Citation
Immature stages of the skipper butterflies (Lepidoptera: hesperiidae) of the United States

Material Information

Title:
Immature stages of the skipper butterflies (Lepidoptera: hesperiidae) of the United States biology, morphology, and descriptions
Creator:
Minno, Marc C., 1956-
Publication Date:
Language:
English
Physical Description:
xxviii, 509 leaves : ill., photos ; 29 cm.

Subjects

Subjects / Keywords:
Zoology thesis Ph.D
Dissertations, Academic -- Zoology -- UF
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1994.
Bibliography:
Includes bibliographical references (leaves 479-507).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Marc C. Minno.

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Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright Marc C. Minno. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
030545795 ( ALEPH )
31736288 ( OCLC )

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Full Text
IMMATURE STAGES OF THE SKIPPER BUTTERFLIES
(LEPIDOPTERA: HESPERIIDAE) OF THE UNITED STATES:
BIOLOGY, MORPHOLOGY, AND DESCRIPTIONS
BY
MARC C. MINNO

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA

1994




ACKNOWLEDGMENTS

Numerous biologists loaned specimens or contributed data for my studies of immature hesperiids. My sincerest appreciation goes to Andrew Atkins, H. David Baggett, Greg Ballmer, Susan S. Borkin, John W. Brown, John M. Burns, Hideyuki Chiba, Frank C. Elia, John F. Emmel, Thomas C. Emmel, Hugh Avery Freeman, Dale H. Habeck, Donald W. Hall, Donald J. Harvey, J. Richard Heitzman, John B. Heppner, Timothy L. McCabe, Noel McFarland, Jacqueline Y. Miller, Steven Passoa, Gordon F. Pratt, Dale F. Schweitzer, and Jeffrey R. Slotten for their contributions. Special thanks go to Roy 0. Kendall, who has been especially supportive and generously loaned specimens, many being unique, from his personal collection. The United States National Museum, Florida State Collection of Arthropods, and the Santa Barbara Natural History Museum also loaned preserved specimens from their collections. Host plant specimens were identified by David W. Hall and Kenneth R. Langdon. Peter J. Eliazar and Andrei Sourakov assisted with the scanning electron microscope. Many thanks go to the librarians at the University of Florida and the Florida Department of Agriculture, Division of Plant Industry and Consumer




Services who helped tremendously in locating a number of obscure references. I also heartily thank my committee members, Lincoln P. Brower, Gary R. Buckingham, Dale H. Habeck, Jonathan Reiskind, and Chairman Thomas C. Emmel for their patient guidance, support, and critique of my graduate work. This research was supported in part by a small grant from the Florida Entomological Society. In conclusion, my greatest appreciation goes to my loving wife, Maria Frances Minno, for her support and help during the course of this research.

iii




TABLE OF CONTENTS
page
ACKNOWLEDGMENTS ...................................... ii
LIST OF TABLES ....................................... vi
LIST OF FIGURES ...................................... vii
ABSTRACT ............................................. xxvi
CHAPTERS
1 INTRODUCTION ................................. 1
2 METHODS ...................................... 5
Study Material ............................... 5
Rearing ...................................... 7
Preservation and Preparation .................... 11
Descriptive Methods .......................... 12
3 BIOLOGY OF NORTH AMERICAN HESPERIIDAE .......... 15
Biology of the Pyrrhopyginae .................... 31
Biology of the Pyrginae ...................... 31
Biology of the Heteropterinae ................... 38
Biology of the Hesperiinae ................... 38
Biology of the Megathyminae .................. 44
4 GENERAL MORPHOLOGY OF THE IMMATURE STAGES ... 50
Eggs ......................................... 50
Larvae ....................................... 51
Pupae ........................................ 66




5 DIAGNOSES AND HOST PLANTS FOR THE UNITED
STATES SKIPPERS ............................ 88
Subfamily Pyrrhopyginae .......................... 88
Subfamily Pyrginae ........................... 90
Subfamily Heteropterinae ..................... 204
Subfamily Hesperiinae ........................ 206
Subfamily Megathyminae ....................... 328
6 TAXONOMIC KEYS TO THE IMMATURE STAGES .......... 453
Preliminary Key to Eggs of U. S.
Hesperiidae (Subfamily and Genus) ............ 453
Preliminary Key to Larvae of U. S.
Hesperiidae (Subfamily and Genus) ............ 455
Preliminary Key to Pupae of U. S.
Hesperiidae (Subfamily and Genus) ............ 463
7 SUMMARY AND CONCLUSIONS ...................... 468
Biology ...................................... 468
Morphology ................................... 471
Taxonomic Insights and Problems ................. 475
REFERENCES ........................................... 479
BIOGRAPHICAL SKETCH .................................. 508




LIST OF TABLES

Table page
1 Number of species and subfamilies of U. S.
Hesperiidae recorded from various host plant
families...................................... 16
2 Maximum number of lenticles observed per
segment and position on Hesperiidae larvae
in the present study. Lenticles may be found in eight possible positions on the
prothorax (Ti), five positions on thoracic
segments 2-3 and abdominal segments 1-9,
and two positions on the last segment (A10). An asterisk indicates 10 to 15
lenticles present............................ 54
3 Maximum number of lenticles observed per
segment and position on Megathyminae larvae
in the present study. Lenticle positions
are labelled 1 for lenticles above the
spiracle line, 2 for lenticles below the
spiracle line, and 3 for ventral lenticles.
Position 1 on abdominal segment 10 refers to lenticles on the lateral margin of the caudal
proleg. An asterisk indicates 10 to 15
lenticles present............................ 58




LIST OF FIGURES
Figure page
1 Larval shelter of Urbanus proteus on
Phaseolus vulgaris (A), Polygonus leo on Piscidia piscipula (B), Erynnis zarucco
on Sesbania macrocarpa (C), Erynnis brizo
on Quercus myrtifolia (D), Wallengrenia
otho on Paspalum sp. (E), and Asbolis
capucinus on Sabal sp. (F)................... 20
2 Larval shelter of Euphyes pilatka on Cladium
jamaicense (A), Euphyes arpa on Sabal etonia
(B), Megathymus yuccae on Yucca aloifolia (C), and an agave leaf cut open to reveal
the tunnels of Agathymus aryxna (D)......... 22
3 SEM micrographs of the egg of Urbanus
proteus (A), Telemiades epicalus (B),
Thorybes pylades (C), Polites vibex (D), Hylephila phyleus (E), Atrytone delaware
(F), Euphyes ruricola (G), Copaeodes minimus
(H), and Megathymus cofaqui (I).............. 70
4 SEM micrographs of the micropylar region of
the egg of Urbanus proteus (A), Telemiades
epicalus (B), Erynnis horatius (C),
Hesperia attalus (D), Polites baracoa (E), Atrytone delaware (F), Problema byssus (G),
Poanes yehl (H), and Euphyes ruricola (I)... 72
5 SEM micrographs of the sculpturing on the
sides of the egg of Atalopedes campestris (A), Polites vibex (B), Atrytone delaware (C), Problema byssus (D), Euphyes ruricola
(E), and Megathymus cofaqui (F)............. 74
6 Comparison of setae on the body of Pholisora
catullus (A), the head of Achalarus casica (B), the body of P. catullus (C), the body
of Urbanus proteus (D), the body of
Ancyloxypha numitor (E), the body of
Hesperia juba (F), the head of Xenophanes
trixus (G), the head of Pyrgus communis (H), the body of Thorybes bathyllus (I), and the
body of Cogia outis (J)...................... 75

vii




Figure

7 Generalized map of the lenticles and other
features of the last instar hesperiid larva. 76
8 Lenticles on the fourth abdominal segment of
Pholisora catullus (A), Problema byssus (B),
and Megathymus yuccae (C).................... 77
9 Color patterns on the fourth abdominal
segment of some U. S. hesperiids: dorsal
view of transverse stripes on Astraptes
fulgerator (A), lateral view of subdorsal
and lateral stripes bounded by white (shaded
portion) on Copaeodes aurantiacus (B),
lateral view of the interrupted subdorsal
stripe of Codatractus arizonensis (C), and
lateral view of patches of black microspines
on Epargyreus clarus (D)..................... 78
10 Relative size of the "neck" of last instar
larvae of Pyrrhopyge araxes (A), Urbanus
teleus (B), Problema byssus (C), and
Megathymus yuccae (D)........................ 79
11 Features of the larval head of hesperiids:
pebbly sculpturing of Polythrix procera (A),
rough sculpturing of Staphylus mazans (B), pitted sculpturing of Hesperia pahaska (C),
ridges on the head of Pyrrhopyge chalybea
(D), and general structures on Amblyscirtes
vialis (E).......... .........o.............. 80
12 Processes on the head of Urbanus procne (A),
Achalarus lyciades (B), and Copaeodes
aurantiacus (C).............................. 81
13 Comparison of the stemmata of Pyrrhopyge
araxes (A), Achlyodes thraso (B), and
Euphyes pilatka (C).......................... 82
14 Ventral view of the hesperiine-type
mandibles of Atalopedes campestris (A) and
the pyrgine-type mandibles of Erynnis
horatius (B)....... ........................ 83
15 Wax gland distribution on Problema byssus
(A), Lerema accius (B), Synapte malitiosa (C), Asbolis capucinus (D), Hesperia juba (E), Ochlodes agricola (F), and Perichares
philetes (G)................................ 84

viii




16 Comparison of the anal comb of Urbanus
proteus (A), Thorybes mexicanus (B), Pyrgus
scriptura (C), Polites themistocles (D),
Pseudocopaeodes eunus (E), and Megathymus
yuccae (F).................................... 85
17 Lenticle distribution on the prothorax of
the pupa of Pseudocopaeodes eunus (A), on
the dorsal abdomen of Hylephila phyleus (B),
and on the proleg scars of Yvretta carus
(C)........................................... 86
18 Lateral view of the pupal cap of Perichares
philetes (A), Atrytone delaware (B), Panoquina panoquin (C), and Calpodes
ethlius (D).................................. 87
19 Last instar larva of Autochton cellus (A),
Epargyreus clarus (B), Thorybes pylades (C),
Urbanus proteus (D), Urbanus dorantes (E),
Polygonus leo (F), Gesta gesta (G), and
Erynnis horatius (H)......................... 343
20 Last instar larva of Achlyodes thraso (A),
Pholisora catullus (B), Pyrgus oileus (C), Staphylus hayhurstii (D), Hesperia attalus
(E), Pompeius verna (F), Wallengrenia
egeremet (G), and Atrytonopsis hianna (H)... 345
21 Last instar larva of Atrytone delaware (A),
Problema byssus (B), Asbolis capucinus (C),
Euphyes arpa (D), Lerema accius (E),
Panoquina panoquin (F), Lerodea eufala (G),
and Megathymus yuccae (H).................... 347
22 Frontal view of the larval head of Phocides
palemon (A), Epargyreus clarus (B),
Polygonus leo (C), Chioides catillus (D),
Typhedanus undulatus (E), and Polythrix
mexicana (F). All drawings are enlarged 12
times........................................ 348
23 Frontal view of the larval head of Polythrix
procera (A), Codatractus alcaeus (B), C.
arizonensis (C), Urbanus proteus (D),
Urbanus dorantes (E), and Astraptes
fulgerator (F). All drawings are enlarged
12 times..................................... 349

Figure

apage




24 Frontal view of the larval head of Astraptes
gilberti (A), Autochton cellus (B),
Achalarus lyciades (C), Thorybes bathyllus
(D), Cabares potrillo (E), and Cogia
hippalus (F). All drawings are enlarged
12 times..................................... 350
25 Frontal view of the larval head of
Nisoniades rubescens (A), Staphylus ceos (B), Carrhenes canescens (C), Xenophanes
trixus (D), Systasea pulverulenta (E),
Achlyodes thraso (F), Timochares
ruptifasciatus (G), and Gesta gesta (H).
All drawings are enlarged 12 times......... 351
26 Frontal view of the larval head of
Ephyriades brunneus (A), Erynnis icelus (B), E. brizo (C), E. juvenalis (D), E.
propertius (E), and E. horatius (F). All
drawings are enlarged 12 times.............. 352
27 Frontal view of the larval head of Erynnis
tristis (A), E. martialis (B), E. pacuvius
(C), E. zarucco (D), E. funeralis (E), E.
baptisiae (F), E. afranius (G), and E.
persius (H). All drawings are enlarged 12
times........................................ 353
28 Frontal view of the larval head of Pyrgus
centaureae (A), Heliopetes ericetorum (B),
Celotes nessus (C), Pholisora catullus (D),
Piruna pirus (E), Synapte malitiosa (F),
Vidius perigenes (G), and Nastra lherminier
(H). All drawings are enlarged 12 times.... 354
29 Frontal view of the larval head of Cymaenes
tripunctus (A), Lerema accius (B),
Perichares philetes (C), Ancyloxypha numitor (D), A. arene (E), Copaeodes aurantiacus (F), C. minimus (G), and
Thymelicus lineola (H). Drawings A-C and H are enlarged 12 times; D-G are enlarged
25 times..................................... 355
30 Frontal view of the larval head of Hylephila
phyleus (A), Yvretta carus (B),
Pseudocopaeodes eunus (C), Hesperia uncas
(D), H. leonardus (E), H. pahaska (F),
Polites coras (G), and P. sabuleti (H).
All drawings are enlarged 12 times.......... 356

Figure

page




31 Frontal view of the larval head of Polites
mardon (A), P. draco (B), P. baracoa (C),
P. themistocles (D), P. origenes (E),
P. mystic (F), P. sonora (G), and P. vibex
(H). All drawings are enlarged 12 times.... 357
32 Frontal view of the larval head of
Wallengrenia otho (A), Pompeius verna (B), Atalopedes campestris (C), Atrytone arogos (D), A. delaware (E), Problema byssus (F),
Ochlodes sylvanoides (G), and Ochlodes
agricola (H). All drawings are enlarged
12 times..................................... 358
33 Frontal view of the larval head of Ochlodes
yuma (A), Poanes hobomok (B), P. zabulon
(C), P. taxiles (D), P. aaroni (E), P. yehl (F), P. viator (G), and Paratrytone melane
(H). All drawings are enlarged 12 times.... 359
34 Frontal view of the larval head of
Choranthus haitensis (A), Euphyes arpa (B), E. pilatka (C), E. alabamae (D), E. dukesi (E), E. bimacula (F), E. ruricola (G), and
Asbolis capucinus (H). All drawings are
enlarged 12 times............................ 360
35 Frontal view of the larval head of
Atrytonopsis hianna (A), Amblyscirtes linda
(B), A. aesculapias (C), A. vialis (D), A.
celia (E), Lerodea eufala (F), Oligoria maculata (G), and Calpodes ethlius (H).
All drawings are enlarged 12 times.......... 361
36 Frontal view of the larval head of Panoquina
panoquin (A), Nyctelius nyctelius (B),
Agathymus valverdiensis (C), and Megathymus
yuccae (D). All drawings are enlarged 12
times........................................ 362
37 Stemmatal pattern of Pyrrhopyge araxes (A),
Phocides pigmalion (B), P. palemon (C),
Epargyreus zestos (D), E. clarus (E), E.
exadeus (F), Polygonus leo (G), Chioides
catillus (H), Typhedanus undulatus (I),
Polythrix mexicana (J), P. procera (K), and
Codatractus alcaeus (L). All drawings are
enlarged 25 times............................ 363

Figure

-age




Figure

38 Stemmatal pattern of Codatractus arizonensis
(A), Urbanus proteus (B), U. esmeraldus (C),
U. dorantes (D), U. teleus (E), U. procne (F), Astraptes fulgerator (G), A. gilberti (H), A. anaphus (I), Autochton cellus (J), Achalarus lyciades (K), and A. casica (L).
All drawings are enlarged 25 times........... 364
39 Stemmatal pattern of Thorybes bathyllus (A),
T. pylades (B), T. diversus (C), T.
mexicanus (D), T. confusis (E), T. drusius
(F), Cabares potrillo (G), Cogia hippalus
(H), C. outis (I), C. caicus (J), Nisoniades
rubescens (K), Staphylus mazans (L), S.
hayhurstii (M), Carrhenes canescens (N),
Xenophanes trixus (0), Systasea pulverulenta (P), S. zampa (Q), and Achlyodes thraso (R).
All drawings are enlarged 25 times........... 365
40 Stemmatal pattern of Timochares
ruptifasciatus (A), Chiomara asychis (B), Gesta gesta (C), Ephyriades brunneus (D),
Erynnis icelus (E), E. brizo (F), E.
juvenalis (G), E. propertius (H), E.
horatius (I), E. tristis (J), E. martialis
(K), E. pacuvius (L), E. zarucco (M), E.
funeralis (N), E. baptisiae (0), E. afranius
(P), E. persius (Q), Pyrgus centaureae (R),
P. ruralis (S), P. scriptura (T), P.
communis (U), P. albescens (V), P. oileus
(W), Heliopetes ericetorum (X), H. lavianus
(Y), H. macaira (Z), H. arsalte (AA),
Celotes nessus (BB), C. limpa (CC), and
Pholisora catullus (DD). All drawings are
enlarged 25 times ........................... 366
41 Stemmatal pattern of Pholisora alpheus (A),
P. gracielae (B), Piruna pirus (C), Synapte malitiosa (D), Vidius perigenes (E), Nastra
lherminier (F), N. julia (G), N. neamathla (H), Cymaemes tripunctus (I), Lerema accius (J), L. liris (K), Ancyloxypha numitor (L),
Perichares philetes (M), A. arene (N),
Copaeodes aurantiacus (0), C. minimus (P), Thymelicus lineola (Q), Hylephila phyleus
(R), Yvretta carus (S), Pseudocopaeodes eunus (T), Hesperia uncas (U), H. juba (V), H. comma
(W), H. woodgatei (X), H. leonardus (Y), H.
pahaska (Z), H. metea (AA), and H. viridis
(BB). All drawings are enlarged 25 times.... 367

xii

aDAGe




42 Stemmatal pattern of Hesperia attalus (A),
H. meskei (B), H. nevada (C), Polites coras
(D), P. sabuleti (E), P. mardon (F), P.
draco (G), P. baracoa (H), P. themistocles
(I), P. origenes (J), P. mystic (K), P.
sonora (L), P. vibex (M), Wallengrenia otho
(N), W. egeremet (0), Pompeius verna (P),
Atalopedes campestris (Q), Atrytone arogos (R), A. delaware (S), Problema byssus (T), Ochlodes sylvanoides (U), O. agricola (V),
O. yuma (W), Poanes hobomok (X), and P.
zabulon (Y). All drawings are enlarged 25
times........................................ 368
43 Stemmatal pattern of Poanes taxiles (A), P.
aaroni (B), P. yehl (C), P. viator (D),
Paratrytone melane (E), Choranthus haitensis
(F), Euphyes arpa (G), E. pilatka (H), E.
alabamae (I), E. dukesi (J), E. macguirei
(K), E. bimacula (L), E. ruricola (M),
Asbolis capucinus (N), Atrytonopsis hianna (0), A. loammi (P), Amblyscirtes linda (Q), A. aesculapias (R), A. vialis (S), A. celia
(T), and Lerodea eufala (U). All drawings
are enlarged 25 times. ....................... 369
44 Stemmatal pattern of Oligoria maculata (A),
Calpodes ethlius (B), Panoquina panoquin (C), P. panoquinoides (D), P. ocola (E),
P. sylvicola (F), Nyctelius nyctelius (G), Agathymus valverdiensis (H), A. stephensi
(I), Megathymus yuccae (J), M. coloradensis (K), M. cofaqui (L), and M. ursus (M). All
drawings are enlarged 25 times.............. 370
45 Dorsal view of the larval terminalia of
Pyrrhopyge araxes (A), Phocides pigmalion (B), Epargyreus zestos (C), and Polygonus
leo (D). All drawings are enlarged six
times........................................ 371
46 Dorsal view of the larval terminalia of
Chioides catillus (A), Typhedanus undulatus
(B), Polythrix mexicana (C), Codatractus
alcaeus (D), Urbanus proteus (E), and
Astraptes fulgerator (F). All drawings
are enlarged six times. ...................... 372

xiii

Figure

page




Figure

47 Dorsal view of the larval terminalia of
Autochton cellus (A), Achalarus lyciades
(B), Thorybes bathyllus (C), Cabares
potrillo (D), Cogia hippalus (E),
Nisoniades rubescens (F), Staphylus
ceos (G), Carrhenes canescens (H), and
Xenophanes trixus (I). All drawings are
enlarged six times. .......................... 373
48 Dorsal view of the larval terminalia of
Achlyodes thraso (A), Timochares
ruptifasciatus (B), Gesta gesta (C),
Ephyriades brunneus (D), Erynnis icelus (E), and Pyrgus centaureae (F). All drawings are
enlarged six times. .......................... 374
49 Dorsal view of the larval terminalia of
Heliopetes ericetorum (A), Celotes nessus (B), Pholisora catullus (C), Piruna pirus
(D), Synapte malitiosa (E), Vidius perigenes
(F), and Nastra lherminier (G). All
drawings are enlarged six times............. 375
50 Dorsal view of the larval terminalia of
Cymaenes tripunctus (A), Lerema accius (B),
Perichares philetes (C), Ancyloxypha numitor
(D), A. arene (E), Copaeodes aurantiacus
(F), and C. minimus (G). Drawings D and F-G
are enlarged 6 times; E is enlarged 12
times...................... ................. 376
51 Dorsal view of the larval terminalia of
Thymelicus lineola (A), Hylephila phyleus
(B), Yvretta carus (C), Pseudocopaeodes eunus
(D), Hesperia uncas (E), Polites coras (F), and
P. sabuleti (G). Drawings A-E are enlarged 6
times; F and G are enlarged 12 times........ 377
52 Dorsal view of the larval terminalia of
Polites sabuleti (A), P. mardon (B), P.
draco (C), P. baracoa (D), P.themistocles
(E), and P. origenes (F). Drawings A-E are
enlarged 12 times; F is enlarged 6 times.... 378
53 Dorsal view of the larval terminalia of
Polites mystic (A), P. sonora (B), P. vibex
(C), Wallengrenia otho (D), Pompeius verna
(E), Atalopedes campestris (F), and Atrytone
arogos (G). Drawings A-C are enlarged 12
times; D-G are enlarged 6 times............. 379

xiv

apage




54 Dorsal view of the larval terminalia of
Atrytone delaware (A), third instar Problema byssus (B), last instar Problema byssus (C),
Ochlodes yuma (D), Poanes hobomok (E), and
Paratrytone melane (F). Drawings A and C-F
are enlarged 6 times; B is enlarged 12
times........................................ 380
55 Dorsal view of the larval terminalia of
Choranthus haitensis (A), Euphyes arpa (B),
E. ruricola (C), Asbolis capucinus (D),
Atrytonopsis hianna (E), and Amblyscirtes linda (F). All drawings are enlarged six
times........................................ 381
56 Dorsal view of the larval terminalia of
Lerodea eufala (A), Oligoria maculata (B),
Panoquina panoquin (C), P. panoquinoides (D), P. ocola (E), and P. sylvicola (F).
Drawings A-D are enlarged 6 times; E-F are
enlarged 12 times. ........................... 382
57 Dorsal view of the larval terminalia of
Nyctelius nyctelius (A), Calpodes ethlius
(B), Agathymus valverdiensis (C), and
Megathymus yuccae (D). All drawings are
enlarged six times. .......................... 383
58 Pupa of Epargyreus clarus (A), Thorybes
bathyllus (B), Urbanus proteus (C), Phocides
pigmalion (D), Polygonus leo (E), Anastrus sempiternus (F), Achlyodes thraso (G), and
Pyrgus communis (H).......................... 385
59 Pupa of Hesperia attalus (A), Pompeius verna
(B), Poanes yehl (C), Oligoria maculata (D),
Cymaenes tripunctus (E), and Synapte
malitiosa (F)................................ 387
60 Pupa of Nastra lherminier (A), Perichares
philetes (B), and Calpodes ethlius (C)...... 389
61 Dorsal view of anterior pupa of Phocides
pigmalion (A), P. palemon (B), Epargyreus
zestos (C), and E. clarus (D). All drawings
are enlarged six times. ...................... 390

xv

Figure

page




Figure

62 Dorsal view of anterior pupa of Polygonus
leo (A), Chioides catillus (B), Polythrix mexicanus (C), Codatractus alcaeus (D), C.
arizonensis (E), and Urbanus proteus (F).
All drawings are enlarged six times......... 391
63 Dorsal view of anterior pupa of Urbanus
esmeraldus (A), U. dorantes (B), U. teleus
(C), U. procne (D), and Astraptes fulgerator
(E). All drawings are enlarged six times... 392
64 Dorsal view of anterior pupa of Astraptes
gilberti (A), Autochton cellus (B),
Achalarus lyciades (C), Thorybes bathyllus
(D), T. pylades (E), and T. confusis (F).
All drawings are enlarged six times......... 393
65 Dorsal view of anterior pupa of Cabares
potrillo (A), Cogia outis (B), Staphylus
ceos (C), S. hayhurstii (D), Carrhenes
canescens (E), Systasea pulverulenta (F),
Achlyodes thraso (G), and Chiomara asychis
(H). All drawings are enlarged six times... 394
66 Dorsal view of anterior pupa of Gesta gesta
(A), Ephyriades brunneus (B), Erynnis icelus
(C), E. brizo (D), E. juvenalis (E), E.
horatius (F), E. tristis (G), and E.
martialis (H). All drawings are enlarged
six times.................................... 395
67 Dorsal view of anterior pupa of Erynnis
zarucco (A), E. funeralis (B), E. baptisiae
(C), E. persius (D), Pyrgus scriptura (E),
P. communis (F), P. oileus (G), and
Heliopetes ericetorum (H). All drawings are
enlarged six times. .......................... 396
68 Dorsal view of anterior pupa of Heliopetes
lavianus (A), Celotes nessus (B), Pholisora
catullus (C), P. alpheus (D), Synapte
malitiosa (E), Nastra lherminier (F), N.
julia (G), and N. neamathla (H). All
drawings are enlarged six times............. 397

xvi

page




69 Dorsal view of anterior pupa of Cymaenes
tripunctus (A), Lerema accius (B), L. liris
(C), Perichares philetes (D), Ancyloxypha
numitor (E), A. arene (F), and Copaeodes
aurantiacus (G). All drawings are enlarged
six times.................................... 398
70 Dorsal view of anterior pupa of Copaeodes
minimus (A), Hylephila phyleus (B), Yvretta
carus (C), Pseudocopaeodes eunus (D),
Hesperia uncas (E), H. comma (F), H. attalus
(G), and Polites coras (H). All drawings
are enlarged six times. ...................... 399
71 Dorsal view of anterior pupa of Polites
sabuleti (A), P. mardon (B), P. draco (C),
P. baracoa (D), P. themistocles (E), P.
origenes (F), P. mystic (G), and P. sonora
(H). All drawings are enlarged six times... 400
72 Dorsal view of anterior pupa of Polites
vibex (A), Wallengrenia otho (B), W.
egeremet (C), Pompeius verna (D),
Atalopedes campestris (E), Atrytone
delaware (F), A. arogos (G), and Problema
byssus (H). All drawings are enlarged
six times.................................... 401
73 Dorsal view of anterior pupa of Ochlodes
sylvanoides (A), Poanes hobomok (B), P.
taxiles (C), P. aaroni (D), P. yehl (E),
P. viator (F), Paratrytone melane (G), and
Choranthus haitensis (H). All drawings are
enlarged six times. .......................... 402
74 Dorsal view of anterior pupa of Euphyes arpa
(A), E. pilatka (B), E. alabamae (C), E.
dukesi (D), E. berryi (E), and E. bimacula
(F). All drawings are enlarged six times... 403
75 Dorsal view of anterior pupa of Euphyes
ruricola (A), Asbolis capucinus (B),
Atrytonopsis loammi (C), Amblyscirtes
aesculapias (D), A. celia (E), Lerodea
eufala (F), and Oligoria maculata (G). All
drawings are enlarged six times............. 404

xvii

Figure

page




76 Dorsal view of anterior pupa of Calpodes
ethlius (A), Panoquina panoquin (B), P.
panoquinoides (C), P. ocola (D), and P.
sylvicola (E). All drawings are enlarged
six times.................................... 405
77 Dorsal view of anterior pupa of Nyctelius
nyctelius (A), Megathymus coloradensis (B),
M. cofaqui (C), and M. ursus (D). Drawing A
is enlarged six times; B-D are enlarged
three times.................................. 406
78 Ventral view of anterior pupa of Phocides
pigmalion (A) and P. palemon (B). All
drawings are enlarged six times............. 407
79 Ventral view of anterior pupa of Epargyreus
zestos (A) and E. clarus (B). All drawings
are enlarged six times. ...................... 408
80 Ventral view of anterior pupa of Polygonus
leo (A) and Chioides catillus (B). All
drawings are enlarged six times............. 409
81 Ventral view of anterior pupa of Polythrix
mexicana (A), Codatractus alcaeus (B), and
C. arizonensis (C). All drawings are
enlarged six times. .......................... 410
82 Ventral view of anterior pupa of Urbanus
proteus (A), U. esmeraldus (B), and U.
dorantes (C). All drawings are enlarged
six times.................................... 411
83 Ventral view of anterior pupa of Urbanus
teleus (A) and U. procne (B). All drawings
are enlarged six times. ...................... 412
84 Ventral view of anterior pupa of Astraptes
fulgerator (A), A. gilberti (B), and
Autochton cellus (C). All drawings are
enlarged six times. .......................... 413
85 Ventral view of anterior pupa of Thorybes
bathyllus (A), T. pylades (B), T. confusis (C), Cabares potrillo (D), and Cogia outis
(E). All drawings are enlarged six times... 414

xviii

Figure

page




86 Ventral view of anterior pupa of Staphylus
ceos (A), S. hayhurstii (B), Carrhenes
canescens (C), Systasea pulverulenta (D),
Achlyodes thraso (E), Chiomara asychis (F),
Gesta gesta (G), and Ephyriades brunneus
(H). All drawings are enlarged six times... 415
87 Ventral view of anterior pupa of Erynnis
icelus (A), E. brizo (B), E. juvenalis (C),
E. horatius (D), E. tristis (E), E.
martialis (F), E. zarucco (G), and E.
funeralis (H). All drawings are enlarged
six times.................................... 416
88 Ventral view of anterior pupa of Erynnis
baptisiae (A), E. persius (B), Pyrgus
scriptura (C), P. communis (D), P. oileus
(E), Heliopetes ericetorum (F), H. lavianus
(G), and Celotes nessus (H). All drawings
are enlarged six times. ...................... 417
89 Ventral view of anterior pupa of Pholisora
catullus (A), P. alpheus (B), Synapte
malitiosa (C), Nastra lherminier (D), N.
julia (E), N. neamathla (F), and Cymaenes
tripunctus (G). All drawings are enlarged
six times.................................... 418
90 Ventral view of anterior pupa of Lerema
accius (A), L. liris (B), Perichares
philetes (C), Ancyloxypha numitor (D), A.
arene (E), Copaeodes aurantiacus (F), and C. minimus (G). Drawings A-B and D-G are
enlarged six times; C is enlarged three
times........................................ 419
91 Ventral view of anterior pupa of Hylephila
phyleus (A), Yvretta carus (B),
Pseudocopaeodes eunus (C), Hesperia uncas
(D), H. comma (E), H. attalus (F), Polites
coras (G), and P. sabuleti (H). All
drawings are enlarged six times............. 420
92 Ventral view of anterior pupa of Polites
mardon (A), P. draco (B), P. baracoa (C),
P. themistocles (D), P. origenes (E), P.
mystic (F), P. sonora (G), and P. vibex (H).
All drawings are enlarged six times......... 421

xix

Figure

page




93 Ventral view of anterior pupa of
Wallengrenia otho (A), W. egeremet (B),
Pompeius verna (C), and Atalopedes
campestris (D). All drawings are enlarged
six times.................................... 422
94 Ventral view of anterior pupa of Atrytone
arogos (A), A. delaware (B), Problema byssus
(C), and Ochlodes sylvanoides (D). All
drawings are enlarged six times............. 423
95 Ventral view of anterior pupa of Poanes
hobomok (A), P. taxiles (B), P. aaroni (C), and P. yehl (D). All drawings are enlarged
six times.................................... 424
96 Ventral view of anterior pupa of Poanes
viator (A), Paratrytone melane (B),
Choranthus haitensis (C), and Euphyes arpa
(D). All drawings are enlarged six times... 425
97 Ventral view of anterior pupa of Euphyes
pilatka (A), E. alabamae (B), E. dukesi (C),
E. berryi (D), E. bimacula (E), and E.
ruricola (F). All drawings are enlarged six
times...................... ................. 426
98 Ventral view of anterior pupa of Asbolis
capucinus (A), Atrytonopsis loammi (B),
Amblyscirtes aesculapias (C), A. celia (D),
Lerodea eufala (E), and Oligoria maculata
(F). All drawings are enlarged six times... 427
99 Ventral view of anterior pupa of Calpodes
ethlius (A), Panoquina panoquin (B), P.
panoquinoides (C), P. ocola (D), and P.
sylvicola (E). All drawings are enlarged
six times.................................... 428
100 Ventral view of anterior pupa of Nyctelius
nyctelius (A), Agathymus valverdiensis (B), Megathymus yuccae (C), M. coloradensis (D),
M. cofaqui (E), and M. ursus (F). All
drawings are enlarged three times........... 429

Figure

pau ac
.L-




101 Lateral view of the cremaster of Phocides
pigmalion (A), P. palemon (B), Epargyreus zestos (C), E. clarus (D), Polygonus leo
(E), Chioides catillus (F), Polythrix mexicana (G), Codatractus alcaeus (H),
and C. arizonensis (I). All drawings are
enlarged 12 times ........................... 430
102 Lateral view of the cremaster of Urbanus
proteus (A), U. esmeraldus (B), U. dorantes (C), U. teleus (D), U. procne (E), Astraptes
fulgerator (F), A. gilberti (G), Autochton
cellus (H), and Achalarus lyciades (I). All
drawings are enlarged 12 times............... 431
103 Lateral view of the cremaster of Thorybes
bathyllus (A), T. pylades (B), T. confusis
(C), Cabares potrillo (D), Cogia hippalus
(E), C. outis (F), C. caicus (G), Nisoniades
rubescens (H), and Staphylus ceos (I). All
drawings are enlarged 12 times............... 432
104 Lateral view of the cremaster of Staphylus
mazans (A), S. hayhurstii (B), Carrhenes
canescens (C), Xenophanes trixus (D),
Systasea pulverulenta (E), Achlyodes thraso (F), Timochares ruptifasciatus (G), Chiomara
asychis (H), Gesta gesta (I), Ephyriades
brunneus (J), Erynnis icelus (K), E. brizo (L), E. juvenalis (M), E. horatius (N), E.
tristis (0), and E. martialis (P). All
drawings are enlarged 12 times............... 433
105 Lateral view of the cremaster of Erynnis
zarucco (A), E. funeralis (B), E. baptisiae
(C), E. persius (D), Pyrgus scriptura (E), P. communis (F), P. oileus (G), Heliopetes
ericetorum (H), H. lavianus (I), Celotes
nessus (J), Pholisora catullus (K), P.
alpheus (L), Synapte malitiosa (M), Nastra
lherminier (N), N. julia (0), and N.
neamathla (P). All drawings are enlarged
12 times .................................... 434

xxi

Figure

page




106 Lateral view of the cremaster of Cymaenes
tripunctus (A), Lerema accius (B), L. liris
(C), Perichares philetes (D), Ancyloxypha
numitor (E), A. arene (F), Copaeodes
aurantiacus (G), C. minimus (H), Hylephila
phyleus (I), Yvretta carus (J),
Pseudocopaeodes eunus (K), and Hesperia
uncas (L). Drawings A-C and E-L are
enlarged 12 times; D is enlarged 6 times.... 435
107 Lateral view of the cremaster of Hesperia
comma (A), H. woodgatei (B), H. viridis (C),
H. attalus (D), Polites coras (E), P.
sabuleti (F), P. mardon (G), P. draco (H),
P. baracoa (I), P. themistocles (J), P.
origenes (K), P. mystic (L), P. sonora (M), P. vibex (N), Wallengrenia otho (0), and W.
egeremet (P). All drawings are enlarged 12
times........................................ 436
108 Lateral view of the cremaster of Pompeius
verna (A), Atalopedes campestris (B), Atrytone arogos (C), A. delaware (D),
Problema byssus (E), Ochlodes sylvanoides
(F), Poanes hobomok (G), P. taxiles (H), P.
aaroni (I), and P. yehl (J). All drawings
are enlarged 12 times. ....................... 437
109 Lateral view of the cremaster of Poanes
viator (A), Paratrytone melane (B),
Choranthus haitensis (C), Euphyes arpa (D), E. pilatka (E), E. alabamae (F), E. dukesi
(G), E. berryi (H), E. macguirei (I), E.
bimacula (J), E. ruricola (K), and Asbolis
capucinus (L). Drawings A-C and H-K are
enlarged 12 times; D-G and L are enlarged
6 times...................................... 438
110 Lateral view of the cremaster of
Atrytonopsis loammi (A), Amblyscirtes
aesculapias (B), A. vialis (C), A. celia
(D), Lerodea eufala (E), Oligoria maculata
(F), Calpodes ethlius (G), Panoquina
panoquin (H), P. panoquinoides (I), and P.
ocola (J). Drawings A-F and H-J are
enlarged 12 times; G is enlarged 6 times.... 439

xxii

Figure

apage




111 Lateral view of the cremaster of Panoquina
sylvicola (A), Nyctelius nyctelius (B), Agathymus valverdiensis (C), Megathymus
yuccae (D), M. coloradensis (E), M. cofaqui
(F), and M. ursus (G). Drawings A-B are
enlarged 12 times; C-G are enlarged 6 times. 440
112 Dorsal view of the cremaster of Phocides
pigmalion (A), P. palemon (B), Epargyreus zestos (C), E. clarus (D), Polygonus leo (E), and Chioides catillus (F). Drawings
A and B are enlarged 6 times; C-F are
enlarged 12 times. ........................... 441
113 Dorsal view of the cremaster of Polythrix
mexicana (A), Codatractus alcaeus (B), C.
arizonensis (C), Urbanus proteus (D), U.
esmeraldus (E), U. dorantes (F), U. teleus
(G), U. procne (H), and Astraptes fulgerator
(I). All drawings are enlarged 12 times.... 442
114 Dorsal view of the cremaster of Astraptes
gilberti (A), Autochton cellus (B),
Achalarus lyciades (C), Thorybes bathyllus
(D), T. pylades (E), T. confusis (F),
Cabares potrillo (G), Cogia hippalus (H),
and C. outis (I). All drawings are
enlarged 12 times. ........................... 443
115 Dorsal view of the cremaster of Cogia caicus
(A), Nisoniades rubescens (B), Staphylus
ceos (C), S. mazans (D), S. hayhurstii (E), Carrhenes canescens (F), Xenophanes trixus
(G), Systasea pulverulenta (H), Achlyodes thraso (I), Timochares ruptifasciatus (J), Chiomara asychis (K), and Gesta gesta (L).
All drawings are enlarged 12 times.......... 444
116 Dorsal view of the cremaster of Ephyriades
brunneus (A), Erynnis icelus (B), E. brizo (C), E. juvenalis (D), E. horatius (E), E.
tristis (F), E. martialis (G), E. zarucco
(H), E. funeralis (I), E. baptisiae (J), E.
persius (K), and Pyrgus scriptura (L). All
drawings are enlarged 12 times.............. 445

xxiii

Figure

page




117 Dorsal view of the cremaster of Pyrgus
communis (A), P. oileus (B), Heliopetes ericetorum (C), H. lavianus (D), Celotes
nessus (E), Pholisora catullus (F), P.
alpheus (G), Synapte malitiosa (H), Nastra lherminier (I), N. julia (J), N. neamathla
(K), and Cymaenes tripunctus (L). All
drawings are enlarged 12 times............... 446
118 Dorsal view of the cremaster of Lerema
accius (A), L. iris (B), Perichares
philetes (C), Ancyloxypha numitor (D), A.
arene (E), Copaeodes aurantiacus (F), C.
minimus (G), Hylephila phyleus (H), Yvretta
carus (I), Pseudocopaeodes eunus (J), and Hesperia uncas (K). Drawings A-B and D-K
are enlarged 12 times; C is enlarged 6
times ....................................... 447
119 Dorsal view of the cremaster of Hesperia
comma (A), H. woodgatei (B), H. viridis (C),
H. attalus (D), Polites coras (E), P.
sabuleti (F), P. mardon (G), P. draco (H), P. baracoa (I), P. themistocles (J), and P.
origenes (K). All drawings are enlarged 12
times ....................................... 448
120 Dorsal view of the cremaster of Polites
mystic (A), P. sonora (B), P. vibex (C),
Wallengrenia otho (D), W. egeremet (E),
Pompeius verna (F), Atalopedes campestris (G), Atrytone arogos (H), A. delaware (I), and Problema byssus (J). Drawings A-I are
enlarged 12 times; J is enlarged 6 times.... 449
121 Dorsal view of the cremaster of Ochlodes
sylvanoides (A), Poanes hobomok (B), P.
taxiles (C), P. aaroni (D), P. yehl (E),
P. viator (F), Paratrytone melane (G),
Choranthus haitensis (H), Euphyes arpa (I), E. pilatka (J), E. alabamae (K), E. dukesi
(L), and E. berryi (M). Drawings A, C-E,
G-H, and M are enlarged 12 times; B, F, and
I-L are enlarged 6 times..................... 450

xxiv

Figure

paq




122 Dorsal view of the cremaster of Euphyes
macguirei (A), E. bimacula (B), E. ruricola
(C), Asbolis capucinus (D), Atrytonopsis loammi (E), Amblyscirtes aesculapias (F),
A. vialis (G), A. celia (H), Lerodea eufala
(I), Oligoria maculata (J), and Calpodes
ethlius (K). Drawings A-C and E-J are
enlarged 12 times; D and K are enlarged 6
times........................................ 451
123 Dorsal view of the cremaster of Panoquina
panoquin (A), P. panoquinoides (B), P. ocola
(C), P. sylvicola (D), Nyctelius nyctelius
(E), Megathymus yuccae (F), M. coloradensis
(G), M. cofaqui (H), and M. ursus (I).
Drawings A-E are enlarged 12 times; F-I are
enlarged 6 times............................. 452

xxv

Figure

page




Abstract of Dissertation to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
IMMATURE STAGES OF THE SKIPPER BUTTERFLIES
(LEPIDOPTERA: HESPERIIDAE) OF THE UNITED STATES:
BIOLOGY, MORPHOLOGY, AND DESCRIPTIONS
By
Marc C. Minno
April 1994
Chairman: Thomas C. Emmel Major Department: Zoology
The purpose of this study was to investigate the biology of as many U. S. skippers as possible, compare morphological features of their immature stages, and produce standardized descriptions. This revision summarizes of our current knowledge of immature stages of North American Hesperiidae, and provides a framework for future taxonomic and systematic studies.
About 300 species of skipper butterflies (Hesperiidae) have been reported from the United States. Most of the fauna is composed of the subfamilies Pyrginae and Hesperiinae, but Pyrrhopyginae, Heteropterinae, and Megathyminae are also represented.
Species of Hesperiidae found in the U. S. feed on 31 families of plants as larvae. These include 8 families of monocots and 23 families of dicots. Most (92%) of these butterflies are limited to only 1 of the 31 families of plants, but a few skipper species feed on plants in 2, or xxvi




very rarely, as many as 6 different families. Species having relatively broad host ranges include Lerema accius (35 spp.), Epargyreus clarus (31 spp.), Thorybes pylades (27 spp.), and Megathymus yuccae (27 spp.). One hundred and thirty-eight new host records are reported for 55 species of hesperiids. Some skippers are important pests of beans, rice, sugarcane, and pasture grasses. Host plants remain to be discovered for 22% of the U. S. Hesperiidae.
The eggs of 49 species, the larvae of 156 species, and the pupae of 120 species of Hesperiidae of the U. S. are described in this thesis. These include the eggs of 16 species, the larvae of 41 species, and the pupae of 35 species that were previously unknown. Twenty-six percent of the U. S. skippers are totally unknown with respect to their egg, larval, and pupal stages. About half of these species with undescribed immatures are breeding residents. The other half are neotropical species that rarely enter the U. S. and probably do not reproduce within its limits.
Larval characters that may prove useful to systematists conducting phylogenetic analyses include the type of mandible articulation, setal types, sculpturing of the head, lenticle distribution on the larvae, wax gland patterns, and pigments. Differences in the sculpturing patterns of hesperiid eggs have been known for some time, but have not been extensively surveyed. Promising characteristics of the pupae include the shape of the pupal cap, lenticle

xxvii




distribution, relative length of the antennae and proboscis, cremaster shape, type of thoracic spiracle guard, and presence of crenulations on the posterior margin of the prothorax.

xxviii




CHAPTER 1
INTRODUCTION
Adult skippers (Hesperiidae) are thick-bodied
butterflies with large compound eyes, wide heads, and conspicuous labial palps. The antennal club is usually bent and has a tapering extension called the apiculus. The radial veins of the forewings all arise from the discal cell, and the second medial vein of the hindwings is absent or weakly developed. All six legs are well-developed and used for perching. Many skippers are drably colored, but some tropical species are bright yellow, red or iridescent blue or green. The wings are frequently dotted with semihyaline whitish spots. The Hesperiidae is a group that is easily recognized as distinct from all other butterfly families.
Skippers occur in all faunal regions of the world
except at the poles (Heppner 1991). Estimates of the number of described species range from 3000 to 3658 (Robbins 1982, Bridges 1983, and Heppner 1991). The Neotropics are especially rich in skippers, with 2,016 described species (Heppner 1991), followed by the Old World tropics, with 570 species in the Oriental region and 440 species in the Ethiopian region. Among temperate regions, the Nearctic has




2
more species (290) than Australia (191) or the Palearctic (155). Overall, about 63% of all described species of skippers occur in the New World.
Within the Hesperiidae, eight subfamilies have been recognized (Heppner 1991). The Heteropterinae is the smallest subfamily, with only about 30 species distributed in the Palearctic, Nearctic, and Neotropical regions. The Megathyminae is another small subfamily of about 50 species. Megathymines are found in xeric areas of the United States, Mexico, and Central America. The Coeliadinae is a group of about 80 species restricted to the Old World tropics. Another Old World group, the Trapezitinae, includes about 85 species endemic to the Australian faunal region. Euschemoninae consists of a single Australian species, Euschemon rafflesia. Some 180 skippers of the subfamily Pyrrhopyginae occur in the Neotropics. The two largest groups, the Pyrginae (1,195 species) and the Hesperiinae (2,044 species), are widely distributed throughout the world.
Some hesperiids are economically important to people. The larvae of one species, Aegiale hesperiaris, known as gusanos de maguey in Spanish, are eaten as a delicacy in Mexico (Blasquez 1870). The Indians of Mexico have long harvested gusanos de maguey for food (Draudt 1924). Today, these hesperiid larvae are collected and sold fresh or canned in Mexican markets (Borror et al. 1954), and are




3
fried before eating (Ancona 1934, Sokolov 1989). Small larvae are sometimes added to bottles of tequila as a garnishment (Sbordoni and Forestiero 1985, Toliver 1987).
Skippers are also pests of crops, forests, and plants used as garden ornamentals. Urbanus proteus is destructive to beans (Comstock 1881, Riley and Howard 1893, Chittenden 1902, Cotton 1918, Watson 1919, Wolcott 1933, Greene 1971, and Heppner 1975), Ancyloxypha numitor, Panoquina ocola, and Nyctelius nyctelius to rice (Bell 1940, Ross and Lambremont 1963), and Panoquina sylvicola, Nyctelius nyctelius, Choranthus haitensis, Choranthus vitellius, Perichares philetes, and others to sugarcane (Smyth 1919, Wolcott 1921, Jones and Wolcott 1922, Wolcott 1933, 1936, and Hayward 1943). Hesperiids that have been listed as occasionally injurious to trees include Epargyreus clarus and Erynnis species (Felt 1918, McDaniel 1933, Craighead 1950, Baker 1972, and Furniss and Carolin 1977). Grass-feeding species such as Hylephila phyleus and Atalopedes campestris may occasionally damage lawns (Bohart 1947, Warren and Roberts 1956, Ross and Lambremont 1963) and Calpodes ethlius frequently will defoliate ornamental cannas (Dyar 1898, Chittenden 1905, 1912, Essig 1926, Weigel and Middleton 1926, and Reinert et al. 1983).
A few skippers accidentally introduced into new areas have become pests. A European species introduced into Canada, Thymelicus lineola, is a major pest of pastures and




4
hay crops (Pengelly 1961, Arthur 1966, and Duchesne and McNeil 1978). Similarly, of two exotic skippers now established in Hawaii, Hylephila phyleus causes damage to lawns (Tashiro and Mitchell 1985) and Erionota thrax to bananas (Riotte and Uchida 1979).
Aside from human economics, skippers have biological value, being important in the pollination of flowering plants and in the cycling of nutrients and flow of energy in food webs. Since many species are restricted to certain habitats or are very local in distribution, they may also prove useful for biological monitoring of the ecological health of nature reserves or for biodiversity studies.
Skippers also have scientific interest, and a large
literature on this group has accumulated over the last few centuries. Many life history descriptions of New World hesperiids have been published, especially by workers in the United States (U. S.), but these accounts sometimes contain errors and are uneven in the choice of characters described. The intent of this study was to investigate the biology of as many U. S. species as possible, compare morphological features of their immature stages, and produce standardized descriptions. This revision will not only serve as a summary of our current knowledge of immature stages of North American Hesperiidae, but will provide a framework for future taxonomic and systematic studies.




CHAPTER 2
METHODS
Study Material
A major problem associated with studying the immature stages of butterflies lies in finding specimens. Butterflies are relatively rare insects compared to groups such as Diptera, Coleoptera, or Hymenoptera, and locating immatures in the field is often a challenging task. Although many collectors and researchers are interested in immature stages, few preserve eggs, larvae, or pupae. Insect collections that contain lepidopteran immatures will often have at least some skipper butterfly larvae, but these are usually common species. Preserved pupae, even of common species, are rare in museum collections. Roy O. Kendall (ROK) has one of the largest (in terms of numbers of individuals as well as species) and best curated collections of North American hesperiids in existence. I borrowed many specimens for study from the Kendall collection as well as from the Florida State Collection of Arthropods (FSCA), Santa Barbara Natural History Museum (SBNHM), United States National Museum (USNM), and the collections of G. R. Ballmer (GRB), T. C. Emmel (TCE), D. H. Habeck (DHH), J. R. Heitzman (JRH), T. L. McCabe (TLM), N. McFarland (NM), and S. Passoa




6
(SP). Several lepidopterists, including H. D. Baggett, S. S. Borkin, J. F. Emmel, D. W. Hall, D. F. Schweitzer, A. M. Shapiro, and J. R. Slotten, also donated specimens for study. In addition, I have conducted my own field research in California, Colorado, Florida, Georgia, Indiana, Maryland, New Mexico, New York, North Carolina, Pennsylvania, South Carolina, and Texas over the last 20 years. Outside of the United States, I have been able to make collections of hesperiid immatures in Colombia, Costa Rica, Dominican Republic, and the Bahamas. Some tropical species represented in these collections have occasionally been recorded from the U. S. Andrew Atkins and H. Chiba sent preserved immatures of interesting species from Australia and Indonesia, which were useful for making comparative studies.
The number of hesperiid eggs, larvae, and pupae that I examined during the course of this study is presented under the species accounts in Chapter 6 of this work. An identifying label, with the format "MCM-Lot XXXX" (where the X's represent a unique number), was placed in each vial or jar of immatures that I examined. The consecutive numbers used began with 0001 and ended with 1561. The code number was also recorded on an accession form listing specimen deposition, identification, locality, host plant, parasites, number of individuals of each stage, type of container, and preservative. Much of this information was entered into a




7
digital database using an IBM personal computer and LOTUS 12-3 software. The accession code may be used to trace specimens for future studies and was used to link database files together.
Rearing
One method of collecting hesperiid immatures is to
search for them in the field. Some species are relatively easy to find with a little practice, since the eggs are laid only on particular plants and the larvae usually fold or tie leaves together to form shelters in which they live. However, many species live such secretive lives that they are best reared in the laboratory.
It has long been known that female butterflies will often lay eggs if confined in a container with the larval host plant. Edwards (1870) got a female zebra swallowtail to lay eggs by placing a nail-keg, from which the bottom had been knocked out, over a pawpaw bush and covering the top with a cloth. Scudder (1870) used tin cans in a similar way and secured eggs of Thorybes pylades, Polites mystic, and a satyrid butterfly. Both Edwards and Scudder were able to rear many butterflies using this technique. Fletcher (1888) recommended not only caging females over hosts in the field, but also rearing larvae on potted hosts covered by wire cages or glass lamp chimneys. Laurent (1908) found that if female skippers were removed from a cyanide killing jar soon after they succumbed, they could easily be transferred to




8
individual pill boxes and that they usually recovered without harm in a short time. He then secured eggs from the female skippers by confining them in cages over pots of the host plants.
Caging females with a known or suspected host does not always work. Some species of skippers seem to lay eggs freely, while others absolutely will not oviposit in confinement. Not only is there wide variation between species, there is also much variation between individuals. Generally, a slightly worn female may be more likely to lay eggs than one that is freshly emerged (and perhaps not mated). It has been my experience that pyrgines are less likely to lay eggs in confinement than hesperiines. Heitzman (1964a) noted that one or two fertile eggs may sometimes be dissected from females that will not otherwise oviposit in captivity.
I have found that females may be transported from the field to the laboratory without harm by placing them individually in the plastic containers in which 35mm film is sold. Either the opaque Kodak or transparent Fuji Film containers work well, but the clear containers have the advantage of the investigator being able to examine the specimens without opening them. Containers with live females should be kept in a cool, dark place such as a field bag or cooler. In the laboratory, the containers may be kept refrigerated for a few days without harm, but it is




9
best to feed and place the females in oviposition containers as soon as possible. I manually fed female skippers a dilute honey solution (1 part honey to 10 parts tap water) twice a day. By grasping the adult's closed wings with one hand, the skipper's proboscis could be uncoiled into a dish of honey water using an insect pin held by the other hand.
To secure eggs, I found it best to place the females in transparent plastic containers containing small sprigs of host plants. The cut ends of the plants may be placed into small vials of water or wrapped with a piece of wet paper towel to prevent the leaves from wilting. The plant should be flattened against the container wall or arranged such that the butterfly will easily come into contact with the leaves and not become trapped in pockets of vegetation. A hole, 2 to 3cm in diameter, cut into the top of the oviposition container and covered with fine netting, will provide ventilation and prevent condensation.
The behavior of adult skippers in cages consists of
periods of rest punctuated by short bursts of activity. The oviposition containers with females and hosts may be placed under incandescent lights to stimulate activity, but need to be monitored frequently, as excessive heat, moisture, or dryness may be a problem. I prefer to place the containers on bright, but not sunny window sills or in a sheltered location out-of-doors where natural light and breezes stimulate the females. Richard Boscoe (pers. comm.)




10
recommends placing the oviposition containers into closed plastic bags or refrigerating females of species reluctant to oviposit in captivity overnight, but I have had limited success with these techniques.
I have not tried to secure eggs from female
megathymines in the laboratory. Megathymus species glue the eggs to the leaves of yuccas, and the eggs are relatively easy to find in the field at the right time of year. Agathymus species either drop the eggs onto the host, or ova laid on the leaves fall off soon after oviposition, making them difficult to find in nature. Comstock and Dammers (1934), Roever (1964), and Stallings and Stallings (1986) were successful in having females of Agathymus species oviposit in cages in the laboratory.
Once eggs have been laid, care must be taken to prevent excessive moisture, which promotes the growth of molds, or dryness which may cause the eggs to desiccate. The plastic containers used for securing eggs also worked well for rearing larvae. Ventilation through the screened hole in the lid, fresh food plants, and maintaining clean conditions within the containers were important in the prevention of disease.
Temperate species that undergo larval hibernation pose special problems, as it is often difficult to provide proper environmental conditions in the laboratory. I have had some success in caging larvae out-of-doors on potted plants, or




11
in rearing the larvae under artificial lights with long-day photoperiods. Other rearing techniques are presented in MacNeill (1964), McFarland (1964), Brown (1965), Kendall (1965), Newcomer (1966), and Tashiro and Mitchell (1985). Wielgus and Wielgus (1973) and Wielgus and Stallings (1974) present methods for rearing Megathymus in the laboratory.
Preservation and Preparation
I preserved hesperiid eggs and first instar larvae by placing them in vials with 70% alcohol. Older larvae and pupae were dropped into hot (near boiling) water to kill them (Peterson 1962), blotted dry with paper toweling, then transferred to 2 or 4 dram vials containing 70% alcohol. After several weeks, the old discolored preservative was drained from the vials and new alcohol added. I used 70% isopropyl alcohol for preserving immatures, because it is inexpensive, readily available, and less hazardous than formulations containing acetic acid, formalin, xylene, or kerosene. Specimens preserved in this manner remained relatively pliable and easy to work with, but lost most of the body colors.
To document colors and the general features of living
caterpillars and pupae, I made color slides using a Canon A1 35mm camera with a 100mm macrolens. A 50mm extension tube was used with this apparatus to magnify specimens between 1 and 3 cm in length. Most specimens were photographed in natural light. Both Kodak Kodachrome 64 and Ektachrome 100




12
color transparency films gave good results, but it was sometimes difficult to get satisfactory slides with the slower film when photographing small specimens under low light conditions.
Descriptive Methods
I chose the checklist by Miller and Brown (1983) as a starting point for my descriptive work. According to these authors, the hesperiid fauna of North America, north of Mexico, contains 1 species of Pyrrhopyginae, 117 species of Pyrginae, 5 species of Heteropterinae, 139 species of Hesperiinae, and 28 species of Megathyminae for a total of 290 species. The Canadian fauna included in this checklist does not contain any species not found in the U. S. (Gregory 1975).
External features of the immature stages were compared with the aid of a Wild M5 stereomicroscope fitted with 20 power eyepieces. A preliminary list of potentially interesting characters was drafted and modified during an initial examination of all of the material. After this review, a new standard character list was produced. The specimens were then examined a second time, new character data were recorded, and drawings of interesting features were made. Abbreviations for segments referred to in the diagnoses are T1-3 for thoracic segments one through three and Al-10 for abdominal segments one through ten.




13
The fine structure of eggs was observed and
photographed using the University of Florida, Department of Zoology scanning electron microscope (SEM) (Hitachi model S415A). To prevent distortion, the eggs were dehydrated through a series of alcohol solutions (70%-99%). The alcohol was subsequently replaced through a series of acetate solutions. The specimens were then subjected to critical point drying using a Denton Vacuum Inc. Apparatus DCP-l. The dry eggs were glued to SEM stubs and coated with gold-palladium using a Denton Vacuum Inc. Desk-l Sputterer, then viewed with the SEM.
I made sketches and measurements of structures with a
camera lucida attached to the stereomicroscope. The drawing tube allowed for superimposing the image of a metric ruler onto the specimen. Then, by dividing by the magnification power, size could be determined. This method worked well for structures greater than 0.1 mm. The heights and diameters of the eggs of 49 species were measured and recorded using this technique. The lengths of larvae and pupae were made simply with a metric ruler. Measurements of the transverse width of the head and abdominal segment four (generally the widest segment) of larvae, and abdominal segment three (usually the widest) of pupae were made with the microscope. A total of 1584 larvae and 623 pupae of 158 species were measured. For measurements of setal lengths and number of crochets, I chose the largest specimens




14
available, and these data are reported as maxima in the diagnoses. In comparing pupal abdomen length among the various species of U. S. hesperiids, the distribution of the ratio of A5-8 length and A1-4 length was divided into three equal groups representing short (0.3 to 0.69mm), moderate (0.7 to 0.89mm), and long (0.9 to 1.4mm) classes.




CHAPTER 3
BIOLOGY
Although the larvae of some specialized moths feed on fungi, lower plants (algae, lichens, liverworts, mosses, ferns, cycads, and gymnosperms), other insects, feces, beeswax, or the keratin-containing parts of dead vertebrates, most butterflies feed only on angiosperm plants. Exceptions include a few neotropical nymphalids (Satyrinae) that eat lycopsids (Singer et al. 1971, Singer and Mallet 1986), a few lycaenids and pierids that eat the leaves of pines, and some lycaenids which feed on the leaves of cycads (Eumaeus spp.) or are predators of aphids (Liphyinae) or ant larvae (some Polyommatinae). Skippers feed only on angiosperms, but utilize a diversity of plants that are defended by a wide variety of secondary plant compounds.
Species of skippers tend to be stenophagous, limiting the host range to just a few species of plants, yet sixty families of higher plants have been recorded as hosts in the New World. The host relationships of U. S. hesperiids is presented in Table 1. Thirty-one families representing 8 Mbnocot groups and 23 dicots have been reported. Hesperiinae, Heteropterinae, and Megathyminae use only




Table 1. Number of species and subfamilies of U.S. Hesperiidae recorded from various host plant families.

HOST PLANT
FAMILY

Monocots:
Agavaceae Arecaceae Cannaceae
Cyperaceae Marantaceae
Musaceae
Poaceae
Zingiberaceae
Dicots:
Amaranthaceae Aquifoliaceae
Asteraceae Betulaceae
Chenopodiaceae
Combretaceae
Convolvulaceae
Fabaceae Fagaceae
Lamiaceae
Malpighiaceae
Malvaceae
Myristicaceae
Myrtaceae
Ranunculaceae
Rhamnaceae
Rhizophoraceae
Rosaceae Rutaceae
Salicaceae
Sterculiaceae Trigoniaceae
Verbenaceae

HESPERIIDAE
# SPECIES SUBFAMILIES

28
2 1 15
1 1
103

Megathyminae Hesperiinae Hesperiinae Hesperiinae Hesperiinae Hesperiinae Heteropterinae Hesperiinae Pyrginae Hesperiinae
Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrrhopyginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae Pyrginae




17
monocots; Pyrginae and Pyrrhopyginae usually feed only on dicots. Three subfamilies, Hesperiinae, Heteropterinae, and Pyrginae, use Poaceae, while both Pyrginae and Pyrrhopyginae eat Fagaceae.
Hesperiid females adopt a characteristic fluttering type of flight pattern when searching for host plants on which to lay eggs. Often, the females will hover about potential hosts, sometimes landing upon the leaves one or more times, before laying eggs. By closely inspecting leaf shape and sensing plant chemistry, the butterflies are able to distinguish hosts of suitable size, age, quality, and position in the landscape before proceeding to lay their eggs (Scott 1986).
Females usually lay their eggs singly on the leaves of the host plant. Most glue the eggs to the under surface of leaves, while others prefer to oviposit on the upper surface or exhibit no preference. The color of the egg changes as development proceeds. Scudder (1889a) recognized four embryological stages of butterfly eggs. Stage one is of short duration and consists of the newly laid egg, which is of homogeneous color. Stage two is marked by either a darkening of the color or the appearance of small, uniformly distributed spots. During the third stage there is a polarity of the contents corresponding to the formation of the germinal band in which a central annulus and a large lateral spot (the head) appear. In the last stage, the head




18
moves upward to just underneath the micropyle at the apex of the egg.
The newly laid eggs of skippers may be brownish, pinkish, white, pale yellow, or green. Development of hesperiid eggs agrees with Scudder's (1889a) observations. The eggs soon darken or change color and may develop a reddish ring. Shortly before the eggs are ready to hatch, the dark head of the larva is usually visible at the apex. Skipper eggs typically hatch in 4 to 15 days. When the larva is ready to emerge, it makes a small hole in the chorion with the mandibles, then cuts a circular opening in the apex slightly larger than the head (Ainslie 1922). This process may take several days (Roever 1964), but once the hole is completed, the larva exits quickly. The newly emerged larva usually does not eat the remains of the egg but seeks out a site in which to build a shelter.
Hesperiid larvae tie leaves or parts of leaves together with silk to form characteristic shelters in which they hide when not feeding (Figures 1 and 2). Although skippers are often known as leaf rollers, they do not roll leaves in the manner of tortricids, pyralids, or other moths, but rather fold leaves or tie the edges of several leaves together and line the inside with silk. Among other groups of butterflies, some nymphalids and papilionids construct silklined shelters. Those built by charaxines (Nymphalidae) such as Anaea andria are the most similar to hesperiid shelters.




Figure 1. Larval shelter of Urbanus proteus on Phaseolus vulgaris (A), Polygonus leo on Piscidia piscipula (B) Erynnis zarucco on Sesbania macrocarpa (C), Erynnis brizo on Quercus myrtifolia (D), Wallengrenia otho on Paspalum sp.
(E), and Asbolis capucinus on Sabal sp. (F).




20
7w,




Figure 2. Larval shelter of Euphyes pilatka on Cladium jamaicense (A), Euphyes arpa on Sabal etonia (B), Megathymus yuccae on Yucca aloifolia (C), and an agave leaf cut open to reveal the tunnels of Agathymus aryxna (D).




IIA
9W AO




23
The only predators that I have observed capturing skipper larvae are Polistes wasps. It is not known if hesperiid larvae are palatable to vertebrate predators such as birds and lizards. Parasitic wasps in the families Mymaridae, Eulophidae, and Braconidae frequently attack the eggs and larvae of skippers. The larvae are also commonly attacked by tachinid flies. Potentially, ants, spiders, lizards, birds, and small mammals may eat skipper butterfly immatures unless repulsed by the defenses noted below.
The larval shelters of hesperiids are primarily thought to provide concealment from predators and parasitoids. However, the shelters are often conspicuous, perhaps raising a flag to predators, and skipper larvae are often heavily parasitized. Another plausible reason for shelters may involve chlorophyll. Chlorophyll which becomes uncoupled from the tightly controlled photosynthetic units within plant cells, as during digestion, is capable of diverting electrons to normally inert oxygen molecules, forming highly destructive oxygen radicals (Hendry 1990). Larger herbivores with opaque bodies have little to fear from chlorophyll as the digestion takes place in the dark inside their guts. However, a rare disease of sheep in which partly digested chlorophyll passes into the blood causes cell destruction in epidermal tissues exposed to sunlight due to the formation of activated oxygen (Hendry 1990). Poulton (1894) demonstrated that the ground color of




24
caterpillars is derived largely from the diet. The green color of hesperiids that feed on leaves is due to chlorophyll. Newly emerged larvae become green only after the first feeding. Hesperiines, which have little cuticular pigmentation, change from green to cream-colored before entering diapause or pupation when they are no longer actively feeding. Leaf-feeding caterpillars which have relatively large amounts of chlorophyll in their blood and guts as well as thin, transparent, or translucent cuticles must have physiological and/or behavioral adaptations that prevent the formation of oxygen radicals. Many authors have noted that skipper larvae hide in shelters during the day and emerge to feed at night (Harris 1862, Lintner 1872, Edwards and Chapman 1879, Panton 1897, Scudder 1889a, and numerous others). In fact, most caterpillars are largely nocturnal feeders (J. Rawlins, pers. comm.). Thus the evolution of shelter-forming behavior in skippers may be grounded in a physiological need for concealment from the sun rather than from predators.
The shelter may also be adaptive in allowing the larva easy access to the leaves of the host. Other caterpillars that do not make shelters or nests rest exposed on the underside of leaves or hide on the branches, under bark, or in leaf litter at the base of the plant. Skipper larvae spin a thin trail of silk with a characteristic side-to-side motion of the head (Fletcher 1889, Scudder 1889a, Ainslie




25
1922, Clark 1936, MacNeill 1964, McAlpine 1973, and Heitzman and Heitzman 1974) as they move about, allowing for firm footing as well as guidance to feeding sites and shelter location. The caterpillars may change shelters three or four times as they outgrow them, but usually stay on the same host or near the oviposition site. Tamburo and Butcher (1955) found that marked larvae of Ephyriades brunneus floridensis mostly stayed at the point of release, but one traveled three host bushes away, or about three meters.
Living in one place for extended periods presents the problem of waste disposal. If frass accumulates near the shelter, it may provide visual or olfactory clues to predators. Skipper larvae propel frass away from the shelter site (Harris 1862, King 1882, Frohawk 1892, Kendall 1965) with the anal comb, sometimes sending the pellets up to a distance of one meter (Scudder 1889a, Ainslie 1922).
Skipper larvae are slow-moving and generally exhibit very subdued behavior. They utilize a variety of defenses against predators, including dropping to the ground and remaining motionless, protective coloration, displaying open jaws, regurgitation of fluids from the mouth, and defecation; they probably also secrete chemicals from a gland on the ventral side of the prothorax (Scott 1986). When disturbed, some notodontid moths, such as Lochmaeus manteo, spray a mixture of noxious chemicals from similar glands (Eisner et al. 1972). Although I have handled many




26
hundreds of skipper larvae, I have never been able to detect any secretions from the prothoracic gland. However, Scudder (1889a) noted that ants turned away (apparently in "great disgust") from the caterpillars of Epargyreus clarus.
Coloration is an important adaptation that may involve crypsis, aposematic colors, and disruptive patterns. The heads of the larvae may be all dark, dark with contrasting false eye patches (the "monkey faces" of Seitz 1924 and Moss 1949), dark with pale patches, all pale, or pale with dark stripes or spots. These patterns may deter predators with high visual acuity such as birds or lizards. A predator peering into a folded leaf may see false faces or disruptive patterns that are not recognized as potential prey items or be tricked into interpreting all dark or all pale heads as representing emptiness. Belwood (1990) noted similar color patterns on tube-dwelling neotropical Tettigoniidae, suggesting that these katydids and skippers may have similar selection pressures.
Combinations of cuticular pigments and internally
derived colors produce a variety of body colors commonly ranging from dark green to pale blue. Some species are brightly patterned with red, black, yellow, orange, or white cuticular pigments, suggesting that they are unpalatable. Most are greenish with yellow, orange, white, or rarely black lateral lines. Very few have transverse stripes. The dorsum is usually lightly pigmented or transparent,




27
revealing the pulsating dorsal heart as a dark line. In larger larvae, the male testes often can be seen as two kidney-shaped bodies located toward the dorsum of abdominal segment six. A few species have caudal false face patterns.
For caterpillars to grow larger, they must shed the
cuticle. Skipper larvae usually have five larval stages or instars. Temperate species that undergo hibernation may sometimes have a few more stages, especially if reared under laboratory conditions that give confusing signals, such as short days but warm temperatures. The developmental time of the early instars ranges from four days to about a week, but lengthens as the larvae grow larger. The duration of the entire larval period typically ranges from 14 to 45 days.
Shortly before hesperiids molt, the head is withdrawn from the old head capsule and shows as a whitish bulge in the prothoracic area. How this is accomplished has not been investigated, but skippers would seem to have special problems making the withdrawal since a key characteristic of the group is the narrow posterior opening of the head. At the moment of molting, the cuticle separates from the head capsule; then the larva walks out of the old skin which remains fixed to a mat of silk threads by the crochets. The old head capsule is cast away with a few lateral movements of the head, and the larva rests while the new cuticle hardens.




28
Interestingly, the final larval molt is somewhat
different from the others, perhaps because the head of the prepupal organism cannot be withdrawn through the posterior opening of the old head capsule. With a few rhythmic struggles, the larval cuticle splits along the dorsum of the thorax and along the frontal suture of the head capsule which remains attached to the skin (Fyles 1895, Ainslie 1922). The exuvium is then slid backwards to the cremaster, and is pushed aside, lying as a crumpled mass at the far end of the cocoon.
Changes in coloration may occur in skipper larvae upon molting or in preparation for hibernation or pupation. Most species have dark heads during the first two or three instars, but develop other patterns in later stages. Body coloration may also change as the larvae mature. The change in species of Phocides is extreme, from red with transverse yellow stripes to white. The body color of temperate species may change from green to brown or take on a pinkish cast without shedding the cuticle before entering hibernation (Minno 1981, Capman 1990). Hesperiines often change from green to a creamy color after they stop feeding and seek out overwintering or pupation sites (Frohawk 1892, Heitzman 1965a). The creamy color is derived from whitish fat bodies and other internal organs which are no longer masked by chlorophyll.




29
Temperate species of hesperiids usually hibernate as partly grown larvae. More rarely, some species overwinter as fully developed larvae within the eggs (Fyles 1895, Pengelly 1961), first instars (Dethier 1939a), last instar larvae (Riley 1876a, Edwards 1885a, Heitzman and Heitzman 1974), or pupae (Scott 1981). After winter has passed, nonmature larvae emerge from their hibernaculae and resume feeding. Those that overwintered as mature larvae (Erynnis and Megathymus species) usually do not feed in the spring, but pupate as directed by environmental cues and soon emerge as adults. Thus species with a pupal or mature larval diapause are among the first to emerge as adults during the spring. Species adapted to temperate climates are generally univoltine or bivoltine, whereas tropical skippers often have three or more generations per year.
Unlike most other butterflies, skippers usually pupate in sealed cocoons. The pupa is supported by a silk thread about the junction of the thorax and abdomen, and is anchored to the substrate by cremaster hooks entangled in a small transverse thread or a pad of silk. The cocoon may be formed by sealing the last larval shelter, or the caterpillar may seek out another nearby site. Scudder (1889a) found Epargyreus clarus pupae under boards, shingles, and rubbish under some large locust trees. Temperate species seem more likely to pupate in leaf litter than tropical species.




30
The pupae of many species are coated with a white or
bluish-white wax which is easily rubbed off with a probe or a finger. In some, the wax is a delicate bloom while others are densely coated. Moss (1949) thought that the wax may help keep off moisture while also deceiving predators with the appearance of a cadaver covered with mold. Another protective coloration of the pupa involves the thoracic spiracles, each of which is shielded by a conspicuous cuticular rise. When viewed from the front, these structures become eyes and other features of the head give the appearance of a small animal.
Although most, if not all, skippers secrete at least some wax through pores in the pupal cuticle; a few groups also produce quantities of wax in the larval stage. Late in the last instar, many hesperiines and megathymines produce wax from glands concentrated in patches on the ventral side of the body. These deposits have a characteristic shape and distribution that often vary between genera. Within a genus, some species may have wax glands, while other congeners do not. The hollow filaments of wax that issue from the glands (Locke 1960) have a flaky or cottony appearance. The larva uses the wax to coat the inside or plug the entrance to the pupal chamber.




31
Biology of the Pyrrhopyginae
Little is known of the biology of the Pyrrhopyginae. I have examined a few preserved larvae of Pyrrhopyge araxes arizonae, Pyrrhopyge chalybea, and an unidentified Brazilian species, but have not had the opportunity to study living material. Host plants include members of the Anacardiaceae, Anonaceae, Clusiaceae, Fagaceae, Flacourtiaceae, Lauraceae, Meliaceace, Myristicaceae, Myrtaceae, Sapindaceae, and Sterculiaceae. The larvae live in pyrgine-type shelters of folded leaves (Moss 1949, Burns 1964a). Larvae described by Moss (1949) and Comstock (1956b) have a reddish ground color and contrastingly colored transverse stripes or spots and some resemble the pyrgine genus Phocides. Pupae are also reddish but are covered with white wax in the manner of many pyrgines. The last shelter of Myscelus pardalina was perforated by a few large holes (Moss 1949), a specialized behavior of certain pyrgines such as Quadrus species. Pupation takes place in the last larval shelter or in leaves on ground. Moss notes that Pyrrhopyge larvae are very pugnacious when disturbed and often leave the shelter when it is cut from the host.
Biology of the Pyrginae
As Scudder (1875) noted, members of the Pyrginae
usually feed on dicotyledonous plants. Many pyrgines eat legumes (Fabaceae). However, there has also been a tremendous radiation away from the use of legumes such that




32
at least 44 other families of plants are eaten by New World pyrgines. Many of these hosts contain strongly scented secondary plant chemicals. Remarkably, there are also a few species of Urbanus that feed on grasses (monocots), while their congeners eat legumes (Kendall 1976)! Twenty-four plant families are utilized by pyrgines reported from the U. S. (Table 1).
Female pyrgines usually lay eggs singly on living
leaves of the host but may also choose flower buds, stems, fruit, or dead leaves (Panton 1897, Tamburo and Butcher 1955, Minno 1981). Pholisora catullus deposits eggs on the upper side of the leaf near the midrib (Edwards 1884, Capman 1990), but most other species seem to prefer the edges of the lower surface. Urbanus dorantes frequently oviposits on the inflorescences of Desmodium species in Florida. One pyrgine, Epargyreus clarus, has been observed to lay eggs on nearby plants after inspecting and touching the true host (Opler and Krizek 1984, MCM observation). Newly laid eggs may be white, yellow, pale green, or brownish.
Urbanus proteus and Autochton cellus females lay stacks of eggs containing from two to six individuals (Comstock 1881, Quaintance 1898, Clark 1934, 1936). The eggs in the stack are offset at an angle (Clark 1936), allowing larvae in the lower eggs to emerge in the normal way from the exposed micropylar area. Quaintance (1898) noted that the most distal egg in the string is the first to hatch.




33
Although these species begin life communally, the newly hatched larvae lead separate lives, each constructing a shelter around the margin of the leaf. Kendall (1976) reported finding ten larvae of Codatractus hyster in a single shelter and this may be a truly gregarious species.
First instar larvae usually have dark brown or black heads. However, some Erynnis species have pale head capsules. Newly emerged pyrgine larvae do not eat the remains of the egg, but usually seek out the margin of a leaf and begin constructing a shelter. The larva eats two converging slits into the leaf margin, except for a narrow portion which acts as a hinge. The resulting flap is then folded over and secured with three or four strands of silk (Lintner 1872, Edwards 1885b, Panton 1897, Quaintance 1898, Hayward 1927a, Clark 1936) in such a way as to form a domeshaped roof. The inside of the shelter is lined with silk. Clark (1934) described the shelters of Autochton cellus as limpet-like. Most species fold the flap over onto the upper surface of the leaf, but Zestusa dorus, Paches polla, and Atarnes sallei tie the flaps to the underside (Klots 1971, Kendall and McGuire 1975, Kendall 1976).
Erynnis species that feed on woody plants often
construct their shelters where a vein reaches the margin (Kendall and McGuire 1975) or at the tip of the leaf (Kendall 1965). The vein adds strength to the hinge. Rather than cutting into the leaf margin, the first instar




34
larva of Nisoniades bessus godma eats a circular channel near the center of a leaf then ties the flap over (Kendall 1976). Newly emerged larvae of Heliopetes laviana eat away the leaf surface, then fold a flap over the weakened spot (Kendall 1965). Young Pyrgus species usually seek out deep wrinkles or overlapping leaves in which to build shelters (Panton 1897, Minno 1981) and Pyrgus scriptura clips off the dense stellate hairs of the host and ties them together with silk to form a covering (Minno 1981). The young larva of Urbanus procne forms a "lean-to" shelter by eating away part of a blade of grass so that the terminal end can be pulled over about 85 degrees, but older larvae simply hide in the detritus at the base of the plant (Kendall 1976).
Older pyrgine larvae may make more box-like shelters by eating channels perpendicular to the leaf margin and folding the rectangular flap over (Clark 1936, Edwards 1885a) or tie the edges of one or more leaves together. Heliopetes macaira may hide in the flower bracts or in dead leaves on the plant or the ground, but often fastens two leaves together, then cuts the petiole of the one which is to be the roof, which withers and dies (Kendall 1965). Similarly, Pyrgus communis often uses a living leaf and detritus (Dethier 1944b, Minno 1981). A few species such as Erynnis persius, Atarnes sallei, and Quadrus species eat small holes in the shelter, then web them over with silk (Scudder 1889a, Kendall 1976, and Young 1991). Before moving to a larger




35
shelter, pyrgine larvae clip the silk threads holding the flap (Scudder 1889a, Clark 1936). Autochton cellus usually makes the first shelter toward the middle or tip of a leaflet and the second more toward the base (Clark 1936).
Pyrgine larvae sometimes construct shelters on plants that are not eaten, but are physically near the true host. Clark (1936) found A. cellus shelters on hickory, sassafras, and grape near to the true host (Amphicarpa bracteata,) and I have found shelters of the legume-feeding U. proteus on blackberry and oak leaves. One fourth instar U. proteus larva, which I placed on a potted Desmodium incanum, built a shelter on a small potted guava adjacent to the host. This behavior suggests that some predators may first direct their hunting toward the food plant rather than the caterpillar.
Pyrgine larvae usually feed on the edges of leaves near to the shelter. Scudder (1889a) noted that the first instar larvae of Erynnis persius feed on the surface layers of the host leaves. Similarly, Pyrgus scriptura larvae feed on the leaf surface within the shelter, rather than the edges of leaves. A few species such as Heliopetes ericetorum and Staphyllus hayhurstii may eat holes in the leaves (Coolidge 1923, Heitzman 1963). Young larvae prefer young tender leaves, but older caterpillars are often able to feed on tougher mature foliage.
Small larvae tend to rest on the top of the shelter in an inverted position (Clark 1936, Kendall 1965). Pyrgine




36
shelters are usually more circular than linear, at least during the early instars, and some species (Achalarus lyciades and Heliopetes ericetorum) assume a coiled resting position, with the head and tail facing the same direction in this unrestricted space (Scudder 1889a, Coolidge 1923).
When disturbed, pyrgine larvae use several defensive behaviors in order to protect themselves. Most will turn the head toward the direction of the stimulus with open jaws, and attempt to bite or thrash the head from side to side. Scudder (1889a) noted that Erynnis juvenalis scrapes the open jaws backwards on the surface of the leaf, making a scratching sound. While they may appear ferocious, I have never actually been bitten by a pyrgine larva. They usually thrash briefly if picked up with fingers, and some such as U. proteus, E. clarus, and A. cellus spit out a watery green-colored fluid if greatly harassed (Scudder 1889a, Clark 1936, MCM observation). Kendall (1976) observed Urbanus teleus, a grass-feeder, to fall to the ground upon being disturbed and lie motionless for long periods of time in the manner of some hesperiines.
The larval stage typically lasts from 15 to 30 days for most pyrgines that do not have diapause requirements. Temperate species usually pass the winter as partly or fully grown larvae; thus for these skippers, the larval stage may extend over several months. A few species diapause as




37
pupae. I do not know of any pyrgines that overwinter as eggs or first instar larvae.
Most pyrgines pupate in cocoons fashioned from the last larval shelter or spun in detritus and debris on the ground. Kendall (1965) noted that although Cogia outis usually pupates on the ground, one larva burrowed under the soil surface and pupated in a silk-lined chamber. Within the cocoon, the pupa is supported by a silk girdle about the middle and is attached to silk threads by the cremaster hooks at the caudal end. Many other butterfly families and some moths use this arrangement, but pyrgines have a specialized feature not known in any other group. The silk girdle supporting the middle of the body also has a median attachment to the cocoon. Thus if the ventral surface of the pupa is toward the bottom of the page, the shape of the girdle is of an inverted letter "Y" (Forbes 1923). Similarly, the cremaster attachment may be Y-shaped (Coolidge 1923).
The pupae of pyrgines may be opaque dark brown, or
greenish with dark markings on the dorsum and wing cases. A few have a short peg-like horn on the frons. The pupae of many pyrgines are thickly coated with wax. The cephalic end of the body is frequently modified to give the appearance of a larger animal with staring eyes. Clark (1936) likened the Autochton cellus pupa to the head of a manatee when viewed




38
from the front. Moss (1926) and Box (1928) describe the frightening appearance of Bungalotis pupae.
Biology of the Heteropterinae
Not much is known of the biology of the Heteropterinae, except for the Arctic Skipper, Carterocephalus palaemon (Fletcher 1889, Frohawk 1892). The morphology, coloration, and behavior of the immature stages of this species are closely similar to that of the Hesperiinae.
Biology of the Hesperiinae
Unlike the pyrgines, Hesperiinae feed on monocots (Scudder 1875). Most New World hesperiines eat grasses (Poaceae), sedges (Cyperaceae), and palms (Arecaceae), but some also feed on the leaves of Bromeliaceae, Cannaceae, Liliaceae, Marantaceae, Musaceae, Orchidaceae, and Zingiberaceae. Host plants of hesperiines recorded from the U. S. include 8 families of monocots (Table 1).
Hesperiine females usually glue the eggs singly to the leaves of the host. Although most hesperiines prefer to deposit their eggs directly on the larval food plant, Hesperia species, Polites species, Wallengrenia otho, Atalopedes campestris, Ochodes yuma, Poanes hobomok, Lerodea eufala, and Panoquina panoquinoides will sometimes lay eggs on dicots or other substrates near the larval host plant (Scott 1986). On two occasions, I have observed Polites vibex, a grass feeder, to lay eggs on dicots after fluttering closely over patches of grass. However, I have




39
also seen females of this species oviposit directly on grass on several occasions. Some populations of Hesperia lindseyi oviposit on lichens growing on fence posts and trees near the grass on which the larvae feed (MacNeill 1964). Dana (1981) discovered that Hesperia ottoe in southwestern Minnesota oviposits on the disk flowers of pale purple coneflower (Echinacea pallida), an adult nectar plant that is not eaten by cattle. After eclosion, the young larvae of these hesperiines must wander, sometimes over relatively long distances, to find a host and begin feeding.
Although the introduced skipper, Thymelicus lineola, always lays eggs in rows inside the sheaths of grasses, there are no known New World hesperiines that consistently lay clusters of eggs. Some such as Calpodes ethlius and Problema byssus may haphazardly deposit eggs in small clusters on occasion (Chittenden 1912, MCM observation). The eggs are usually white or green, but a few species have yellow or pinkish eggs. Green or yellowish eggs frequently develop a bright red ring and apical spot after a few days (Dethier 1942e, Heitzman 1964a, 1966, Brown and McGuire 1983, MCM observation).
The newly emerged larvae of many hesperiines first eat the remains of the egg, leaving only the shiny circular base, before building a shelter (Fyles 1895, Chittenden 1905, Ainslie 1922, Coolidge 1922, 1925, Dethier 1942d, Nielsen 1958, Heitzman 1964b, 1965a, 1966, Heitzman and




40
Heitzman 1970b, MCM observation). Shelters are often formed at the tip of a leaf by tying the edges together with silk or by eating a slit part way into the leaf and folding over the flap. Depending upon the species, the flap may be folded onto the upper or the lower surface of the leaf. Some hesperiines, such as Euphyes species and Atrytonopsis hianna, tie the tips of two overlapping leaves together (Heitzman and Heitzman 1974, MCM observation). Calpodes ethlius cuts two slits parallel to the leaf margin and folds the flap over.
Older larvae typically live in tubular shelters formed by tying the edges of several leaves together and lining the inside with silk. The distal end of the shelter may be tightly closed. Lerema accius feeds below the shelter, nearly severing the midvein, so that the shelter dangles down from the cut. The older larvae of Asbolis capucinus fold the edges of a palm leaflet under and eat the leaf tissue near the tip, except for the midrib, which protrudes like a long spine. Wallengrenia species are most unusual in that they form a case from a piece of leaf. They drag the case to feeding sites, temporarily attaching it to the host with silk (Kendall 1960, MCM observations).
Although most hesperiines tend to live upon the leaves of the host, Hesperia species and their relatives usually construct horizontal shelters of silk and detritus near the base of the plant. Sometimes the shelters may extend below




41
ground (Cockayne 1952, MacNeill 1964, Heitzman and Heitzman 1970b). McCabe and Post (1977) found larvae of Hesperia comma assiniboia in silk-lined burrows under dried cattle droppings. Older larvae of a few hesperiines such as Panoquina species, Nastra species, Oarisma powesheik, Pompeius pompeius, and Decinea percosius, do not make shelters, but rest exposed on leaves or hide in natural recesses (Dyar 1892, McAlpine 1973, Kendall 1976, Kendall and Rickard 1976, MCM observations). Although Kendall (1966b) mentioned that Poanes viator does not make a shelter, I have found older larvae in typical shelters as well as in recesses at the bases of the leaves in Alachua County, Florida. As Shapiro (1971) noted, the larva usually reinforces the recess with some silk.
First instar larvae of hesperiines usually have a dark head. The only exception that I have observed is Poanes viator which has a cream-colored head with some small dark spots. Species that live on leaves are mostly green with longitudinal stripes or reticulate patterns of white or pale yellow. The head capsule frequently bears dark stripes or spots on a pale ground. Hesperiines that live near the ground surface are dully colored brown or dark green and have heads that are plain black or black with pale patches.
For some species, the caudal end may be modified to provide a defense from predators. During the act of defecation, the larva must expose the terminalia which may




attract the attention of lizards or other predators. Some hesperiines (such as Atrytone delaware and Asbolis capucinus) have a flattened, pale-colored suranal plate rimmed or patterned with black. In addition, the last pair of spiracles is often black, contrasting with the pale body color. When viewed from behind, the spiracles become eyes and the suranal plate a broad bill.
Many hesperiines, especially those that do not build shelters, rest with head flattened ventrally against the substrate and the mouth protracted forward (Forbes 1960, MCM observation). Although most hesperiine larvae will rapidly retreat into the shelter if threatened, they tend to be relatively docile. When disturbed, they may drop from the plant, curl ventrally, and remain motionless for a long time (Panton 1897, Heitzman 1964a, MacNeill 1964, Heitzman and Heitzman 1970b). Hesperia species and Amblyscirtes belli thrash violently within the shelter if disturbed (MacNeill 1964, Heitzman 1965b).
Hesperiines that diapause during the winter usually do so as partly grown larvae (Scott 1981). Less commonly, they may pass the winter as fully developed larvae within the eggs, first instar larvae, last instar larvae, or pupae. Although some species may complete the larval stage in about two weeks (Comstock and Dammers 1931, Kendall 1965), one month or longer is more typical. During the last instar




43
many species develop ventral wax glands on the abdominal segments.
Like the pyrgines, hesperiines usually pupate in sealed cocoons. However, Lerema accius and Cymaenes tripunctus pupate exposed, but under a leaf with the edges slightly drawn together. Panoquina panoquin pupates exposed on the upperside of leaves. Amblyscirtes belli, Niconiades nikko, and Quinta cannae cut the last shelter from the host and form the cocoon on the ground (Kendall and McGuire 1975).
Hesperiine pupae are supported by a simple silk girdle about the middle and are attached to a pad of silk by the cremaster. Species of Atrytone and Euphyes lack these supports and pupate loosely in the cocoon in a vertical position. The cremaster of these species is modified into one or two spines and lacks hooks.
The pupal cuticle of hesperiines tends to be lightly pigmented and thin. Those that pupate exposed are pale green and may have yellow or white stripes on the abdomen. Most have a cream colored or slightly greenish ground color with dark patches or lines on the head, wing cases, and dorsum. Some groups of hesperiines have an elongate point on the frons of the pupa (a "beak") that may be upturned or T-shaped at the tip. Atrytone delaware and Poanes viator have a bulbous frons reminiscent of the Australian Trapezitinae, some of which are wildly modified into false face patterns resembling the jaws of termite soldiers. As




44
development proceeds, the eyes become reddish, the wings darken, and just before emergence the adult is clearly visible through the transparent cuticle.
When the adult emerges, the pupa splits along the
dorsum of the thorax, the edges of the head, and the costal margin of the wing cases. The pupal cap (the cuticle covering of the head) is often torn away completely. The exuvium frequently contains a clear or yellowish meconium, which is expelled during emergence (Panton 1897, MCM observation).
Biology of the Megathyminae
I have closely observed the biology of Megathymus yuccae and Megathymus cofaqui in Florida, but have no experience with other Megathyminae. Fortunately, there is a large literature base available on this interesting group. Riley (1876a,b, 1877), Freeman (1951a), Tinkham (1954), Remington (1959), Roever (1964), and Wielgus and Stallings (1974) provided much of the summarized information presented below. Not only do the Megathyminae have a highly specialized biology that differs considerably from other skippers, but the three tribes within this subfamily have each evolved their own peculiar specializations.
Megathymines feed only on plants in the Agavaceae. Megathymus species feed in the stem and roots of yuccas. Members of the closely related genus Stallingsia bore into the stem of Polianthes species. Although Yucca species have




45
large, starchy roots, they also contain saponins and some are commonly known as soap plant. The sap of many yuccas and agaves is highly irritating to human skin and causes a severe poison ivy-like dermatitis. The specialized host preferences of many megathymines seems due, in part, to female choice, as the larvae of Megathymus yuccae can be reared in the laboratory on starchy roots of quite different chemistry such as potatoes.
Megathymus species (Megathymini) glue their eggs singly to the leaves of the host but sometimes select nearby dead leaves or other plants (Riley 1876a,b, 1877, MCM observation). Females often select small or isolated plants in favor of those that are more robust or clumped (Oslar 1900, Tinkham 1954, Stallings and Turner 1956, Remington 1959, McCabe and Post 1977). The eggs are greenish when first laid but later turn white, brownish, or pinkish. They hatch in 7 to 16 days (Wielgus and Stallings 1974).
Newly emerged larvae are reddish. The remains of the egg are not eaten. The young larva of Megathymus yuccae usually constructs a covering of silk and plant fibers over itself near the tip of a leaf and feeds on the epidermal tissue. Later, the larva ties together the youngest leaves at the center of the plant, then bores into the stem. Most other Megathymus species bore directly into the stem after emerging from the egg.




46
Unlike other skippers, larvae of Megathymus species make a cigar-shaped "tent" of silk, plant fibers, and detritus. Some species such as M. yuccae make the tent when they begin boring into the stem. As the larva grows, the tent is enlarged, and may be seven or eight centimeters long by pupation time. All during the feeding period, frass is expelled from the tip of the tent, but rather than being propelled away, the pellets are simply dropped from the opening.
Megathymus streckeri and Megathymus cofaqui are
somewhat different in larval behavior. These species bore directly into the stem after emerging from the egg, and pack the frass behind as they feed. When ready to pupate, the larvae burrow from the roots upward through the soil and make a short tent of silk, detritus, and soil.
Mature larvae of Megathymus species are brownish or
cream-colored with reddish brown or black heads. Wax glands on the ventral side of abdominal segments 7 and 8 supply copious quantities of wax to line the tunnel. Overwintering occurs in the larval stages. Megathymus yuccae hibernates as a mature larva and pupates in the spring without further feeding.
The tunnels may extend several feet into the stem and roots of the plant and although the terminal bud is killed, the damage done by the larva usually has a pruning effect, causing the plant to produces new shoots (Remington 1959).




47
Large species such as Megathymus ursus may inflict enough damage to kill the host (Stallings and Turner 1956).
The pupae of Megathymus species are brown and lack silk supports. The tip of the abdomen is freely movable, allowing the pupa to move up and down in the tunnel. When disturbed, they quickly retreat to the bottom of the burrow. Before emergence, they usually position themselves just below the tent. Tinkham (1954) speculated that these movements may serve to regulate temperature as well as retreat from disturbances. Megathymus larvae defend themselves by biting with the jaws, locking themselves in the burrow, defecating, and expelling a brownish fluid from the mouth (Wielgus and Stallings 1974).
After the adult emerges, it crawls upward and pushes its way through the loosely sealed opening to the tent. A yellowish meconium is expelled into the pupal exuvium. The adult usually perches on the tent while expanding and drying the wings. Megathymus species have only one or two generations per year, depending on the species.
Aegiale hesperiaris (Aegialini) and Agathymus species (Agathymini) feed only on century plants (Agave species). Aegiale hesperiaris glues small clusters of eggs to the host leaves (Dampf 1924). Agathymus species either perch on the host or nearby and drop or flick the eggs haphazardly (Comstock and Dammers 1934, Freeman 1951a) or lay the eggs on a host leaf. However, since no adhesive is applied, the




48
eggs fall to the ground (Roever 1964). Stallings et al. (1985) noted that females of Agathymus estelleae dropped between five and ten eggs at one time. Egg color changes from green to reddish in Agathymus species (Comstock and Dammers 1934, Roever 1975). One species, Agathymus estelleae, produces a range of egg colors varying from green to beige to pinkish (Stallings and Stallings 1986). The duration of the egg stage is from 22 to 52 days (Roever 1964).
Newly emerged larvae do not eat the remains of the egg, but crawl toward the distal half of a leaf and make a short burrow directed toward the tip (Roever 1964). Agathymus larvae spin a trail of silk when moving about on the leaf surface. The burrow is also lined with silk to prevent sap from entering, which is not always successful, forcing the larva to move to another site or another plant. The frass produced by young larvae is relatively dry and green (Roever 1964).
After several molts, the larva passes the winter within the burrow, then moves to the base of the leaf and makes a new tunnel directed toward the stem. This tunnel is also lined with silk except at the distal end, where the larva feeds primarily on sap. The entrance may be on the upper side, lower side or either surface of the leaf, depending upon the species. Defecation occurs at the tunnel entrance, after which the entry is partially barred by a few strands




49
of silk. The oily, dark brown frass accumulates around the entrance to the tunnel during the summer feeding period. Aegiale hesperiaris, Agathymus evansi, and Agathymus stephensi cause harm to the plant, as evidenced by necrotic tissue near the point of entry.
Agathymus larvae have dark heads and greenish or bluish white bodies. When feeding is completed, the entire tunnel is lined with silk and powdered with wax, except for Aegiale hesperiaris and Turnerina hazelae, which do not develop wax glands. Instead of constructing a tent over the tunnel entrance as do Megathymus, Agathymus exude a pliable silk from the mouth and fashion it into an opaque finger naillike door. The silk door completely seals the tunnel entrance.
Pupation occurs in the bottom of the tunnel. The brown pupa is not supported by silk and is oriented with the head toward the entry. At the time of emergence, the adult crawls from the exuvium and pushes on the trap door, which swings open on a hinge along the bottom edge. The adult then exits the tunnel, expands and dries the wings, and flies away.




CHAPTER 4
GENERAL MORPHOLOGY
This chapter describes external morphological
characteristics of the eggs, larvae, and pupae of U. S. skippers. Many structures exhibit a wide range of variation between the various species. Morphological variation is further noted in the species diagnoses of Chapter 5.
The eggs of hesperiids found within the U. S. vary from
0.4 to 1.8mm in height and 0.6 to 3mm in diameter. Hesperiid eggs are wider than high. Shape varies from hemispherical (vertical sides) to somewhat conical (sloping sides) (Figure 3). Numerous minute holes (aeropyles) occur on the surface of the chorion. The micropylar area at the top of the egg is surrounded by rosette patterns of sculpturing (Figure 4). Scudder (1875) first noted that the sculpturing of hesperiid eggs seemed to vary consistently between subfamilies. Pyrgines have ribbed eggs (Figure 3) while hesperiines have smooth eggs or eggs with polygonal patterns of sculpturing (Figure 5). While this pattern generally holds true, some Pyrginae, such as Cogia outis, have polygonal sculpturing. Other pyrgines have vertical rows of short spines. Megathymines, like hesperiines, have




51
non-ribbed eggs with polygonal sculpturing patterns. Heteropterine eggs are described as having vertical grooves (Higgins 1975); however, a Butleria species has ribbed eggs (Herrera et al. 1991), and the egg of Carterocephalus palaemon is smooth with polygonal sculpturing (Brooks and Knight 1982). I could not find specimens, detailed descriptions, or illustrations of pyrrhopygine eggs.
Larvae
Hesperiid larvae, like the caterpillars of other
lepidopterans, have a head, three thoracic segments, and ten abdominal segments. Each thoracic segment bears a segmented pair of legs. In addition, abdominal segments three through six and ten each have a pair of unsegmented prolegs. Spiracles are located on the prothorax and abdominal segments one through eight. The spiracles on the prothorax and abdominal segment eight are usually slightly larger than the others. The body segmentation of hesperiids tends to be ill defined and confused by folds or wrinkles of the cuticle (annuli or plicae).
Mature larvae of U. S. hesperiids range in length from
8 to 86mm. The prothorax is larger than the head in Megathyminae, about the same size as the head in Pyrrhopyginae, and generally smaller than the head in the Pyrginae and Hesperiinae. Body width tends to be more or less uniform. However, the thorax of Megathymus species is slightly enlarged. These plant boring larvae are able to




52
expand the thoracic region and thereby lock themselves into their tunnels upon being disturbed by a predator or human observer. Other Megathyminae are somewhat wider in the area of abdominal segments three to five. The transverse width of the U. S. hesperiids varies from 1.6 to 11.7mm. Hesperiines tend to be more slender than pyrgines.
The setae occurring on skipper larvae exhibit variation in shape (Figure 6). Although simple setae are the most abundant type, some species also have modified setae. Modified setae vary in the shape of the tip, which may be blunt, slightly expanded, or greatly expanded. Pyrgines and some hesperiines frequently have setae with expanded tips. Branching setae are another modified type found only on the head of certain pyrgines.
The length of the setae on the body and head varies between species. On an individual caterpillar, the setae are usually longest toward the ventral side of the body and shortest on the dorsum. The length of the setae on the dorsum of abdominal segment four varies from < 0.1 to 6.4mm for U. S. species. On the head, setae range from < 0.1 to 5mm near the apex to 0.1 to 5.7 on the ventral side. Pyrrhopygines have very long setae. Most other hesperiids have short or very short setae, but a few pyrgines and hesperiines have long or moderately long setae.
Lenticles are small sclerotized plates that seem to represent modified setae. Franzl et al. (1984) found the




53
lenticles of Calpodes ethlius to be innervated secretory structures that probably produce lipids. The positions of the lenticles change with each larval molt (Franzl et al. 1984), alternating between two basic patterns. The expression of lenticles at a particular location may not always occur or they may be present in excess. Not only is there variation between individuals, the number of lenticles often varies from side to side of a single larva. Several larvae usually need to be examined in order to determine the pattern for a particular species. A generalized map of lenticle distribution is presented in Figure 7. Table 2 lists the maximum number of lenticles observed in the present study at eight positions on the prothorax, five positions on thoracic segments two, three, and abdominal segments one through nine, and two positions on abdominal segment ten for subfamilies Pyrrhopyginae, Pyrginae, Heteropterinae, and Hesperiinae. Megathymines also have lenticles, but usually in a supraspiracular cluster or in small groups on the ventral side of abdominal segments one, two, seven, or eight (Figure 8). Table 3 presents the number of lenticles present on ten megathymine taxa.
Pigments that may be present in the cuticle of
hesperiid larvae are red, yellow, orange, white, or rarely black. Unfortunately, most of these pigments are destroyed in preserved larvae. Literature descriptions are often unreliable as they may refer to wax deposits, cuticular




Table 2. Maximum number of lenticles observed per segment and position on Hesperiidae larvae in the present study. Lenticles may be found in eight possible positions on the prothorax (Tl), five positions on thoracic segments 2-3 and abdominal segments 1-9, and two positions on the last segment (AO10). An asterisk indicates 10 to 15 lenticles present.
MAXIMUM # LENTICLES PER POSITION
T1 T2-3 A1-2 A3-6 A7-9 AO10

SPECIES

12345678 12345 12345 12345 12345 12

PYRRHOPYGINAE: P. araxes
PYRGINAE: P. pigmalion P. palemon E. zestos E. clarus E. exadeus P. leo C. catillus T. undulatus P. mexicanus P. procerus C. alcaeus C. arizonensis U. proteus U. esmeraldus U. dorantes U. teleus U. procne A. fulgerator A. gilberti A. anaphus A. cellus A. lyciades A. casica T. bathyllus T. pylades T. diversus T. mexicanus T. confusis T. drusius C. potrillo C. hippalus C. outis C. caicus

10001010 00100 00114 10120 00122 10

10001020 10001140 10001120 10001120 10001120 10001130 10001120 10001020 10100213 10001244 10001020 10001020 10001120 10001120 10001120 10001122 10001110 10001120 10001120 10001120 10001120 10001120 10001120 10001120 10001120 10000120 10001100 10001122 10001121 10001020 10001020 10101020 10001120

01010 01120 11110
10100 11014 1300* 11000
10014 1110*
1210* 11100
01100 11000 11100 11100
01003 11110 11100
01000 01100 11100 11000 11000
01110 12113 01000 01000 12104 11012 11000
10000 10002 00000

10112 10110 10111 10111 10111 10111 10111 10021 10111 10112 10113 10113 10111 10022 10111 10111 10011 10121 10121 10121 10121 10011 10111 10111 10112 10011 00011 10111 10111 10111 10011 10012 10011

10110 10110 10121 10021 10111 10111 10111 10021 10122 10113 10122 10113 10111 10121 10111 10112 10111 10121 10121 10121 10121 10011 10111 10111 10111 10111 10011 10111 10111 10111 10011 10122 10021

00110 10110 10111
10111 10111 10111 10011
10021 10121 10111 10122 10112 10111 10011 10111 10112 10111 10121 10121 10111 10121 10011 10111 10111 10112 10111 10111 10111 10111 10111 10012 10012 10011




Table 2--continued.

MAXIMUM # LENTICLES PER POSITION T1 T2-3 A1-2 A3-6 A7-9 A10 12345678 12345 12345 12345 12345 12

N. rubescens
S. ceos S. mazans S. hayhurstii C. canescens X. trixus S. pulverulenta S. zampa A. thraso T. ruptifasciatus G. gesta E. brunneus E. icelus E. brizo E. juvenalis E. propertius E. horatius E. tristis E. martialis E. pacuvius E. zarucco E. funeralis E. baptisiae E. persius E. afranius P. centaureae P. ruralis P. scriptura P. communis P. albescens P. oileus H. ericetorum H. lavianus H. macaira H. arsalte C. nessus C. limpia P. catullus P. alpheus
HETEROPTERINAE: P. pirus

10001111 10001110 10001112 10001113 10001110 10001110 10002120 10001120 00001110 00001120 10001120 10001100 10001110 10001110 10001010 10001110 10001110 10001110 10001010 10001010 00001110 10001110 10001110 10001110 10001110 10001120 10001110 10000110 10001110 10001110 10001110 10001110 10001110 10001110 10001110 10001120 10001120 10001111 00001110

00110110 03100 00112 00117 00121 10

SPECIES

12010 00100 02101 02100 11010
01110 11000
11010 10000 01110 12100 11010
12010 12010 12110 12010 11010
12110 02100 12100 12110 02100 12100 12100 12100 12110 11000 11010 11110
12010 02010 02010 00010 00000 12010 12010 12110 01100 01000

10021 10111 00111 00121 10121 10021 10122 10021 10021 10122 10112 10012 10122 10123 10122 10122 10122 10122 10122 10122 10122 10122 10122 10122 10122 10021 10121 10111 10111 10111 10111 10111 10021 00111 10111 00111 10111 10111 00110

10022 10111 10111 10111 10121 10021 10110 10120 10111 10121 10111 10021 10121 10121 10121 10120 10121 10121 10121 10121 10111 10121 10111 10121 10111 10121 10011 10121 10111 10111 10111 10111 10021 10111 10111 10111 10111 10111 10111

10021 10111 00121 10121 10121 10021 10120 10120 10111 10121 10111 10022 10121 10111 10111 10110 10110 10111 10111 10111 10111 10111 10111 10121 10111 10121 10111 10121 10111 10111 10111 10111 10121 10111 10111 10111 10111 10111 10100




Table 2--continued.

MAXIMUM # LENTICLES PER POSITION T1 T2-3 Al-2 A3-6 A7-9 A10 12345678 12345 12345 12345 12345 12

HESPERIINAE: S. malitiosa V. perigenes N. lherminier N. julia N. neamathla C. tripunctus L. accius L. liris P. philetes A. numitor A. arene C. aurantiaca C. minimus T. lineola H. phyleus Y. carus P. eunus H. uncas H. juba H. comma H. woodgatei H. leonardus H. pahaska H. viridis H. attalus H. meskei P. coras P. sabuleti P. mardon P. draco P. baracoa P. themistocles P. origenes P. mystic P. sonora P. vibex W. otho W. egeremet P. verna A. campestris A. arogos A. delaware P. byssus

00100010 02000 01112 01118 00112 11 00100000 10110 10011 1001* 10111 01 00000010 02100 01111 11116 11111 10 00000110 02100 00101 00101 00112 10 00000110 02100 01121 01116 00111 10 00100110 03100 01112 01117 01111 01 00100110 03010 01112 01119 01111 01 00101110 03010 01112 01118 01111 01 00000010 02010 01112 01113 01112 11 00101020 12010 01121 10123 11111 17 00001120 02000 01111 01111 01111 14 00000000 01000 00102 00104 00110 01 00101020 01000 00011 00103 00111 11 00101020 01000 00111 01116 01111 01 00101120 12010 01121 01113 01111 10 00101120 02000 01111 01114 01111 10 00001010 11010 10011 10013 10011 01 00101010 11010 10011 10018 10011 12 10120010 02010 01111 11116 10111 11 10110120 02100 01111 01119 00111 11 10121110 10010 10011 10015 10011 11 00101120 02100 01111 01116 01111 11 00101120 12010 11111 11116 11111 11 00100020 11010 10011 10015 10111 11 00101010 11010 10011 10017 10111 11 00100010 02010 01111 01113 01111 11 10101110 12010 10111 11118 11111 11 00101010 01010 11111 10123 11021 11 00101010 13010 11111 11113 11111 01 10110120 02000 10011 1111* 01011 11 00000120 03000 01111 01115 01111 10 10121120 03100 01111 01115 01111 10 10140020 02100 01111 11117 01111 10 10120020 03000 01111 01110 01111 10 10121020 12110 01011 11110 01111 10 10110120 02100 01111 01114 11111 10 10101110 13010 11111 11116 11111 11 00000120 02000 01111 01117 01111 10 11111010 11010 10011 10018 10111 11 11121120 02000 01111 01116 01111 10 00001110 12100 11112 11117 11112 00 10111120 02000 00112 00113 10111 01 00101120 02110 01112 0111* 01112 10

SPECIES




Table 2--continued.

MAXIMUM # LENTICLES PER POSITION T1 T2-3 A1-2 A3-6 A7-9 AO10 12345678 12345 12345 12345 12345 12

O. sylvanoides
0. agricola
0. yuma P. hobomok P. zabulon P. taxiles P. aaroni P. yehl P. viator P. melane C. haitensis E. alabamae E. arpa E. pilatka E. dukesi E. bimacula E. ruricola A. capucinus A. hianna A. loammi A. aesculapias A. vialis A. celia L. eufala
0. maculata C. ethlius P. panoquin P. panoquinoides P. ocola P. sylvicola N. nyctelius

10150020 03100 01122 01117 01111 11 10130020 02100 01111 01119 01111 11 10140120 02000 01111 01110 01111 11 11141110 11010 10001 1001* 10011 11 10130000 03000 01111 01118 01111 11 10101220 11010 10021 10029 10021 11 00131110 11010 11011 1101* 11111 11 12141110 12010 11111 1111* 11111 11 031*1120 12110 11111 11118 10111 01 10141020 02110 01111 01117 01111 11 10110120 01100 11111 11116 01111 01 00101130 02100 01011 01119 01111 11 00100030 12110 11112 1111* 11111 11 00101130 12110 11111 11118 11011 11 00101030 12110 11111 11117 11111 01 00101130 02100 01112 01115 01111 11 00101130 02100 01112 01116 01111 11 10111010 11010 11123 11119 11232 01 00100110 02100 01212 0111* 01112 01 10130120 02100 01212 0111* 01112 01 00111010 11000 10011 10010 10111 11 10120110 03100 00112 00119 00112 10 00100110 02100 00122 00116 00111 10 00100110 03000 01112 01115 01111 11 00001110 12110 11112 11118 11111 11 00102020 05000 01112 01117 01112 01 00101000 11110 11111 1111* 10111 10 00101020 01010 01012 01114 01112 11 00101020 01010 01111 01113 01111 11 00101010 01110 01111 01115 01111 11 11121020 02000 01111 01111 01111 11

SPECIES




Table 3. Maximum number of lenticles observed per segment and position on Megathyminae larvae in the present study. Lenticle positions are labelled 1 for lenticles above the spiracle line, 2 for lenticles below the spiracle line, and 3 for ventral lenticles. Position 1 on abdominal segment 10 refers to lenticles on the lateral margin of the caudal proleg. An asterisk indicates 10 to 15 lenticles present.
MAXIMUM # LENTICLES PER POSITION
Tl T2-3 A1-2 A3-6 A7-9 A10

SPECIES

123 123 123 123 123 1

A. valverdiensis A. stephensi M. y. yuccae M. y. martini M. y. reinthali M. coloradensis
kendalli
M. coloradensis
navajo
M. cofaqui M. u. ursus M. u. violae

300 200 100 100
000

*00 800 300 120 200

700 400 612 813 515

400 400 910 610 800

500 301
*02 402 802

000 200 503 700 802 0

000 000 010 000

300 900 300 210

903 304 602 302

800
*00 500 900

601
*01 402 702




59
thickenings, internally derived colors, or may not have been included in the description. Megathymines lack bright cuticular pigments, but may be cream-colored, bluish or greenish from chlorophyll and other pigments inside the body. Red colors seem to occur only in pyrgines and pyrrhopygines. The ventral prothorax of some pyrgines such as Epargyreus clarus is often bright red or reddish-brown in color. Yellow or orange pigments occur in many pyrgines and some hesperiines. White or cream-colored cuticular pigments are more broadly distributed in hesperiines. The only skipper larva that I have seem with extensive black pigment is Codatractus hyster.
Colors on the body of hesperiid larvae are most
commonly distributed as stripes (Figure 9). Usually the stripes occur longitudinally, but Pyrrhopygines and some pyrgines have transverse stripes of yellow or white. Transverse stripes seem to occur on unpalatable lepidopteran larvae such as the sphingid, Pseudosphinx tetrio.
Longitudinal stripes typically occur only in certain locations such as a heart line, a subdorsal line, or a lateral line. The heart line is always present because the cuticle above the heart tends to be less pigmented than elsewhere on the body. The subdorsal line is usually a narrow stripe that may sometimes be interrupted to form a line of spots. The lateral stripe is less frequently present, but may form a broad band across the side of the




60
body. In some species the body is covered with tiny white or yellowish spots and the stripes are formed by the absence of these colors. In other cases the stripes are outlined with white or yellow, forming double bands (Figure 9B). Some lines or spots on pyrgines such as Epargyreus and Urbanus may also be present due to patches of black microspines (Figure 9D). The texture of the cuticle of most hesperiids is rough, particular on pyrgines.
The larval head consists of several sclerites. The
head capsule is divided into two major sections (epicrania) by the frontal and adfrontal sclerites and the epicranial suture along the vertex. The front varies in shape from nearly an equilateral to an isosceles triangle, usually the latter. Below the frontal sclerite lies the clypeus and next the labrum. The labrum of hesperiids is small with a shallow mesal notch. Along the posterior margin of the head is the postocciput. This sclerite is normally a very narrow band in most Lepidoptera, but is frequently very wide or moderately wide in the Hesperiidae. In pyrgines, the postocciput is usually very wide to moderately wide, lending in part to the "narrow neck" often referred to by authors (Figure 10). The postocciput is narrow to occasionally moderately wide in hesperiines (Orthos lycortas, Paratrytone melane) as well. Megathymines have a narrow postocciput coupled with a large prothorax, creating a "wide neck"




61
(Figure 10D). The transverse width of the head capsule of U. S. skippers varies from 1.5 to 7.2mm.
The head capsule of hesperiids usually has a rough texture due to polygonal patterns of sculpturing (Figure 11). The sculpturing may form small pebbly knobs in pyrgines. These knobs are sometimes enlarged and modified into small spines at the apex of the epicranium (Figure 12). The ridges of the polygonal sculpturing may also be greatly widened, except for the center, resulting in a pattern of pits in hesperiines. Pyrrhopygines have a unique and characteristic series of low vertical ridges on the face (Figure 11D). The head capsule of Copaeodes aurantiacus, a hesperiine, has two forward pointing processes, but its close relative, Copaeodes minimus, does not.
Klots (1966) named and described paraclypeal hooks on the larva of Amblyscirtes samoset. The hooks are downward projecting spines, one on each side of the clypeus (Figure 11E). Their function remains unknown. I have seen this structure only in Amblyscirtes species (Hesperiinae) and Piruna pirus (Heteropterinae).
Hesperiids have six simple eyes or stemmata on the lower part of each epicranium (Figure 13). The relative size and spacing of the stemmata often varies between species. Stemma one through four lie in an arc and are usually equally spaced. Stemma five and six are usually farther apart.




62
The labial-submental complex occupies a relatively large portion of the ventral head of most lepidopteran larvae. In hesperiids, however, the labial-submental complex is small relative to the size of the head, except for megathymines, which have large labial-submental complexes.
The mandibles of hesperiines and heteropterines are articulated somewhat differently from other skippers. Hesperiine mandibles are longer in the dimension between the labrum and labium than in the dimension between the antennae (Figure 14). The articulation supporting this longer mandible is also elongate. Pyrgine mandibles are subequal in these dimensions. Most hesperiids lack teeth or at most have shallow lobes on the mandibles. Well developed-teeth are present in some pyrgines and megathymines.
Hesperiid larvae have a pair of short antennae that
arise from a membranous area adjacent to the mandibles. The antenna consists of three segments, the distal segment being very small. A long sensory seta occurs on the tip of the second segment.
Color patterns on the heads of skipper larvae are often striking. The pattern may be light spots or stripes on a dark ground, dark spots or stripes on a light ground, or variegated. Pyrgines frequently have spots on the face, and sometimes a lateral line. Hesperiines often have stripes on the face. The stripes usually occur in specific areas. The




63
medial stripe proceeds from the vertex along the epicranial suture, then branches and proceeds partway along the adfrontal sclerites. The epicranial line proceeds from near the apex to the lower part of the face or may be broadly joined with the medial stripe along the adfrontals. The lateral stripe proceeds from the vertex, along the lateral margin of the head to the stemmatal area. Two skipper groups have uniformly colored heads, megathymines which are mostly concealed in stems, leaves, or roots, and pyrrhopygines, which have dense long setae on the head. In addition, pyrgines sometimes have two brightly colored (yellow, orange, or red) false eye patches on the lower face between the stemmata and the mandibles. Eye patches also occur in hesperiines, but are smaller and less colorful.
The dorsum of the prothorax is often sclerotized and darkened in larval hesperiids. The prothoracic shield may be relatively wide, covering much of the dorsum, reduced to a narrow band between the major annuli, or indistinct. The shield usually extends laterally onto a small hump. The hump is often darkened, but separated from the dorsal part of the shield in third and fourth instar larvae. The hump also frequently bears a lenticle and a pinaculum, the seta of which is longer and more slender than the surrounding setae. Another pinaculum is present in a more dorsal position on the mesothorax. A small cluster of lenticles usually accompanies the mesothoracic pinaculum as well. On




64
the ventral side of the prothorax lies the opening to a specialized gland, thought to produce defensive chemicals (Eisner et al. 1972). Most hesperiids have the ventral prothoracic gland, but I cannot find the opening in Megathyminae. The prothoracic spiracle often varies in shape between hesperiid species. Typically the spiracle is oval, but the anterior rim is frequently elongate in some species.
The color of the thoracic legs is often useful in
distinguishing species. In most skippers the legs are all pale or all dark. Others have only the first or second legs darkened. The thoracic leg length of hesperiines and pyrgines is usually moderate, but some species have relatively shorter or longer legs. Leg length seems to be partly correlated to larval ecology. Hesperiines that live near the ground, such as Hesperia species, often have long thoracic legs, whereas epiphytic species, such as Lerodea eufala, have very short legs. A few pyrgines have a row or small clusters of lenticles anterior to the legs.
Abdominal segments one and two of the larvae lack legs and are transitional to the thoracic segments. Segments three through six are similar and bear a pair of prolegs. Segments seven and eight lack legs and are transitional to the terminalia. Segment nine is reduced to a small annulus between segment eight and ten. Segment ten bears the anal prolegs and is modified for disposal of the frass.




65
The prolegs of segments three through six bear small hooks (crochets) in a circle, a near circle, a mesal penellipse, or transverse bands. The crochets are sometimes uniordinal or biordinal, but most often are in more or less three ranks (triordinal). The crochet band is always uniordinal near the lateral margin and quickly becomes biordinal and triordinal toward the meson. In some pyrgine genera such as Achalarus, Thorybes, and Cabares, the posterior or mesal crochets are nearly twice as large as the anterior series of each proleg. Only one species that I have examined, Polygonus leo, has ventral lenticles in small clusters anterior of the prolegs.
The last stage larvae of some hesperiines and
megathymines develop wax glands prior to pupation (Dethier 1942c). The wax produced by the glands is used to coat the inside of the cocoon. The gland pattern on the ventral surface of the larva varies considerably (Figure 15). The U. S. species with wax glands usually develop longitudinal or transverse patches on the ventral side of abdominal segments seven and eight (Figure 15A and B). The wax glands of Perichares philetes form a transverse patch on the ventral side of abdominal segment one. Synapte malitiosa has transverse patches on abdominal segments three through six. In Choranthus haitensis and Asbolis capucinus, these patches are reduced to small spots posterior of the prolegs.




66
Abdominal segment ten usually bears an anal comb. The anal comb is a fan-shaped, sclerotized structure located caudad and dorsal to the anus (Figure 16). During the act of defecation, the anal comb functions to propel the fecal pellet away from the caterpillar. This is accomplished with some force and the pellets may be launched several feet away. The base of the anal comb is wider in pyrgines than hesperiines. The anal comb is degenerate, but usually present, in megathymines, consisting of up to about dozen spines.
The suranal plate of most U. S. hesperiids is rounded, but the plate is pointed in a few hesperiines (Copaeodes spp.). Other hesperiines (Atrytone delaware, Polites spp.) have dark lines or markings on the suranal plate. The broad, bill-like plate of Asbolis capucinus is edged with black and the terminal spiracles are black, forming a false face.
Pupae
The pupae of U. S. hesperiids range in length from 10.5 to 52mm and in width from 2.1 to 10.3mm. The length of the abdomen varies between groups of hesperiids, being very short in species such as Epargyreus clarus and very long in Megathymus spp. The difference is due mostly to a change in the size of abdominal segments five through ten, as these lie beyond the wings. Most skipper pupae are brown or cream-colored, but some are green. Like their larvae,




67
hesperiid pupae have a head, three thoracic segments, and ten abdominal segments. The antennae, legs, and wings are visible, but are glued to the body. The prothorax and abdominal segments two through eight bear spiracles. The thoracic spiracle is usually guarded by a cluster of microspines. In most pyrgines, the spiracle guard is on a small cuticular rise, giving the pupa a false face appearance when viewed from the front. The distal tip of the abdomen is modified into the cremaster, a structure used to anchor the pupa to the substrate. The cremaster shows considerable variation between species and higher groups.
The setae on hesperiid pupae are usually short and
simple. A few pyrgines, however, have some bifurcate setae. A neotropical species, Arteurotia tractipennis, has long plumose setae on the head. Species with long larval setae generally have long setae on the pupae as well. Many hesperiines have very short or inconspicuous setae. Lenticles occur only on hesperiine pupae. The prothorax, dorsal abdomen, and ventral proleg scars of hesperiine pupae typically have lenticles (Figure 17).
The head of most hesperiid pupae is rounded; however,
some modifications do occur (Figure 18). A few pyrgines and many hesperiines have pointed processes on the cap. The point may be straight and relatively thick at the base (Lerema accius), slender and upturned (Calpodes ethlius), or T-shaped at the tip (Justinia phaetusa, a neotropical




68
species). Instead of being pointed, the cap may be bulbous as in Atrytone delaware and Poanes viator. The head also bears a pair of sclerites (pilifers) which represent the mandible remnants. Pyrrhopygines, pyrgines, and megathymines always have the pilifers separated, whereas, most hesperiines have the pilifers touching.
The position of the antennal tip relative to the tip of the middle leg on the pupae of hesperiids shows a good deal of variation. The antennae extend to the tip of the middle leg in Pyrrhopygines, and many pyrgines. In other pyrgines, most hesperiines, and megathymines the distal tip of the antenna lies far cephalad of the tip of the middle leg. Likewise, the proboscis usually reaches to the wing tips in pyrrhopygines and pyrgines, extends far beyond the wing tips in hesperiines, and lies far cephalad of the wing tips in megathymines. The distal part of the proboscis case is not glued to the body if this structure extends beyond the wings.




Figure 3. SEM micrographs of the egg of Urbanus proteus
(A), Telemiades epicalus (B), Thorybes pylades (C), Polites vibex (D), Hylephila phyleus (E), Atrytone delaware (F), Euphyes ruricola (G), Copaeodes minimus (H), and Megathymus cofaqui (I).




70
. 3 8 ....... ........... ..................... ..i....i..
3 i m i ii .43iiiiiiiii~~i~l~l m m.
is:!iil .38 mm I 8




Figure 4. SEM micrographs of the micropylar region of the egg of Urbanus proteus (A), Telemiades epicalus (B), Erynnis horatius (C), Hesperia attalus (D), Polites baracoa (E), Atrytone delaware (F), Problema byssus (G), Poanes yehl (H), and Euphyes ruricola (I).




72
A D
30 microns
30 microns E
30 microns




Full Text

PAGE 1

IMMATURE STAGES OF THE SKIPPER BUTTERFLIES (LEPIDOPTERA: HESPERIIDAE) OF THE UNITED STATES; BIOLOGY, MORPHOLOGY, AND DESCRIPTIONS BY MARC C. MINNO A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1994

PAGE 2

ACKNOWLEDGMENTS Numerous biologists loaned specimens or contributed data for my studies of immature hesperiids. My sincerest appreciation goes to Andrew Atkins, H. David Baggett, Greg Ballmer, Susan S. Borkin, John W. Brown, John M. Burns, Hideyuki Chiba, Frank C. Elia, John F. Emmel Thomas C. Emmel Hugh Avery Freeman, Dale H. Habeck, Donald W. Hall, Donald J. Harvey, J. Richard Heitzman, John B. Heppner, Timothy L. McCabe, Noel McFarland, Jacqueline Y. Miller, Steven Passoa, Gordon F. Pratt, Dale F. Schweitzer, and Jeffrey R. Slotten for their contributions. Special thanks go to Roy 0. Kendall, who has been especially supportive and generously loaned specimens, many being unique, from his personal collection. The United States National Museum, Florida State Collection of Arthropods, and the Santa Barbara Natural History Museum also loaned preserved specimens from their collections. Host plant specimens were identified by David W. Hall and Kenneth R. Langdon. Peter J. Eliazar and Andrei Sourakov assisted with the scanning electron microscope. Many thanks go to the librarians at the University of Florida and the Florida Department of Agriculture, Division of Plant Industry and Consumer ii

PAGE 3

Services who helped tremendously in locating a number of obscure references. I also heartily thank my committee members, Lincoln P. Brower, Gary R. Buckingham, Dale H. Habeck, Jonathan Reiskind, and Chairman Thomas C. Emmel for their patient guidance, support, and critigue of my graduate work. This research was supported in part by a small grant from the Florida Entomological Society. In conclusion, my greatest appreciation goes to my loving wife, Maria Frances Minno, for her support and help during the course of this research. 111

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TABLE OF CONTENTS Page ACKNOWLEDGMENTS ii LIST OF TABLES v i LIST OF FIGURES v ii ABSTRACT xxvi CHAPTERS 1 INTRODUCTION x 2 METHODS 5 Study Material 5 Rearing 7 Preservation and Preparation 11 Descriptive Methods 12 3 BIOLOGY OF NORTH AMERICAN HESPERIIDAE 15 Biology of the Pyrrhopyginae 31 Biology of the Pyrginae 31 Biology of the Heteropterinae 38 Biology of the Hesperiinae 38 Biology of the Megathyminae 44 4 GENERAL MORPHOLOGY OF THE IMMATURE STAGES ... 50 Eggs 50 Larvae 51 Pu P ae .....*!!!!!.*!!!!!! 66 IV

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5 DIAGNOSES AND HOST PLANTS FOR THE UNITED STATES SKIPPERS 88 Subfamily Pyrrhopyginae 88 Subfamily Pyrginae 90 Subfamily Heteropterinae 204 Subfamily Hesperiinae 206 Subfamily Megathyminae 328 6 TAXONOMIC KEYS TO THE IMMATURE STAGES 453 Preliminary Key to Eggs of U. S. Hesperiidae (Subfamily and Genus) 453 Preliminary Key to Larvae of U. S. Hesperiidae (Subfamily and Genus) 455 Preliminary Key to Pupae of U. S. Hesperiidae (Subfamily and Genus) 463 7 SUMMARY AND CONCLUSIONS 468 Biology 468 Morphology 471 Taxonomic Insights and Problems 475 REFERENCES 47g BIOGRAPHICAL SKETCH 508

PAGE 6

Table LIST OF TABLES page Number of species and subfamilies of U. S. Hesperiidae recorded from various host plant families 16 Maximum number of lenticles observed per segment and position on Hesperiidae larvae in the present study. Lenticles may be found in eight possible positions on the prothorax (Tl), five positions on thoracic segments 2-3 and abdominal segments 1-9, and two positions on the last segment (A10). An asterisk indicates 10 to 15 lenticles present 54 Maximum number of lenticles observed per segment and position on Megathyminae larvae in the present study. Lenticle positions are labelled 1 for lenticles above the spiracle line, 2 for lenticles below the spiracle line, and 3 for ventral lenticles. Position 1 on abdominal segment 10 refers to lenticles on the lateral margin of the caudal proleg. An asterisk indicates 10 to 15 lenticles present 58 VI

PAGE 7

LIST OF FIGURES Figure page 1 Larval shelter of Urbanus proteus on Phaseolus vulgaris (A) Polygonus leo on Piscidia piscipula (B), Erynnis zarucco on Sesbania macrocarpa (C), Erynnis brizo on Quercus myrtifolia (D), Wallengrenia otho on Paspalum sp. (E), and Asbolis capucinus on Sabal sp. (F) 20 2 Larval shelter of Euphyes pilatka on Cladium jamaicense (A), Euphyes arpa on Sabal etonia (B), Megathymus yuccae on Fucca aloifolia (C), and an agave leaf cut open to reveal the tunnels of Agathymus aryxna (D) 22 3 SEM micrographs of the egg of Urbanus proteus (A), Telemiades epicalus (B), rhoryhes pylades (C), Polites vibex (D), Hylephila phyleus (E), Atrytone delaware (F), Euphyes ruricola (G), Copaeodes minimus (H), and Megathymus cofaqui (I) 70 4 SEM micrographs of the micropylar region of the egg of Urbanus proteus (A), Telemiades epicalus (B) Erynnis horatius (C), Hesperia attalus (D), Polites baracoa (E), Atrytone deiai/are (F), Problema byssus (G), Poanes yehl (H), and Euphyes ruricola (I)... 72 5 SEM micrographs of the sculpturing on the sides of the egg of Atalopedes campestris (A), Polites vibex (B) Atrytone delaware (C), Problema byssus (D) Euphyes ruricola (E) and Megathymus cofaqui (F) 74 6 Comparison of setae on the body of Pholisora catullus (A), the head of Achalarus casica (B), the body of P. catullus (C), the body of Urbanus proteus (D), the body of Ancyloxypha numitor (E), the body of Hesperia juba (F), the head of Xenophanes trixus (G), the head of Pyrgus communis (H), the body of Thorybes bathyllus (I), and the body of Cogia outis (J) 75 vii

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Figure page 7 Generalized map of the lenticles and other features of the last instar hesperiid larva. 76 8 Lenticles on the fourth abdominal segment of Pholisora catullus (A), Problema byssus (B), and Megathymus yuccae (C) 77 9 Color patterns on the fourth abdominal segment of some U. S. hesperiids: dorsal view of transverse stripes on Astraptes fulgerator (A), lateral view of subdorsal and lateral stripes bounded by white (shaded portion) on Copaeodes aurantiacus (B), lateral view of the interrupted subdorsal stripe of Codatr actus arizonensis (C) and lateral view of patches of black microspines on Epargyreus clarus (D) 78 10 Relative size of the "neck" of last instar larvae of Pyrrhopyge araxes (A), Urbanus teleus (B), Problema byssus (C), and Megathymus yuccae (D) 79 11 Features of the larval head of hesperiids: pebbly sculpturing of Polythrix procera (A), rough sculpturing of Staphylus mazans (B), pitted sculpturing of Hesperia pahaska (C), ridges on the head of Pyrrhopyge chalybea (D), and general structures on Amblyscirtes vialis (E) 80 12 Processes on the head of Urbanus procne (A), Achalarus lyciades (B), and Copaeodes aurantiacus (C) 81 13 Comparison of the stemmata of Pyrrhopyge araxes (A), Achlyodes thraso (B), and Euphyes pilatka (C) 82 14 Ventral view of the hesperiine-type mandibles of Atalopedes campestris (A) and the pyrgine-type mandibles of Erynnis horatius (B) 83 15 Wax gland distribution on Problema byssus (A), Lerema accius (B) Synapte malitiosa (C), Asbolis capucinus (D), Hesperia juba (E), Oc hi odes agricola (F), and Per ic hares philetes (G) 84 vm

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Figure page 16 Comparison of the anal comb of Urbanus proteus (A), Thorybes mexicanus (B), Pyrgus scriptura (C) Polites themistocles (D) Pseudocopaeodes eunus (E), and Megathymus yuccae (F) 85 17 Lenticle distribution on the prothorax of the pupa of Pseudocopaeodes eunus (A), on the dorsal abdomen of Hylephila phyleus (B), and on the proleg scars of Yvretta carus (C) 86 18 Lateral view of the pupal cap of Perichares philetes (A), Atrytone delaware (B), Panoquina panoquin (C) and Calpodes ethlius (D) 87 19 Last instar larva of Autochton cellus (A) Epargyreus clarus (B), Thorybes py lades (C), Urbanus proteus (D), Urbanus dor antes (E), Polygonus leo (F) Gesta gesta (G) and Erynnis horatius (H) 343 20 Last instar larva of Achlyodes thraso (A) Pholisora catullus (B), Pyrgus oileus (C), Staphylus hayhurstii (D) Hesperia attalus (E), Pompeius verna (F), Wallengrenia egeremet (G), and Atrytonopsis hianna (H) 345 21 Last instar larva of Atrytone delaware (A), Problema byssus (B), Asbolis capucinus (C), Euphyes arpa (D), Lerema accius (E) Panoquina panoquin (F), Lerodea eufala (G), and Megathymus yuccae (H) 347 22 Frontal view of the larval head of Phocides palemon (A), Epargyreus clarus (B) Polygonus leo (C) Chioides catillus (D), Typhedanus undulatus (E), and Polythrix mexicana (F). All drawings are enlarged 12 times 348 23 Frontal view of the larval head of Polythrix procera (A), Codatractus alcaeus (B), C. arizonensis (C), Urbanus proteus (D) Urbanus dorantes (E), and Astraptes fulgerator (F). All drawings are enlarged 12 times 34 9 IX

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Figure page 24 Frontal view of the larval head of Astraptes gilberti (A), Autochton cellus (B), Achalarus lyciades (C), Thoryjbes bathyllus (D), Cabares potrillo (E), and Cogia hippalus (F). All drawings are enlarged 12 times 350 25 Frontal view of the larval head of Nisoniades rubescens (A), Staphylus ceos ( B ) Carrhenes canescens ( C ) Xenophanes trixus (D), Systasea pulverulenta (E), Achlyodes thraso (F), Timochares ruptifasciatus (G), and Gesta gesta (H). All drawings are enlarged 12 times 351 26 Frontal view of the larval head of Ephyriades brunneus (A), Erynnis icelus (B), E. brizo (C), E. juvenalis (D), E. propertius (E), and E. horatius (F). All drawings are enlarged 12 times 352 27 Frontal view of the larval head of Erynnis tristis (A), E. martialis (B), E. pacuvius (C) E. zarucco (D), £. funeralis (E), £. baptisiae (F), E. afranius (G) and E. persius (H) All drawings are enlarged 12 times 353 28 Frontal view of the larval head of Pyrgus centaureae (A), Heliopetes ericetorum (B), Celotes nessus (C) Pholisora catullus (D), Piruna pirus (E), Synapte malitiosa (F), Vidius perigenes (G), and Nastra lherminier (H). All drawings are enlarged 12 times.... 354 29 Frontal view of the larval head of Cymaenes tripunctus (A), Lerema accius (B), Perichares philetes (C), Ancyloxypha numitor (D), A. arene (E) Copaeodes aurantiacus (F) C. minimus (G) and Thymelicus lineola (H) Drawings A-C and H are enlarged 12 times; D-G are enlarged 25 times 30 355 Frontal view of the larval head of Hylephila phyleus (A), Yvretta carus (B), Pseudocopaeodes eunus (C), Hesperia uncas (D), H. leonardus (E), H. pahaska (F), Polites cor as (G), and P. sabuleti (H) All drawings are enlarged 12 times 356

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Figure page 31 Frontal view of the larval head of Polites mardon (A), P. draco (B) P. baracoa (C) P. themistocles (D), P. origenes (E), P. mystic (F), P. sonora (G) and P. vibex (H). All drawings are enlarged 12 times.... 357 3 2 Frontal view of the larval head of Wallengrenia otho (A), Pompeius verna (B), Atalopedes campestris (C), Atrytone arogos (D) A. delaware (E), Problema byssus (F), Ochlodes sylvanoides (G), and Ochlodes agricola (H) All drawings are enlarged 12 times 358 3 3 Frontal view of the larval head of Ochlodes yuma (A), Poanes hobomok (B), P. zabulon (C), P. taxiles (D), P. aaroni (E), P. yehl (F), P. viator (G) and Paratrytone melane (H). All drawings are enlarged 12 times.... 359 3 4 Frontal view of the larval head of Choranthus haitensis (A), Euphyes arpa (B) E. pilatka (C), E. alabamae (D), E. dukesi (E), E. bimacula (F), E. ruricola (G), and Asbolis capucinus (H) All drawings are enlarged 12 times 360 35 Frontal view of the larval head of Atrytonopsis hianna (A), Amjbiyscirtes linda (B), A. aesculapias (C), A. vialis (D) A. celia (E), Lerodea eufala (F), Oligoria maculata (G) and Calpodes ethlius (H) All drawings are enlarged 12 times 361 36 Frontal view of the larval head of Panoquina panoquin (A), Nyctelius nyctelius (B), Agathymus valverdiensis (C), and Megathymus yuccae (D). All drawings are enlarged 12 times 362 37 Stemmatal pattern of Pyrrhopyge araxes (A), Phocides pigmalion (B), P. palemon (C), Epargyreus zestos (D) E. clarus (E), E. exadeus (F), Polygonus leo (G) Chioides catillus (H), Typhedanus undulatus (I), Polythrix mexicana (J), p. procera (K), and Codatractus alcaeus (L) All drawings are enlarged 25 times 363 XI

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Figure page 38 Stemmatal pattern of Codatractus arizonensis (A), Urbanus proteus (B) U. esmeraldus (C), U. dorantes (D) U. teleus (E) U. procne (F), Astraptes fulgerator (G), A. gilberti (H), A. anaphus (I), Autochton cellus (J), Achalarus lyciades (K) and A. casica (L). All drawings are enlarged 25 times 364 39 Stemmatal pattern of Thorybes bathyllus (A) T. pylades (B), r. diversus (C) T. mexicanus (D), r. confusis (E), T. drusius (F), Ca£>ares potrillo (G) Cogia hippalus (H), C. outis (I), C. caicus (J), Nisoniades rubescens (K), Staphylus mazans (L), S. hayhurstii (M) Carrhenes canescens (N), Xenophanes trixus (0) Systasea pulverulenta (P), S. zampa (Q), and Achiyodes thraso (R) All drawings are enlarged 25 times 365 40 Stemmatal pattern of Timochares ruptifasciatus (A), Chiomara asychis (B), Gesta gesta (C), Ephyriades brunneus (D), Erynnis icelus (E), E. jbrizo (F), E. juvenalis (G), E. propertius (H), E. horatius (I), E. tristis (J), E. martialis (K), E. pacuvius (L), E. zarucco (M) E. funeralis (N) E. baptisiae (0), E. afranius (P), E. persius (Q), Pyrgus centaureae (R) P. ruralis (S), P. scriptura (T) P. communis (U), P. albescens (V) P. oileus (W) Heliopetes ericetorum (X), H. lavia/ius (Y), ff. macaira (Z), tf. arsalte (AA), Celotes nessus (BB), C. limpa (CC), and Pholisora catullus (DD). All drawings are enlarged 25 times 366 41 Stemmatal pattern of Pholisora alpheus (A), P. gracielae (B), Piruna pirus (C), Synapte malitiosa (D), Vidius perigenes (E), Nastra lherminier (F), N. julia (G), W. neamathla (H), Cyinaemes tripunctus (I), Lerema accius (J), L. liris (K), Ancyloxypha numitor (L), Perichares philetes (M) A. arene (N), Copaeodes aurantiacus (0), C. minimus (P), Thymelicus lineola (Q), Hylephila phyleus (R), Yvretta carus (S), Pseudocopaeodes eunus (T), Hesperia uncas (U) tf. jujba (V), if. conuna (W), H. v/oodgatei (X), H. leonardus (Y), if. pahas/ca (Z), tf. metea (AA), and tf. viridis (BB). All drawings are enlarged 25 times 367 xii

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Figure page 42 Stemmatal pattern of Hesperia attalus (A) H. meskei (B), H. nevada (C) Polites cor as (D), P. sabuleti (E), P. mardon (F), P. draco (G) P. jbaracoa (H), P. themistocles (I), P. origenes (J), P. mystic (K) P. sonora (L), P. vibex (M) Wallengrenia otho (N) W. egeremet (0) Pompeius verna (P), Atalopedes campestris (Q), Atrytone arogos (R) A. delaware (S), Problema byssus (T), Ochlodes sylvanoides (U), O. agricola (V), O. yuma (W) Poanes hobomok (X), and P. zajbuJon (Y). All drawings are enlarged 25 times 368 43 Stemmatal pattern of Poanes taxiles (A) P. aaroni (B), P. yehl (C) P. viator (D), Paratrytone melane (E), Choranthus haitensis (F), Euphyes arpa (G), £. pilatka (H) £. alabamae (I), E. dukes i (J), £. macguirei (K), £. bimacula (L), £. ruricola (M) Asbolis capucinus (N), Atrytonopsis hianna (0), A. loammi (P), Amjblyscirtes linda (Q), A. aesculapias (R) A. vialis (S), A. celia (T), and Lerodea eufala (U). All drawings are enlarged 25 times 369 44 Stemmatal pattern of Oligoria maculata (A), Calpodes ethlius (B), Panoquina panoquin (C), P. panoquinoides (D), P. ocola (E), P. sylvicola (F), Nyctelius nyctelius (G) Agathymus valverdiensis (H), Astephensi (I), Megathymus yuccae (J), M. coloradensis (K), M. cofaqui (L) and M. ursus (M) All drawings are enlarged 25 times 370 45 Dorsal view of the larval terminalia of Pyrrhopyge araxes (A), Phocides pigmalion (B), £pargyreus zestos (C), and Polygonus leo (D). All drawings are enlarged six times 371 46 Dorsal view of the larval terminalia of Chioides catillus (A) Typhedanus undulatus (B), Polythrix mexicana (C) Codatractus alcaeus (D), Urbanus proteus (E), and Astraptes fulgerator (F) All drawings are enlarged six times 372 Xlll

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Figure page 47 Dorsal view of the larval terminalia of Autochton cellus (A) Achalarus lyciades (B), Thorybes bathyllus (C), Cabares potrillo (D), Cogia hippalus (E), Nisoniades rubescens (F), Staphylus ceos (G) Carrhenes canescens (H) and Xenophanes trixus (I). All drawings are enlarged six times 373 48 Dorsal view of the larval terminalia of Achlyodes thraso (A) Timochares ruptifasciatus (B), Gesta gesta (C), Ephyriades brunneus (D), Erynnis icelus (E), and Pyrgus centaureae (F). All drawings are enlarged six times 374 49 Dorsal view of the larval terminalia of Heliopetes ericetorum (A) Celotes nessus (B), Pholisora catullus (C) Piruna pirus (D), Synapte malitiosa (E), Vidius perigenes (F), and Nastra Iherminier (G). All drawings are enlarged six times 375 50 Dorsal view of the larval terminalia of Cymaenes tripunctus (A), Lerema accius (B), Perichares philetes (C), Ancyloxypha numitor (D), A. arene (E), Copaeodes aurantiacus (F), and C. minimus (G). Drawings D and F-G are enlarged 6 times; E is enlarged 12 times 376 51 Dorsal view of the larval terminalia of Thymelicus lineola (A), Hylephila phyleus (B), Yvretta carus (C) Pseudocopaeodes eunus (D), Hesperia uncas (E), Polites coras (F), and P. sabuleti (G). Drawings A-E are enlarged 6 times; F and G are enlarged 12 times 377 52 Dorsal view of the larval terminalia of Polites sabuleti (A), P. mardon (B), P. draco (C), P. baracoa (D), P.themistocles (E), and P. origenes (F). Drawings A-E are enlarged 12 times; F is enlarged 6 times 378 53 Dorsal view of the larval terminalia of Polites mystic (A), P. sonora (B) P. vibex (C), Wallengrenia otho (D), Pompeius verna (E), Atalopedes campestris (F), and Atrytone arogos (G) Drawings A-C are enlarged 12 times; D-G are enlarged 6 times 379 xiv

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Figure page 54 Dorsal view of the larval terminalia of Atrytone delaware (A) third instar Problema byssus (B), last instar Problema byssus (C) Ochlodes yuma (D), Poanes hobomok (E), and Paratrytone melane (F). Drawings A and C-F are enlarged 6 times; B is enlarged 12 times 380 55 Dorsal view of the larval terminalia of Choranthus haitensis (A), Euphyes arpa (B) E. ruricola (C) Asbolis capucinus (D), Atrytonopsis hianna (E), and Amblyscirtes linda (F). All drawings are enlarged six times 381 56 Dorsal view of the larval terminalia of Lerodea eufala (A), Oligoria maculata (B), Panoquina panoquin (C) P. panoquinoides (D), P. ocola (E), and P. sylvicola (F). Drawings A-D are enlarged 6 times; E-F are enlarged 12 times 382 57 Dorsal view of the larval terminalia of Nyctelius nyctelius (A), Calpodes ethlius (B), Agathymus valverdiensis (C), and Megathymus yuccas (D) All drawings are enlarged six times 383 58 Pupa of Epargyreus clarus (A) Thorybes bathyllus (B), Urbanus proteus (C), Phocides pigmalion (D), Polygonus leo (E), Anastrus sempiternus (F), Achlyodes thraso (G) and Pyrgus communis (H) 385 59 Pupa of Hesperia attalus (A) Pompeius verna (B), Poanes yehl (C) Oligoria maculata (D), Cymaenes tripunctus (E), and Synapte malitiosa (F) 387 60 Pupa of Nastra Iherminier (A) Perichares philetes (B), and Calpodes ethlius (C) 389 61 Dorsal view of anterior pupa of Phocides pigmalion (A), P. palemon (B), Epargyreus zestos (C), and £. clarus (D). All drawings are enlarged six times 390 xv

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Figure page 62 Dorsal view of anterior pupa of Polygonus leo (A), Chioides catillus (B) Polythrix mexicanus (C), Codatractus alcaeus (D), C. arizonensis (E), and Urbanus proteus (F). All drawings are enlarged six times 391 6 3 Dorsal view of anterior pupa of Urbanus esmeraldus (A), U. dorantes (B), U. teleus (C), U. procne (D) and Astraptes fulgerator (E). All drawings are enlarged six times... 392 64 Dorsal view of anterior pupa of Astraptes gilberti (A), Autochton cellus (B) Achalarus lyciades (C), Thorybes bathyllus (D), T. pylades (E), and T. confusis (F). All drawings are enlarged six times 393 65 Dorsal view of anterior pupa of Cabares potrillo (A), Cogia outis (B), Staphylus ceos (C), S. hayhurstii (D), Carrhenes canescens (E), Systasea pulverulenta (F), Achlyodes thraso (G) and Chiomara asychis (H). All drawings are enlarged six times... 394 66 Dorsal view of anterior pupa of Gesta gesta (A), Ephyriades brunneus (B), Erynnis icelus (C), E. brizo (D) E. juvenalis (E), E. horatius (F), E. tristis (G) and E. martialis (H). All drawings are enlarged six times 395 67 Dorsal view of anterior pupa of Erynnis zarucco (A), E. funeralis (B), E. baptisiae (C), E. persius (D), Pyrgus scriptura (E), P. communis (F), P. oileus (G), and Heliopetes ericetorum (H) All drawings are enlarged six times 396 68 Dorsal view of anterior pupa of Heliopetes lavianus (A), Celotes nessus (B), Pholisora catullus (C), P. alpheus (D), Synapte malitiosa (E), Nastra lherminier (F), W. Julia (G), and W. neamathla (H). All drawings are enlarged six times 397 xvi

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Figure page 69 Dorsal view of anterior pupa of Cymaenes tripunctus (A), Lerema accius (B), L. liris (C), Perichares philetes (D), Ancyloxypha numitor (E), A. arene (F), and Copaeodes aurantiacus (G) All drawings are enlarged six times 398 70 Dorsal view of anterior pupa of Copaeodes minimus (A), Hylephila phyleus (B), Yvretta carus (C) Pseudocopaeodes eunus (D) Hesperia uncas (E), if. comma (F), H. attalus (G) and Polites coras (H) All drawings are enlarged six times 399 71 Dorsal view of anterior pupa of Polites sabuleti (A), P. mardon (B), P. draco (C) P. jbaracoa (D), P. themistocles (E), P. origenes (F), P. mystic (G) and P. sonora (H) All drawings are enlarged six times... 400 72 Dorsal view of anterior pupa of Polites vibex (A), Wallengrenia otho (B), W. egeremet (C) Pompeius verna (D), Atalopedes campestris (E), Atrytone del aware (F), A. arogos (G), and Problema byssus (H). All drawings are enlarged six times 401 73 Dorsal view of anterior pupa of Ochlodes sylvanoides (A), Poanes hobomok (B), P. taxiles (C), P. aaroni (D), P. yehl (E), P. viator (F), Paratrytone melane (G) and Choranthus haitensis (H). All drawings are enlarged six times 402 74 Dorsal view of anterior pupa of Euphyes arpa (A), E. pilatka (B), E. alabamae (C), E. dukesi (D) E. berryi (E), and E. bimacula (F). All drawings are enlarged six times... 403 75 Dorsal view of anterior pupa of Euphyes ruricola (A), Asbolis capucinus (B), Atrytonopsis loammi (C) Amblyscirtes aesculapias (D), A. celia (E), Lerodea eufala (F), and Oligoria maculata (G). All drawings are enlarged six times 404 XVI 1

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Figure page 76 Dorsal view of anterior pupa of Calpodes ethlius (A), Panoquina panoquin (B), P. panoquinoides (C), P. ocola (D), and P. sylvicola (E). All drawings are enlarged six times 405 77 Dorsal view of anterior pupa of Nyctelius nyctelius (A), Megathymus coloradensis (B), M. cofaqui (C) and M. ursus (D). Drawing A is enlarged six times; B-D are enlarged three times 406 78 Ventral view of anterior pupa of Phocides pigmalion (A) and P. palemon (B). All drawings are enlarged six times 407 79 Ventral view of anterior pupa of Epargyreus zestos (A) and E. clarus (B). All drawings are enlarged six times 408 80 Ventral view of anterior pupa of Polygonus leo (A) and Chioides catillus (B). All drawings are enlarged six times 409 81 Ventral view of anterior pupa of Polythrix mexicana (A), Codatractus alcaeus (B), and C. arizonensis (C). All drawings are enlarged six times 410 82 Ventral view of anterior pupa of Urbanus proteus (A), U. esmeraldus (B), and U. dorantes (C). All drawings are enlarged six times 411 8 3 Ventral view of anterior pupa of Urbanus teleus (A) and U. procne (B). All drawings are enlarged six times 412 84 Ventral view of anterior pupa of Astraptes fulgerator (A), A. gilbert i (B), and Autochton cellus (C). All drawings are enlarged six times 413 85 Ventral view of anterior pupa of Thorybes bathyllus (A), T. pylades (B), T. confusis (C), Cabares potrillo (D), and Cogia outis (E). All drawings are enlarged six times... 414 XVI 11

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Figure page 86 Ventral view of anterior pupa of Staphylus ceos (A), S. hayhurstii (B), Carrhenes canescens (C), Systasea pulverulenta (D), Achlyodes thraso (E), Chiomara asychis (F), Gesta gesta (G) and Ephyriades brunneus (H) All drawings are enlarged six times... 415 87 Ventral view of anterior pupa of Erynnis icelus (A), E. brizo (B), E. juvenalis (C), E. horatius (D) E. tristis (E), E. martialis (F), E. zarucco (G), and E. funeralis (H) All drawings are enlarged six times 416 88 Ventral view of anterior pupa of Erynnis baptisiae (A), E. persius (B), Pyrgus scriptura (C), P. communis (D), P. oileus (E), Heliopetes ericetorum (F), H. lavianus (G), and Celotes nessus (H). All drawings are enlarged six times 417 89 Ventral view of anterior pupa of Pholisora catullus (A), P. alpheus (B), Synapte malitiosa (C) Nastra lherminier (D), N. julia (E), N. neamathla (F), and Cymaenes tripunctus (G) All drawings are enlarged six times 418 90 Ventral view of anterior pupa of Lerema accius (A), L. liris (B), Perichares philetes (C), Ancyloxypha numitor (D), A. arene (E), Copaeodes aurantiacus (F), and C. minimus (G). Drawings A-B and D-G are enlarged six times; C is enlarged three times 419 91 Ventral view of anterior pupa of Hylephila phyleus (A), Yvretta carus (B), Pseudocopaeodes eunus (C), Hesperia uncas (D), H. comma (E), H. attalus (F), Polites coras (G), and P. sabuleti (H). All drawings are enlarged six times 420 92 Ventral view of anterior pupa of Polites mardon (A), P. draco (B), P. haracoa (C), P. themistocles (D), P. origenes (E), P. mystic (F), P. sonora (G), and P. vijbex (H). All drawings are enlarged six times 421 xix

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Figure page 93 Ventral view of anterior pupa of Wallengrenia otho (A), W. egeremet (B), Pompeius verna (C), and Atalopedes campestris (D). All drawings are enlarged six times 422 94 Ventral view of anterior pupa of Atrytone arogos (A), A. delaware (B), Problema byssus (C), and Ochlodes sylvanoides (D). All drawings are enlarged six times 423 95 Ventral view of anterior pupa of Poanes hobomok (A), P. taxiles (B), P. aaroni (C), and P. yehl (D) All drawings are enlarged six times 424 96 Ventral view of anterior pupa of Poanes viator (A), Paratrytone melane (B), Choranthus haitensis (C) and Euphyes arpa (D). All drawings are enlarged six times... 425 97 Ventral view of anterior pupa of Euphyes pilatka (A), E. alabamae (B), E. dukesi (C) E. berryi (D), E. bimacula (E), and E. ruricola (F). All drawings are enlarged six times 426 98 Ventral view of anterior pupa of Asbolis capucinus (A), Atrytonopsis loammi (B), Amblyscirtes aesculapias (C), A. celia (D), Lerodea eufala (E), and Oligoria maculata (F). All drawings are enlarged six times... 427 99 Ventral view of anterior pupa of Calpodes ethlius (A), Panoquina panoquin (B), P. panoquinoides (C) P. ocola (D), and P. sylvicola (E). All drawings are enlarged six times 428 100 Ventral view of anterior pupa of Nyctelius nyctelius (A), Agathymus valverdiensis (B), Megathymus yuccae (C) M. coloradensis (D), M. cofaqui (E), and M. ursus (F). All drawings are enlarged three times 429 xx

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Figure page 101 Lateral view of the cremaster of Phocides pigmalion (A), P. palemon (B), Epargyreus zestos (C), E. clarus (D), Polygonus leo (E), Chioides catillus (F), Polythrix mexicana (G) Codatr actus alcaeus (H), and C. arizonensis (I). All drawings are enlarged 12 times 430 102 Lateral view of the cremaster of Urbanus proteus (A), U. esmeraldus (B) U. dorantes (C), U. teleus (D), U. procne (E), Astraptes fulgerator (F), A. gilberti (G), Autochton cellus (H) and Achalarus lyciades (I). All drawings are enlarged 12 times 431 103 Lateral view of the cremaster of Thorybes bathyllus (A), T. pylades (B), T. confusis (C), Cabares potrillo (D), Cogia hippalus (E), C. outis (F), C. caicus (G) Nisoniades rubescens (H) and Staphylus ceos (I). All drawings are enlarged 12 times 432 104 Lateral view of the cremaster of Staphylus mazans (A), S. hayhurstii (B), Carrhenes canescens ( C ) Xenophanes trixus ( D ) Systasea pulverulenta (E), Achlyodes thraso (F), Timochares ruptifasciatus (G) Chiomara asychis (H) Gesta gesta (I), Ephyriades brunneus (J), Erynnis icelus (K), £. jbrizo (L), £. juvenalis (M) ; £. horatius (N) £. tristis (0), and £. martialis (P). All drawings are enlarged 12 times 433 105 Lateral view of the cremaster of Erynnis zarucco (A), £. funeralis (B), E. baptisiae (C), E. persius (D), Pyrgrus scriptura (E), P. communis (F), P. oileus (G), Heliopetes ericetorum (H), H. lavianus (I), Celotes nessus (J), Pholisora catullus (K) P. alpheus (L), Synapte malitiosa (M) Nastra lherminier (N) W. julia (0) and W. neamathla (P). All drawings are enlarged 12 times 434 xxi

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Figure page 106 Lateral view of the cremaster of Cymaenes tripunctus (A), Lerema accius (B), L. liris (C), Perichares philetes (D), Ancyloxypha numitor (E), A. arene (F), Copaeodes aurantiacus (G) C minimus (H) Hylephila phyleus (I), Yvretta carus (J), Pseudocopaeodes eunus (K), and Hesperia uncas (L). Drawings A-C and E-L are enlarged 12 times; D is enlarged 6 times.... 435 107 Lateral view of the cremaster of Hesperia comma (A), H. woodgatei (B), H. viridis (C) if. attalus (D), Polites coras (E), P. sabuleti (F), P. mardon (G), P. draco (H), P. baracoa (I), P. themistocles (J), P. origenes (K) P. mystic (L) P. sonora (M) P. vijbex (N) Wallengrenia otho (0) and IV. egeremet (P). All drawings are enlarged 12 times 436 108 Lateral view of the cremaster of Pompeius verna (A), Atalopedes campestris (B), Atrytone arogos (C), A. delav/are (D), Problema byssus (E), Ochlodes sylvanoides (F), Poanes hobomok (G), P. taxiles (H), P. aaroni (I), and P. yehi (J). All drawings are enlarged 12 times 437 109 Lateral view of the cremaster of Poanes viator (A), Paratrytone melane (B), Choranthus haitensis (C), Euphyes arpa (D), £. pilatka (E), £. alabamae (F), i?. dukesi (G) £. jberryi (H), E. macguirei (I), £. bimacula (J), £. ruricola (K), and Asbolis capucinus (L). Drawings A-C and H-K are enlarged 12 times; D-G and L are enlarged 6 times 438 110 Lateral view of the cremaster of Atrytonopsis loammi (A) AnLblyscirtes aesculapias (B), A. vialis (C) A. celia (D) Lerodea eufala (E), Oligoria maculata (F) Calpodes ethlius (G) Panoquina panoquin (H), P. panoquinoides (I), and P. ocola ( J) Drawings A-F and H-J are enlarged 12 times; G is enlarged 6 times.... 439 xxii

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Figure page 111 Lateral view of the cremaster of Panoquina sylvicola (A), Nyctelius nyctelius (B), Agathymus valverdiensis (C) Megathymus yuccae (D), M. coloradensis (E), M. cofaqui (F), and M. ursus (G) Drawings A-B are enlarged 12 times; C-G are enlarged 6 times. 440 112 Dorsal view of the cremaster of Phocides pigmalion (A), P. palemon (B), Epargyreus zestos (C), E. clarus (D), Polygonus leo (E), and Chioides catillus (F). Drawings A and B are enlarged 6 times; C-F are enlarged 12 times 441 113 Dorsal view of the cremaster of Polythrix mexicana (A), Codatractus alcaeus (B), C. arizonensis (C), Urbanus proteus (D), U. esmeraldus (E), £7. dorantes (F), (7. teleus (G), £7. procne (H) and Astraptes fulgerator (I). All drawings are enlarged 12 times.... 442 114 Dorsal view of the cremaster of Astraptes gilberti (A), Autochton cellus (B), Achalarus lyciades (C), ThoryJbes bathyllus (D), T. pylades (E), T. confusis (F) Cajbares potrillo (G), Cogia hippalus (H) and C. outis (I). All drawings are enlarged 12 times 443 115 Dorsal view of the cremaster of Cogia caicus (A), Nisoniades rubescens (B), Staphylus ceos (C), S. mazans (D), S. hayhurstii (E), Carrhenes canescens (F), Xenophanes trixus (G) Systasea pulverulenta (H) AcMyodes thraso (I), Timochares ruptifasciatus (J), Chiomara asychis (K) and Gesta gesta (L). All drawings are enlarged 12 times 444 116 Dorsal view of the cremaster of Ephyriades brunneus (A), Erynnis icelus (B), E. brizo (C), E. juvenalis (D), E. horatius (E), E. tristis (F), E. martialis (G), £. zarucco (H) jE. funeralis (I), £. baptisiae (J), £. persius (K) and Pyrgus scriptura (L). All drawings are enlarged 12 times 445 xxm

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Figure page 117 Dorsal view of the cremaster of Pyrgus communis (A), P. oileus (B), Heliopetes ericetorum (C) H. lavianus (D), Celotes nessus (E), Pholisora catullus (F), P. alpheus (G) Synapte malitiosa (H), Nastra Iherminier (I), N. julia (J), N. neamathla (K) and Cyinaenes tripunctus (L). All drawings are enlarged 12 times 446 118 Dorsal view of the cremaster of Lerema accius (A), L. liris (B), Perichares philetes (C), Ancyloxypha numitor (D), A. arene (E), Copaeodes aurantiacus (F), C. minimus (G), Hylephila phyleus (H), yvretta carus (I), Pseudocopaeodes eunus (J), and Hesperia uncas (K) Drawings A-B and D-K are enlarged 12 times; C is enlarged 6 times 447 119 Dorsal view of the cremaster of Hesperia comma (A), H. woodgatei (B), H. viridis (C), H. attalus (D), Polites coras (E), P. sabuleti (F), P. mardon (G) P. draco (H), P. baracoa (I), P. themistocles (J), and P. origenes (K) All drawings are enlarged 12 times 448 120 Dorsal view of the cremaster of Polites mystic (A), P. sonora (B), P. vi&ex (C), Wallengrenia otho (D), IV. egeremet (E), Pompeius verna (F), Atalopedes campestris (G) Atrytone arogos (H) A. delaware (I), and Problema byssus ( J) Drawings A-I are enlarged 12 times; J is enlarged 6 times.... 449 121 Dorsal view of the cremaster of Ochlodes sylvanoides (A), Poanes hobomok (B), P. taxiles (C), P. aaroni (D), P. yehl (E), P. viator (F), Paratrytone melane (G), Choranthus haitensis (H) Euphyes arpa (I), £. pilatka (J), £. alabamae (K), £. dukesi (L), and 2?. i>erryi (M) Drawings A, C-E, G-H, and M are enlarged 12 times; B, F, and I-L are enlarged 6 times 450 xxiv

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Figure page 122 Dorsal view of the cremaster of Euphyes macguirei (A), E. bimacula (B), E. ruricola (C), Asbolis capucinus (D), Atrytonopsis loammi (E), Amblyscirtes aesculapias (F), A. vialis (G), A. celia (H) Lerodea eufala (I), Oligoria maculata (J), and Calpodes ethlius (K). Drawings A-C and E-J are enlarged 12 times; D and K are enlarged 6 times 451 123 Dorsal view of the cremaster of Panoquina panoquin (A), P. panoquinoides (B), P. ocola (C) P. sylvicola (D) Nyctelius nyctelius (E), Megathymus yuccae (F), M. coloradensis (G), Mcofaqui (H), and M. ursus (I). Drawings A-E are enlarged 12 times; F-I are enlarged 6 times 452 xxv

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Abstract of Dissertation to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy IMMATURE STAGES OF THE SKIPPER BUTTERFLIES (LEPIDOPTERA: HESPERIIDAE) OF THE UNITED STATES: BIOLOGY, MORPHOLOGY, AND DESCRIPTIONS By Marc C. Minno April 1994 Chairman: Thomas C. Emmel Major Department: Zoology The purpose of this study was to investigate the biology of as many U. S. skippers as possible, compare morphological features of their immature stages, and produce standardized descriptions. This revision summarizes of our current knowledge of immature stages of North American Hesperiidae, and provides a framework for future taxonomic and systematic studies. About 300 species of skipper butterflies (Hesperiidae) have been reported from the United States. Most of the fauna is composed of the subfamilies Pyrginae and Hesperiinae, but Pyrrhopyginae, Heteropterinae, and Megathyminae are also represented. Species of Hesperiidae found in the U. S. feed on 31 families of plants as larvae. These include 8 families of monocots and 23 families of dicots. Most (92%) of these butterflies are limited to only 1 of the 31 families of plants, but a few skipper species feed on plants in 2 or xxvi

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very rarely, as many as 6 different families. Species having relatively broad host ranges include Lerema accius (35 spp.), Epargyreus clarus (31 spp.), Thorybes py lades (27 spp.)/ and Megathymus yuccae (27 spp. ) • One hundred and thirty-eight new host records are reported for 55 species of hesperiids. Some skippers are important pests of beans, rice, sugarcane, and pasture grasses. Host plants remain to be discovered for 22% of the U. S. Hesperiidae. The eggs of 49 species, the larvae of 156 species, and the pupae of 120 species of Hesperiidae of the U. S. are described in this thesis. These include the eggs of 16 species, the larvae of 41 species, and the pupae of 35 species that were previously unknown. Twenty-six percent of the U. S. skippers are totally unknown with respect to their egg, larval, and pupal stages. About half of these species with undescribed immatures are breeding residents. The other half are neotropical species that rarely enter the U. S. and probably do not reproduce within its limits. Larval characters that may prove useful to systematists conducting phylogenetic analyses include the type of mandible articulation, setal types, sculpturing of the head, lenticle distribution on the larvae, wax gland patterns, and pigments. Differences in the sculpturing patterns of hesperiid eggs have been known for some time, but have not been extensively surveyed. Promising characteristics of the pupae include the shape of the pupal cap, lenticle xxvii

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distribution, relative length of the antennae and proboscis, cremaster shape, type of thoracic spiracle guard, and presence of crenulations on the posterior margin of the prothorax. xxvm

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CHAPTER 1 INTRODUCTION Adult skippers (Hesperiidae) are thick-bodied butterflies with large compound eyes, wide heads, and conspicuous labial palps. The antennal club is usually bent and has a tapering extension called the apiculus. The radial veins of the forewings all arise from the discal cell, and the second medial vein of the hindwings is absent or weakly developed. All six legs are well-developed and used for perching. Many skippers are drably colored, but some tropical species are bright yellow, red or iridescent blue or green. The wings are freguently dotted with semihyaline whitish spots. The Hesperiidae is a group that is easily recognized as distinct from all other butterfly families. Skippers occur in all faunal regions of the world except at the poles (Heppner 1991). Estimates of the number of described species range from 3000 to 3658 (Robbins 1982, Bridges 1983, and Heppner 1991). The Neotropics are especially rich in skippers, with 2,016 described species (Heppner 1991), followed by the Old World tropics, with 570 species in the Oriental region and 440 species in the Ethiopian region. Among temperate regions, the Nearctic has

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2 more species (290) than Australia (191) or the Palearctic (155). Overall, about 63% of all described species of skippers occur in the New World. Within the Hesperiidae, eight subfamilies have been recognized (Heppner 1991). The Heteropterinae is the smallest subfamily, with only about 30 species distributed in the Palearctic, Nearctic, and Neotropical regions. The Megathyminae is another small subfamily of about 50 species. Megathymines are found in xeric areas of the United States, Mexico, and Central America. The Coeliadinae is a group of about 80 species restricted to the Old World tropics. Another Old World group, the Trapezitinae, includes about 85 species endemic to the Australian faunal region. Euschemoninae consists of a single Australian species, Euschemon rafflesia. Some 180 skippers of the subfamily Pyrrhopyginae occur in the Neotropics. The two largest groups, the Pyrginae (1,195 species) and the Hesperiinae (2,044 species), are widely distributed throughout the world. Some hesperiids are economically important to people. The larvae of one species, Aegriale hesperiaris known as gusanos de maguey in Spanish, are eaten as a delicacy in Mexico (Blasguez 1870). The Indians of Mexico have long harvested gusanos de maguey for food (Draudt 1924). Today, these hesperiid larvae are collected and sold fresh or canned in Mexican markets (Borror et al 1954), and are

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3 fried before eating (Ancona 1934, Sokolov 1989). Small larvae are sometimes added to bottles of tequila as a garnishment (Sbordoni and Forestiero 1985, Toliver 1987). Skippers are also pests of crops, forests, and plants used as garden ornamentals. Urbanus proteus is destructive to beans (Comstock 1881, Riley and Howard 1893, Chittenden 1902, Cotton 1918, Watson 1919, Wolcott 1933, Greene 1971, and Heppner 1975), Ancyloxypha numitor Panoquina ocola, and Nyctelius nyctelius to rice (Bell 1940, Ross and Lambremont 1963), and Panoquina sylvicola, Nyctelius nyctelius Choranthus haitensis Choranthus vitellius Perichares philetes and others to sugarcane (Smyth 1919, Wolcott 1921, Jones and Wolcott 1922, Wolcott 1933, 1936, and Hayward 1943). Hesperiids that have been listed as occasionally injurious to trees include Epargyreus clarus and Erynnis species (Felt 1918, McDaniel 1933, Craighead 1950, Baker 1972, and Furniss and Carolin 1977). Grass-feeding species such as Hylephila phyleus and Atalopedes campestris may occasionally damage lawns (Bohart 1947, Warren and Roberts 1956, Ross and Lambremont 1963) and Calpodes ethlius frequently will defoliate ornamental cannas (Dyar 1898, Chittenden 1905, 1912, Essig 1926, Weigel and Middleton 1926, and Reinert et al 1983). A few skippers accidentally introduced into new areas have become pests. A European species introduced into Canada, Thymelicus lineola is a major pest of pastures and

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4 hay crops (Pengelly 1961, Arthur 1966, and Duchesne and McNeil 1978). Similarly, of two exotic skippers now established in Hawaii, Hylephila phyleus causes damage to lawns (Tashiro and Mitchell 1985) and Erionota thrax to bananas (Riotte and Uchida 1979). Aside from human economics, skippers have biological value, being important in the pollination of flowering plants and in the cycling of nutrients and flow of energy in food webs. Since many species are restricted to certain habitats or are very local in distribution, they may also prove useful for biological monitoring of the ecological health of nature reserves or for biodiversity studies. Skippers also have scientific interest, and a large literature on this group has accumulated over the last few centuries. Many life history descriptions of New World hesperiids have been published, especially by workers in the United States (U. S.), but these accounts sometimes contain errors and are uneven in the choice of characters described. The intent of this study was to investigate the biology of as many U. S. species as possible, compare morphological features of their immature stages, and produce standardized descriptions. This revision will not only serve as a summary of our current knowledge of immature stages of North American Hesperiidae, but will provide a framework for future taxonomic and systematic studies.

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CHAPTER 2 METHODS Study Material A major problem associated with studying the immature stages of butterflies lies in finding specimens. Butterflies are relatively rare insects compared to groups such as Diptera, Coleoptera, or Hymenoptera, and locating immatures in the field is often a challenging task. Although many collectors and researchers are interested in immature stages, few preserve eggs, larvae, or pupae. Insect collections that contain lepidopteran immatures will often have at least some skipper butterfly larvae, but these are usually common species. Preserved pupae, even of common species, are rare in museum collections. Roy 0. Kendall (ROK) has one of the largest (in terms of numbers of individuals as well as species) and best curated collections of North American hesperiids in existence. I borrowed many specimens for study from the Kendall collection as well as from the Florida State Collection of Arthropods (FSCA), Santa Barbara Natural History Museum (SBNHM), United States National Museum (USNM), and the collections of G. R. Ballmer (GRB), T. C. Emmel (TCE) D. H. Habeck (DHH), J. R. Heitzman (JRH), T. L. McCabe (TLM), N. McFarland (NM), and S. Passoa

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6 (SP). Several lepidopterists including H. D. Baggett, S. S. Borkin, J. F. Emmel D. w. Hall, D. F. Schweitzer, A. M. Shapiro, and J. R. Slotten, also donated specimens for study. In addition, I have conducted my own field research in California, Colorado, Florida, Georgia, Indiana, Maryland, New Mexico, New York, North Carolina, Pennsylvania, South Carolina, and Texas over the last 20 years. Outside of the United States, I have been able to make collections of hesperiid immatures in Colombia, Costa Rica, Dominican Republic, and the Bahamas. Some tropical species represented in these collections have occasionally been recorded from the U. S. Andrew Atkins and H. Chiba sent preserved immatures of interesting species from Australia and Indonesia, which were useful for making comparative studies. The number of hesperiid eggs, larvae, and pupae that I examined during the course of this study is presented under the species accounts in Chapter 6 of this work. An identifying label, with the format "MCM-Lot XXXX" (where the X's represent a unigue number) was placed in each vial or jar of immatures that I examined. The consecutive numbers used began with 0001 and ended with 1561. The code number was also recorded on an accession form listing specimen deposition, identification, locality, host plant, parasites, number of individuals of each stage, type of container, and preservative. Much of this information was entered into a

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7 digital database using an IBM personal computer and LOTUS 12-3 software. The accession code may be used to trace specimens for future studies and was used to link database files together. Rearing One method of collecting hesperiid immatures is to search for them in the field. Some species are relatively easy to find with a little practice, since the eggs are laid only on particular plants and the larvae usually fold or tie leaves together to form shelters in which they live. However, many species live such secretive lives that they are best reared in the laboratory. It has long been known that female butterflies will often lay eggs if confined in a container with the larval host plant. Edwards (1870) got a female zebra swallowtail to lay eggs by placing a nail-keg, from which the bottom had been knocked out, over a pawpaw bush and covering the top with a cloth. Scudder (1870) used tin cans in a similar way and secured eggs of Thorybes pylades Polites mystic, and a satyr id butterfly. Both Edwards and Scudder were able to rear many butterflies using this technique. Fletcher (1888) recommended not only caging females over hosts in the field, but also rearing larvae on potted hosts covered by wire cages or glass lamp chimneys. Laurent (1908) found that if female skippers were removed from a cyanide killing jar soon after they succumbed, they could easily be transferred to

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8 individual pill boxes and that they usually recovered without harm in a short time. He then secured eggs from the female skippers by confining them in cages over pots of the host plants. Caging females with a known or suspected host does not always work. Some species of skippers seem to lay eggs freely, while others absolutely will not oviposit in confinement. Not only is there wide variation between species, there is also much variation between individuals. Generally, a slightly worn female may be more likely to lay eggs than one that is freshly emerged (and perhaps not mated). It has been my experience that pyrgines are less likely to lay eggs in confinement than hesperiines. Heitzman (1964a) noted that one or two fertile eggs may sometimes be dissected from females that will not otherwise oviposit in captivity. I have found that females may be transported from the field to the laboratory without harm by placing them individually in the plastic containers in which 35mm film is sold. Either the opague Kodak or transparent Fuji Film containers work well, but the clear containers have the advantage of the investigator being able to examine the specimens without opening them. Containers with live females should be kept in a cool, dark place such as a field bag or cooler. In the laboratory, the containers may be kept refrigerated for a few days without harm, but it is

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9 best to feed and place the females in oviposition containers as soon as possible. I manually fed female skippers a dilute honey solution (1 part honey to 10 parts tap water) twice a day. By grasping the adult's closed wings with one hand, the skipper's proboscis could be uncoiled into a dish of honey water using an insect pin held by the other hand. To secure eggs, I found it best to place the females in transparent plastic containers containing small sprigs of host plants. The cut ends of the plants may be placed into small vials of water or wrapped with a piece of wet paper towel to prevent the leaves from wilting. The plant should be flattened against the container wall or arranged such that the butterfly will easily come into contact with the leaves and not become trapped in pockets of vegetation. A hole, 2 to 3cm in diameter, cut into the top of the oviposition container and covered with fine netting, will provide ventilation and prevent condensation. The behavior of adult skippers in cages consists of periods of rest punctuated by short bursts of activity. The oviposition containers with females and hosts may be placed under incandescent lights to stimulate activity, but need to be monitored freguently, as excessive heat, moisture, or dryness may be a problem. I prefer to place the containers on bright, but not sunny window sills or in a sheltered location out-of-doors where natural light and breezes stimulate the females. Richard Boscoe (pers. comm. )

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10 recommends placing the oviposition containers into closed plastic bags or refrigerating females of species reluctant to oviposit in captivity overnight, but I have had limited success with these techniques. I have not tried to secure eggs from female megathymines in the laboratory. Megathymus species glue the eggs to the leaves of yuccas, and the eggs are relatively easy to find in the field at the right time of year. Agathymus species either drop the eggs onto the host, or ova laid on the leaves fall off soon after oviposition, making them difficult to find in nature. Comstock and Dammers (1934), Roever (1964), and Stallings and Stallings (1986) were successful in having females of Agathymus species oviposit in cages in the laboratory. Once eggs have been laid, care must be taken to prevent excessive moisture, which promotes the growth of molds, or dryness which may cause the eggs to desiccate. The plastic containers used for securing eggs also worked well for rearing larvae. Ventilation through the screened hole in the lid, fresh food plants, and maintaining clean conditions within the containers were important in the prevention of disease. Temperate species that undergo larval hibernation pose special problems, as it is often difficult to provide proper environmental conditions in the laboratory. I have had some success in caging larvae out-of-doors on potted plants, or

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11 in rearing the larvae under artificial lights with long-day photoperiods. Other rearing techniques are presented in MacNeill (1964), McFarland (1964), Brown (1965), Kendall (1965), Newcomer (1966), and Tashiro and Mitchell (1985). Wielgus and Wielgus (1973) and Wielgus and Stallings (1974) present methods for rearing Megathymus in the laboratory. Preservation and Preparation I preserved hesperiid eggs and first instar larvae by placing them in vials with 70% alcohol. Older larvae and pupae were dropped into hot (near boiling) water to kill them (Peterson 1962), blotted dry with paper toweling, then transferred to 2 or 4 dram vials containing 70% alcohol. After several weeks, the old discolored preservative was drained from the vials and new alcohol added. I used 70% isopropyl alcohol for preserving immatures, because it is inexpensive, readily available, and less hazardous than formulations containing acetic acid, formalin, xylene, or kerosene. Specimens preserved in this manner remained relatively pliable and easy to work with, but lost most of the body colors. To document colors and the general features of living caterpillars and pupae, I made color slides using a Canon A1 35mm camera with a 100mm macrolens. A 50mm extension tube was used with this apparatus to magnify specimens between 1 and 3 cm in length. Most specimens were photographed in natural light. Both Kodak Kodachrome 64 and Ektachrome 100

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12 color transparency films gave good results, but it was sometimes difficult to get satisfactory slides with the slower film when photographing small specimens under low light conditions. Descriptive Methods I chose the checklist by Miller and Brown (1983) as a starting point for my descriptive work. According to these authors, the hesperiid fauna of North America, north of Mexico, contains 1 species of Pyrrhopyginae, 117 species of Pyrginae, 5 species of Heteropterinae, 139 species of Hesperiinae, and 28 species of Megathyminae for a total of 290 species. The Canadian fauna included in this checklist does not contain any species not found in the U. S. (Gregory 1975) External features of the immature stages were compared with the aid of a Wild M5 stereomicroscope fitted with 20 power eyepieces. A preliminary list of potentially interesting characters was drafted and modified during an initial examination of all of the material. After this review, a new standard character list was produced. The specimens were then examined a second time, new character data were recorded, and drawings of interesting features were made. Abbreviations for segments referred to in the diagnoses are Tl-3 for thoracic segments one through three and Al-10 for abdominal segments one through ten.

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13 The fine structure of eggs was observed and photographed using the University of Florida, Department of Zoology scanning electron microscope (SEM) (Hitachi model S415A) To prevent distortion, the eggs were dehydrated through a series of alcohol solutions (70%-99%). The alcohol was subseguently replaced through a series of acetate solutions. The specimens were then subjected to critical point drying using a Denton Vacuum Inc. Apparatus DCP-1 The dry eggs were glued to SEM stubs and coated with gold-palladium using a Denton Vacuum Inc. Desk-1 Sputterer, then viewed with the SEM. I made sketches and measurements of structures with a camera lucida attached to the stereomicroscope. The drawing tube allowed for superimposing the image of a metric ruler onto the specimen. Then, by dividing by the magnification power, size could be determined. This method worked well for structures greater than 0.1 mm. The heights and diameters of the eggs of 49 species were measured and recorded using this technique. The lengths of larvae and pupae were made simply with a metric ruler. Measurements of the transverse width of the head and abdominal segment four (generally the widest segment) of larvae, and abdominal segment three (usually the widest) of pupae were made with the microscope. A total of 1584 larvae and 623 pupae of 158 species were measured. For measurements of setal lengths and number of crochets, I chose the largest specimens

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14 available, and these data are reported as maxima in the diagnoses. In comparing pupal abdomen length among the various species of U. S. hesperiids, the distribution of the ratio of A5-8 length and Al-4 length was divided into three egual groups representing short (0.3 to 0.69mm), moderate (0.7 to 0.89mm), and long (0.9 to 1.4mm) classes.

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CHAPTER 3 BIOLOGY Although the larvae of some specialized moths feed on fungi, lower plants (algae, lichens, liverworts, mosses, ferns, cycads and gymnosperms ) other insects, feces, beeswax, or the keratin-containing parts of dead vertebrates, most butterflies feed only on angiosperm plants. Exceptions include a few neotropical nymphalids (Satyrinae) that eat lycopsids (Singer et al 1971, Singer and Mallet 1986), a few lycaenids and pierids that eat the leaves of pines, and some lycaenids which feed on the leaves of cycads (Eumaeus spp. ) or are predators of aphids (Liphyinae) or ant larvae (some Polyommatinae) Skippers feed only on angiosperms, but utilize a diversity of plants that are defended by a wide variety of secondary plant compounds Species of skippers tend to be stenophagous, limiting the host range to just a few species of plants, yet sixty families of higher plants have been recorded as hosts in the New World. The host relationships of U. S. hesperiids is presented in Table 1. Thirty-one families representing 8 ttbnocot groups and 23 dicots have been reported. Hesperiinae, Heteropterinae, and Megathyminae use only 15

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16 Table 1. Number of species and subfamilies of U.S. Hesperiidae recorded from various host plant families HOST PLANT FAMILY HESPERIIDAE # SPECIES SUBFAMILIES Monocots : Agavaceae Arecaceae Cannaceae Cyperaceae Marantaceae Musaceae Poaceae Zingiberaceae 28 2 1 15 1 1 103 Megathyminae Hesperiinae Hesperiinae Hesperiinae Hesperiinae Hesperiinae Heteropterinae Hesperiinae Pyrginae Hesperiinae Dicots: Amaranthaceae 4 Pyrginae Aquif oliaceae 1 Pyrginae Asteraceae 1 Pyrginae Betulaceae 2 Pyrginae Chenopodiaceae 8 Pyrginae Combretaceae 1 Pyrginae Convolvulaceae 1 Pyrginae Fabaceae 43 Pyrginae Fagaceae 9 Pyrginae Pyrrhopyginae Lamiaceae 1 Pyrginae Malpighiaceae 4 Pyrginae Malvaceae 16 Pyrginae Myristicaceae 1 Pyrginae Myrtaceae 1 Pyrginae Ranunculaceae 1 Pyrginae Rhamnaceae 3 Pyrginae Rhizophoraceae 1 Pyrginae Rosaceae 4 Pyrginae Rutaceae 1 Pyrginae Salicaceae 1 Pyrginae Sterculiaceae 2 Pyrginae Trigoniaceae 1 Pyrginae Verbenaceae 2 Pyrginae

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17 monocots; Pyrginae and Pyrrhopyginae usually feed only on dicots. Three subfamilies, Hesperiinae, Heteropterinae, and Pyrginae, use Poaceae, while both Pyrginae and Pyrrhopyginae eat Fagaceae. Hesperiid females adopt a characteristic fluttering type of flight pattern when searching for host plants on which to lay eggs. Often, the females will hover about potential hosts, sometimes landing upon the leaves one or more times, before laying eggs. By closely inspecting leaf shape and sensing plant chemistry, the butterflies are able to distinguish hosts of suitable size, age, quality, and position in the landscape before proceeding to lay their eggs (Scott 1986). Females usually lay their eggs singly on the leaves of the host plant. Most glue the eggs to the under surface of leaves, while others prefer to oviposit on the upper surface or exhibit no preference. The color of the egg changes as development proceeds. Scudder (1889a) recognized four embryological stages of butterfly eggs. Stage one is of short duration and consists of the newly laid egg, which is of homogeneous color. Stage two is marked by either a darkening of the color or the appearance of small, uniformly distributed spots. During the third stage there is a polarity of the contents corresponding to the formation of the germinal band in which a central annulus and a large lateral spot (the head) appear. In the last stage, the head

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18 moves upward to just underneath the micropyle at the apex of the egg. The newly laid eggs of skippers may be brownish, pinkish, white, pale yellow, or green. Development of hesperiid eggs agrees with Scudder's (1889a) observations. The eggs soon darken or change color and may develop a reddish ring. Shortly before the eggs are ready to hatch, the dark head of the larva is usually visible at the apex. Skipper eggs typically hatch in 4 to 15 days. When the larva is ready to emerge, it makes a small hole in the chorion with the mandibles, then cuts a circular opening in the apex slightly larger than the head (Ainslie 1922). This process may take several days (Roever 1964), but once the hole is completed, the larva exits quickly. The newly emerged larva usually does not eat the remains of the egg but seeks out a site in which to build a shelter. Hesperiid larvae tie leaves or parts of leaves together with silk to form characteristic shelters in which they hide when not feeding (Figures 1 and 2). Although skippers are often known as leaf rollers, they do not roll leaves in the manner of tortricids, pyralids, or other moths, but rather fold leaves or tie the edges of several leaves together and line the inside with silk. Among other groups of butterflies, some nymphalids and papilionids construct silklined shelters. Those built by charaxines (Nymphalidae) such as Anaea andria are the most similar to hesperiid shelters

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19 Figure 1. Larval shelter of Urbanus proteus on Phaseolus vulgaris (A), Polygonus leo on Piscidia piscipula (B) Erynnis zarucco on Sesbania macrocarpa (C), Erynnis brizo on Quercus myrtifolia (D), Wallengrenia otho on Paspalum sp. (E), and Asbolis capucinus on Sabal sp. (F).

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20

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21 Figure 2. Larval shelter of Euphyes pilatka on Cladium jamaicense (A), Euphyes arpa on Sabal etonia (B), Megathymus yuccae on Yucca aloi folia (C) and an agave leaf cut open to reveal the tunnels of Agathymus aryxna (D).

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23 The only predators that I have observed capturing skipper larvae are Polistes wasps. It is not known if hesperiid larvae are palatable to vertebrate predators such as birds and lizards. Parasitic wasps in the families Mymaridae, Eulophidae, and Braconidae frequently attack the eggs and larvae of skippers. The larvae are also commonly attacked by tachinid flies. Potentially, ants, spiders, lizards, birds, and small mammals may eat skipper butterfly immatures unless repulsed by the defenses noted below. The larval shelters of hesperiids are primarily thought to provide concealment from predators and parasitoids. However, the shelters are often conspicuous, perhaps raising a flag to predators, and skipper larvae are often heavily parasitized. Another plausible reason for shelters may involve chlorophyll. Chlorophyll which becomes uncoupled from the tightly controlled photosynthetic units within plant cells, as during digestion, is capable of diverting electrons to normally inert oxygen molecules, forming highly destructive oxygen radicals (Hendry 1990). Larger herbivores with opaque bodies have little to fear from chlorophyll as the digestion takes place in the dark inside their guts. However, a rare disease of sheep in which partly digested chlorophyll passes into the blood causes cell destruction in epidermal tissues exposed to sunlight due to the formation of activated oxygen (Hendry 1990). Poulton (1894) demonstrated that the ground color of

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24 caterpillars is derived largely from the diet. The green color of hesperiids that feed on leaves is due to chlorophyll. Newly emerged larvae become green only after the first feeding. Hesperiines, which have little cuticular pigmentation, change from green to cream-colored before entering diapause or pupation when they are no longer actively feeding. Leaf -feeding caterpillars which have relatively large amounts of chlorophyll in their blood and guts as well as thin, transparent, or translucent cuticles must have physiological and/or behavioral adaptations that prevent the formation of oxygen radicals. Many authors have noted that skipper larvae hide in shelters during the day and emerge to feed at night (Harris 1862, Lintner 1872, Edwards and Chapman 1879, Panton 1897, Scudder 1889a, and numerous others). In fact, most caterpillars are largely nocturnal feeders (J. Rawlins, pers. comm.). Thus the evolution of shelter-forming behavior in skippers may be grounded in a physiological need for concealment from the sun rather than from predators. The shelter may also be adaptive in allowing the larva easy access to the leaves of the host. Other caterpillars that do not make shelters or nests rest exposed on the underside of leaves or hide on the branches, under bark, or in leaf litter at the base of the plant. Skipper larvae spin a thin trail of silk with a characteristic side-to-side motion of the head (Fletcher 1889, Scudder 1889a, Ainslie

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25 1922, Clark 1936, MacNeill 1964, McAlpine 1973, and Heitzman and Heitzman 1974) as they move about, allowing for firm footing as well as guidance to feeding sites and shelter location. The caterpillars may change shelters three or four times as they outgrow them, but usually stay on the same host or near the oviposition site. Tamburo and Butcher (1955) found that marked larvae of Ephyriades brunneus floridensis mostly stayed at the point of release, but one traveled three host bushes away, or about three meters. Living in one place for extended periods presents the problem of waste disposal. If frass accumulates near the shelter, it may provide visual or olfactory clues to predators. Skipper larvae propel frass away from the shelter site (Harris 1862, King 1882, Frohawk 1892, Kendall 1965) with the anal comb, sometimes sending the pellets up to a distance of one meter (Scudder 1889a, Ainslie 1922). Skipper larvae are slow-moving and generally exhibit very subdued behavior. They utilize a variety of defenses against predators, including dropping to the ground and remaining motionless, protective coloration, displaying open jaws, regurgitation of fluids from the mouth, and defecation; they probably also secrete chemicals from a gland on the ventral side of the prothorax (Scott 1986). When disturbed, some notodontid moths, such as Lochmaeus manteo spray a mixture of noxious chemicals from similar glands (Eisner et al 1972). Although I have handled many

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26 hundreds of skipper larvae, I have never been able to detect any secretions from the prothoracic gland. However, Scudder (1889a) noted that ants turned away (apparently in "great disgust") from the caterpillars of Epargyreus clarus. Coloration is an important adaptation that may involve crypsis, aposematic colors, and disruptive patterns. The heads of the larvae may be all dark, dark with contrasting false eye patches (the "monkey faces" of Seitz 1924 and Moss 1949), dark with pale patches, all pale, or pale with dark stripes or spots. These patterns may deter predators with high visual acuity such as birds or lizards. A predator peering into a folded leaf may see false faces or disruptive patterns that are not recognized as potential prey items or be tricked into interpreting all dark or all pale heads as representing emptiness. Belwood (1990) noted similar color patterns on tube-dwelling neotropical Tettigoniidae, suggesting that these katydids and skippers may have similar selection pressures. Combinations of cuticular pigments and internally derived colors produce a variety of body colors commonly ranging from dark green to pale blue. Some species are brightly patterned with red, black, yellow, orange, or white cuticular pigments, suggesting that they are unpalatable. Most are greenish with yellow, orange, white, or rarely black lateral lines. Very few have transverse stripes. The dorsum is usually lightly pigmented or transparent,

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27 revealing the pulsating dorsal heart as a dark line. In larger larvae, the male testes often can be seen as two kidney-shaped bodies located toward the dorsum of abdominal segment six. A few species have caudal false face patterns. For caterpillars to grow larger, they must shed the cuticle. Skipper larvae usually have five larval stages or instars. Temperate species that undergo hibernation may sometimes have a few more stages, especially if reared under laboratory conditions that give confusing signals, such as short days but warm temperatures. The developmental time of the early instars ranges from four days to about a week, but lengthens as the larvae grow larger. The duration of the entire larval period typically ranges from 14 to 45 days. Shortly before hesperiids molt, the head is withdrawn from the old head capsule and shows as a whitish bulge in the prothoracic area. How this is accomplished has not been investigated, but skippers would seem to have special problems making the withdrawal since a key characteristic of the group is the narrow posterior opening of the head. At the moment of molting, the cuticle separates from the head capsule; then the larva walks out of the old skin which remains fixed to a mat of silk threads by the crochets. The old head capsule is cast away with a few lateral movements of the head, and the larva rests while the new cuticle hardens.

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28 Interestingly, the final larval molt is somewhat different from the others, perhaps because the head of the prepupal organism cannot be withdrawn through the posterior opening of the old head capsule. With a few rhythmic struggles, the larval cuticle splits along the dorsum of the thorax and along the frontal suture of the head capsule which remains attached to the skin (Fyles 1895, Ainslie 1922). The exuvium is then slid backwards to the cremaster, and is pushed aside, lying as a crumpled mass at the far end of the cocoon. Changes in coloration may occur in skipper larvae upon molting or in preparation for hibernation or pupation. Most species have dark heads during the first two or three instars, but develop other patterns in later stages. Body coloration may also change as the larvae mature. The change in species of Phocides is extreme, from red with transverse yellow stripes to white. The body color of temperate species may change from green to brown or take on a pinkish cast without shedding the cuticle before entering hibernation (Minno 1981, Capman 1990). Hesperiines often change from green to a creamy color after they stop feeding and seek out overwintering or pupation sites (Frohawk 1892, Heitzman 1965a). The creamy color is derived from whitish fat bodies and other internal organs which are no longer masked by chlorophyll.

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29 Temperate species of hesperiids usually hibernate as partly grown larvae. More rarely, some species overwinter as fully developed larvae within the eggs (Fyles 1895, Pengelly 1961), first instars (Dethier 1939a), last instar larvae (Riley 1876a, Edwards 1885a, Heitzman and Heitzman 1974), or pupae (Scott 1981). After winter has passed, nonmature larvae emerge from their hibernaculae and resume feeding. Those that overwintered as mature larvae (Erynnis and Megathymus species) usually do not feed in the spring, but pupate as directed by environmental cues and soon emerge as adults. Thus species with a pupal or mature larval diapause are among the first to emerge as adults during the spring. Species adapted to temperate climates are generally univoltine or bivoltine, whereas tropical skippers often have three or more generations per year. Unlike most other butterflies, skippers usually pupate in sealed cocoons. The pupa is supported by a silk thread about the junction of the thorax and abdomen, and is anchored to the substrate by cremaster hooks entangled in a small transverse thread or a pad of silk. The cocoon may be formed by sealing the last larval shelter, or the caterpillar may seek out another nearby site. Scudder (1889a) found Epargyreus clarus pupae under boards, shingles, and rubbish under some large locust trees. Temperate species seem more likely to pupate in leaf litter than tropical species.

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30 The pupae of many species are coated with a white or bluish-white wax which is easily rubbed off with a probe or a finger. In some, the wax is a delicate bloom while others are densely coated. Moss (1949) thought that the wax may help keep off moisture while also deceiving predators with the appearance of a cadaver covered with mold. Another protective coloration of the pupa involves the thoracic spiracles, each of which is shielded by a conspicuous cuticular rise. When viewed from the front, these structures become eyes and other features of the head give the appearance of a small animal. Although most, if not all, skippers secrete at least some wax through pores in the pupal cuticle; a few groups also produce guantities of wax in the larval stage. Late in the last instar, many hesperiines and megathymines produce wax from glands concentrated in patches on the ventral side of the body. These deposits have a characteristic shape and distribution that often vary between genera. Within a genus, some species may have wax glands, while other congeners do not. The hollow filaments of wax that issue from the glands (Locke 1960) have a flaky or cottony appearance. The larva uses the wax to coat the inside or plug the entrance to the pupal chamber.

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31 Biology of the Pyrrhopyainae Little is known of the biology of the Pyrrhopyginae. I have examined a few preserved larvae of Pyrrhopyge araxes arizonae, Pyrrhopyge chalybea, and an unidentified Brazilian species, but have not had the opportunity to study living material. Host plants include members of the Anacardiaceae, Anonaceae, Clusiaceae, Fagaceae, Flacourtiaceae, Lauraceae, Meliaceace, Myristicaceae, Myrtaceae, Sapindaceae, and Sterculiaceae. The larvae live in pyrgine-type shelters of folded leaves (Moss 1949, Burns 1964a). Larvae described by Moss (1949) and Comstock (1956b) have a reddish ground color and contrastingly colored transverse stripes or spots and some resemble the pyrgine genus Phocides. Pupae are also reddish but are covered with white wax in the manner of many pyrgines. The last shelter of Myscelus pardalina was perforated by a few large holes (Moss 1949), a specialized behavior of certain pyrgines such as Quadrus species. Pupation takes place in the last larval shelter or in leaves on ground. Moss notes that Pyrrhopyge larvae are very pugnacious when disturbed and often leave the shelter when it is cut from the host. Biology of the Pyrainae As Scudder (1875) noted, members of the Pyrginae usually feed on dicotyledonous plants. Many pyrgines eat legumes (Fabaceae). However, there has also been a tremendous radiation away from the use of legumes such that

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32 at least 44 other families of plants are eaten by New World pyrgines. Many of these hosts contain strongly scented secondary plant chemicals. Remarkably, there are also a few species of Urbanus that feed on grasses (monocots), while their congeners eat legumes (Kendall 1976)! Twenty-four plant families are utilized by pyrgines reported from the U. S. (Table 1). Female pyrgines usually lay eggs singly on living leaves of the host but may also choose flower buds, stems, fruit, or dead leaves (Panton 1897, Tamburo and Butcher 1955, Minno 1981). Pholisora catullus deposits eggs on the upper side of the leaf near the midrib (Edwards 1884, Capman 1990), but most other species seem to prefer the edges of the lower surface. Urbanus dorantes freguently oviposits on the inflorescences of Desmodium species in Florida. One pyrgine, Epargyreus clarus has been observed to lay eggs on nearby plants after inspecting and touching the true host (Opler and Krizek 1984, MCM observation). Newly laid eggs may be white, yellow, pale green, or brownish. Urbanus proteus and Autochton cellus females lay stacks of eggs containing from two to six individuals (Comstock 1881, Quaintance 1898, Clark 1934, 1936). The eggs in the stack are offset at an angle (Clark 1936), allowing larvae in the lower eggs to emerge in the normal way from the exposed micropylar area. Quaintance (1898) noted that the most distal egg in the string is the first to hatch.

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33 Although these species begin life communally, the newly hatched larvae lead separate lives, each constructing a shelter around the margin of the leaf. Kendall (1976) reported finding ten larvae of Codatractus hyster in a single shelter and this may be a truly gregarious species. First instar larvae usually have dark brown or black heads. However, some Erynnis species have pale head capsules. Newly emerged pyrgine larvae do not eat the remains of the egg, but usually seek out the margin of a leaf and begin constructing a shelter. The larva eats two converging slits into the leaf margin, except for a narrow portion which acts as a hinge. The resulting flap is then folded over and secured with three or four strands of silk (Lintner 1872, Edwards 1885b, Panton 1897, Quaintance 1898, Hayward 1927a, Clark 1936) in such a way as to form a domeshaped roof. The inside of the shelter is lined with silk. Clark (1934) described the shelters of Autochton cellus as limpet-like. Most species fold the flap over onto the upper surface of the leaf, but Zestusa dorus Paches polla, and Atarnes sallei tie the flaps to the underside (Klots 1971, Kendall and McGuire 1975, Kendall 1976). Erynnis species that feed on woody plants often construct their shelters where a vein reaches the margin (Kendall and McGuire 1975) or at the tip of the leaf (Kendall 1965). The vein adds strength to the hinge. Rather than cutting into the leaf margin, the first instar

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34 larva of Nisoniades bessus godma eats a circular channel near the center of a leaf then ties the flap over (Kendall 1976). Newly emerged larvae of Heliopetes laviana eat away the leaf surface, then fold a flap over the weakened spot (Kendall 1965). Young Pyrgus species usually seek out deep wrinkles or overlapping leaves in which to build shelters (Panton 1897, Minno 1981) and Pyrgus scriptura clips off the dense stellate hairs of the host and ties them together with silk to form a covering (Minno 1981). The young larva of Urbanus procne forms a "lean-to" shelter by eating away part of a blade of grass so that the terminal end can be pulled over about 85 degrees, but older larvae simply hide in the detritus at the base of the plant (Kendall 1976). Older pyrgine larvae may make more box-like shelters by eating channels perpendicular to the leaf margin and folding the rectangular flap over (Clark 1936, Edwards 1885a) or tie the edges of one or more leaves together. Heliopetes macaira may hide in the flower bracts or in dead leaves on the plant or the ground, but often fastens two leaves together, then cuts the petiole of the one which is to be the roof, which withers and dies (Kendall 1965). Similarly, Pyrgus communis often uses a living leaf and detritus (Dethier 1944b, Minno 1981). A few species such as Erynnis persius, Atarnes sallei, and Quadrus species eat small holes in the shelter, then web them over with silk (Scudder 1889a, Kendall 1976, and Young 1991). Before moving to a larger

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35 shelter, pyrgine larvae clip the silk threads holding the flap (Scudder 1889a, Clark 1936). Autochton cellus usually makes the first shelter toward the middle or tip of a leaflet and the second more toward the base (Clark 1936). Pyrgine larvae sometimes construct shelters on plants that are not eaten, but are physically near the true host. Clark (1936) found A. cellus shelters on hickory, sassafras, and grape near to the true host (Amphicarpa bracteata,) and I have found shelters of the legume-feeding U. proteus on blackberry and oak leaves. One fourth instar U. proteus larva, which I placed on a potted Desmodium incanum, built a shelter on a small potted guava adjacent to the host. This behavior suggests that some predators may first direct their hunting toward the food plant rather than the caterpillar. Pyrgine larvae usually feed on the edges of leaves near to the shelter. Scudder (1889a) noted that the first instar larvae of Erynnis persius feed on the surface layers of the host leaves. Similarly, Pyrgus scriptura larvae feed on the leaf surface within the shelter, rather than the edges of leaves. A few species such as Heliopetes ericetorum and Staphyllus hayhurstii may eat holes in the leaves (Coolidge 1923, Heitzman 1963). Young larvae prefer young tender leaves, but older caterpillars are often able to feed on tougher mature foliage. Small larvae tend to rest on the top of the shelter in an inverted position (Clark 1936, Kendall 1965). Pyrgine

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36 shelters are usually more circular than linear, at least during the early instars, and some species (Achalarus lyciades and Heliopetes ericetorum) assume a coiled resting position, with the head and tail facing the same direction in this unrestricted space (Scudder 1889a, Coolidge 1923). When disturbed, pyrgine larvae use several defensive behaviors in order to protect themselves. Most will turn the head toward the direction of the stimulus with open jaws, and attempt to bite or thrash the head from side to side. Scudder (1889a) noted that Erynnis juvenalis scrapes the open jaws backwards on the surface of the leaf, making a scratching sound. While they may appear ferocious, I have never actually been bitten by a pyrgine larva. They usually thrash briefly if picked up with fingers, and some such as U. proteus E. clarus and A. cellus spit out a watery green-colored fluid if greatly harassed (Scudder 1889a, Clark 1936, MCM observation). Kendall (1976) observed Urbanus teleus a grass-feeder, to fall to the ground upon being disturbed and lie motionless for long periods of time in the manner of some hesperiines. The larval stage typically lasts from 15 to 30 days for most pyrgines that do not have diapause reguirements. Temperate species usually pass the winter as partly or fully grown larvae; thus for these skippers, the larval stage may extend over several months. A few species diapause as

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37 pupae. I do not know of any pyrgines that overwinter as eggs or first instar larvae. Most pyrgines pupate in cocoons fashioned from the last larval shelter or spun in detritus and debris on the ground. Kendall (1965) noted that although Cogia outis usually pupates on the ground, one larva burrowed under the soil surface and pupated in a silk-lined chamber. Within the cocoon, the pupa is supported by a silk girdle about the middle and is attached to silk threads by the cremaster hooks at the caudal end. Many other butterfly families and some moths use this arrangement, but pyrgines have a specialized feature not known in any other group. The silk girdle supporting the middle of the body also has a median attachment to the cocoon. Thus if the ventral surface of the pupa is toward the bottom of the page, the shape of the girdle is of an inverted letter "Y" (Forbes 1923). Similarly, the cremaster attachment may be Y-shaped (Coolidge 1923) The pupae of pyrgines may be opaque dark brown, or greenish with dark markings on the dorsum and wing cases. A few have a short peg-like horn on the frons. The pupae of many pyrgines are thickly coated with wax. The cephalic end of the body is frequently modified to give the appearance of a larger animal with staring eyes. Clark (1936) likened the Autochton cellus pupa to the head of a manatee when viewed

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38 from the front. Moss (1926) and Box (1928) describe the frightening appearance of Bungalotis pupae. Biology of the Heteropterinae Not much is known of the biology of the Heteropterinae, except for the Arctic Skipper, Carter ocephalus palaemon (Fletcher 1889, Frohawk 1892). The morphology, coloration, and behavior of the immature stages of this species are closely similar to that of the Hesperiinae. Biology of the Hesperiinae Unlike the pyrgines, Hesperiinae feed on monocots (Scudder 1875). Most New World hesperiines eat grasses (Poaceae), sedges (Cyperaceae) and palms (Arecaceae), but some also feed on the leaves of Bromeliaceae, Cannaceae, Liliaceae, Marantaceae, Musaceae, Orchidaceae, and Zingiberaceae. Host plants of hesperiines recorded from the U. S. include 8 families of monocots (Table 1). Hesperiine females usually glue the eggs singly to the leaves of the host. Although most hesperiines prefer to deposit their eggs directly on the larval food plant, Hesperia species, Polites species, Wallengrenia otho Atalopedes campestris Ochodes yuma, Poanes hobomok Lerodea eufala, and Panoquina panoquinoides will sometimes lay eggs on dicots or other substrates near the larval host plant (Scott 1986). On two occasions, I have observed Polites vibex, a grass feeder, to lay eggs on dicots after fluttering closely over patches of grass. However, I have

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39 also seen females of this species oviposit directly on grass on several occasions. Some populations of Hesperia lindseyi oviposit on lichens growing on fence posts and trees near the grass on which the larvae feed (MacNeill 1964). Dana (1981) discovered that Hesperia ottoe in southwestern Minnesota oviposits on the disk flowers of pale purple conef lower (Echinacea pallida), an adult nectar plant that is not eaten by cattle. After eclosion, the young larvae of these hesper lines must wander, sometimes over relatively long distances, to find a host and begin feeding. Although the introduced skipper, Thymelicus lineola, always lays eggs in rows inside the sheaths of grasses, there are no known New World hesperiines that consistently lay clusters of eggs. Some such as Calpodes ethlius and Problema byssus may haphazardly deposit eggs in small clusters on occasion (Chittenden 1912, MCM observation). The eggs are usually white or green, but a few species have yellow or pinkish eggs. Green or yellowish eggs frequently develop a bright red ring and apical spot after a few days (Dethier I942e, Heitzman 1964a, 1966, Brown and McGuire 1983, MCM observation). The newly emerged larvae of many hesperiines first eat the remains of the egg, leaving only the shiny circular base, before building a shelter (Fyles 1895, Chittenden 1905, Ainslie 1922, Coolidge 1922, 1925, Dethier 1942d, Nielsen 1958, Heitzman 1964b, 1965a, 1966, Heitzman and

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40 Heitzman 1970b, MCM observation). Shelters are often formed at the tip of a leaf by tying the edges together with silk or by eating a slit part way into the leaf and folding over the flap. Depending upon the species, the flap may be folded onto the upper or the lower surface of the leaf. Some hesperiines, such as Euphyes species and Atrytonopsis hianna, tie the tips of two overlapping leaves together (Heitzman and Heitzman 1974, MCM observation). Calpodes ethlius cuts two slits parallel to the leaf margin and folds the flap over. Older larvae typically live in tubular shelters formed by tying the edges of several leaves together and lining the inside with silk. The distal end of the shelter may be tightly closed. Lerema accius feeds below the shelter, nearly severing the midvein, so that the shelter dangles down from the cut. The older larvae of Asbolis capucinus fold the edges of a palm leaflet under and eat the leaf tissue near the tip, except for the midrib, which protrudes like a long spine. Wallengrenia species are most unusual in that they form a case from a piece of leaf. They drag the case to feeding sites, temporarily attaching it to the host with silk (Kendall 1960, MCM observations). Although most hesperiines tend to live upon the leaves of the host, Hesperia species and their relatives usually construct horizontal shelters of silk and detritus near the base of the plant. Sometimes the shelters may extend below

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41 ground (Cockayne 1952, MacNeill 1964, Heitzman and Heitzman 1970b). McCabe and Post (1977) found larvae of Hesperia comma assiniboia in silk-lined burrows under dried cattle droppings. older larvae of a few hesperiines such as Panoquina species, Nastra species, Oarisma powesheik, Pompeius pompeius, and Decinea percosius do not make shelters, but rest exposed on leaves or hide in natural recesses (Dyar 1892, McAlpine 1973, Kendall 1976, Kendall and Rickard 1976, MCM observations). Although Kendall (1966b) mentioned that Poanes viator does not make a shelter, I have found older larvae in typical shelters as well as in recesses at the bases of the leaves in Alachua County, Florida. As Shapiro (1971) noted, the larva usually reinforces the recess with some silk. First instar larvae of hesperiines usually have a dark head. The only exception that I have observed is Poanes viator which has a cream-colored head with some small dark spots. Species that live on leaves are mostly green with longitudinal stripes or reticulate patterns of white or pale yellow. The head capsule frequently bears dark stripes or spots on a pale ground. Hesperiines that live near the ground surface are dully colored brown or dark green and have heads that are plain black or black with pale patches. For some species, the caudal end may be modified to provide a defense from predators. During the act of defecation, the larva must expose the terminalia which may

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42 attract the attention of lizards or other predators. Some hesperiines (such as Atrytone delaware and Asbolis capucinus) have a flattened, pale-colored suranal plate rimmed or patterned with black. In addition, the last pair of spiracles is often black, contrasting with the pale body color. When viewed from behind, the spiracles become eyes and the suranal plate a broad bill. Many hesperiines, especially those that do not build shelters, rest with head flattened ventrally against the substrate and the mouth protracted forward (Forbes 1960, MCM observation). Although most hesperiine larvae will rapidly retreat into the shelter if threatened, they tend to be relatively docile. When disturbed, they may drop from the plant, curl ventrally, and remain motionless for a long time (Panton 1897, Heitzman 1964a, MacNeill 1964, Heitzman and Heitzman 1970b). Hesperia species and Amblyscirtes belli thrash violently within the shelter if disturbed (MacNeill 1964, Heitzman 1965b). Hesperiines that diapause during the winter usually do so as partly grown larvae (Scott 1981). Less commonly, they may pass the winter as fully developed larvae within the eggs, first instar larvae, last instar larvae, or pupae. Although some species may complete the larval stage in about two weeks ( Corns tock and Dammers 1931, Kendall 1965), one month or longer is more typical. During the last instar

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43 many species develop ventral wax glands on the abdominal segments Like the pyrgines, hesperiines usually pupate in sealed cocoons. However, Lerema accius and Cymaenes tripunctus pupate exposed, but under a leaf with the edges slightly drawn together. Panoquina panoquin pupates exposed on the upperside of leaves. Amblyscirtes belli, Niconiades nikko and Quinta cannae cut the last shelter from the host and form the cocoon on the ground (Kendall and McGuire 1975). Hesperiine pupae are supported by a simple silk girdle about the middle and are attached to a pad of silk by the cremaster. Species of Atrytone and Euphyes lack these supports and pupate loosely in the cocoon in a vertical position. The cremaster of these species is modified into one or two spines and lacks hooks. The pupal cuticle of hesperiines tends to be lightly pigmented and thin. Those that pupate exposed are pale green and may have yellow or white stripes on the abdomen. Most have a cream colored or slightly greenish ground color with dark patches or lines on the head, wing cases, and dorsum. Some groups of hesperiines have an elongate point on the frons of the pupa (a "beak") that may be upturned or T-shaped at the tip. Atrytone del aware and Poanes viator have a bulbous frons reminiscent of the Australian Trapezitinae, some of which are wildly modified into false face patterns resembling the jaws of termite soldiers. As

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44 development proceeds, the eyes become reddish, the wings darken, and just before emergence the adult is clearly visible through the transparent cuticle. When the adult emerges, the pupa splits along the dorsum of the thorax, the edges of the head, and the costal margin of the wing cases. The pupal cap (the cuticle covering of the head) is often torn away completely. The exuvium freguently contains a clear or yellowish meconium, which is expelled during emergence (Panton 1897, MCM observation) Biology of th e Meaathvminae I have closely observed the biology of Megathymus yuccae and Megathymus cofaqui in Florida, but have no experience with other Megathyminae. Fortunately, there is a large literature base available on this interesting group. Riley (1876a,b, 1877), Freeman (1951a), Tinkham (1954), Remington (1959), Roever (1964), and Wielgus and Stallings (1974) provided much of the summarized information presented below. Not only do the Megathyminae have a highly specialized biology that differs considerably from other skippers, but the three tribes within this subfamily have each evolved their own peculiar specializations. Megathymines feed only on plants in the Agavaceae. Megathymus species feed in the stem and roots of yuccas. Members of the closely related genus Stallingsia bore into the stem of Polianthes species. Although Yucca species have

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45 large, starchy roots, they also contain saponins and some are commonly known as soap plant. The sap of many yuccas and agaves is highly irritating to human skin and causes a severe poison ivy-like dermatitis. The specialized host preferences of many megathymines seems due, in part, to female choice, as the larvae of Megathymus yuccae can be reared in the laboratory on starchy roots of guite different chemistry such as potatoes. Megathymus species (Megathymini ) glue their eggs singly to the leaves of the host but sometimes select nearby dead leaves or other plants (Riley 1876a, b, 1877, MCM observation). Females often select small or isolated plants in favor of those that are more robust or clumped (Oslar 1900, Tinkham 1954, Stallings and Turner 1956, Remington 1959, McCabe and Post 1977). The eggs are greenish when first laid but later turn white, brownish, or pinkish. They hatch in 7 to 16 days (Wielgus and Stallings 1974). Newly emerged larvae are reddish. The remains of the egg are not eaten. The young larva of Megathymus yuccae usually constructs a covering of silk and plant fibers over itself near the tip of a leaf and feeds on the epidermal tissue. Later, the larva ties together the youngest leaves at the center of the plant, then bores into the stem. Most other Megathymus species bore directly into the stem after emerging from the egg.

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46 Unlike other skippers, larvae of Megathymus species make a cigar-shaped "tent" of silk, plant fibers, and detritus. Some species such as M. yuccae make the tent when they begin boring into the stem. As the larva grows, the tent is enlarged, and may be seven or eight centimeters long by pupation time. All during the feeding period, frass is expelled from the tip of the tent, but rather than being propelled away, the pellets are simply dropped from the opening. Megathymus streckeri and Megathymus cofaqui are somewhat different in larval behavior. These species bore directly into the stem after emerging from the egg, and pack the frass behind as they feed. When ready to pupate, the larvae burrow from the roots upward through the soil and make a short tent of silk, detritus, and soil. Mature larvae of Megathymus species are brownish or cream-colored with reddish brown or black heads. Wax glands on the ventral side of abdominal segments 7 and 8 supply copious quantities of wax to line the tunnel. Overwintering occurs in the larval stages. Megathymus yuccae hibernates as a mature larva and pupates in the spring without further feeding. The tunnels may extend several feet into the stem and roots of the plant and although the terminal bud is killed, the damage done by the larva usually has a pruning effect, causing the plant to produces new shoots (Remington 1959).

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47 Large species such as Megathymus ursus may inflict enough damage to kill the host (Stallings and Turner 1956). The pupae of Megathymus species are brown and lack silk supports. The tip of the abdomen is freely movable, allowing the pupa to move up and down in the tunnel. When disturbed, they guickly retreat to the bottom of the burrow. Before emergence, they usually position themselves just below the tent. Tinkham (1954) speculated that these movements may serve to regulate temperature as well as retreat from disturbances. Megathymus larvae defend themselves by biting with the jaws, locking themselves in the burrow, defecating, and expelling a brownish fluid from the mouth (Wielgus and Stallings 1974). After the adult emerges, it crawls upward and pushes its way through the loosely sealed opening to the tent. A yellowish meconium is expelled into the pupal exuvium. The adult usually perches on the tent while expanding and drying the wings. Megathymus species have only one or two generations per year, depending on the species. Aegiale hesperiaris (Aegialini) and Agathymus species (Agathymini) feed only on century plants {Agave species). Aegiale hesperiaris glues small clusters of eggs to the host leaves (Dampf 1924). Agathymus species either perch on the host or nearby and drop or flick the eggs haphazardly (Comstock and Dammers 1934, Freeman 1951a) or lay the eggs on a host leaf. However, since no adhesive is applied, the

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48 eggs fall to the ground (Roever 1964). Stallings et al (1985) noted that females of Agathymus estelleae dropped between five and ten eggs at one time. Egg color changes from green to reddish in Agathymus species (Comstock and Dammers 1934, Roever 1975). One species, Agathymus estelleae, produces a range of egg colors varying from green to beige to pinkish (Stallings and Stallings 1986). The duration of the egg stage is from 22 to 52 days (Roever 1964) Newly emerged larvae do not eat the remains of the egg, but crawl toward the distal half of a leaf and make a short burrow directed toward the tip (Roever 1964). Agathymus larvae spin a trail of silk when moving about on the leaf surface. The burrow is also lined with silk to prevent sap from entering, which is not always successful, forcing the larva to move to another site or another plant. The frass produced by young larvae is relatively dry and green (Roever 1964) After several molts, the larva passes the winter within the burrow, then moves to the base of the leaf and makes a new tunnel directed toward the stem. This tunnel is also lined with silk except at the distal end, where the larva feeds primarily on sap. The entrance may be on the upper side, lower side or either surface of the leaf, depending upon the species. Defecation occurs at the tunnel entrance, after which the entry is partially barred by a few strands

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49 of silk. The oily, dark brown frass accumulates around the entrance to the tunnel during the summer feeding period. Aegiale hesperiaris Agathymus evansi and Agathymus stephensi cause harm to the plant, as evidenced by necrotic tissue near the point of entry. Agathymus larvae have dark heads and greenish or bluish white bodies. When feeding is completed, the entire tunnel is lined with silk and powdered with wax, except for Aegiale hesperiaris and Turnerina hazel ae which do not develop wax glands. Instead of constructing a tent over the tunnel entrance as do Megathymus Agathymus exude a pliable silk from the mouth and fashion it into an opaque finger naillike door. The silk door completely seals the tunnel entrance. Pupation occurs in the bottom of the tunnel The brown pupa is not supported by silk and is oriented with the head toward the entry. At the time of emergence, the adult crawls from the exuvium and pushes on the trap door, which swings open on a hinge along the bottom edge. The adult then exits the tunnel expands and dries the wings and flies away.

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CHAPTER 4 GENERAL MORPHOLOGY This chapter describes external morphological characteristics of the eggs, larvae, and pupae of U. S. skippers. Many structures exhibit a wide range of variation between the various species. Morphological variation is further noted in the species diagnoses of Chapter 5. Eggs The eggs of hesperiids found within the U. S. vary from 0.4 to 1.8mm in height and 0.6 to 3mm in diameter. Hesperiid eggs are wider than high. Shape varies from hemispherical (vertical sides) to somewhat conical (sloping sides) (Figure 3). Numerous minute holes (aeropyles) occur on the surface of the chorion. The micropylar area at the top of the egg is surrounded by rosette patterns of sculpturing (Figure 4). Scudder (1875) first noted that the sculpturing of hesperiid eggs seemed to vary consistently between subfamilies. Pyrgines have ribbed eggs (Figure 3) while hesperiines have smooth eggs or eggs with polygonal patterns of sculpturing (Figure 5). While this pattern generally holds true, some Pyrginae, such as Cogia out is have polygonal sculpturing. Other pyrgines have vertical rows of short spines. Megathymines, like hesperiines, have 50

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51 non-ribbed eggs with polygonal sculpturing patterns. Heteropterine eggs are described as having vertical grooves (Higgins 1975); however, a Butleria species has ribbed eggs (Herrera et al 1991), and the egg of Carter ocephalus palaemon is smooth with polygonal sculpturing (Brooks and Knight 1982). I could not find specimens, detailed descriptions, or illustrations of pyrrhopygine eggs. Larvae Hesperiid larvae, like the caterpillars of other lepidopterans, have a head, three thoracic segments, and ten abdominal segments. Each thoracic segment bears a segmented pair of legs. In addition, abdominal segments three through six and ten each have a pair of unsegmented prolegs. Spiracles are located on the prothorax and abdominal segments one through eight. The spiracles on the prothorax and abdominal segment eight are usually slightly larger than the others. The body segmentation of hesperiids tends to be ill defined and confused by folds or wrinkles of the cuticle (annuli or plicae). Mature larvae of U. S. hesperiids range in length from 8 to 86mm. The prothorax is larger than the head in Megathyminae, about the same size as the head in Pyrrhopyginae, and generally smaller than the head in the Pyrginae and Hesperiinae. Body width tends to be more or less uniform. However, the thorax of Megathymus species is slightly enlarged. These plant boring larvae are able to

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52 expand the thoracic region and thereby lock themselves into their tunnels upon being disturbed by a predator or human observer. Other Megathyminae are somewhat wider in the area of abdominal segments three to five. The transverse width of the U. S. hesperiids varies from 1.6 to 11.7mm. Hesperiines tend to be more slender than pyrgines. The setae occurring on skipper larvae exhibit variation in shape (Figure 6). Although simple setae are the most abundant type, some species also have modified setae. Modified setae vary in the shape of the tip, which may be blunt, slightly expanded, or greatly expanded. Pyrgines and some hesperiines frequently have setae with expanded tips. Branching setae are another modified type found only on the head of certain pyrgines. The length of the setae on the body and head varies between species. On an individual caterpillar, the setae are usually longest toward the ventral side of the body and shortest on the dorsum. The length of the setae on the dorsum of abdominal segment four varies from < 0.1 to 6.4mm for U. S. species. On the head, setae range from < 0.1 to 5mm near the apex to 0.1 to 5.7 on the ventral side. Pyrrhopygines have very long setae. Most other hesperiids have short or very short setae, but a few pyrgines and hesperiines have long or moderately long setae. Lenticles are small sclerotized plates that seem to represent modified setae. Franzl et al (1984) found the

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53 lenticles of Calpodes ethlius to be innervated secretory structures that probably produce lipids. The positions of the lenticles change with each larval molt (Franzl et al 1984), alternating between two basic patterns. The expression of lenticles at a particular location may not always occur or they may be present in excess. Not only is there variation between individuals, the number of lenticles often varies from side to side of a single larva. Several larvae usually need to be examined in order to determine the pattern for a particular species. A generalized map of lenticle distribution is presented in Figure 7. Table 2 lists the maximum number of lenticles observed in the present study at eight positions on the prothorax, five positions on thoracic segments two, three, and abdominal segments one through nine, and two positions on abdominal segment ten for subfamilies Pyrrhopyginae Pyrginae, Heteropterinae, and Hesperiinae. Megathymines also have lenticles, but usually in a supraspiracular cluster or in small groups on the ventral side of abdominal segments one, two, seven, or eight (Figure 8). Table 3 presents the number of lenticles present on ten megathymine taxa. Pigments that may be present in the cuticle of hesperiid larvae are red, yellow, orange, white, or rarely black. Unfortunately, most of these pigments are destroyed in preserved larvae. Literature descriptions are often unreliable as they may refer to wax deposits, cuticular

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54 Table 2. Maximum number of lenticles observed per segment and position on Hesperiidae larvae in the present study. Lenticles may be found in eight possible positions on the prothorax (Tl), five positions on thoracic segments 2-3 and abdominal segments 1-9, and two positions on the last segment (A10). An asterisk indicates 10 to 15 lenticles present. MAXIMUM # LENTICLES PER POSITION PYRRHOPYGINAE : P. araxes PYRGINAE: P. pigmalion P. palemon E. zestos E. clarus E. exadeus P. leo C. catillus T. undulatus P. mexicanus P. procerus C. alcaeus c. arizonensis u. proteus u. esmeraldus u. dorantes u. teleus u. procne A. fulgerator A. gilberti A. anaphus A. cellus A. lyciades A. casica T. bathyllus T. pyl ades T. diversus T. mexicanus T. confusis T. drusius C. potrillo C. hippalus C. outis C. caicus Tl T2-3 Al-2 A3-6 A7-9 A10 SPECIES 12345678 12345 12345 12345 12345 12 10001010 00100 00114 10120 00122 10 10001020 01010 10112 10110 00110 01 10001140 01120 10110 10110 10110 01 10001120 11110 10111 10121 10111 10 10001120 10100 10111 10021 10111 11 10001120 11014 10111 10111 10111 10 10001130 1300* 10111 10111 10111 11 10001120 11000 10111 10111 10011 11 10001020 10014 10021 10021 10021 01 10100213 1110* 10111 10122 10121 11 10001244 1210* 10112 10113 10111 11 10001020 11100 10113 10122 10122 11 10001020 01100 10113 10113 10112 11 10001120 11000 10111 10111 10111 11 10001120 11100 10022 10121 10011 11 10001120 11100 10111 10111 10111 11 10001122 01003 10111 10112 10112 01 10001110 11110 10011 10111 10111 11 10001120 11100 10121 10121 10121 11 10001120 01000 10121 10121 10121 11 10001120 01100 10121 10121 10111 11 10001120 11100 10121 10121 10121 11 10001120 11000 10011 10011 10011 11 10001120 11000 10111 10111 10111 11 10001120 OHIO 10111 10111 10111 11 10001120 12113 10112 10111 10112 11 10000120 01000 10011 10111 10111 11 10001100 01000 00011 10011 10111 10 10001122 12104 10111 10111 10111 11 10001121 11012 10111 10111 10111 11 10001020 11000 10111 10111 10111 11 10001020 10000 10011 10011 10012 01 10101020 10002 10012 10122 10012 01 10001120 00000 10011 10021 10011 01

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Table 2 — continued. 55 MAXIMUM # LENTICLES PER POSITION Tl T2-3 Al-2 A3-6 A7-9 A10 SPECIES N. rubescens S. ceos S. mazans S. hayhurstii C. canescens X. trixus S. pulverulenta S. zampa A. thraso T. ruptifasciatus G. gesta E. brunneus E. icelus E. brizo E. juvenalis E. propertius E. horatius E. tristis E. martialis E. pacuvius E. zarucco E. funeralis E. baptisiae E. persius E. afranius P. centaureae P. rural is P. scriptura P. communis P. albescens P. oileus H. ericetorum H. lavianus H. macaira H. arsalte C. nessus C. limpia P. catullus P. alpheus 12345678 12345 12345 12345 12345 12 10001111 12010 10021 10022 10021 01 10001110 00100 10111 10111 10111 11 10001112 02101 00111 10111 00121 11 10001113 02100 00121 10111 10121 11 10001110 11010 10121 10121 10121 11 10001110 OHIO 10021 10021 10021 11 10002120 11000 10122 10110 10120 01 10001120 11010 10021 10120 10120 10 00001110 10000 10021 10111 10111 11 00001120 OHIO 10122 10121 10121 11 10001120 12100 10112 10111 10111 11 10001100 11010 10012 10021 10022 01 10001110 12010 10122 10121 10121 11 10001110 12010 10123 10121 10111 11 10001010 12110 10122 10121 10111 11 10001110 12010 10122 10120 10110 11 10001110 11010 10122 10121 10110 01 10001110 12110 10122 10121 10111 01 10001010 02100 10122 10121 10111 10 10001010 12100 10122 10121 10111 11 00001110 12110 10122 10111 10111 11 10001110 02100 10122 10121 10111 11 10001110 12100 10122 10111 10111 11 10001110 12100 10122 10121 10121 11 10001110 12100 10122 10111 10111 12 10001120 12110 10021 10121 10121 11 10001110 11000 10121 10011 10111 11 10000110 11010 10111 10121 10121 11 10001110 11110 10111 10111 10111 11 10001110 12010 10111 10111 10111 11 10001110 02010 10111 10111 10111 11 10001110 02010 10111 10111 10111 11 10001110 00010 10021 10021 10121 11 10001110 00000 00111 10111 10111 11 10001110 12010 10111 10111 10111 11 10001120 12010 00111 10111 10111 11 10001120 12110 10111 10111 10111 11 10001111 01100 10111 10111 10111 10 00001110 01000 00110 10111 10100 11 HETEROPTERINAE : P. pirus 00110110 03100 00112 00117 00121 10

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Table 2 — continued. 56 MAXIMUM # LENTICLES PER POSITION Tl T2-3 Al-2 A3-6 A7-9 A10 SPECIES HESPERIINAE: S. malitiosa V. perigenes N. lherminier N. julia N. neamathla C. tripunctus L. accius L. liris P. philetes A. numitor A. arene C. aurantiaca C. minimus T. lineola H. phyleus Y carus P. eunus H. uncas H. juba H comma H. woodgatei H. leonardus H. pahaska H. viridis H. attalus H. meskei P cor as P. sabuleti P mar don P draco P baracoa P. themistocles P. origenes P. mystic P sonora P. vibex W. otho W. egeremet P verna A. campestris A arogos A. del aware P. byssus 12345678 12345 12345 12345 12345 12 00100010 02000 01112 01118 00112 11 00100000 10110 10011 1001* 10111 01 00000010 02100 01111 11116 11111 10 00000110 02100 00101 00101 00112 10 00000110 02100 01121 01116 00111 10 00100110 03100 01112 01117 01111 01 00100110 03010 01112 01119 01111 01 00101110 03010 01112 01118 01111 01 00000010 02010 01112 01113 01112 11 00101020 12010 01121 10123 11111 17 00001120 02000 01111 01111 01111 14 00000000 01000 00102 00104 00110 01 00101020 01000 00011 00103 00111 11 00101020 01000 00111 01116 01111 01 00101120 12010 01121 01113 01111 10 00101120 02000 01111 01114 01111 10 00001010 11010 10011 10013 10011 01 00101010 11010 10011 10018 10011 12 10120010 02010 01111 11116 10111 11 10110120 02100 01111 01119 00111 11 10121110 10010 10011 10015 10011 11 00101120 02100 01111 01116 01111 11 00101120 12010 11111 11116 11111 11 00100020 11010 10011 10015 10111 11 00101010 11010 10011 10017 10111 11 00100010 02010 01111 01113 01111 11 10101110 12010 10111 11118 11111 11 00101010 01010 11111 10123 11021 11 00101010 13010 11111 11113 11111 01 10110120 02000 10011 1111* 01011 11 00000120 03000 01111 01115 01111 10 10121120 03100 01111 01115 01111 10 10140020 02100 01111 11117 01111 10 10120020 03000 01111 OHIO 01111 10 10121020 12110 01011 11110 01111 10 10110120 02100 01111 01114 11111 10 10101110 13010 11111 11H6 11111 H 00000120 02000 01111 01117 01111 10 11111010 11010 10011 10018 10111 11 11121120 02000 01111 01116 01111 10 00001110 12100 11112 11117 11112 00 10111120 02000 00112 00113 10111 01 00101120 02110 01112 0111* 01112 10

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Table 2 — continued. 57 MAXIMUM # LENTICLES PER POSITION SPECIES Tl T2-3 Al-2 A3-6 A7-9 A10 12345678 12345 12345 12345 12345 12 O. sylvanoides O. agricola O yuma P hobomok P. zabulon P. taxiles P. aaroni P. yehl P. viator P. melane C. haitensis E. alabamae E arpa E. pilatka E. dukes i E. bimacula E. ruricola A. capucinus A. hianna A. loammi A. aesculapias A. vialis A. celia L. eufala O. maculata C. ethlius P. panoquin P. panoquinoides P. ocola P. sylvicola N. nyctelius 10150020 03100 01122 01117 01111 11 10130020 02100 01111 01119 01111 11 10140120 02000 01111 OHIO 01111 11 11141110 11010 10001 1001* 10011 11 10130000 03000 01111 01118 01111 11 10101220 11010 10021 1O029 10021 11 00131110 11010 11011 1101* 11111 11 12141110 12010 11111 1111* 11111 11 031*1120 12110 11111 11118 10111 01 10141020 02110 01111 01117 01111 11 10110120 01100 11111 11116 01111 01 00101130 02100 01011 01119 01111 11 00100030 12110 11112 1111* 11111 11 00101130 12110 11111 11118 11011 11 00101030 12110 11111 11117 11111 01 00101130 02100 01112 01115 01111 11 00101130 02100 01112 01116 01111 11 10111010 11010 11123 11119 11232 01 00100110 02100 01212 0111* 01112 01 10130120 02100 01212 0111* 01112 01 00111010 11000 10011 10010 10111 11 10120110 03100 00112 00119 00112 10 00100110 02100 00122 00116 00111 10 00100110 03000 01112 01115 01111 11 00001110 12110 11112 11118 11111 11 00102020 05000 01112 01117 01112 01 00101000 11110 11111 1111* 10111 10 00101020 01010 01012 01114 01112 11 00101020 01010 01111 01113 01111 11 00101010 OHIO 01111 01115 01111 11 11121020 02000 01111 01111 01111 11

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58 Table 3 Maximum number of lenticles observed per segment and position on Megathyminae larvae in the present study. Lenticle positions are labelled 1 for lenticles above the spiracle line, 2 for lenticles below the spiracle line, and 3 for ventral lenticles. Position 1 on abdominal segment 10 refers to lenticles on the lateral margin of the caudal proleg. An asterisk indicates 10 to 15 lenticles present. MAXIMUM # LENTICLES PER POSITION Tl T2-3 Al-2 A3-6 A7-9 A10 SPECIES 123 123 123 123 123 A. valverdiensis A. stephensi M. y. yuccae M. y. martini M. y. reinthali M. coloradensis kendalli 000 200 503 700 802 M. coloradensis navajo M. cofaqui M. u. ursus M. u. violae 300 *00 700 400 500 200 800 400 400 301 100 300 612 910 *02 100 120 813 610 402 1 000 200 515 800 802 000 300 903 800 601 000 900 304 *00 *01 010 300 602 500 402 000 210 302 900 702

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59 thickenings, internally derived colors, or may not have been included in the description. Megathymines lack bright cuticular pigments, but may be cream-colored, bluish or greenish from chlorophyll and other pigments inside the body. Red colors seem to occur only in pyrgines and pyrrhopygines. The ventral prothorax of some pyrgines such as Epargyreus clarus is often bright red or reddish-brown in color. Yellow or orange pigments occur in many pyrgines and some hesperiines. White or cream-colored cuticular pigments are more broadly distributed in hesperiines. The only skipper larva that I have seem with extensive black pigment is Codatr actus hyster. Colors on the body of hesperiid larvae are most commonly distributed as stripes (Figure 9). Usually the stripes occur longitudinally, but Pyrrhopygines and some pyrgines have transverse stripes of yellow or white. Transverse stripes seem to occur on unpalatable lepidopteran larvae such as the sphingid, Pseudosphinx tetrio. Longitudinal stripes typically occur only in certain locations such as a heart line, a subdorsal line, or a lateral line. The heart line is always present because the cuticle above the heart tends to be less pigmented than elsewhere on the body. The subdorsal line is usually a narrow stripe that may sometimes be interrupted to form a line of spots. The lateral stripe is less frequently present, but may form a broad band across the side of the

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60 body. In some species the body is covered with tiny white or yellowish spots and the stripes are formed by the absence of these colors. In other cases the stripes are outlined with white or yellow, forming double bands (Figure 9B). Some lines or spots on pyrgines such as Epargyreus and Urbanus may also be present due to patches of black microspines (Figure 9D) The texture of the cuticle of most hesperiids is rough, particular on pyrgines. The larval head consists of several sclerites. The head capsule is divided into two major sections (epicrania) by the frontal and adfrontal sclerites and the epicranial suture along the vertex. The front varies in shape from nearly an eguilateral to an isosceles triangle, usually the latter. Below the frontal sclerite lies the clypeus and next the labrum. The labrum of hesperiids is small with a shallow mesal notch. Along the posterior margin of the head is the postocciput. This sclerite is normally a very narrow band in most Lepidoptera, but is freguently very wide or moderately wide in the Hesperiidae. In pyrgines, the postocciput is usually very wide to moderately wide, lending in part to the "narrow neck" often referred to by authors (Figure 10). The postocciput is narrow to occasionally moderately wide in hesperiines (Orthos lycortas Paratrytone melane) as well. Megathymines have a narrow postocciput coupled with a large prothorax, creating a "wide neck"

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61 (Figure 10D) The transverse width of the head capsule of U. S. skippers varies from 1.5 to 7.2mm. The head capsule of hesperiids usually has a rough texture due to polygonal patterns of sculpturing (Figure 11). The sculpturing may form small pebbly knobs in pyrgines. These knobs are sometimes enlarged and modified into small spines at the apex of the epicranium (Figure 12). The ridges of the polygonal sculpturing may also be greatly widened, except for the center, resulting in a pattern of pits in hesperiines. Pyrrhopygines have a unique and characteristic series of low vertical ridges on the face (Figure 11D) The head capsule of Copaeodes aurantiacus a hesperiine, has two forward pointing processes, but its close relative, Copaeodes minimus, does not. Klots (1966) named and described paraclypeal hooks on the larva of Amblyscirtes samoset. The hooks are downward projecting spines, one on each side of the clypeus (Figure HE). Their function remains unknown. I have seen this structure only in Amblyscirtes species (Hesperiinae) and Piruna pirus (Heteropterinae) Hesperiids have six simple eyes or stemmata on the lower part of each epicranium (Figure 13). The relative size and spacing of the stemmata often varies between species. Stemma one through four lie in an arc and are usually egually spaced. Stemma five and six are usually farther apart.

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62 The labial-submental complex occupies a relatively large portion of the ventral head of most lepidopteran larvae. In hesperiids, however, the labial-submental complex is small relative to the size of the head, except for megathymines which have large labial-submental complexes. The mandibles of hesperiines and heteropterines are articulated somewhat differently from other skippers. Hesperiine mandibles are longer in the dimension between the labrum and labium than in the dimension between the antennae (Figure 14). The articulation supporting this longer mandible is also elongate. Pyrgine mandibles are subequal in these dimensions. Most hesperiids lack teeth or at most have shallow lobes on the mandibles. Well developed-teeth are present in some pyrgines and megathymines. Hesperiid larvae have a pair of short antennae that arise from a membranous area adjacent to the mandibles. The antenna consists of three segments, the distal segment being very small. A long sensory seta occurs on the tip of the second segment. Color patterns on the heads of skipper larvae are often striking. The pattern may be light spots or stripes on a dark ground, dark spots or stripes on a light ground, or variegated. Pyrgines frequently have spots on the face, and sometimes a lateral line. Hesperiines often have stripes on the face. The stripes usually occur in specific areas. The

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63 medial stripe proceeds from the vertex along the epicranial suture, then branches and proceeds partway along the adfrontal sclerites. The epicranial line proceeds from near the apex to the lower part of the face or may be broadly joined with the medial stripe along the adfrontals. The lateral stripe proceeds from the vertex, along the lateral margin of the head to the stemmatal area. Two skipper groups have uniformly colored heads, megathymines which are mostly concealed in stems, leaves, or roots, and pyrrhopygines which have dense long setae on the head. In addition, pyrgines sometimes have two brightly colored (yellow, orange, or red) false eye patches on the lower face between the stemmata and the mandibles. Eye patches also occur in hesperiines, but are smaller and less colorful. The dorsum of the prothorax is often sclerotized and darkened in larval hesperiids. The prothoracic shield may be relatively wide, covering much of the dorsum, reduced to a narrow band between the major annuli, or indistinct. The shield usually extends laterally onto a small hump. The hump is often darkened, but separated from the dorsal part of the shield in third and fourth instar larvae. The hump also freguently bears a lenticle and a pinaculum, the seta of which is longer and more slender than the surrounding setae. Another pinaculum is present in a more dorsal position on the mesothorax. A small cluster of lenticles usually accompanies the mesothoracic pinaculum as well. On

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64 the ventral side of the prothorax lies the opening to a specialized gland, thought to produce defensive chemicals (Eisner et al 1972). Most hesperiids have the ventral prothoracic gland, but I cannot find the opening in Megathyininae. The prothoracic spiracle often varies in shape between hesperiid species. Typically the spiracle is oval, but the anterior rim is freguently elongate in some species. The color of the thoracic legs is often useful in distinguishing species. In most skippers the legs are all pale or all dark. Others have only the first or second legs darkened. The thoracic leg length of hesperiines and pyrgines is usually moderate, but some species have relatively shorter or longer legs. Leg length seems to be partly correlated to larval ecology. Hesperiines that live near the ground, such as Hesperia species, often have long thoracic legs, whereas epiphytic species, such as Lerodea eufala, have very short legs. A few pyrgines have a row or small clusters of lenticles anterior to the legs. Abdominal segments one and two of the larvae lack legs and are transitional to the thoracic segments. Segments three through six are similar and bear a pair of prolegs. Segments seven and eight lack legs and are transitional to the terminalia. Segment nine is reduced to a small annulus between segment eight and ten. Segment ten bears the anal prolegs and is modified for disposal of the frass.

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65 The prolegs of segments three through six bear small hooks (crochets) in a circle, a near circle, a mesal penellipse, or transverse bands. The crochets are sometimes uniordinal or biordinal, but most often are in more or less three ranks (triordinal) The crochet band is always uniordinal near the lateral margin and quickly becomes biordinal and triordinal toward the meson. In some pyrgine genera such as Achalarus Thorybes and Cabares the posterior or mesal crochets are nearly twice as large as the anterior series of each proleg. Only one species that I have examined, Polygonus leo has ventral lenticles in small clusters anterior of the prolegs. The last stage larvae of some hesperiines and megathymines develop wax glands prior to pupation (Dethier 1942c) The wax produced by the glands is used to coat the inside of the cocoon. The gland pattern on the ventral surface of the larva varies considerably (Figure 15). The U. S. species with wax glands usually develop longitudinal or transverse patches on the ventral side of abdominal segments seven and eight (Figure 15A and B) The wax glands of Perichares philetes form a transverse patch on the ventral side of abdominal segment one. Synapte malitiosa has transverse patches on abdominal segments three through six. In Choranthus haitensis and Asbolis capucinus these patches are reduced to small spots posterior of the prolegs.

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66 Abdominal segment ten usually bears an anal comb. The anal comb is a fan-shaped, sclerotized structure located caudad and dorsal to the anus (Figure 16). During the act of defecation, the anal comb functions to propel the fecal pellet away from the caterpillar. This is accomplished with some force and the pellets may be launched several feet away. The base of the anal comb is wider in pyrgines than hesperiines. The anal comb is degenerate, but usually present, in megathymines consisting of up to about dozen spines. The suranal plate of most U. S. hesperiids is rounded, but the plate is pointed in a few hesperiines (Copaeodes spp.). Other hesperiines (Atrytone delaware, Polites spp.) have dark lines or markings on the suranal plate. The broad, bill-like plate of Asbolis capucinus is edged with black and the terminal spiracles are black, forming a false face. Pupae The pupae of U. S. hesperiids range in length from 10.5 to 52mm and in width from 2.1 to 10.3mm. The length of the abdomen varies between groups of hesperiids, being very short in species such as Epargyreus clarus and very long in Megathymus spp. The difference is due mostly to a change in the size of abdominal segments five through ten, as these lie beyond the wings. Most skipper pupae are brown or cream-colored, but some are green. Like their larvae,

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67 hesperiid pupae have a head, three thoracic segments, and ten abdominal segments. The antennae, legs, and wings are visible, but are glued to the body. The prothorax and abdominal segments two through eight bear spiracles. The thoracic spiracle is usually guarded by a cluster of microspines. In most pyrgines, the spiracle guard is on a small cuticular rise, giving the pupa a false face appearance when viewed from the front. The distal tip of the abdomen is modified into the cremaster, a structure used to anchor the pupa to the substrate. The cremaster shows considerable variation between species and higher groups. The setae on hesperiid pupae are usually short and simple. A few pyrgines, however, have some bifurcate setae. A neotropical species, Arteurotia tractipennis has long plumose setae on the head. Species with long larval setae generally have long setae on the pupae as well. Many hesperiines have very short or inconspicuous setae. Lenticles occur only on hesperiine pupae. The prothorax, dorsal abdomen, and ventral proleg scars of hesperiine pupae typically have lenticles (Figure 17). The head of most hesperiid pupae is rounded; however, some modifications do occur (Figure 18). A few pyrgines and many hesperiines have pointed processes on the cap. The point may be straight and relatively thick at the base (Lerema accius) slender and upturned (Calpodes ethlius) or T-shaped at the tip (Justinia phaetusa, a neotropical

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68 species). Instead of being pointed, the cap may be bulbous as in Atrytone delaware and Poanes viator. The head also bears a pair of sclerites (pilifers) which represent the mandible remnants. Pyrrhopygines, pyrgines, and megathymines always have the pilifers separated, whereas, most hesperiines have the pilifers touching. The position of the antennal tip relative to the tip of the middle leg on the pupae of hesperiids shows a good deal of variation. The antennae extend to the tip of the middle leg in Pyrrhopygines, and many pyrgines. In other pyrgines, most hesperiines, and megathymines the distal tip of the antenna lies far cephalad of the tip of the middle leg. Likewise, the proboscis usually reaches to the wing tips in pyrrhopygines and pyrgines, extends far beyond the wing tips in hesperiines, and lies far cephalad of the wing tips in megathymines. The distal part of the proboscis case is not glued to the body if this structure extends beyond the wings.

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69 Figure 3 SEM micrographs of the egg of Urbanus proteus (A), Telemiades epicalus (B), Thorybes pylades (C) Polites vibex (D) Hylephila phyleus (E), Atrytone delaware (F), Euphyes ruricola (G) Copaeodes minimus (H) and Megathymus cofaqui ( I )

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70

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71 Figure 4. SEM micrographs of the micropylar region of the egg of Urbanus proteus (A), Telemiades epicalus (B), Erynnis horatius (C), Hesperia attalus (D), Polites baracoa (E), Atrytone del aware (F), Problema byssus (G) Poanes yehl (H), and Euphyes ruricola (I).

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72

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' 73 Figure 5. SEM micrographs of the sculpturing on the sides of the egg of Atalopedes campestris (A), Polites vibex (B), Atrytone delaware (C), Problema byssus (D), Euphyes ruricola (E), and Megathymus cofaqui (F).

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74

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75 1 B I X I D E F \y H ci?uI?us"rA?Th ri rV f Set f e n the bod ^ of Pholisora I > catullus '(A %h 6a S 2 A ^ aJ ™ ^^a (B), the body of P. catuiius (C), the body of Urbanus proteus (D), the bodv of A^cyloxyp^a numitor (E), the body of Hesperia iubt (F) the head ofXe.opha.es trixus (G), the hea^of ?yrguf ( >' "fc^ja TutlTZ^ Th rybeS bathyUUS (I) Ld the

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76 Tl T2-3 Al-2 DORSAL SUBDORSAL SUPRASPIRACULAR SUBSPIRACULAR VENTRAL PROTHORACIC GLAND • 1 • '2 • i • I* PINACULUM I PROTHORACIC •' SHIELD SPIRACLE • 6 D. LEG A3-6 A7-8 DORSAL SUBDORSAL SUPRASPIRACULAR A9-10 SUBSPIRACULAR VENTRAL SPIRACLE Figure 7. Generalized map of the lenticles and other features of the last instar hesperiid larva.

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77 Figure 8. Lenticles on Pholisora catullus (A) yuccae (C) the fourth abdominal segment of Problema byssus (B), and Megathymus

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78 SUBDORSAL STRIPE LATERAL STRI PE MICROSPINES somTu 9 S SinL? a H tern 3 n the fourth abdominal segment of some U S. hespernds: dorsal view of transverse strioe^ on T a t r J^ e "/ Ul9e T r J A) lateral View of sub5orsafand Co™oL StripeS bOUnded by white (shaded portion) on Copaeodes aurantiacus (B), lateral view of the interrupted viewof a pa?cher o ? f b^cf raCtUS *"— W .nllTttral view or patches of black microspmes on Epargyreus clarus

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79 postocciput Figure 10. Relative larvae of Pyrrhopyge Problema byssus (C), size of the "neck" of last instar araxes (A), Urbanus teleus (B), and Megathymus yuccae (D)

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80 jirv^: %$& RIDGES EPICRANIAL SUTURE CLYPEUS LABRUM EPICRANIUM STEMMATA MANDIBLE" PARACLYPEAL HOOK ANTENNA Figure 11. Features of the larval head of hesperiids: pebbly sculpturing of Polythrix procera (A) rough sculpturing of Staphylus mazans (B), pitted sculpturing of Hesperia pahaska (C), ridges on the head of Pyrrhopyge chalybea (D) and general structures on Amblyscirtes vialis (E).

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81 Figure 12. Processes on the head of Urbanus procne (A), Achalarus lyciades (B), and Copaeodes aurantiacus (C).

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STEMMA 1 STEMMA 2 o o STEMMA 3 *-\ STEMMA 4 o o o 82 STEMMA 6 o o o o STEMMA 5 B Figure 13. Comparison of the stemmata of Pyrrhopyge araxes (A), Achlyodes thraso (B), and Euphyes pilatka (C).

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83 Figure 14. Ventral view of the hesperiine-type mandibles of Atalopedes campestris (A) and the pyrgine-type mandibles of Erynnis horatius (B).

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84 l<£3^> .@_J5t .dQ ^o-' ^ D Figure Lerema 15,

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85 A B D E a? Ur ?^^S OInPariSOn f the anal c ™ h of Urbanus (A) Thorybes mexicanus (B), Pyrqus scri ntnr* themzstocles (D) Pseudocopaeodes yuccae (F). eunus (E) proteus (C), Polites and Megathymus

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86 B ^a of pie^co^^ S*f ?? <£ Pth f* the

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87 B Figure 18. Lateral view of the pupal cap of Perichares philetes (A), Atrytone delaware (B), Panoquina panoquin (C) and Calpodes ethlius (D).

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CHAPTER 5 DIAGNOSES AND HOST PLANTS FOR THE UNITED STATES SKIPPERS In the following chapter, I present a diagnosis of the immature stages for each subfamily and for each species of Hesperiidae recorded for the United States. If personal observations were lacking and a thorough search of the literature turned up no reference to previous description, that negative information is listed under the species name so that future workers can concentrate their efforts on those taxa to complete our picture of the North American skipper fauna. Subfamily Pvrrhopvainae Diagnosis EGG: no specimens available, undescribed in literature. LAST INSTAR LARVA: BODY: length to ca. 50mm, A4 transverse width to ca. lOmm; brightly colored with red, yellow, or white, freguently with transverse stripes; setae simple, long; spiracles tan, Tl and A8 spiracles largest. HEAD: uniformly dark; transverse width ca. 7mm; sculpturing rough, with several low ridges extending from the upper to lower face; mandibles without teeth, pyrgine type articulation; labial-submental complex relatively small; setae simple, long; stemmata subequal; postocciput narrow. THORAX: prothorax slightly larger than head; legs tan; 88

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89 shield narrow; ventral prothoracic gland present. ABDOMEN: A4 proleg with ca. 140 crochets, irregularly triordinal, arranged in a circle, posterior prolegs always with a mesal penellipse; suranal plate rounded, unmarked; anal comb well developed; wax glands absent. PUPA: robust; thoracic spiracle guard distinct; setae relatively long; pilifers separated; head rounded; antennal tip extending to slightly beyond tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster blunt with numerous hooked setae in a cluster at the tip. 1. Pyrrhopyge araxes Diagnosis LAST INSTAR LARVA: BODY: length 47mm, A4 transverse width 9.7mm; preserved specimen brownish red with narrow transverse yellow stripes on T1-A9; setae simple, to 6.4mm long on A4 dorsum; spiracles tan. HEAD: dark brown; transverse width 7.2mm; sculpturing rough, with ridges on face; mandibles without teeth; setae simple, to 5mm long at apex, a few ventral setae to 5.7mm; stemmata subegual (Figure 37A) ; postocciput narrow. THORAX: legs tan; shield narrow, a dark brown band between the annul i. ABDOMEN: prolegs each with about 140 crochets, irregularly triordinal, arranged in nearly a circle; suranal plate rounded, unmarked (Figure 45A). Other Descrip tions. Dyar 1906 (larva), Comstock 1956b (larva, pupa).

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90 Host Plants FAGACEAE: Quercus arizonica (Burns 1964a), Quercus emoryi (Bailowitz and Brock 1991, in lab), Quercus oblongifolia (Bailowitz and Brock 1991, in lab). MYRTACEAE Psidium guajava (Beutelspacher 1980) [questionable]. Specimens Examined LARVAE: ARIZONA: Lot 002, 1 larva (instar 5) (NM); Lot 552, 1 larva ( instar 4) (NM) .subfamily Pyrainae Diagnosis EGG: usually with 10 to 30 vertical ribs, occasionally with polygonal sculpturing or with 30 to 50 vertical rows of short spines, height 0.5-1. 3mm, diameter 0.6-1. 8mm; white, yellow, orange, or green, occasionally red, gray, or brown. LAST INSTAR LARVA: BODY: length 849mm, A4 transverse width 1.8-9.8ram; often with red, yellow, or orange markings, rarely with transverse stripes; some setae often modified with blunt or expanded tips, < 0.11.6mm long; spiracles pale to dark, Tl and A8 spiracles largest. HEAD: pale, dark, or patterned, sometimes with colorful eye patches; transverse width 2-6. 4mm; sculpturing rough to pebbly, occasionally with small spines or knobs at apex; mandibles with or without teeth, pyrgine type articulation; labial-submental complex relatively small; setae simple, branching, or feathery, < 0.1-2mm long; stemmata usually subequal; postocciput wide or moderately wide. THORAX: prothorax about the same size as head or smaller; legs pale to black; shield narrow to broad; ventral

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91 prothoracic gland present. ABDOMEN: A4 proleg with 30 to 230 crochets, uniordinal biordinal, or irregularly triordinal, arranged in a circle, a near circle, or a mesal penellipse, posterior prolegs always with a mesal penellipse; suranal plate rounded, unmarked; anal comb well developed; wax glands absent. PUPA: length 10.5-32mm, A3 transverse width 3. 2-10. 3mm; green or brown; thoracic spiracle guard distinct; setae < 0.1-lmm; pilifers separated; head usually rounded, occasionally with a short pointed process on cap; antennal tip may lie slightly caudad of tip of middle leg or far cephalad of middle leg tip; proboscis extending to the wing tips, occasionally slightly beyond; abdomen short or moderately long; cremaster blunt with numerous hooked setae in a cluster at the tip. 2. Phocides pigmalion (Cramer) Diagnosis LAST INSTAR LARVA: BODY: length 2 3-3 2mm, A4 transverse width 5. 3-7. 7mm; white; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: reddish brown, darker around the mouth, large yellow eye patches present; transverse width 5-5. 3mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.7mm; stemmata subegual (Figure 37B) ; postocciput wide. THORAX: legs tan; shield broad, tan. ABDOMEN: prolegs each with about 230 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 45B) PUPA: Figures 58D, 61A and 78A,

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92 length 25-32mm, A3 transverse width 7. 5-9. 8mm; greenish white; thoracic spiracle guard absent; setae simple, to 0.5mm long on head; head rounded, cap bulbous with a slight mesal indentation; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 2.5mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101A and 112A) Other Descriptions. Dyar 1890 (egg, larva, pupa), Strohecker 1938 (larva, pupa), Minno and Emmel 1993 (larva, pupa ) Host Plants RHIZOPHORACEAE: Rhizophora mangle (Dyar 1890) Specimens Examined LARVAE: FLORIDA: BROWARD CO.: Lot 364, 1 larva (instar 5) (MCM) DADE CO.: Lot 1272, 1 larva (instar 2) (DHH). MONROE CO.: Lot 365, 1 larva (instar 5) (MCM); Lot 728, 1 larva (instar 4) (MCM); Lot 729, 1 larva (instar 4) (MCM). PUPAE COLLIER CO.: Lot 363, 1 pupa (MCM); Lot 365, 2 pupae (MCM). 3. Phocides palemon (Cramer) Diagnosis EGG: height 1.6-1. 8mm, width 1-1. 3mm, 17-19 ribs, red. LAST INSTAR LARVA: BODY: length 21.5-33mm, A4 transverse width 5. 8-9. 1mm; preserved specimens pale; setae simple, to 0.3mm long on A4 dorsum; spiracles tan. HEAD: Figure 22A, pale reddish brown, darker around the mouth, large yellow eye patches present; transverse width 4.7-

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93 5.5mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.7mm; stemmata subequal (Figure 37C); postocciput wide. THORAX: legs tan; shield broad, tan. ABDOMEN: prolegs each with about 165 crochets, irregularly triordinal, arranged in a circle. PUPA: Figures 61B and 78B, length 24-26mm, A3 transverse width 8.3-10mm; preserved specimens pale; thoracic spiracle guard absent; setae simple, to 0.5mm long on head; head rounded, cap bulbous with a slight mesal indentation; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 2.2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101B and 112B). Other Descriptions Comstock and Vazquez 1960 (larva, pupa ) Host Plants MYRTACEAE: Eugenia unifora (Neck 1983), Psidium cattleianum (Neck 1978), Psidium guajava (Comstock and Vazquez 1960). Specimens Examined EGGS: TEXAS: CAMERON CO.: Lot 962, 12 eggs (ROK) LARVAE: MEXICO: GUERRERO: Lot 866, 7 larvae (instars 4, 5) (USNM) TEXAS: CAMERON CO.: Lot 777, 1 larva (instar 5) (USNM); Lot 883, 1 larva (instar 5) (USNM); Lot 962, 3 larvae (instar 5) (ROK). PUPAE: TEXAS: CAMERON CO.: Lot 962, 4 pupae (ROK).

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94 4. Phocides urania (Westwood) Nothing is known of the biology of this neotropical species. 5. Proteides mercurius (Fabricius) This species rarely enters the U. S. from tropical America. Its biology is poorly known. No immature specimens were located for study. Descriptions Draudt 1924 (larva, pupa), Comstock and Vazguez 1960 (larva, pupa). Host Plants. FABACEAE: Cassia species (Draudt 1924), Derris eliptica (Wolcott 1941), Ecastophyllum species (MacNeill 1975), Muellera moniliformis (Moss 1949), Rhynchosia species (MacNeill 1975), Vigna species (MacNeill 1975). 6. Epargyreus zestos (Geyer) Diagnosis LAST INSTAR LARVA: BODY: length 31-48mm, A4 transverse width 6. 2-8. 2mm; green with numerous narrow transverse yellow stripes, prothorax reddish ventrally, prolegs orange; some setae with expanded tips, to 0.1 mm long on A4 dorsum; spiracles dark brown. HEAD: black, large yellow eye patches present; transverse width 5.25.9mm; sculpturing pebbly; mandibles without teeth; setae simple, to O.lmm long at apex, a few ventral setae to 0.7mm; stemmata subegual (Figure 37D); postocciput wide. THORAX: legs tan; shield broad, brown. ABDOMEN: prolegs each with about 180 crochets, irregularly triordinal, arranged in a

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95 circle; suranal plate rounded, unmarked (Figure 45C) PUPA: Figures 61C and 79A, length 24.5-28mm, A3 transverse width 8.8mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; some setae with slightly expanded tips, to 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101C and 112C) Other Descriptions Minno and Emmel 1993 (larva, pupa). Host Plants FABACEAE: Galactia striata (Scott 1986). Specimens Examined LARVAE: FLORIDA: MONROE CO.: Lot 137, 1 larva ( instar 5) (MCM) ; Lot 138, 1 larva (instar 5) (MCM); Lot 139, 3 larvae ( instars 3, 5) (MCM); Lot 141, 2 larvae (instar 5) (MCM); Lot 142, 1 larva (instar 2) (MCM); Lot 1085, 3 larvae (instars 2, 5) (DHH). PUPAE: FLORIDA: MONROE CO.: Lot 137, 1 pupa (MCM); Lot 140, 1 pupa (MCM). 7. Epargyreus clarus (Cramer) Diagnosis LAST INSTAR LARVA: BODY: Figure 19B, length 19-47iwn, A4 transverse width 3. 5-9. 8mm; green with numerous narrow transverse yellow stripes, a few lines and patches of black microspines on sides, ventral prothorax red, prolegs orange; setae simple, to 0.1mm long on A4 dorsum; spiracles black. HEAD: Figure 22B, dark reddish brown, large yellow eye patches present; transverse width 4. 4-5. 8mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.1mm long

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96 at apex, a few ventral setae to 0.7mm; stemmata subequal (Figure 37E); postocciput wide. THORAX: Tl legs red, T2-3 legs tan; shield broad, tan. ABDOMEN: prolegs each with about 145 crochets, irregularly triordinal, arranged in a circle. PUPA: Figures 58A, 61D and 79B, length 18-25. 5mm, A3 transverse width 6. 3-9. 2mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.3mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101D and 112D) Other Descriptions Harris 1862 (larva, pupa), Scudder 1881 (larva, pupa), French 1886 (egg, larva, pupa), Scudder 1889a (egg, larva, pupa), Comstock and Comstock 1895 (larva), Mosher 1916 (pupa), Comstock 1927b (egg, larva, pupa), Klots 1951 (larva, pupa), Peterson 1962 (larva), Emmel and Emmel 1973 (egg, larva, pupa), Opler and Krizek 1984 (egg, larva, pupa), Scott 1986 (egg, larva, pupa). Host Plants FABACEAE: Acacia species (Edwards 1889) [erroneous], Amorpha californica (Emmel and Emmel 1973), Amorpha fruticosa (Walsh and Riley 1869), Amphicarpa bracteata (Scudder 1889a, b), Apios americana (Scudder 1889a, b), Astragalus species (Tietz 1952), Cassia marilandica (Heitzman and Heitzman 1987), Clitoria mariana (Hayward 1947), Desmodium canadense (Scudder 1889a, b),

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97 Desmodium "dillenii" (Clark 1936) [this name is currently placed under both D. glabellum and D. perplexum) Desmodium marilandicum (Scudder 1889a, b), Desmodium "michauxii" (Clark 1936) [I cannot find this name listed in floras of the region], Desmodium nudiflorum (Scudder 1889a, b), Desmodium paniculatum (Clark 1936), Desmodium rotundifolium (Scott 1986), Erythrina herbacea (Kendall 1965, oviposition but no larval feeding), Galactia regularis (Minno 1992), Galactia volubilis (MCM collection), Gleditsia triacanthos (Walsh and Riley 1869), Glycine max (DHH collection), Glycyrrhiza lepidota (McCabe and Post 1977), Lathyrus palustris (Moffat 1882), Lespedeza capitata (Scudder 1889a, b), Lotus crassifolius (Emmel and Emmel 1973), Lotus grandiflorus (Lembert 1894), Lupinus species (DHH collection), Phaseolus polystachios (Fiske 1896), Phaseolus vulgaris (Fiske 1896, probably; Lenczewski 1980), Pueraria lobata (Sim 1936), Rhynchosia minima (Kendall 1965, oviposition but no larval feeding), Robinia hispida (French 1878), Robinia neomexicana (Scudder 1889a, b), Robinia pseudoacacia (Harris 1862), Robinia viscosa (Harris 1862), Wisteria frutescens (Scudder 1869), Wisteria sinensis (Kendall 1965). Specimens Examined LARVAE: NO DATA: Lot 1193, 3 larvae (instar 5) (DHH). ARIZONA: COCHISE CO.: Lot 536, 2 larvae (instars 4, 5) (TLM) CALIFORNIA: RIVERSIDE CO.: Lot 583, 1 larva (instar 5) (GRB) FLORIDA: Lot 1172, 1 larva (instar 5) (DHH). ALACHUA CO.: Lot 147, 2 larvae (instar

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98 5) (MCM) ; Lot 722, 2 larvae ( instars 3, 5) (MCM) ; Lot 723, 1 larva ( instar 5) (MCM); Lot 725, 1 larva ( instar 5) (MCM); Lot 1173, 1 larva (instar 5) (DHH); Lot 1174, 1 larva (instar 5) (DHH); Lot 1175, 1 larva (instar 4) (DHH); Lot 1176, 1 larva (instar 4) (DHH); Lot 1177, 1 larva (instar 2) (DHH); Lot 1179, 2 larvae (instars 2, 4) (DHH); Lot 1180, 2 larvae (instar 5) (DHH); Lot 1181, 1 larva (instar 5) (DHH); Lot 1185, 1 larva (instar 5) (DHH); Lot 1186, 9 larvae (instars 3, 4, 5) (DHH); Lot 1187, 1 larva (instar 5) (DHH); Lot 1188, 3 larvae (instars 2, 3, 4) (DHH); Lot 1190, 1 larva (instar 2) (DHH); Lot 1192, 1 larva (instar 5) (DHH); Lot 1194, 1 larva (instar 5) (DHH); Lot 1195, 1 larva (instar 5) (DHH); Lot 1197, 1 larva (instar 5) (DHH); Lot 1198, 1 larva (instar 5) (DHH); Lot 1200, 1 larva (instar 5) (DHH); Lot 1201, 1 larva (instar 5) (DHH); Lot 1203, 1 larva (instar 5) (DHH); Lot 1204, 1 larva (instar 2) (DHH); Lot 1205, 1 larva (instar 5) (DHH); Lot 1207, 1 larva (instar 4) (DHH). DUVAL CO.: Lot 721, 1 larva (instar 3) (MCM). GADSDEN CO.: Lot 1170, 2 larvae (instar 5) (DHH). HIGHLANDS CO.: Lot 143, 1 larva (instar 5) (MCM). PUTNAM CO.: Lot 724, 1 larva (instar 5) (MCM); Lot 726, 1 larva (instar 2) (MCM); Lot 1184, 2 larvae (instar 5) (DHH). VOLUSIA CO.: Lot 144, 1 larva (instar 5) (MCM). GEORGIA: TIFT CO.: Lot 1169, 1 larva (instar 5) (DHH). ILLINOIS: Lot 617, 1 larva (instar 5) (SP) INDIANA: KOSCIUSKO CO.: Lot 3, 1 larva (instar 5) (MCM). KANSAS: DOUGLAS CO.: Lot

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99 633, 4 larvae ( instars 3, 5) (TCE) MINNESOTA: BECKER CO.: Lot 534, 1 larva ( instar 3) (TLM); Lot 537, 1 larva ( instar 5) (TLM). MISSOURI: BOONE CO.: Lot 1178, 1 larva (instar 5) (DHH); Lot 1183, 2 larvae (instar 5) (DHH); Lot 1191, 1 larva (instar 5) (DHH). COOPER CO.: Lot 1189, 5 larvae (instars 4, 5) (DHH); Lot 1202, 1 larva (instar 5) (DHH). JACKSON CO.: Lot 556, 3 larvae (instar 5) ( JRH) ; Lot 1196, 3 larvae (instar 5) (DHH); Lot 1199, 4 larvae (instar 5) (DHH). NEW YORK: TOMKINS CO.: Lot 535, 1 larva (instar 5) (TLM). NORTH DAKOTA: SLOPE CO.: Lot 533, 1 larva (instar 5) (TLM). SOUTH CAROLINA: PICKENS CO.: Lot 1206, 1 larva (instar 3) (DHH). VIRGINIA: GILES CO.: Lot 1182, 1 larva (instar 3) (DHH). PUPAE: FLORIDA: ALACHUA CO.: Lot 145, 1 pupa (MCM). KANSAS: DOUGLAS CO.: Lot 63 3, 1 pupa (TCE). MARYLAND: PRINCE GEORGE'S CO.: Lot 146, 2 pupae (MCM). MISSOURI: JACKSON CO.: Lot 564, 2 pupae (JRH). NEW MEXICO: SANDINAL CO.: Lot 727, 1 pupa (MCM). 8. Epargyreus exadeus (Cramer) Diagnosis LAST INSTAR LARVA: BODY: length 47mm, A4 transverse width 6.3mm; green with numerous narrow transverse yellow stripes, a few lines and patches of black microspines on sides, ventral prothorax red, prolegs orange; setae simple, to 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: dark reddish brown, large yellow eye patches present; transverse width 5.2mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.1mm long at

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100 apex, a few ventral setae to 0.7mm; stemmata subequal (Figure 37F); postocciput wide. THORAX: legs tan; shield broad, tan. ABDOMEN: prolegs each with about 175 crochets, irregularly triordinal arranged in a circle. Other Descriptions Draudt 1924 (larva), Moss 1949 (larva), Comstock and Vazquez 1960 (larva, pupa). Host Plants FABACEAE: Cassia species (Comstock and Vazquez 1960), Lecythis paraensis (Moss 1949, probably), Phaseolus species (Moss 1949), Rhynchosia minima (MCM collection) Specimens Examined LARVAE: COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 148, 2 larvae ( instars 3, 5) (MCM). 9. Polygonus leo (Gmelen) Diagnosis LAST INSTAR LARVA: BODY: Figure 19F, length 23-42mm, A4 transverse width 4-7. 5mm; green with numerous tiny yellow spots; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 22C, pale with two black eye spots on upper face, lateral line black; transverse width 4. 3-5. 3mm; sculpturing pebbly; mandibles without teeth; setae simple, < 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subequal (Figure 37G); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 180 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 45D) PUPA: Figures 58E, 62A, and 80A, length 22-24. 5mm, A3 transverse width 7.2-

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101 8.7mm; pale green; thoracic spiracle guard distinct; setae simple, to 0.1mm long on head; head with a short pointed process on cap, anterior margin of eyes with a short point; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 2.2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101E and 112E) Other Descriptions Dyar 1897 (larva), Wolcott 1923 (larva), Dethier 1942b (larva), MacNeill 1975 (egg, larva), Minno and Emmel 1993 (larva, pupa). Host Plants FABACEAE: Derris eliptica (Wolcott 1941), Lonchocarpus sericeus (Bates 1935), Piscidia piscipula (Dyar 1897), Pongamia pinnata (Kimball 1965). Specimens Examined LARVAE: FLORIDA: BROWARD CO.: Lot 426, 4 larvae (instars 2, 4, 5) (MCM) ; Lot 1274, 2 larvae (instars 3, 5) (DHH) MONROE CO.: Lot 423, 1 larva ( instar 5) (MCM); Lot 425, 3 larvae (instar 5) (MCM); Lot 427, 1 larva (instar 5) (MCM); Lot 428, 2 larvae (instar 5) (MCM); Lot 730, 1 larvae (instar 5) (MCM); Lot 731, 1 larva (instar 5) (MCM); Lot 732, 1 larva (instar 5) (MCM); Lot 733, 1 larva (instar 5) (MCM); Lot 734, 2 larvae (instar 5) (MCM); Lot 735, 1 larva (instar 5) (MCM), Lot 1273, 1 larva (instar 3) (DHH); Lot 1275, 1 larva (instar 5) (DHH); Lot 1276, 1 larva (instar 5) (DHH). PUPAE: FLORIDA: BROWARD CO.: Lot 426, 1 pupa (MCM). MONROE CO.: Lot 424, 1 pupa (MCM); Lot 425, 1 pupa (MCM)

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102 10. Polygonus manueli Bell and W. P. Comstock No specimens of the immature stages of this neotropical species were located for study. This skipper has rarely been recorded from the U. S. Other Descriptions Moss 1949 (larva, pupa). Host Plants FABACEAE: Muellera moniliformis (Moss 1949). 11. Chioides catillus (Cramer) Diagnosis LAST INSTAR LARVA: BODY: length 18-3 3. 5mm, A4 transverse width 4. 7-7. 2mm; preserved specimens pale with some small clusters of black microspines, ventral prothorax red, prolegs orange; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 2 2D, tan, front black, a black W-shaped mark on upper face, stemmatal area black, two black eye spots on lower face, large orange eye patches present; transverse width 4. 7-5. 6mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.8mm; stemmata subegual (Figure 37H); postocciput wide. THORAX: legs reddish; shield broad, dark brown. ABDOMEN: prolegs each with about 140 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 46A) PUPA: Figures 62B and 80B, length 18. 5-23. 5mm, A3 transverse width 5. 8-7. 7mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.3mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing

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103 tips; abdomen short; cremaster to 2.2mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 101F and 112F). Other Descriptions Hayward 1927a (egg, larva), Comstock and Vazquez 1960 (larva, pupa). Host Plants CHENOPODIACEAE: Chenopodium album (Hayward 1947) [erroneous]. FABACEAE: Galactia canescens (ROK collection), Glycine max (Hayward 1941), Mimosa species (MacNeill 1975), Phaseolus atropurpureus (Kendall 1965), Rhynchosia minima (Kendall 1965), Rhynchosia senna (Hayward 1927a), Tephrosia lindheimeri (Kendall 1965). LAMIACEAE: Monarda punctata (Hayward 1947) [erroneous], Origanum vulgare (Hayward 1947) [erroneous]. Specimens Examined LARVAE: TEXAS: CAMERON CO.: Lot 921, 69 larvae (instar 5) (ROK). SAN PATRICIO CO.: Lot 922, 13 larvae ( instars 2, 3, 4, 5) (ROK). PUPAE: TEXAS: CAMERON CO.: Lot 921, 6 pupae (ROK). SAN PATRICIO CO.: Lot 922, 1 pupa (ROK) 12. Chioides zilpa (Butler) The immature stages of this species are undescribed. No specimens were located for study. Host Plants FABACEAE: Nissolia schottii (Bailowitz and Brock 1991, possibly), Phaseolus species (Beutelspacher 1980), Rhynchosia species (Beutelspacher 1980), Tephrosia species (Beutelspacher 1980).

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104 13. Aguna asander (Hewitson) Little is known of the biology of this neotropical species, except that the larvae probably feed on leguminous plants (da Costa Lima 1936). 14. Aguna claxon Evans Nothing is known of the biology of this neotropical species. 15. Typhedanus undulatus (Hewitson) Diagnosis LAST INSTAR LARVA: BODY: length 18-2 4mm, A4 transverse width 3. 8-6. 2mm; preserved specimens brownish; some setae with expanded tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 22E, black, large yellow eye patches present; transverse width 4. 7-5. 6mm; sculpturing pebbly; mandibles without teeth; setae branching, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 371); postocciput wide. THORAX: legs tan; shield narrow, a brown line between the annuli. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 46B) Other Descriptions Hayward 1931 (egg, larva, pupa), Hoffmann 1932 (larva, pupa), Moss 1949 (larva, pupa). Host Plants FABACEAE: Cassia alata (Moss 1949), Cassia bicapsularis (FSCA collection), Cassia corymbosa (Hayward 1931), Cassia occidentalis (Moss 1949), Cassia reticulata (Moss 1949), Cassia tora (Moss 1949).

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105 Specimens Examined LARVAE: TEXAS: HIDALGO CO.: Lot 1003, 2 larvae ( instar 5) (FSCA) 16. Polythrix mexicana H. A. Freeman Diagnosis LAST INSTAR LARVA: BODY: length 19-2 5. 5mm, A4 transverse width 4.6-5mm; preserved specimens pale, reticulated with yellow, subdorsal line a chain of orange spots connected by a narrow yellow line on T3-A9; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 22F, pale, lateral stripe yellow anteriorly, black posteriorly, two black spots on upper face, posterior black; transverse width 3.7-4. 3mm; sculpturing pebbly; mandibles without teeth; setae simple, < 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 37J) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 175 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 46C) PUPA: Figures 62C and 81A, length 19mm, A3 transverse width 4. 8-5. 7mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.1mm long on head; head with a short pointed process on cap; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 2.1mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101G and 113A)

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106 Host Plants FABACEAE: Amerimnon granadillo (Kendall and McGuire 1975), Ichtyomethia communis (Kendall and McGuire 1975) Specimens Examined LARVAE: MEXICO: TAMAULIPAS: Lot 967, 4 larvae (instar 5) (ROK) ; Lot 968, 2 larvae ( instar 5) (ROK). PUPAE: MEXICO: TAMAULIPAS: Lot 967, 3 pupae (ROK) 17. Polythrix octomaculata (Sepp) No immature specimens of this species were located for study. The larva and pupa stages were briefly described by Draudt (1924) and Moss (1949). Host Plants FABACEAE: Muellera moniliformis (Moss 1949), Pterocarpus draco (Riley 1975), Pterocarpus indicus (Draudt 1924), Toluifera perreirae (Scott 1986). 18. Polythrix procera (Plotz) Diagnosis EGG: height 1.7mm, width 0.8mm, 14 ribs. LAST INSTAR LARVA: BODY: length 24.5mm, A4 transverse width 6.5mm; preserved specimens pale, subdorsal line broken into a row of red spots on Al-7, large red patches around the spiracles, Al-8 red ventral ly; some setae with expanded tips, to 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 23A, reddish brown, paler toward the apex, large yellowish eye patches present, stemmatal area black, vertex deeply cleft; transverse width 4.5mra; sculpturing pebbly; mandibles without teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual

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107 (Figure 37K) ; postocciput wide. THORAX: legs brown; shield broad, tan. ABDOMEN: crochets irregularly triordinal, arranged in a circle. Host Plants COMBRETACEAE : Laguncularia racemosa (Kendall and McGuire 1975), Terminalia catappa (Kendall and McGuire 1975) Specimens Examined EGGS: MEXICO: TAMAULIPAS: Lot 969, 1 egg (ROK) LARVAE: MEXICO: TAMAULIPAS: Lot 969, 2 larvae (instars 4, 5) (ROK). 19. Zestusa dorus (Edwards) No immature specimens of this southwestern species were located for study. Klots (1971) briefly described the egg, larva, and pupa. Host Plants FAGACEAE: Quercus alba (Klots 1971, in lab), Quercus arizonica (Klots 1971, associated with; MacNeill 1975), Quercus emoryi (Klots 1971), Quercus gambelii (Klots 1971, in lab) 20. Codatractus alcaeus (Hewitson) Diagnosis LAST INSTAR LARVA: BODY: length 27-38. 5mm, A4 transverse width 5. 5-8. 5mm; preserved specimens pale with numerous tiny yellow spots, subdorsal line broken into a row of round yellow spots on Al-8; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 23B, reddish brown, lower half of face black, large yellow eye patches present; transverse width 5. 7-6. 4mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at

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108 apex, a few ventral setae to 0.7mm; stemmata subequal (Figure 37L) ; postocciput moderately wide. THORAX: legs tan; shield broad, tan. ABDOMEN: prolegs each with about 140 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked (Figure 46D). PUPA: Figures 62D and 81B, length 22mm, A3 transverse width 7.7mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.9mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 101H and 113B) Host Plants FABACEAE: Amerimnon granadillo (Kendall and McGuire 1975), Ichtyomethia communis (Kendall and McGuire 1975) Specimens Examined LARVAE: MEXICO: TAMAULIPAS: Lot 924, 2 larvae (instars 3, 5) (ROK) SAN LUIS POTOSI : Lot 925, 2 larvae (instar 5) (ROK). PUPAE: MEXICO: TAMAULIPAS: Lot 924, 1 pupa (ROK). SAN LUIS POTOSI: Lot 925, 1 pupa (ROK). 21. Codatractus melon (Godman and Salvin) Nothing is known of the biology of this neotropical species 22. Codatractus arizonensis (Skinner) Diagnosis LAST INSTAR LARVA: BODY: length 2 72 7. 5mm, A4 transverse width 6.4mm; preserved specimens pale with

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109 numerous tiny yellow spots, subdorsal line broken into a row of rectangular yellow spots on T2-A8; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 23C, pale brown, front black, large yellow eye patches present; transverse width 4.8-5mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 38A) ; postocciput moderately wide. THORAX: legs tan; shield broad, tan. ABDOMEN: prolegs each with about 80 crochets, irregularly triordinal, arranged in a near circle. PUPA: Figures 62E and 81C, length 18mm, A3 transverse width 5.8mm; dark reddish brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.4mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 1011 and 113C). Host Plants FABACEAE: Eysenhardtia orthocarpa (Bailowitz and Brock 1991), Eysenhardtia texana (ROK collection). Specimens Examined LARVAE: TEXAS: BREWSTER CO.: Lot 926, 11 larvae (instars 3, 5) (ROK). PUPAE: TEXAS: BREWSTER CO.: Lot 926, 2 pupae (ROK). 23. Urbanus proteus (Linnaeus) Diagnosis EGG: Figures 3A and 4A, height 0.9-lmm, width 0.8-0. 9mm, 11-12 ribs, yellow. LAST INSTAR LARVA: BODY:

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110 Figure 19D, length 17-41mm, A4 transverse width 3. 5-7. 8mm; green, heart line darkened with black microspines, subdorsal line narrow, yellow, widening and becoming orange on A8-10, small scattered patches of black microspines present, ventral prothorax red, prolegs orange; some setae with blunt tips, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 23D, light brown, lower face blackish, large orange or reddish eye patches present, posterior black; transverse width 4-5. 3mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 38B); postocciput wide. THORAX: Tl legs dark brown, T2-3 legs tan; shield broad, dark brown. ABDOMEN: prolegs each with about 115 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 46E). PUPA: Figures 58C, 62F and 82A, length 18. 5-26. 5mm, A3 transverse width 5.4-8mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.3mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.5mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102A and 113D) Other Descriptions Edwards and Chapman 1879 (larva, pupa), Comstock 1881 (larva, pupa), Scudder 1889a (egg, larva, pupa), Quaintance 1898 (egg, larva, pupa), Chittenden 1902 (egg, larva, pupa), Watson 1919 (larva), Comstock 193 2b

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Ill (egg), Moss 1949 (larva, pupa), Emmel and Emmel 1973 (larva, pupa), Heppner 1975 (larva), Pyle 1981 (egg, larva, pupa), Young 1985 (egg, larva, pupa), Toliver 1987 (larva), Minno and Emmel 1993 (egg, larva, pupa). Host Plants ASTERACEAE: Camara amarella (Moss 1949) [guestionable] Clibadium erosum (USNM collection) [guestionable] BRASSICACEAE: Brassica oleracea (Comstock 1881) [erroneous], Brassica rapa (Comstock 1881) [erroneous], Brassica species (Scudder 1889b) [erroneous], Raphanus sativus (MCM observation, oviposition and larvae cut shelters, but refused to eat this plant in the lab). CANNACEAE: Canna species (Hayward 1941) [erroneous]. FABACEAE: Alysicarpus vaginalis (MCM collection), Amphicarpa bracteata (Opler and Krizek 1984), Bauhinia species (Hayward 1941), Centrosema floridanum (Minno 1992), Centrosema pubescens (USNM collection), Centrosema virginianum (MCM collection), Clitoria mariana (Scudder 1872), Clitoria ternatea (Scudder 1889a, b), Crotalaria spectabilis (MCM observation, oviposition and larvae cut shelters, but refused to eat this plant), Crotalaria species (Fennah 1947) [questionable], Desmodium batocaulon (Bailowitz and Brock 1991), Desmodium canescens (Chittenden 1902), Desmodium floridanum (MCM observation), Desmodium incanum (Minno 1992), Desmodium neomexicanum (Kendall 1976), Desmodium paniculatum (MCM collection) Desmodium tortuosum (Quaintance 1898), Desmodium viridiflorum (Scudder 1889a, b),

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112 Galactia regularis (MCM collection), Galactia striata (Minno and Emmel 1993), Glycine max (Tietz 1952), Macroptilium lathyroides (Wolcott 1923), Mucuna species (Young 1985), Phaseolus aureus (Bruner et al. 1945), Phaseolus limensis (Wolcott 1936), Phaseolus polystachios (Edwards and Chapman 1879), Phaseolus semierectus (Hayward 1947), Phaseolus vulgaris (Comstock 1881), Pisum sativum (Comstock 1944), Prosopis glandulosa var. torreyana (Tietz 1952), Prosopis species (Comstock 1927b), Pueraria lobata (MCM collection), Rhynchosia minima (MCM collection), Vigna luteola (Minno 1992), Vigna sinensis (Tietz 1952), Vigna species (Chittenden 1902), Wisteria floribunda (Hayward 1941), Wisteria frutescens (Scudder 1889a, b), Wisteria sinensis (Gerberg and Arnett 1989). LAMIACEAE: (Kimball 1965) [erroneous]. MALPIGHIACEAE : Stigmaphyllon lingulatum (Wolcott 1923) [erroneous]. ROSACEAE: Prunus species (Kimball 1965) [erroneous]. Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1322, 24 eggs (DHH); Lot 1417, 14 eggs (DHH) ; Lot 1531, 1 egg (MCM); Lot 1540, 2 eggs (MCM); Lot 1541, 1 egg (MCM). LARVAE: COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 502, 2 larvae (instar 5) (MCM). COSTA RICA: Lot 1484, 1 larva (instar 4) (DHH). HONDURAS: DEPT. COMAYAGUA: Lot 619, 1 larva (instar 3) (SP). JAMAICA: Lot 1081, 8 larvae (instars 2, 3, 4) (DHH). PUERTO RICO: Lot 861, 1 larva (instar 5) (USNM); Lot 862, 1 larva (instar 5) (USNM)

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113 CALIFORNIA: RIVERSIDE CO.: Lot 584, 1 larva ( instar 5) (GRB). FLORIDA: Lot 494, 2 larvae (instar 5) (MCM) ; Lot 1372, 1 larva (instar 5) (DHH); Lot 1454, 6 larvae (instar 5) (DHH); Lot 1479, 1 larva (instar 5) (DHH). ALACHUA CO.: Lot 489, 1 larva (instar 5) (MCM); Lot 498, 2 larvae (instar 5) (MCM); Lot 500, 1 larva (instar 5) (MCM); Lot 503, 1 larva (instar 5) (MCM); Lot 511, 1 larva (instar 5) (MCM); Lot 757, 1 larva (instar 5) (MCM); Lot 761, 3 larvae (instars 4, 5) (MCM); Lot 762, 1 larva (instar 5) (MCM); Lot 764, 1 larva (instar 5) (MCM); Lot 1073, 2 larvae (instar 3) (DHH); Lot 1074, 2 larvae (instar 2) (DHH); Lot 1077, 1 larva (instar 3) (DHH); Lot 1287, 3 larvae (instar 3) (DHH); Lot 1292, 1 larva (instar 4) (DHH); Lot 1293, 8 larvae (instars 4, 5) (DHH) Lot 1294, 2 larvae (instar 5) (DHH); Lot 1295, 16 larvae (instars 3, 4, 5) (DHH); Lot 1296, 18 larvae (instars 4, 5) (DHH); Lot 1297, 8 larvae (instars 4, 5) (DHH); Lot 1298, 2 larvae (instar 5) (DHH); Lot 1299, 18 larvae (instars 1, 2, 3, 4, 5) (DHH); Lot 1300, 11 larvae (instars 3, 4, 5) (DHH); Lot 1301, 2 larvae (instar 4) (DHH); Lot 1304, 5 larvae (instar 5) (DHH); Lot 1307, 1 larva (instar 4) (DHH); Lot 1308, 1 larva (instar 5) (DHH); Lot 1309, 5 larvae (instars 4, 5) (DHH); Lot 1310, 1 larva (instar 5) (DHH); Lot 1311, 5 larvae (instars 4, 5) (DHH); Lot 1312, 1 larva (instar 5) (DHH); Lot 1313, 7 larvae (instars 1, 2, 3) (DHH); Lot 1314, 1 larva (instar 5) (DHH); Lot 1315, 1 larva (instar 5) (DHH); Lot 1317, 1 larva

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114 (instar 5) (DHH); Lot 1318, 2 larvae (instar 2) (DHH); Lot 1319, 9 larvae ( instars 4, 5) (DHH); Lot 1322, 9 larvae (instars 1, 2, 3, 4) (DHH); Lot 1323, 2 larvae (instars 3, 5) (DHH); Lot 1324, 1 larva (instar 5) (DHH); Lot 1325, 5 larvae (instars 4, 5) (DHH); Lot 1326, 1 larva (instar 5) (DHH); Lot 1327, 2 larvae (instar 5) (DHH); Lot 1328, 4 larvae (instars 4, 5) (DHH); Lot 1329, 1 larva (instar 5) (DHH); Lot 1330, 3 larvae (instar 3) (DHH); Lot 1333, 1 larva (instar 5) (DHH); Lot 13 34, 1 larva (instar 5) (DHH); Lot 1335, 2 larvae (instar 5) (DHH); Lot 1336, 1 larva (instar 5) (DHH); Lot 13 37, 3 larvae (instar 5) (DHH); Lot 1338, 1 larva (instar 5) (DHH); Lot 1339, 1 larva (instar 4) (DHH); Lot 1340, 1 larva (instar 4) (DHH); Lot 1341, 1 larva (instar 5) (DHH); Lot 1342, 1 larva (instar 5) (DHH); Lot 1344, 1 larva (instar 5) (DHH); Lot 1345, 1 larva (instar 3) (DHH); Lot 1346, 3 larvae (instars 2, 3) (DHH); Lot 1347, 1 larva (instar 5) (DHH); Lot 1348, 1 larva (instar 5) (DHH); Lot 1349, 2 larvae (instar 5) (DHH); Lot 1350, 2 larvae (instar 5) (DHH); Lot 1352, 1 larva (instar 3) (DHH); Lot 1353, 1 larva (instar 5) (DHH); Lot 1355, 1 larva (instar 5) (DHH); Lot 1356, 1 larva (instar 5) (DHH); Lot 1358, 1 larva (instar 5) (DHH); Lot 1360, 1 larva (instar 5) (DHH); Lot 1361, 1 larva (instar 5) (DHH); Lot 1362, 1 larva (instar 5) (DHH); Lot 1363, 2 larvae (instars 4, 5) (DHH); Lot 1364, 1 larva (instar 5) (DHH); Lot 1366, 2 larvae (instar 5) (DHH) Lot 1367, 1 larva (instar 5) (DHH); Lot 1368, 3 larvae

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115 (instars 4, 5) (DHH); Lot 1369, 3 larvae (instar 5) (DHH); Lot 1370, 5 larvae (instars 4, 5) (DHH); Lot 1371, 1 larva (instar 4) (DHH); Lot 1373, 4 larvae (instars 3, 5) (DHH); Lot 1374, 2 larvae (instar 4) (DHH); Lot 1375, 1 larva (instar 5) (DHH); Lot 1376, 1 larva (instar 5) (DHH); Lot 1377, 2 larvae (instar 5) (DHH); Lot 1379, 4 larvae (instars 2, 3, 4) (DHH); Lot 1380, 1 larva (instar 4) (DHH); Lot 1381, 1 larva (instar 3) (DHH); Lot 1382, 3 larvae (instars 4, 5) (DHH); Lot 1383, 1 larva (instar 5) (DHH); Lot 1384, 1 larva (instar 5) (DHH); Lot 1385, 2 larvae (instar 5) (DHH); Lot 1386, 1 larva (instar 5) (DHH); Lot 1387, 1 larva (instar 5) (DHH); Lot 1388, 1 larva (instar 5) (DHH); Lot 1389, 1 larva (instar 5) (DHH); Lot 1390, 1 larva (instar 5) (DHH); Lot 1391, 10 larvae (instars 2, 3, 4) (DHH); Lot 1392, 1 larva (instar 5) (DHH); Lot 1393, 1 larva (instar 5) (DHH); Lot 1394, 4 larvae (instars 1, 2, 3) (DHH); Lot 1395, 1 larva (instar 4) (DHH); Lot 1396, 4 larvae (instar 2) (DHH); Lot 1397, 2 larvae (instar 4) (DHH); Lot 1398, 2 larvae (instar 3) (DHH); Lot 1400, 4 larvae (instar 5) (DHH); Lot 1403, 1 larva (instar 5) (DHH); Lot 1404, 2 larvae (instar 3) (DHH); Lot 1406, 1 larva (instar 5) (DHH); Lot 1407, 1 larva (instar 4) (DHH); Lot 1409, 1 larva (instar 4) (DHH); Lot 1410, 7 larvae (instar 4) (DHH); Lot 1411, 1 larva (instar 5) (DHH); Lot 1412, 3 larvae (instars 4, 5) (DHH); Lot 1414, 5 larvae (instars 4, 5) (DHH); Lot 1415, 1 larva (instar 5) (DHH); Lot 1416, 2 larvae (instars

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116 4, 5) (DHH); Lot 1417, 8 larvae ( instars 2, 3, 4, 5) (DHH); Lot 1418, 5 larvae (instars 3, 4, 5) (DHH); Lot 1421, 1 larva ( instar 5) (DHH); Lot 1422, 1 larva ( instar 5) (DHH); Lot 1423, 4 larvae (instars 4, 5) (DHH); Lot 1424, 1 larva (instar 5) (DHH); Lot 1426, 2 larvae (instar 5) (DHH); Lot 1429, 1 larva (instar 5) (DHH); Lot 1432, 1 larva (instar 4) (DHH); Lot 1433, 1 larva (instar 5) (DHH); Lot 1434, 1 larva (instar 5) (DHH); Lot 1436, 1 larva (instar 5) (DHH); Lot 1438, 1 larva (instar 5) (DHH); Lot 1439, 3 larvae (instar 5) (DHH); Lot 1440, 3 larvae (instars 3, 4) (DHH); Lot 1441, 1 larva (instar 5) (DHH); Lot 1442, 1 larva (instar 4) (DHH); Lot 1443, 5 larvae (instars 3, 5) (DHH); Lot 1445, 4 larvae (instars 3, 5) (DHH); Lot 1448, 2 larvae (instar 5) (DHH); Lot 1449, 2 larvae (instar 5) (DHH); Lot 1451, 1 larva (instar 4) (DHH); Lot 1452, 1 larva (instar 2) (DHH); Lot 1453, 2 larvae (instars 4, 5) (DHH); Lot 1455, 5 larvae (instars 4, 5) (DHH); Lot 1456, 2 larvae (instar 5) (DHH); Lot 1457, 2 larvae (instar 5) (DHH); Lot 1458, 3 larvae (instar 5) (DHH); Lot 1459, 1 larva (instar 5) (DHH); Lot 1460, 1 larva (instar 5) (DHH); Lot 1462, 1 larva (instar 5) (DHH); Lot 1463, 1 larva (instar 5) (DHH); Lot 1465, 1 larva (instar 5) (DHH); Lot 1466, 3 larvae (instars 3, 5) (DHH); Lot 1469, 1 larva (instar 2) (DHH); Lot 1470, 2 larvae (instars 3, 5) (DHH); Lot 1471, 1 larva (instar 5) (DHH); Lot 1472, 4 larvae (instars 2, 3) (DHH); Lot 1473, 1 larva (instar 5) (DHH); Lot 1474, 1 larva (instar 3) (DHH); Lot

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117 1475, 1 larva ( instar 4) (DHH); Lot 1476, 2 larvae ( instars 3, 4) (DHH); Lot 1477, 5 larvae (instars 3, 4, 5) (DHH); Lot 1478, 2 larvae (instar 3) (DHH); Lot 1480, 7 larvae (instars 4, 5) (DHH); Lot 1481, 2 larvae (instars 4, 5) (DHH); Lot 1485, 1 larva (instar 4) (DHH); Lot 1487, 3 larvae (instars 2, 4) (DHH); Lot 1488, 2 larvae (instars 3, 5) (DHH); Lot 1489, 1 larva (instar 4) (DHH); Lot 1490, 4 larvae (instars 4, 5) (DHH); Lot 1493, 2 larvae (instar 4) (DHH); Lot 1507, 1 larva (instar 5) (MCM); Lot 1531, 2 larvae (instar 2) (MCM); Lot 1540, 4 larvae (instar 1) (MCM); Lot 1542, 8 larvae (instars 1, 2) (MCM). BRADFORD CO.: Lot 1425, 1 larva (instar 5) (DHH); Lot 1447, 3 larvae (instars 4, 5) (DHH); Lot 1467, 8 larvae (instars 4, 5) (DHH); Lot 1468, 4 larvae (instar 5) (DHH). BROWARD CO.: Lot 505, 6 larvae (instar 3, 5) (MCM); Lot 507, 2 larvae (instar 5) (MCM); Lot 1357, 1 larva (instar 5) (DHH). COLUMBIA CO.: Lot 1430, 1 larva (instar 4) (DHH). DADE CO.: Lot 546, 2 larvae (instar 5) (TLM); Lot 1076, 1 larva (instar 3) (DHH); Lot 1303, 2 larvae (instar 5) (DHH); Lot 1401, 5 larvae (instar 3, 4, 5) (DHH); Lot 1428, 1 larva (instar 5) (DHH); Lot 1483, 1 larva (instar 4) (DHH). GADSDEN CO.: Lot 1331, 1 larva (instar 5) (DHH); Lot 1354, 2 larvae (instars 4, 5) (DHH); Lot 1359, 1 larva (instar 4) (DHH); Lot 1378, 1 larva (instar 5) (DHH); Lot 1431, 1 larva (instar 5) (DHH); Lot 1435, 1 larva (instar 4) (DHH); Lot 1437, 1 larva (instar 5) (DHH). HARDEE CO.: Lot 1302, 2 larvae (instars 2, 3)

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118 (DHH). HIGHLANDS CO.: Lot 491, 2 larvae (instars 3, 5) (MCM); Lot 492, 2 larvae ( instar 5) (MCM) ; Lot 510, 2 larvae (instar 5) (MCM). HILLSBOROUGH CO.: Lot 1461, 1 larva (instar 5) (DHH). LEON CO.: Lot 755, 2 larvae (instar 5) (MCM). LEVY CO.: Lot 1316, 3 larvae (instar 5) (DHH); Lot 1320, 1 larva (instar 3) (DHH); Lot 1343, 1 larva (instar 3) (DHH); Lot 1351, 1 larva (instar 5) (DHH); Lot 1399, 2 larvae (instar 5) (DHH). LIBERTY CO.: Lot 1413, 3 larvae (instar 5) (DHH). MADISON CO.: Lot 1075, 1 larva (instar 3) (DHH); Lot 1332, 1 larva (instar 5) (DHH). MANATEE CO.: Lot 1365, 2 larvae (instar 2) (DHH); Lot 1405, 3 larvae (instars 3, 4, 5) (DHH); Lot 1419, 6 larvae (instars 1, 2, 3) (DHH); Lot 1420, 2 larvae (instar 5) (DHH); Lot 1491, 4 larvae (instar 4) (DHH). MARION CO.: Lot 495, 1 larva (instar 4) (MCM); Lot 1305, 1 larva (instar 4) (DHH); Lot 1444, 1 larva (instar 3) (DHH). MONROE CO.: Lot 504, 1 larva (instar 5) (MCM); Lot 1486, 1 larva (instar 4) (DHH). NASSAU CO.: Lot 760, 1 larva (instar 5) (MCM). ORANGE CO.: Lot 1408, 1 larva (instar 4) (DHH). POLK CO.: Lot 497, 3 larvae (instars 3, 5) (MCM); Lot 506, 2 larvae (instar 5) (MCM); Lot 1482, 1 larva (instar 5) (DHH). PUTNAM CO.: Lot 496, 1 larva (instar 5) (MCM); Lot 501, 2 larvae (instar 5) (MCM); Lot 509, 1 larva (instar 5) (MCM); Lot 512, 1 larva (instar 5) (MCM); Lot 753, 2 larvae (instar 5) (MCM); Lot 758, 1 larva (instar 5) (MCM); Lot 759, 1 larva (instar 5) (MCM); Lot 1321, 4 larvae (instars 4, 5) (DHH); Lot 1450, 1

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119 larva (instar 4) (DHH). ST. JOHNS CO.: Lot 490, 1 larva (instar 5) (MCM) ; Lot 499, 2 larvae (instar 5) (MCM) ST. LUCIE CO.: Lot 784, 1 larva (instar 3) (USNM) GEORGIA: COFFEE CO.: Lot 1446, 1 larva (4 instar) (DHH). PUPAE: FLORIDA: Lot 1402, 1 pupa (DHH). ALACHUA CO.: Lot 508, 2 pupae (MCM); Lot 754, 2 pupae (MCM); Lot 1304, 1 pupa (DHH); Lot 1306, 1 pupa (DHH); Lot 1362, 1 pupa (DHH); Lot 1417, 1 pupa (DHH); Lot 1438, 1 pupa (DHH); Lot 1477, 1 pupa (DHH). DADE CO.: Lot 1401, 1 pupa (DHH). HIGHLANDS CO.: Lot 492, 2 pupae (MCM); Lot 630, 3 pupae (MCM). MANATEE CO.: Lot 1464, 1 pupa (DHH). PUTNAM CO.: Lot 756, 1 pupa (MCM); Lot 763, 1 pupa (MCM). ST. JOHNS CO.: Lot 490, 1 pupa (MCM); Lot 493, 1 pupa (MCM). 24. Urbanus pronta Evans A single specimen of this neotropical species has been reported from Texas, but Scott (1986) corrected the identification to Urbanus pronus. Nothing is known of the biology of either species. 25. Urbanus esmeraldus (Butler) Diagnosis EGG: height 1mm, width 0.9mm, 12 ribs. LAST INSTAR LARVA: BODY: length 27mm, A4 transverse width 5.1mm; preserved specimen pale, subdorsal line faintly red, scattered patches of black microspines present, ventral prothorax and prolegs red; some setae with blunt tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: brown, paler toward the apex, large orange eye patches present;

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120 transverse width 4.6mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 38C) ; postocciput wide. THORAX: legs tan; shield broad, tan. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a circle. PUPA: Figure 63A and 82B, length 20.5-23mm, A3 transverse width 6.77.2mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.4mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.7mm long, blunt with numerous hooked setae in a cluster at the tip (Figure 102B and 113E) Other Descriptions Moss 1949 (larva). Host Plants FABACEAE: Desmodium neomexicanum (Kendall 1976) Specimens Examined EGGS: MEXICO: TAMAULIPAS: Lot 985, 1 egg (ROK) LARVAE: MEXICO: TAMAULIPAS: Lot 985, 3 larvae (instars 2, 4, 5) (ROK). PUPAE: MEXICO: TAMAULIPAS: Lot 985, 2 pupae (ROK) 26. Urbanus dorantes (Stoll) Diagnosis EGG: height 0.8-0. 9mm, width 0.7-0. 8mm, 12-13 ribs, green. LAST INSTAR LARVA: BODY: Figure 19E, length 20. 5-36. 5mm, A4 transverse width 5-6. 5mm; color variable, green to pinkish brown, heart line dark, subdorsal line outlined by dark bands, constricted at the segmental

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121 boundaries, small scattered patches of black microspines present, ventral prothorax red; some setae with slightly expanded tips, to 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 23E, black; transverse width 4.14.9mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.3mm long at apex, a few ventral setae to 0.5mm; stemmata subequal (Figure 38D) ; postocciput wide. THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a circle. PUPA: Figures 63B and 82C, length 18.5-25mm, A3 transverse width 5. 9-7. 2mm; brown with darker spots and markings; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.2mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.9mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102C and 113F) Other Descriptions Dethier 1942b (egg, larva), Moss 1949 (larva, pupa), Comstock and Vazquez 1960 (egg, larva), Heppner 1975 (larva), Minno and Emmel 1993 (egg, larva, pupa) Host Plants FABACEAE: Clitoria mariana (Bailowitz and Brock 1991), Desmodium batocaulon (Bailowitz and Brock 1991), Desmodium incanum (Minno 1992), Desmodium

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122 neomexicanum (Bailowitz and Brock 1991), Desmodium tortuosum (Heppner 1975), Phaseolus vulgaris (Dethier 1942b), Vigna luteola (MCM observation). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1532, 4 eggs (MCM). LARVAE: FLORIDA: ALACHUA CO.: Lot 751, 3 larvae (instar 3, 5) (MCM). BROWARD CO.: Lot 517, 1 larva (instar 5) (MCM). DADE CO.: Lot 516, 1 larva (instar 5) (MCM); Lot 750, 1 larva (instar 5) (MCM). HIGHLANDS CO.: Lot 513, 2 larvae (instar 5) (MCM); Lot 514, Lot 515, 1 larva (instar 5) (MCM). SARASOTA CO.: Lot 1492, 1 larva (instar 5) (DHH). PUPAE: FLORIDA: ALACHUA CO.: Lot 752, 2 pupae (MCM). BROWARD CO.: Lot 515, 1 pupa (MCM). HIGHLANDS CO.: Lot 513, 2 pupae (MCM); Lot 514, 2 pupae (MCM) 27. Urbanus teleus (Hiibner) Diagnosis EGG: height 1-1. lmm, width 0.6-0. 8mm, 15-16 ribs. LAST INSTAR LARVA: BODY: length 20-4 3mm, A4 transverse width 5. 1-7. 8mm; preserved specimens brownish; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: dark brown; transverse width 4. 3-4. 8mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.3mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 38E); postocciput wide. THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 100 crochets, irregularly triordinal, arranged in a circle.

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123 PUPA: Figures 63C and 83A, length 18-23. 5mm, A3 transverse width 5. 8-8. 3mm; brown with darker spots and markings; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, < 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 2.3mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102D and 113G) Other Descriptions Moss 1949 (larva). Host Plants POACEAE: Panicum maximum (Kendall 1976), Paspalum ciliatifolium (Scott 1986), Sorghum halepense (Kendall 1976). Specimens Examined EGGS: MEXICO: SAN LUIS POTOSI : Lot 987, 10 eggs (ROK) LARVAE: MEXICO: SAN LUIS POTOSI: Lot 987, 16 larvae ( instars 3, 4, 5) (ROK). PUPAE: MEXICO: SAN LUIS POTOSI: Lot 987, 13 pupae (ROK). 28. Urbanus tanna Evans Nothing is known of the biology of this neotropical species. 29. Urbanus simplicius (Stoll) No immature specimens of this tropical species were located for study. Moss (1949) briefly described the larva. Host Plants CANNACEAE: Canna species (da Costa Lima 1936) [erroneous]. FABACEAE: Phaseolus vulgaris (Garth and Tilden 1986), Schrankia species (Moss 1949).

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124 30. Urbanus procne (Plotz) Diagnosis EGG: height 0.9-lmm, width 0.7-0. 8mm, 13-15 ribs. LAST INSTAR LARVA: BODY: length 2 4. 53 6mm, A4 transverse width 4. 3-6. 3mm; preserved specimens brownish; some setae with expanded tips, to 0.3mm long on A4 dorsum; spiracles dark brown. HEAD: dark reddish brown; transverse width 4. 3-4. 8mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.4mm long at apex, ventral setae to 0.4mm; stemmata subegual (Figure 38F); postocciput wide. THORAX: legs brown; shield broad, tan. ABDOMEN: prolegs each with about 80 crochets, irregularly triordinal, arranged in a near circle. PUPA: Figures 63D and 83B, length 21.5-22mm, A3 transverse width 7. 6-7. 8mm; brown with darker markings; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, < 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.9mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102E and 113H) Host Plants POACEAE: Cynodon dactylon (Kendall 1966a), Lolium perenne (Kendall 1976, in lab), Panicum maximum (Kendall 1976, in lab), Sorghum halepense (Scott 1986), Stenotaphrum secundatum (Kendall 1976, in lab). Specimens Examined EGGS: TEXAS: CAMERON CO.: Lot 986, 10 eggs (ROK). LARVAE: COLOMBIA: DEPT. VALLE DEL CAUCA:

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125 Lot 518, 1 larva ( instar 5) (MCM) TEXAS: CAMERON CO.: Lot 986, 5 larvae ( instars 4, 5) (ROK) PUPAE: TEXAS: CAMERON CO.: Lot 986, 4 pupae (ROK). 31. Urbanus doryssus (Swainson) Nothing is known of the biology of this neotropical species. 32. Urbanus albimargo (Mabille) Nothing is known of the biology of this neotropical species. 33. Astraptes fulgerator (Reakirt) Diagnosis EGG: height 1 2-1 4nun, width 0.9-1. 2mm, 15-17 ribs. LAST INSTAR LARVA: BODY: length 29-49mm, A4 transverse width 5.1-8. 3mm; preserved specimens brownish red with transverse yellow stripes on T2-A9, ventral side bright red; setae simple, to 1.6mm long on A4 dorsum; spiracles tan. HEAD: Figure 23F, light brown with darker brown patches on upper face; transverse width 5. 3-6. 2mm; sculpturing pebbly; mandibles without teeth; setae simple, to 2mm long at apex, ventral setae to 2mm; stemmata subegual (Figure 38G); postocciput wide. THORAX: legs red basally, tan distal ly; shield broad, brown. ABDOMEN: prolegs each with about 150 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 46F). PUPA: Figures 63E and 84A, length 22-28. 5mm, A3 transverse width 7. 710. 3mm; brown to dark brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated;

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126 setae simple, to 0.3mm long on head; head rounded, a short point on the anterior margin of eyes; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 2.4mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102F and 1131) Other Descriptions Moss 1949 (larva, pupa), Comstock and Vazguez 1960 (larva, pupa), Young 1985 (egg, larva). Host Plants AQUIFOLIACEAE: Jlex paraguariensis (Hayward 1941). ASTERACEAE: Calea urticifolia (Young 1985). FABACEAE: Cassia hoffmanseggii (Moss 1949), Cassia species (da Costa Lima 1936), Erythrina species (Young 1985), Inga species (Moss 1949). RHAMNACEAE: Karwinskia humboldtiana (Kendall and McGuire 1975). TRIGONIACEAE : Trigonia rugosa (Young 1985). VERBENACEAE: Vitex mollis (Comstock and Vazguez 1960) Specimens Examined EGGS: MEXICO: TAMAULIPAS: Lot 910, 4 eggs (ROK) USA: TEXAS: HIDALGO CO.: Lot 911, 3 eggs (ROK). LARVAE: MEXICO: TAMAULIPAS: Lot 910, 3 larvae (instars 3, 5) (ROK). TEXAS: HIDALGO CO.: Lot 909, 1 larva (instar 5) (ROK); Lot 911, 1 larva (instar 5) (ROK). PUPAE: USA: TEXAS: HIDALGO CO.: Lot 909, 1 pupa (ROK) ; Lot 910, 3 pupae (ROK). 34. Astraptes egregius (Butler) Nothing is known of the biology of this neotropical species.

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127 35. Astraptes alardus (Stoll) Moss (1949) briefly described the larva and pupa of this neotropical species, but I could not find specimens of the immature stages for study. This skipper has rarely been recorded from the U. S. 36. Astraptes gilbert i H. A. Freeman Diagnosis LAST INSTAR LARVA: BODY: length 18.5-32mm, A4 transverse width 3. 7-6. 6mm; preserved specimens red with numerous tiny yellow spots, the sides yellow, a rectangular orange spot above the spiracle of A8; setae simple, to 2mm long on A4 dorsum; spiracles tan. HEAD: Figure 24A, light brown with darker around the mouth, large yellow eye patches present; transverse width 4. 6-5. 6mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.3mm long at apex, a few ventral setae to 0.8mm; stemmata subequal (Figure 38H) ; postocciput wide. THORAX: legs red basally, tan distally; shield broad, tan. ABDOMEN: prolegs each with about 140 crochets, irregularly triordinal, arranged in a circle. PUPA: Figures 64A and 84B, length 18mm, A3 transverse width 6.2mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, < 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.6mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102G and 114A)

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128 Host Plants FABACEAE: Bauhinia divaricata (Kendall and McGuire 1975) Specimens Examined LARVAE: MEXICO: SAN LUIS POTOSI : Lot 912, 18 larvae ( instars 2, 3, 4, 5) (ROK) PUPAE: MEXICO: SAN LUIS POTOSI: Lot 912, 4 pupae (ROK). 37. Astraptes galesus (Mabille) Nothing is known of the biology of this neotropical species. 38. Astraptes anaphus (Cramer) Diagnosis LAST INSTAR LARVA: BODY: length 24mm, A4 transverse width 5.5mm; preserved specimen brownish; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: reddish brown, large yellow eye patches present; transverse width 5mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.3mm long at apex, a few ventral setae to 0.7mm; stemmata subegual (Figure 381); postocciput wide. THORAX: legs tan; shield broad, dark brown. ABDOMEN: prolegs each with about 100 crochets, irregularly triordinal, arranged in a circle. Host Plants FABACEAE: (Moss 1949), Phaseolus species (Pyle 1981), Pueraria lobata (Kendall 1976). Specimens Examined LARVAE: COLOMBIA: VALLE DEL CAUCA PROV. : Lot 83, 1 larva (instar 5) (MCM) 39. Autochton cellus (Boisduval and Leconte) Diagnosis LAST INSTAR LARVA: BODY: Figure 19A, length 17. 5-30. 5mm, A4 transverse width 4. 2-7. lmm; green with

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129 numerous tiny yellow spots, subdorsal line wide, lemon yellow, ventral prothorax bright red; some setae with slightly expanded tips, < 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 24B, brown, paler toward apex, large pale orange eye patches present; transverse width 4. 2-7. lmm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.7mm; stemmata subegual (Figure 38J) ; postocciput wide. THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about 100 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 47A) PUPA: Figures 64B and 84C, length 16-21mm, A3 transverse width 5. 3-7. 7mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.5mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.6mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 102H and 114B) Other Descriptions Beutenmiiller 1893 (larva, pupa), Clark 1934, 1936 (egg, larva, pupa). Host Plants CONVOLVULACEAE : Bonamia aquatica (Scudder 1889a) [erroneous]. FABACEAE: Amphicarpa bracteata (Clark 1934), Clitoria mariana (Burns 1984, probably; Scott 1986), Desmodium batocaulon (Bailowitz and Brock 1991, possibly), Phaseolus grayanus (Burns 1984), Phaseolus ritensis

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130 (Bailowitz and Brock 1991, possibly), Phaseolus wrightii (Burns 1984), Pueraria lobata (MCM collection), Robinia neomexicana (Bailowitz and Brock 1991), Vigna species (Burns 1984). SAXIFRAGACEAE: Hydrangea species (Clark 1936) [erroneous] Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 48, 1 larva ( instar 3) (MCM); Lot 49, 3 larvae ( instars 3, 5) (MCM); Lot 50, 4 larvae (instar 5) (MCM); Lot 51, 10 larvae (instars 3, 4, 5). TEXAS: BRESTER CO.: Lot 914, 6 larvae (instar 5) (ROK). PUPAE: FLORIDA: ALACHUA CO.: Lot 50, 1 pupa (MCM). TEXAS: BREWSTER CO.: Lot 914, 2 pupae (ROK) 40. Autochton pseudocellus (Coolidge and Clemence) No immature specimens of this tropical species were located for study. This skipper is rarely found in the U. S. Host Plants FABACEAE: Desmodium batocaulon (Bailowitz and Brock 1991) 41. Achalarus lyciades (Geyer) Diagnosis EGG: height 1.1mm, width 0.9-lmm, 14-16 ribs. LAST INSTAR LARVA: BODY: length 15-29mm, A4 transverse width 4-6. 7mm; brown with small scattered patches of black microspines; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 24C, black; transverse width 4. 5-5. 4mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.3mm

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131 long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 38K) ; postocciput wide. THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 47B) PUPA: Figure 64C, length 17.5mm, A3 transverse width 5.3mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.2mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.8mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 1021 and 114C) Other Descriptions Scudder 1889a (egg, larva, pupa) Host Plants CONVOLVULACEAE : Ipomoea pandurata (Scudder 1889a, b) [erroneous], Ipomoea species (Davenport and Dethier 1938) [erroneous]. FABACEAE: Amorpha fruticosa (Tietz 1972), Baptisia species (Scudder 1889a, b), Clitoria mariana (MCM collection), Desmodium canadense (Shapiro 1974a), Desmodium ciliare (Kendall 1965), Desmodium cuspidatum (Shapiro 1974a), Desmodium glabellum (Tietz 1972), Desmodium "dillenii" (Scudder 1889a, b) [this name is currently placed under both D. glabellum and D. perplexum] Desmodium nudiflorum (Shapiro 1974a), Desmodium nuttallii (Shapiro 1974a), Desmodium paniculatum (Scudder 1889a, b), Desmodium rotundi folium (Shapiro 1974a), Hedysarum species (Scudder

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132 1869), Indigofera caroliniana (Tietz 1952), Indigofera species ( Scudder 1889a, b), Lespedeza hirta (Tietz 1952), Lespedeza texana (Kendall 1965). FUMARIACEAE: Corydalis sempervirens (Scudder 1889a, b, possibly; Tietz 1952) [erroneous ] Specimens Examined EGGS: TEXAS: WALKER CO.: Lot 896, 10 eggs (ROK) LARVAE: FLORIDA: ALACHUA CO.: Lot 45, 1 larva (instar 5) (MCM) GEORGIA: HARRIS CO.: Lot 47, 2 larvae ( instars 2,4) (MCM). NORTH CAROLINA: IREDELL CO.: Lot 46, 1 larva (instar 5) (MCM). TEXAS: WALKER CO.: Lot 896, 21 larvae (instars 1,2,4,5) (ROK). PUPAE: TEXAS: WALKER CO.: Lot 896, 1 pupa (ROK). 42. Achalarus casica (Herrich-Schaf f er ) Diagnosis LAST INSTAR LARVA: BODY: length 163 lmm, A4 transverse width 4. 7-7. 2mm; preserved specimens brownish with small scattered patches of black microspines; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: dark reddish brown; transverse width 5-5. 4mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.7mm; stemmata subequal (Figure 3 8L); postocciput wide. THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs irregularly triordinal, arranged in a circle. Host Plants FABACEAE: Clitoria mariana (Bailowitz and Brock 1991), Desmodium batocaulon (Bailowitz and Brock

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133 1991), Desmodium cinerascens (Bailowitz and Brock 1991), Desmodium grahamii (ROK collection). Specimens Examined LARVAE: TEXAS: JEFF DAVIS CO.: Lot 895, 6 larvae ( instars 4,5) (ROK). 43. Achalarus albociliatus (Mabille) Nothing is known of the biology of this neotropical species. 44. Achalarus toxeus (Plotz) No immature specimens of this neotropical species were located for study. Host Plants FABACEAE: Acacia farnesiana (Kendall 1965, possibly), Pithecellobium flexicaule (Kendall 1965), Prosopis glandulosa (Kendall 1965, possibly). 45. Achalarus jalapus (Plotz) Nothing is known of the biology of this neotropical species. 46. Thorybes bathyllus (J. E. Smith) Diagnosis LAST INSTAR LARVA: BODY: length 20.5-37mm, A4 transverse width 3.6-7mm; color variable, green or brown, heart line dark, subdorsal line narrow, pale yellow or pinkish; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 24D, black; transverse width 4. 3-5. 5mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.8mm; stemmata subegual (Figure 39A) ; postocciput wide. THORAX: legs dark brown; shield broad,

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134 dark brown. ABDOMEN: prolegs each with about 75 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked (Figure 47C) PUPA: Figures 58B, 64D and 85A, length 19. 5-20. 5mm, A3 transverse width 6. 2-6. 8mm; pale green or cream-colored with brown spots and markings; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.3mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.6mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 103A and 114D) Other Descriptions Harris 1862 (pupa), Scudder 1889a (egg, larva, pupa) Host Plants FABACEAE: Apios americana (Beutenmiiller 1893), Astragalus engelmannii (Kendall 1965), Centrosema virginianum (Scudder 1889a, b), Clitoria mariana (MCM collection), Desmodium ciliare (Kendall 1965), Desmodium "dillenii" (Middleton 1881) [this name is currently placed under both D. glabellum and D. perplexum] Desmodium paniculatum (Kendall 1965), Desmodium rotundifolium (Shapiro 1968), Desmodium striatum (MCM collection), Desmodium viridiflorum (MCM collection) Glycine reticulata (Harris 1972), Glycine species (Harris 1862), Hedysarum species (Harris 1862), Lespedeza capitata (Grossbeck 1917), Lespedeza hirta (Scudder 1889a, b, probably; Tietz 1952),

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135 Lespedeza texana (Kendall 1965, in lab), Lespedeza virginica (Tietz 1952), Rhynchosia tomentosa (Scudder 1889a, b), Strophostyles helveola (Huber 1966), Strophostyles leiosperma (Huber 1966), Strophostyles species (Klots 1951), Tephrosia florida (Scudder 1889a, b) Tephrosia virginiana (Harris 1972), Trifolium pratense (Tietz 1972). Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 475, 1 larva ( instar 5) (MCM) ; Lot 478, 1 larva ( instar 5) (MCM); Lot 1277, 1 larva (instar 5) (DHH); Lot 1279, 1 larva (instar 5) (DHH); Lot 1280, 1 larva (instar 5) (DHH). LEON CO.: Lot 1278, 1 larva (instar 5) (DHH). MARION CO.: Lot 742, 1 larva (instar 5) (MCM). PUTNAM CO.: Lot 473, 1 larva (instar 5) (MCM); Lot 474, 5 larvae ( instars 3, 4, 5) (MCM); Lot 476, 4 larvae (instars 3, 5) (MCM); Lot 481, 1 larva (instar 5) (MCM); Lot 740, 2 larvae (instar 3) (MCM); Lot 741, 1 larva (instar 5) (MCM); Lot 744, 2 larvae (instar 5) (MCM); Lot 745, 1 larva (instar 5) (MCM); Lot 746, 1 larva (instar 5) (MCM); Lot 747, 1 larva (instar 5) (MCM). INDIANA: TIPPECANOE CO.: Lot 477, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: PUTNAM CO.: Lot 743, 2 pupae (MCM); Lot 749, 1 pupa (MCM). 47. Thorybes py lades (Scudder) Diagnosis EGG: Figure 3C, height 0.6-0. 9mm, width 0.81.1mm, ribs 12-17, white. LAST INSTAR LARVA: BODY: Figure 19C, length 18-41mm, A4 transverse width 4. 2-6. 9mm; color variable, green or pinkish brown, heart line dark, subdorsal

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136 line narrow, pale yellow or pinkish, ventral prothorax reddish; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: black; transverse width 4. 3-5. 5mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 39B) ; postocciput wide. THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a near circle. PUPA: Figures 64E and 85B, length 20-28mm, A3 transverse width 6. 2-7. 2mm; pale green or cream-colored with brown spots and markings; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.2mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.6mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 103B and 114E) Other Descriptions Scudder 1881 (egg), Scudder 1889a (egg, larva, pupa), Comstock and Dammers 1933b (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants FABACEAE: Amorpha calif ornica (Comstock and Dammers 1933b), Apios americana (MCM collection), Astragalus nuttallianus (Kendall 1965), Centrosema floridanum (MCM collection), Centrosema virginianum (MCM collection), Cologania angustifolia (Bailowitz and Brock 1991), Desmodium

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137 batocaulon (Bailowitz and Brock 1991), Desmodium canadense (Shapiro 1974a), Desmodium glabellum (Tietz 1972), Desmodium "dillenii" (Scudder 1889a, b) [this name is currently placed under both D. glabellum and D. perplexum] Desmodium nudiflorum (Tietz 1952), Desmodium paniculatum (Kendall 1965), Desmodium rotundi folium (Shapiro 1974a), Galactia elliottii (MCM collection), Galactia regularis (Minno 1992), Hedysarum species (Edwards 1870, in lab), Lathyrus eucosmus (Scott 1986), Lathyrus jepsonii ssp. californicus (Shapiro et al. 1981, associated with), Lathyrus ochroleucus (Scott 1986), Lespedeza capitata (Scudder 1889a, b), Lespedeza hirta (Scudder 1889a, b), Lespedeza intermedia (Shapiro 1974a), Lotus crassifolius (Scott 1986), Lotus douglasii (Newcomer 1964), Medicago sativa (Shapiro 1966), Rhynchosia difformis (Minno 1992), Rhynchosia texana (Kendall 1965), Trifolium pratense (Scudder 1889a, b), Trifolium repens (Scudder 1889a, b), Vicia americana (Scott 1986). Specimens Examined EGGS: TEXAS: JEFF DAVIS CO.: Lot 981, 3 eggs (ROK) FLORIDA: NASSAU CO.: Lot 1535, 1 egg (MCM). PUTNAM CO.: Lot 1536, 1 egg (MCM). LARVAE: CALIFORNIA: RIVERSIDE CO.: Lot 594, 1 larva ( instar 5) (GRB). FLORIDA: HIGHLANDS CO.: Lot 482, 1 larva (instar 5) (MCM); Lot 485, 1 larva (instar 5) (MCM); Lot 486, 3 larvae (instar 5) (MCM); Lot 487, 2 larvae (instar 5) (MCM). LEVY CO.: Lot 484, 3 larvae (instar 5) (MCM). MARION CO.: Lot 483, 2 larvae (instar 5) (MCM). TEXAS: JEFF DAVIS CO.:

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138 Lot 981, 6 larvae (instars 3, 5) (ROK). PUPAE: FLORIDA: HIGHLANDS CO.: Lot 487, 2 pupae (MCM) TEXAS: JEFF DAVIS CO.: Lot 981, 2 pupae (ROK). 48. Thorybes diversus Bell Diagnosis LAST INSTAR LARVA: BODY: length 31. 5-32. 5mm, A4 transverse width 6. 7-6. 8mm; preserved specimen brownish; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: black; transverse width 4.44.5mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 39C); postocciput wide. THORAX: Tl-2 legs dark brown, T3 legs brown; shield broad, dark brown. ABDOMEN: prolegs each with about 55 crochets, irregularly triordinal arranged in a circle or near circle. Other Descriptions MacNeill 1975 (egg, larva). Host Plants FABACEAE: Trifolium wormskjoldii (MacNeill 1975) Specimens Examined LARVAE: CALIFORNIA: DEL NORTE CO.: Lot 488, 1 larva (instar 5) (MCM); Lot 595; 1 larva (instar 5) (GRB). 49. Thoryjbes mexicanus (Herrich-Schaf f er ) Diagnosis LAST INSTAR LARVA: BODY: length 26mm, A4 transverse width 5.5mm; preserved specimen brownish; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: black; transverse width 3.7mm; sculpturing pebbly; mandibles without teeth; setae simple,

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139 to 0.2mm long at apex, a few ventral setae to 0.8mm; stemmata subequal (Figure 39D) ; postocciput wide. THORAX: Tl legs dark brown, T2-3 legs brown; shield broad, dark brown. ABDOMEN: crochets, irregularly triordinal arranged in a circle. Other Descriptions Emmel et al 1992 (larva). Host Plants FABACEAE: Amorpha californica ( Corns tock and Dammers 1933b) [erroneous, MacNeill 1975], Astragalus species (Beutelspacher 1980), Desmodium species (Beutelspacher 1980), Lathyrus arizonicus (Bailowitz and Brock 1991), Lathyrus leucanthus (Shields et al 1970), Lathyrus species (Dornfeld 1980), Lespedeza species (Beutelspacher 1980), Rhynchosia species (Beutelspacher 1980), Trifolium longipes (Scott 1986), Trifolium monanthum (Lembert 1894), Vicia americana (Scott 1986). POACEAE: Sitanion species (Dornfeld 1980) [erroneous]. Specimens Examined LARVAE: CALIFORNIA: INYO CO.: Lot 579, 1 larva ( instar 5) (GRB). 50. Thorybes confusis Bell Diagnosis LAST INSTAR LARVA: BODY: length 18-3 7mm, A4 transverse width 3. 6-6. 3mm; pinkish brown, heart line dark, subdorsal line narrow, pale pink, ventral prothorax reddish; some setae with expanded tips, to 0.3mra long on A4 dorsum; spiracles dark brown. HEAD: black; transverse width 4.24.9mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.3mm;

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140 stemmata subequal (Figure 39E); postocciput wide. THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a circle or near circle. PUPA: Figures 64F and 85C, length 19.5mm, A3 transverse width 5. 7-6. 4mm; pale green or cream-colored with brown spots and markings; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.3mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.5mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 103C and 114F) Host Plants FABACEAE: Lespedeza species (Shapiro 1966, possibly) Tephrosia florida (MCM collection) Trifolium repens (MCM collection, in lab). Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 1494, 1 larva ( instar 5) (DHH) PUTNAM CO.: Lot 479, 1 larva (instar 5) (MCM); Lot 748, 5 larvae ( instars 3, 5) (MCM). PUPAE: FLORIDA: PUTNAM CO.: Lot 748, 2 pupae (MCM) 51. Thorybes drusius (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 2 0-3 0.5mm, A4 transverse width 4. 6-5. 8mm; preserved specimens pinkish brown, heart line dark, subdorsal line pink, ventral prothorax pale red; some setae with expanded tips, to 0.1mm

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141 long on A4 dorsum; spiracles dark brown. HEAD: dark brown; transverse width 4-4. 5mm; sculpturing pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 39F); postocciput wide. THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 50 crochets, irregularly triordinal, arranged in a circle or near circle. Host Plants FABACEAE: Cologania angustifolia (Bailowitz and Brock 1991) Specimens Examined LARVAE: MEXICO: CHIHUAHUA: Lot 980, 10 larvae (instars 3, 5) (ROK) probably this species. 52. Thorybes valerianus (Plotz) Nothing is known of the biology of this species, except that the larvae feed on leguminous plants (Scott 1986). 53. Cabares potrillo (Lucas) Diagnosis EGG: height 0.8-0. 9mm, width 0.5-0. 6mm, ribs 10-11. LAST INSTAR LARVA: BODY: length 25mm, A4 transverse width 5mm; preserved specimen pale; some setae with expanded tips, to 0.1mm long on A4 dorsum; spiracles brown. HEAD: Figure 24E, dark reddish brown; transverse width 3.8mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subequal (Figure 3 9G) ; postocciput wide. THORAX: legs dark brown; shield pale dorsally, brown laterally. ABDOMEN: crochets, irregularly triordinal, arranged in a circle; suranal plate

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142 rounded, unmarked (Figure 47D) PUPA: Figures 65A and 85D, length 18.5mm, A3 transverse width 5.2mm; brown; thoracic spiracle guard distinct; posterior margin of prothorax crenulated; setae simple, to 0.3mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.5mm long, tapering, blunt with numerous hooked setae in a cluster at the tip (Figures 103D and 114G) Other Descriptions Dethier 1940a (egg), MacNeill 1975 (egg, larva, pupa). Host Plants LAMIACEAE: Salvia species (ROK collection). VERBENACEAE: Priva lappulacea (Kendall and Rickard 1976). Specimens Examined EGGS: TEXAS: HIDALGO CO.: Lot 915, 10 eggs (ROK). LARVAE: TEXAS: HIDALGO CO.: Lot 915, 2 larvae (instars 2, 5) (ROK). PUPAE: TEXAS: HIDALGO CO.: Lot 915, 1 pupa (ROK). 54. Celaenorrhinus fritztgaertneri (Bailey) Nothing is known of the biology of this neotropical species. 55. Celaenorrhinus stallingsi H. A. Freeman Nothing is known of the biology of this neotropical species. 56. Dyscophellus euribates (Stoll) Moss (1949) gave a description of the larva and pupa of this neotropical species, and listed the host plant as

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143 Virola spp. (Myristicaceae) This skipper is rarely recorded from the U. S. I could not locate specimens of the immature stages for study. 57. Spathilepia clonius (Cramer) No immature specimens of this tropical species were located for study. Moss (1949) briefly described the larva and pupa. Host Plants FABACEAE: Inga edulis (Moss 1949), Phaseolus species (Moss 1949). 58. Cogia calchas (Herrich-Schaf fer) No immature specimens of this neotropical species were located for study. Moss (1949) briefly described the larva. Host Plants FABACEAE: Indigofera species (Moss 1949), Malicia species (Brown and Heineman 1972), Mimosa pigra var. berlandieri (Kendall 1966a), Schrankia species (Moss 1949). 59. Cogia hippalus (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 16.5-22mm, A4 transverse width 3.6-5mm; preserved specimen pale with numerous tiny yellow spots; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 24F, light brown, front dark brown, posterior black, yellowish eye patches present; transverse width 4-4. 3mm; sculpturing pebbly; mandibles without teeth; setae branching, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 39H); postocciput wide. THORAX: legs pale; shield indistinct. ABDOMEN: Prolegs each with about

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144 45 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 47E) PUPAL EXUVIUM: cremaster blunt with numerous hooked setae at tip (Figures 103E and 114H) Host Plants FABACEAE: Acacia angustissima (Pyle 1981). Specimens Examined LARVAE: MEXICO: TAMAULIPAS: Lot 928, 3 larvae ( instar 5) (ROK) 60. Cogia outis (Skinner) Diagnosis EGG: height 0.6-0. 7mm, width 0.5-0. 6mm, polygonal sculpturing, no ribs. LAST INSTAR LARVA: BODY: length 13.5-24mm, A4 transverse width 3. 8-5. 8mm; preserved specimens pale; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: light brown, posterior black, faint yellowish eye patches present; transverse width 3.7-4. 3mm; sculpturing pebbly; mandibles without teeth; setae branching, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 391); postocciput wide. THORAX: legs tan to pale; shield indistinct. ABDOMEN: prolegs each with about 45 crochets, irregularly triordinal, arranged in a near circle or mesal penellipse. PUPA: Figures 65B and 85E, length 15-19mm, A3 transverse width 4.8-6mm; light brown with darker spots and markings; thoracic spiracle guard distinct; setae simple, < 0.1mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.7mm long, tapering, blunt with

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145 numerous hooked setae in a cluster at the tip (Figures 103F and 1141) Host Plants FABACEAE: Acacia angustissima var. hirta (Kendall 1965), Acacia texensis (Kendall 1965). Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 930, 4 eggs (ROK) LARVAE: TEXAS: BEXAR CO.: Lot 930, 10 larvae (instars 2, 3, 5) (ROK). PUPAE: TEXAS: BEXAR CO.: Lot 930, 14 pupae (ROK) 61. Cogia caicus (Herrich-Schaf fer) Diagnosis LAST INSTAR LARVA: BODY: length 20-37. 5mm, A4 transverse width 4.1-7. 3mm; preserved specimens pale with numerous tiny yellow spots; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: light brown, front darker brown, posterior black, yellow eye patches present; transverse width 3. 7-4. 7mm; sculpturing pebbly; mandibles without teeth; setae branching, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 39J) ; postocciput wide. THORAX: legs tan to pale; shield indistinct. ABDOMEN: prolegs each with about 30 crochets, irregularly triordinal, arranged in a mesal penellipse. PUPAL EXUVIUM: cremaster blunt with numerous hooked setae at tip (Figures 103G and 115A) Host Plants FABACEAE: Acacia angustissima (Kendall and McGuire 1975)

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146 Specimens Examined LARVAE: MEXICO: TAMAULIPAS: Lot 929, 1 larva ( instar 5) (ROK) ARIZONA: Lot 553, 3 larvae (instar 5) (NM) 62. Nisoniades rubescens (Moschler) Diagnosis LAST INSTAR LARVA: BODY: length 16-24mm, A4 transverse width 3. 6-5. 3mm; preserved specimens pale, subdorsal yellow spots on T2-3; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 25A, dark reddish brown; transverse width 3. 5-3. 6mm; sculpturing pebbly, small spines at apex; mandibles with teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 39K) ; postocciput moderately wide. THORAX: legs tan to pale; shield broad, dark brown. ABDOMEN: prolegs each with about 110 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 47F) PUPAL EXUVIUM: cremaster blunt with numerous hooked setae at tip (Figures 103H and 115B) Host Plants CONVOLVULACEAE : Ipomoea batatas (Kendall 1976), Ipomoea luteum (Kendall 1976), Ipomoea pandurata (ROK collection, probably), Ipomoea trichocarpa (Kendall 1976, possibly) Specimens Examined LARVAE: MEXICO: TAMAULIPAS: Lot 955, 1 larva (instar 5) (ROK); Lot 956, 2 larvae (instar 5) (ROK).

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147 63 Pellicia angra Evans Nothing is known of the biology of this neotropical species. 64. Pellicia arina Evans Nothing is known of the biology of this species from tropical America. 65. Bolla clytius (Godman and Salvin) Nothing is known of the biology of this neotropical species. 66. Bolla brennus (Godman and Salvin) Nothing is known of the biology of this species from tropical America. 67. Staphylus ceos (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 19.5mm, A4 transverse width 3.9mm; preserved specimen pale; some setae with blunt tips, to 0.2mm long on A4 dorsum; spiracles pale, HEAD: Figure 25B, dark reddish brown; transverse width 2.6mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.5mm long at apex, ventral setae to 0.5mm; stemmata subegual; postocciput wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 55 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 47G) PUPA: Figures 65C and 86A, length 12.5mm, A3 transverse width 3.8mm; light brown; thoracic spiracle guard distinct; setae simple, to 0.3mm long on head; head rounded; antennae

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148 extend to nearly the tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 1031 and 115C) Host Plants CHENOPODIACEAE : Chenopodium fremontii (Bailowitz and Brock 1991), Chenopodium species (Bailowitz and Brock 1991) Specimens Examined LARVAE: ARIZONA: COCHISE CO.: Lot 1010, 1 larva (instar 5) (SDNHM). PUPAE: ARIZONA COCHISE CO.: Lot 1010, 1 pupa (SDNHM). 68. Staphylus mazans (Reakirt) Diagnosis LAST INSTAR LARVA: BODY: length 14-16mm, A4 transverse width 3.3mm; preserved specimen pale; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: dark reddish brown; transverse width 3.3mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.4mm long at apex, ventral setae to 0.4mm; stemmata subegual (Figure 39L); postocciput wide. THORAX: legs pale; shield broad, tan. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a near circle. PUPAL EXUVIUM: cremaster blunt with numerous hooked setae at tip (Figures 104A and 115D) Host Plants AMARANTHACEAE : Amaranthus aspera (Scott 1986), Amaranthus retroflexus (Kendall 1965). CHENOPODIACEAE: Chenopodium album (Kendall 1965), Chenopodium ambrosioides (Kendall 1965).

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149 Specimens Examined LARVAE: TEXAS: HIDALGO CO.: Lot 975, 2 larvae (instar 5) (ROK) 69. Staphylus hayhurstii (Edwards) Diagnosis LAST INSTAR LARVA: BODY: Figure 20D, length 15.5-26mm, A4 transverse width 2.8-4. 3mm; dark green with numerous tiny white spots; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles pale. HEAD: black; transverse width 2.7-3mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.3mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 39M) ; postocciput wide. THORAX: legs pale; shield broad, tan. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a circle. PUPA: Figures 65D and 85B, length 13.5-16mm, A3 transverse width 3. 8-4. 3mm; brown; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 104B and 115E) Other Descriptions Dyar 1891 (larva, pupa), Heitzman 1963 (egg, larva, pupa), Minno and Emmel 1993 (larva, pupa). Host Plants AMARANTHACEAE : Alternanthera species (Grossbeck 1917) [erroneous], Amaranthus species (Forbes 1906), Celosia nitida (ROK collection), Iresine diffusa (MCM

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150 collection), Iresine flavescens (Dyar 1891). CHENOPODIACEAE: Chenopodium album (Opler and Krizek 1984). Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 467, 3 larvae (instar 5) (MCM) ; Lot 468, 3 larvae ( instar 5) (MCM); Lot 469, 1 larva (instar 5) (MCM); Lot 738, 3 larvae (instar 5) (MCM); Lot 739, 4 larvae (instar 5) (MCM); Lot 1281, 1 larva (instar 5) (DHH) PUPAE: FLORIDA: ALACHUA CO.: Lot 467, 1 pupa (MCM); Lot 468, 1 pupa (MCM); Lot 739, 2 pupae (MCM) 70. Gorgythion begga (Kirby) Nothing is known of the biology of this species from tropical America. 71. Sostrata bifasciata (Menetries) Nothing is known of the biology of this neotropical species. 72. Carrhenes canescens (R. Felder) Diagnosis LAST INSTAR LARVA: BODY: length 18-27mm, A4 transverse width 4-4. 6mm; preserved specimens pale; some setae with blunt tips, to 0.3mm long on A4 dorsum; spiracles pale. HEAD: Figure 25C, dark reddish brown; transverse width 3. 4-3. 7mm; sculpturing rough; mandibles without teeth; setae simple, to 0.4mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 39N) ; postocciput wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 80 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked i

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151 (Figure 47H) PUPA: Figures 65E and 86C, length 14mm, A3 transverse width 4.7mm; light brown; thoracic spiracle guard distinct; setae simple, to 0.8mm long on head; head rounded; antennae extend slightly caudad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.5mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 104C and 115F). Host Plants MALVACEAE: Hibiscus species (Scott 1986), Malache rosae (ROK collection). Specimens Examined LARVAE: MEXICO: SAN LUIS POTOSI : Lot 916, 9 larvae ( instars 2, 3, 5) (ROK). PUPAE: MEXICO: SAN LUIS POTOSI: Lot 916, 2 pupae (ROK). 73. Xenophanes trixus (R. Felder) Diagnosis LAST INSTAR LARVA: BODY: length 29mm, A4 transverse width 4.8mm; preserved specimens pale with numerous tiny white spots; some setae with slightly expanded tips, to 0.5mm long on A4 dorsum; spiracles pale. HEAD: Figure 25D, brown; transverse width 3.7mm; sculpturing rough; mandibles without teeth; some setae with expanded tips, to 0.3mm long at apex, a few ventral setae to 0.6mm; stemma 5 slightly small (Figure 390); postocciput wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked (Figure 471). PUPAL EXUVIUM: cremaster blunt with numerous hooked setae at tip (Figures 104D and 115G)

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152 Other Descriptions Moss 1949 (larva, pupa). Host Plants MALVACEAE: Malachra fasciata (Moss 1949), Malvaviscus drummondii (Kendall and Rickard 1976). Specimens Examined LARVAE: TEXAS: CAMERON CO.: Lot 991, 1 larva (instar 5) (ROK). 74. Systasea pulverulenta (R. Felder) Diagnosis EGG: height 0.9-1. lmm, width 0.5-0. 8mm, 44-52 vertical rows of short spines. LAST INSTAR LARVA: BODY: length 11-29. 5mm, A4 transverse width 3. 2-4. 7mm; preserved specimens pale; setae simple, to 0.5mm long on A4 dorsum; spiracles pale. HEAD: Figure 25E, light brown, pale eye patches present or dark brown with two paler lines parallel to vertex, light brown around the mouth; transverse width 2. 7-3. 4mm; sculpturing rough; mandibles with teeth; setae simple, to 0.5mm long at apex, a few ventral setae to 0.8mm; stemmata subegual (Figure 39P); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 30 crochets, uniordinal, arranged in a mesal penellipse. PUPA: Figures 65F and 86D, length 15-17mm, A3 transverse width 4.8mm; preserved specimens pale; thoracic spiracle guard distinct; setae simple, to lmm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 1.2mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 104E and 115H)

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153 Host Plants MALVACEAE: Abutilon abutiloides (Kendall 1965), Abutilon incanum (Kendall 1965), Abutilon pedunculare (Hayward 1947), Abutilon sonorae (Bailowitz and Brock 1991), Abutilon wrightii (Kendall 1961b), Pseudabutilon lozani (Hayward 1947), Sphaeralcea angustifolia (Kendall 1965), Wissadula amplissima (Kendall 1965), Wissadula holosericea (Kendall 1960). Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 976, 25 eggs (ROK) LARVAE: TEXAS: BEXAR CO.: Lot 976, 8 larvae (instar 5) (ROK). COMAL CO.: Lot 1002, 6 larvae ( instars 3, 4, 5) (FSCA). PUPAE: TEXAS: BEXAR CO.: Lot 976, 2 pupae (ROK). 75. Systasea zampa (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 20.5-22mm, A4 transverse width 4. 3-4. 4mm; preserved specimens pale; setae simple, to 0.5mm long on A4 dorsum; spiracles pale. HEAD: light brown; transverse width 2. 9-3. 2mm; sculpturing rough; mandibles with teeth; setae simple, to 0.4mm long at apex, a few ventral setae to 0.7mm; stemmata subegual (Figure 39Q) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 30 crochets, uniordinal, arranged in a mesal penellipse. Host Plants MALVACEAE: Abutilon abutiloides (Bailowitz and Brock 1991), Abutilon incanum (Bailowitz and Brock 1991), Abutilon parishii (Bailowitz and Brock 1991), Abutilon reventum (Bailowitz and Brock 1991), Abutilon

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154 sonorae (Bailowitz and Brock 1991), Herissantia crispa (Bailowitz and Brock 1991), Hibiscus denudatus (Emmel and Emmel 1973, possibly). Specimens Examined LARVAE: CALIFORNIA: SAN DIEGO CO.: Lot 580, 1 larva (instar 5) (GRB) TEXAS: PRESIDIO CO.: Lot 977, 1 larva (instar 5) (ROK) 76. Achlyodes thraso (Hiibner) Diagnosis EGG: height 0.8-lmm, width 0.6-0.7iam, 14-16 ribs, pale yellow. LAST INSTAR LARVA: BODY: Figure 20A, length 15-32. 5mm, A4 transverse width 3. 5-6. 8mm; green with numerous tiny yellow spots, heart line dark, subdorsal line moderately wide, bright yellow; some setae with slightly expanded tips, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 25F, light brown; transverse width 3. 8-5. 2mm; sculpturing pebbly; mandibles with a few small teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.7mm; stemma 1 very large, 6 large (Figure 39R) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 190 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 48A) PUPA: Figures 58G, 65G, and 86E, length 17-23mm, A3 transverse width 4. 5-6. 5mm; pale green, process on cap dark, anterior pronotum dark, pilifers with a black spot; thoracic spiracle guard a small black knob; setae simple, to 0.4mm long on head; head with a short, peglike process on cap; antennae extend slightly caudad of tip

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155 of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.7mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 104F and 1151) Other Descriptions Panton 1897 (egg, larva, pupa), Wolcott 1923 (larva, pupa), Hayward 1927b (larva, pupa), Moss 1949 (larva, pupa) Host Plants ROSACEAE: Prunus occidental is (MCM observation). RUTACEAE: Citrus aurantifolia (Bruner et al 1945), Citrus aurantium (Comstock 1944), Citrus limon (Martorell 1945), Citrus paradisi (Wolcott 1923), Citrus sinensis (Wolcott 1951), Fagara hyemale (Hayward 1927b), Fagara naranjillo (Hayward 1941), Fagara rhoifolia (Moss 1949), Zanthoxylum fagara (Kendall 1965), Zanthoxylum martinicensis (Bruner et al 1945), Zanthoxylum monophyllum (Wolcott 1923), Zanthoxylum pterota (ROK collection). Specimens Examined EGGS: DOMINICAN REPUBLIC: PUERTO PLATA PROV.: Lot 40, 1 egg (MCM). USA: TEXAS: SAN PATRICIO CO.: Lot 898, 24 eggs (ROK). LARVAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 5, 2 larvae ( instar 5) (MCM). PUERTO PLATA PROV.: Lot 40, 2 larvae (instars 3, 5) (MCM); Lot 41, 1 larva (instar 5) (MCM); Lot 856, 2 larvae (instar 5) (USNM); Lot 864, 2 larvae (instar 5) (USNM) JAMAICA: Lot 857, 1 larva (instar 5) (USNM). MEXICO: LEON GUANAJUATA: Lot 853, 1 larva (instar 5) (USNM). PUERTO RICO: Lot 847, 2 larvae (instar 2) (USNM); Lot 848, 1 larva

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156 (instar 3) (USNM); Lot 852, 1 larva (instar 5) (USNM); Lot 855, 1 larva (instar 5) (USNM); Lot 858, 1 larva (instar 4) (USNM); Lot 1082, 2 larvae (instar 4) (DHH) TEXAS: SAN PATRICIO CO.: Lot 898, 3 2 larvae ( instars 1, 3, 4, 5) (ROK). WEST INDIES: Lot 859, 2 larvae (instar 5) (USNM). PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV. : Lot 5, 1 pupa (MCM). PUERTO PLATA PROV.: Lot 40, 1 pupa (MCM) ; Lot 41, 1 pupa (MCM). PUERTO RICO: Lot 854, 1 pupa (USNM). TEXAS: SAN PATRICIO CO.: Lot 898, 9 pupae (ROK). 77. Grais stigmatica (Mabille) Nothing is known of the biology of this neotropical species. 78. Timochares ruptifasciatus (Plotz) Diagnosis LAST INSTAR LARVA: BODY: length 22.5mm, A4 transverse width 4.8mm; preserved specimens pale; setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 25G, pale with a black patch at vertex and a black U-shaped mark on face, lateral line brown; transverse width 4.8mm; sculpturing rough; mandibles with teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemma 5 and 6 small (Figure 40A) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 100 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 48B) PUPAL EXUVIUM: cremaster

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157 blunt with numerous hooked setae in a cluster at the tip (Figures 104G and 115J) Other Descriptions Comstock 1953 (egg, larva, pupa). Host Plants MALPIGHIACEAE: (Comstock 1953), Malpighia glabra (Kendall and Rickard 1976). Specimens Examined LARVAE: TEXAS: HIDALGO CO.: Lot 982, 1 larva ( instar 5) (ROK). 79. Chiomara asychis (Stoll) Diagnosis LARVAL EXUVIUM: stemmata subegual (Figure 40B) PUPA: Figures 65H and 86F, length 10.5mm, A3 transverse width 3.2mm; pale; thoracic spiracle guard distinct; setae simple, to 0.1mm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 104H and 115K) Other Descriptions Moss 1949 (larva, pupa). Host Plants MALPIGHIACEAE: (Moss 1949), Aspicarpa hirtella (Bailowitz and Brock 1991, possibly), Gaudichaudia pentrandra (Kendall 1976), Janusia gracilis (Kendall and Rickard 1976, possibly), Malpighia glabra (Kendall and Rickard 1976), Thryallis angustifolia (Kendall and Rickard 1976, possibly). Specimens Examined PUPAE: TEXAS: HIDALGO CO.: Lot 923, 1 pupa ( ROK )

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158 80. Gesta gesta (Herrich-Schaf f er) Diagnosis LAST INSTAR LARVA: BODY: Figure 19G, length 819mm, A4 transverse width 1.8-4. 4mm; grayish green, heart line dark subdorsal line a chain of yellow spots on T3-A8; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 25H, tan with six black spots on face, posterior black, area around mouth and stemmata black, occasionally mostly black; transverse width 2.63.2mm; sculpturing rough; mandibles without teeth, or at most a few shallow lobes; setae simple, < 0.1mm long at apex, a few ventral setae to 0.3mm; stemmata subegual (Figure 40C) ; postocciput moderately wide. THORAX: legs brown; shield indistinct. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 48C). PUPA: Figures 66A and 86G, length 13-15mm, A3 transverse width 3.7-4mm; pale green; thoracic spiracle guard a black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.1mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 1041 and 115L) Other Descriptions Comstock and Vazguez 1960 (larva, pupa), Comstock and Kendall 1967 (egg, larva, pupa). Host Plants FABACEAE: Cassia occidentalis (da Costa Lima 1936), Cassia species (Comstock and Vazguez 1960),

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159 Indigofera leptosepala (Kendall 1965, in lab), Indigofera lindheimeriana (Kendall 1965), Indigofera suffruticosa (Moss 1949), Indigofera tinctoria (Bruner et al. 1945). Specimens Examined LARVAE: COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 241, 1 larva (instar 5) (MCM) ; Lot 242, 1 larva (instar 5) (MCM); Lot 243, 4 larvae ( instars 4, 5) (MCM). DOMINICAN REPUBLIC: LA VEGA PROV. : Lot 240, 8 larvae (instar 5) (MCM). TEXAS: ARANSAS CO.: Lot 561, 6 larvae (instars 2, 3, 4, 5) ( JRH) ; Lot 562, 2 larvae (instars 2, 5) (JRH). PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 240, 6 pupae (MCM) 81. Ephyriades brunneus (Herrich-Schaf fer) Diagnosis LAST INSTAR LARVA: BODY: length 14.5-21mm, A4 transverse width 3. 6-4. 8mm; green with numerous tiny yellow spots, heart line dark outlined with yellow, subdorsal line very narrow, yellow; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan to pale. HEAD: Figure 26A, dark brown with orange patches at apex and laterally, orange eye patches present; transverse width 3. 2-3. 6mm; sculpturing rough to pebbly at apex; mandibles without teeth, or at most a few shallow lobes; setae simple, to 0.1mm long at apex, a few ventral setae to 1mm; stemmata subegual (Figure 40D); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 130 crochets, irregularly triordinal, arranged in a circle, suranal plate rounded, unmarked (Figure 48D) PUPA:

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160 Figures 66B and 86H, length 16-18mm, A3 transverse width 4. 8-5. 4mm; pale green; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 104 J and 116A) Other Descriptions Tamburo and Butcher 1955 (larva), Brown and Heineman 1972 (egg, larva, pupa), Minno and Emmel 1993 (larva, pupa) Host Plants MALPIGHIACEAE: Byrsonima lucida (Kimball 1965), Malpighia glabra (Butcher 1951). ROSACEAE: Prunus species (Brown and Heineman 1972) [erroneous]. Specimens Examined LARVAE: FLORIDA: DADE CO.: Lot 779, 2 larvae ( instars 4, 5) (USNM). MONROE CO.: Lot 149, 18 larvae (instar 5) (MCM) ; Lot 150, 8 larvae (instars 2, 4, 5) (MCM). PUPAE: FLORIDA: MONROE CO.: Lot 149, 8 pupae (MCM); Lot 150, 3 pupae (MCM). 82. Erynnis icelus (Scudder and Burgess) Diagnosis LAST INSTAR LARVA: BODY: length 12-2 3. 5mm, A4 transverse width 4. 1-5. lmm; green with numerous tiny white spots, heart line dark, subdorsal line narrow, yellow; setae simple, to 0.2mm long on A4 dorsum; spiracles tan to pale. HEAD: Figure 26B, brown, blackish around mouth, stemmata, vertex, and posterior; transverse width 3. 3-3. 5mm; sculpturing rough; mandibles without teeth; setae simple, to

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161 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subequal (Figure 40E) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 75 crochets, irregularly triordinal arranged in a circle; suranal plate rounded, unmarked (Figure 48E). PUPA: Figures 66C and 87A, length 16-18mm, A3 transverse width 4. 8-5. 4mm; pale brown; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal and middle leg tips subequal; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1mm long, tapering, blunt with numerous hooked setae in a cluster at the tip (Figures 104K and 116B) Other Descriptions Mead 1875 (egg), Edwards 1885a (larva, pupa), French 1886 (egg), Fletcher 1888 (egg), Scudder 1889a (egg, larva, pupa), Harrington 1892 (larva), Emmel et al. 1992 (larva, pupa). Host Plants BETULACEAE: Betula lenta (DHH collection), Betula populifolia (Shapiro 1966), Corylus species (Scudder 1889a, b, questionable; Burns 1964a, erroneous). FABACEAE: Baptisia tinctoria (Folsom 1896) [erroneous], Baptisia species (Scudder 1889a, b) [erroneous], Robinia pseudoacacia (Schaffner and Griswold 1934). FAGACEAE: Quercus ilicifolia (Craighead 1950) [erroneous], Quercus rubra (Craighead 1950) [erroneous], Quercus velutina (Craighead 1950) [erroneous]. HAMAMELIDACEAE: Hamamelis virginiana (Scudder 1889a, b, possibly) [questionable]. SALICACEAE:

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162 Populus balsamifera (Burns 1964a), Populus grandidentata (Shapiro and Shapiro 1973), Populus nigra (Shapiro 1966), Populus tremuloides (Edwards 1885a), Salix bebbiana (McCabe 1991), Salix cordata (Fletcher 1888). Specimens Examined LARVAE: COLORADO: TELLER CO.: Lot 158, 5 larvae (instar 5) (MCM) INDIANA: BROWN CO.: Lot 209, 1 larva (instar 5) (MCM). NEW YORK: TOMPKINS CO.: Lot 547, 1 larva (instar 5) (TLM) VIRGINIA: GILES CO.: Lot 1242, 1 larva (instar 3) (DHH) PUPAE: COLORADO: TELLER CO.: Lot 158, 2 pupae (MCM). 83. Erynnis brizo (Boisduval and Leconte) Diagnosis LAST INSTAR LARVA: BODY: length 15-28mm, A4 transverse width 3.6-6mm; green with numerous tiny yellow spots, heart line dark, subdorsal line narrow, yellow; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan to pale. HEAD: Figure 26C, brown, yellowish eye patches present; transverse width 3. 6-4. 4mm; sculpturing rough; mandibles without teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.6mm; stemma 5 slightly small, 6 slightly large (Figure 40F) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a near circle. PUPA: Figures 66D and 87B, length 18.5mm, A3 transverse width 4.7mm; pale brown; thoracic spiracle guard a small black knob; setae simple, < 0.1mm long on head; head rounded; antennal tip lies slightly

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163 cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen long; cremaster to 1.3mm long, tapering, blunt with numerous hooked setae in a cluster at the tip (Figures 104L and 116C) Other Descriptions Scudder 1889a (egg, larva, pupa), Scott 1986 (larva, pupa). Host Plants BORAGINACEAE: Cynoglossum morrisoni (Edwards 1885a) [erroneous], Cynoglossum officiale (Duffy and Garland 1978) [erroneous], FABACEAE: Baptisia tinctoria (Harris 1972) [erroneous], Baptisia species (Scudder 1872) [erroneous], Galactia regularis (Scudder 1881) [erroneous], Lespedeza species (Klots 1951, dubious; Duffy and Garland 1978) [erroneous], Sesbania longifolia (Mather and Mather 1958) [erroneous]. FAGACEAE: Castanea dentata (Shapiro 1966), Quercus chapmanii (MCM collection), Quercus dumosa (MacNeill 1975, possibly), Quercus durata (Burns 1964a), Quercus fusiformis (Kendall 1966b, in lab), Quercus gambelii (Burns 1964a), Quercus harvardii (ROK collection), Quercus ilicifolia (Scudder 1889a, b), Quercus inopina (Minno 1992), Quercus laevis (MCM collection), Quercus lobata (Burns 1964a, in lab), Quercus macrocarpa (McCabe and Post 1977), Quercus myrtifolia (Minno 1992), Quercus turbinella (Burns 1964a), Quercus undulata (Burns 1964a). Specimens Examined LARVAE: FLORIDA: HIGHLANDS CO.: Lot 157, 5 larvae ( instars 2, 5) (MCM). LEVY CO.: Lot 154, 2 larvae (instar 5) (MCM). MARION CO.: Lot 155, 4 larvae

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164 (instars 4, 5) (MCM) ; Lot 156, 4 larvae (instars 3, 4). PUTNAM CO.: Lot 709, 1 larva (instar 4) (MCM). TEXAS: MOTLEY CO.: Lot 935, 4 larvae (instars 3, 4, 5) (ROK) PUPAE: FLORIDA: HIGHLANDS CO.: Lot 157, 1 pupa (MCM). 84. Erynnis juvenalis (Fabricius) Diagnosis LAST INSTAR LARVA: BODY: length 172 5mm, A4 transverse width 4.1-6mm; green with numerous tiny white spots, heart line dark, subdorsal line narrow, yellow; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 26D, brown with orange patches at apex and laterally, blackish around the mouth, orangish eye patches present; transverse width 4.1-6mm; sculpturing rough, small spines at apex; mandibles without teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 40G) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 90 crochets, multiordinal ca. 2-3 ranks, arranged in a near circle. PUPA: Figures 66E and 87C, length 16.5-19mm, A3 transverse width 4. 5-5. 8mm; pale green; thoracic spiracle guard a small black knob; setae simple, to 0.3mm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.3mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 104M and 116D)

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165 Other Descriptions Harris 1862 (larva, pupa), Saunders 1869b (larva), Scudder 1881 (larva, pupa), Scudder 1889a (egg, larva, pupa), Scott 1986 (egg, larva, pupa). Host Plants BETULACEAE: Corylus americana (McCabe and Post 1977) [possibly], Corylus species (Saunders 1869b). FABACEAE: Apios americana (French 1886) [erroneous], Baptisia species (Middleton 1881) [erroneous], Galactia regularis (Scudder 1889a, b) [erroneous], Galactia volubilis (Scudder 1889a, b) [erroneous], Glycine species (Harris 1862) [erroneous], Lathyrus species (Harris 1862) [erroneous], Lotus species (Basinger 1926) [erroneous]. FAGACEAE: Fagus americana (Scott 1986, probably), Quercus agrifolia (Burns 1964a, in lab), Quercus alba (Scudder 1889a, b), Quercus arizonica (Burns 1964a), Quercus chapmanii (MCM collection), Quercus emoryi (Burns 1964a, probably; MacNeill 1975), Quercus falcata (MCM collection) Quercus fusiformis (ROK collection), Quercus gambelii (Burns 1964a, possibly), Quercus hypoleucoides (Burns 1964a, probably; MacNeill 1975), Quercus ilicifolia (Scudder 1889a, b), Quercus inopina (Minno 1992), Quercus laevis (MCM collection), Quercus lobata (Burns 1964a, in lab), Quercus macrocarpa (McCabe and Post 1977), Quercus marilandica (Kendall 1966a), Quercus mohriana (Burns 1964a, probably; MacNeill 1975), Quercus muehlenbergii (Klots 1951), Quercus nigra (MacNeill 1975), Quercus phellos (Scudder 1889a, b, probably; Tietz 1952), Quercus prinus (Scudder 1889a, b), Quercus rubra (Craighead

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166 1950), Quercus stellata (Kendall 1966a), Quercus velutina (Craighead 1950) Specimens Examined LARVAE: MEXICO: NUEVO LEON: Lot 937, 1 larva ( instar 5) (ROK) INDIANA: PULASKI CO.: Lot 178, 7 larvae (instar 5) (MCM) ; Lot 179, 2 larvae (instar 3) (MCM); Lot 181, 4 larvae (instar 2) (MCM); Lot 182, 5 larvae (instars 4, 5) (MCM); Lot 183, 7 larvae (instar 4) (MCM); Lot 184, 1 larva (instar 5) (MCM); Lot 185, 2 larvae (instar 5) (MCM); Lot 186, 1 larva (instar 5) (MCM); Lot 559, 1 larva (instar 5) (JRH); Lot 714, 1 larva (instar 4) (MCM); Lot 715, 1 larva (instar 5) (MCM); Lot 782, 1 larva (instar 5) (USNM) PUPAE: MEXICO: NUEVO LEON: Lot 937, 1 pupa (ROK). INDIANA: PULASKI CO.: Lot 180, 6 pupae (MCM). 85. Erynnis telemachus Burns The immature stages of this species are undescribed, and I could not locate specimens for study. Burns (1964a) found larvae on Quercus gambelii and reared these in the laboratory on Quercus lobata. 86. Erynnis propertius (Scudder and Burgess) Diagnosis LAST INSTAR LARVA: BODY: length 32mm, A4 transverse width 6mm; preserved specimen pale; some setae with blunt tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 26E, brown with pale orange patches at apex and laterally, pale orange eye patches present; transverse width 4.4mm; sculpturing pebbly, small spines at apex; mandibles without teeth; setae simple, to 0.1mm long at

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167 apex, a few ventral setae to 0.7mm; stemmata subequal (Figure 40H) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 105 crochets, irregularly triordinal, arranged in a circle. Other Descriptions Emmel and Emmel 1973 (pupa), Pyle 1981 (egg, larva, pupa). Host Plants FAGACEAE: Quercus agrifolia (Burns 1964a), Quercus garryana (Burns 1964a), Quercus virginiana (Tietz 1972) Specimens Examined LARVAE: OREGON: BENTON CO.: Lot 636, 1 larva (instar 5) (TCE). 87. Erynnis meridianus Bell The immature stages of this species are undescribed, and I could not locate specimens for study. Host Plants FAGACEAE: Quercus alba (Burns 1964a, in lab), Quercus arizonica (Burns 1964), Quercus fusiformis (Burns 1964a). 88. Erynnis scudderi (Skinner) Nothing is known of the biology of this species. 89. Erynnis horatius (Scudder and Burgess) Diagnosis EGG: Figure 4C, height 0.9mm, width 0.7-0. 8mm, 13 ribs, yellow to orange. LAST INSTAR LARVA: BODY: Figure 19H, length 12-30. 5mm, A4 transverse width 4-6. 2mm; green with numerous tiny white spots; some setae with blunt tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 26F, brown with orange patches at apex and laterally,

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168 blackish around the mouth and stemmata, orangish eye patches present; transverse width 3. 7-4. 8mm; sculpturing rough to pebbly; mandibles without teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 401); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 105 crochets, multiordinal ca. 2-3 ranks, arranged in a circle or near circle. PUPA: Figures 66F and 87D, length 17-21mm, A3 transverse width 5. 7-6. 3mm; pale green, often with a black patch on the anterior margin of the proboscis; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.4mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 104N and 116E) Other Descriptions Scudder 1889a (larva, pupa). Host Plants FABACEAE: Wisteria frutescens (Scudder 1889a, b) [erroneous]. FAGACEAE: Quercus alba (Burns 1964a, in lab), Quercus chapmanii (MCM collection), Quercus falcata var. pagodaefolia (Scott 1986), Quercus fusiformis (Burns 1964a), Quercus gambelii (Burns 1964a, associated with; Kendall 1965, in lab), Quercus geminata (MCM collection), Quercus hemisphaerica (Kendall 1965), Quercus ilicifolia (Burns 1964a), Quercus inopina (Minno 1992), Quercus laurifolia (Kendall 1965), Quercus laevis (MCM collection),

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169 Quercus marilandica (Kendall 1965), Quercus muehlenbergii (Shapiro 1966), Quercus myrtifolia (Minno 1992), Quercus nigra (Ross and Lambremont 1963), Quercus phellos (Kendall 1965), Quercus rubra (Scott 1986), Quercus shumardii (Burns 1964a), Quercus stellata (Burns 1964a), Quercus stellata var. margaretta (MCM collection), Quercus texana (Burns 1964a), Quercus velutina (Opler and Krizek 1984), Quercus virginiana (Burns 1964a). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1533, 2 eggs (MCM). LARVAE: FLORIDA: ALACHUA CO.: Lot 168, 3 larvae ( instar 5) (MCM); Lot 169, 1 larva (instar 5) (MCM); Lot 170, 1 larva (instar 5) (MCM); Lot 172, 2 larvae (instar 5) (MCM); Lot 173, 2 larvae (instar 5) (MCM); Lot 175, 2 larvae (instar 5) (MCM); Lot 176, 3 larvae (instars 3, 5) (MCM); Lot 177, 1 larva (instar 4) (MCM); Lot 711, 1 larva (instar 5) (MCM). BROWARD CO.: Lot 164, 1 larva (instar 5) (MCM); Lot 165, 1 larva (instar 4) (MCM); Lot 166, 2 larvae (instar 5) (MCM). HIGHLANDS CO.: Lot 6, 2 larvae (instar 5) (MCM); Lot 712, 1 larva (instar 5) (MCM). MARION CO.: Lot 171, 1 larva (instar 4) (MCM). PUTNAM CO.: Lot 710, 2 larvae (instar 5) (MCM); Lot 713, 1 larva (instar 5) (MCM); Lot 1508, 1 larva (instar 4) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 168, 1 pupa (MCM); Lot 173, 1 pupa (MCM). BROWARD CO.: Lot 163, 1 pupa (MCM); Lot 166, 1 pupa (MCM)

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170 90. Erynnis tristis (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 152 9mm, A4 transverse width 3. 3-5. 8mm; green, heart line dark, subdorsal line very narrow, pale yellow; some setae with blunt or slightly expanded tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 27A, pale brown with pale orange patches at apex and laterally, posterior dark brown, small pale orange eye patches present; transverse width 3.84.8mm; sculpturing rough, small spines at apex; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.8mm; stemmata subegual (Figure 40 J) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 90 crochets, multiordinal ca. 2-3 ranks, arranged in a circle or near circle. PUPA: Figures 66G and 87E, length 14.5-16mm, A3 transverse width 4. 4-4. 7mm; preserved specimen pale; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.5mm long, slightly constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 1040 and 116F). Other Descriptions Corns tock and Dammers 19 3 2a (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants FABACEAE: Lotus species (Wright 1905) [erroneous]. FAGACEAE: Quercus agrifolia (Burns 1964a),

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171 Quercus devia (Burns 1964a, associated with), Quercus douglasii (Burns 1964a), Quercus dumosa (Johnson 1982), Quercus emoryi (Burns 1964a, associated with; ROK collection), Quercus grisea (Burns 1964a, associated with; Bailowitz and Brock 1991), Quercus hemisphaerica (MCM collection, in lab), Quercus idonea (Burns 1964a, associated with), Quercus lobata (Burns 1964a), Quercus suber (Comstock and Dammers 1932a), Quercus wislizenii (Shapiro 1974b). Specimens Examined LARVAE: CALIFORNIA: SACRAMENTO CO.: Lot 187, 6 larvae ( instar 5) (MCM). SAN DIEGO CO.: Lot 589, 2 larvae (instar 5) (GRB). TEXAS: BREWSTER CO.: Lot 938, 34 larvae ( instars 2, 3, 4, 5) (ROK). PUPAE: TEXAS: BREWSTER CO.: Lot 938, 3 pupae (ROK). 91. Erynnis martialis (Scudder) Diagnosis LAST INSTAR LARVA: BODY: length 19-23.5ram, A4 transverse width 4. 7-4. 9mm; green, heart lined dark, subdorsal line narrow, pale yellow; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 27B, dark brown with pale orange patches at apex and laterally, posterior black, pale orange eye patches present; transverse width 3. 3-3. 7mm; sculpturing rough, small knobs at apex; mandibles with teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 40K) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 60 crochets, multiordinal, ca. 2-3 ranks, arranged in a circle. PUPA:

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172 Figures 66H and 87F, length 16mm, A3 transverse width 5.3mm; green; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; creraaster to 1.3mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 104P and 116G) Other Descriptions Scudder 1889a (egg, larva, pupa). Host Plants AMARANTHACEAE : Amaranthus retro flexus (Tietz 1952) [erroneous], Amaranthus species (Holland 1898) [erroneous]. FABACEAE: Amorpha species (Klots 1951) [erroneous], Baptisia species (Forbes 1906, possibly) [erroneous], Indigofera caroliniana (Scudder 1889a, b) [erroneous]. HAEMODORACEAE : Lachnanthes caroliniana (Scudder 1889a, b, possibly) [erroneous]. RHAMNACEAE: Ceanothus americanus (Scudder 1889a, b), Ceanothus cordulatus (Scott 1986), Ceanothus fendleri (Scott and Scott 1980), Ceanothus herbaceus var. pubescens (Opler and Krizek 1984, associated with), Ceanothus ovatus (Scott 1986). ROSACEAE: Adenostoma fasciculatum (Tietz 1952) [erroneous]. ULMACEAE: Trema micrantha (Tietz 1952) [erroneous]. Specimens Examined LARVAE: INDIANA: JASPER CO.: Lot 992, 2 larvae ( instar 5)(MCM). PENNSYLVANIA: CHESTER CO.: Lot 1241, 1 larva (instar 5) (DHH). PUPAE: INDIANA: JASPER CO.: Lot 992, 1 pupa (MCM)

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173 92. Erynnis pacuvius (Lintner) Diagnosis LAST INSTAR LARVA: BODY: length 30mm, A4 transverse width 6.3mm; preserved specimen pale; some setae with blunt tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 27C, light brown with a darker brown patch across face, posterior dark brown; transverse width 3.3mm; sculpturing rough, small knobs at apex; mandibles with teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 40L) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 70 crochets, multiordinal ca. 2-3 ranks, arranged in a circle or near circle. Other Descriptions Comstock and Dammers 1932b (egg), Pyle 1981 (egg, larva, pupa). Host Plants FAGACEAE: Quercus species (Beutelspacher 1980) [erroneous]. RHAMNACEAE: Ceanothus cordulatus (Burns 1964a), Ceanothus fendleri (Eff 1955), Ceanothus oliganthus (Burns 1964a) Specimens Examined LARVAE: CALIFORNIA: LOS ANGELES CO.: Lot 590, 1 larva (instar 5) (GRB). 93. Erynnis zarucco (Lucas) Diagnosis EGG: height 0.7-0. 8mm, width 0.6-0. 7mm, yellow to orange. LAST INSTAR LARVA: BODY: length 12-30mm, A4 transverse width 2. 9-6. 2mm; green with numerous tiny yellow spots, heart line dark, subdorsal line narrow, pale yellow; some setae with expanded tips, < 0.1mm long on A4 dorsum;

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174 spiracles tan. HEAD: Figure 27D, light brown with orange patches, a dark W-shape on face, posterior black, occasionally mostly black or light brown; transverse width 3. 1-4. lmm; sculpturing rough, small knobs at apex; mandibles with teeth; setae feathery, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 40M) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a circle or near circle. PUPA: Figures 67A and 87G, length 14. 5-19. 5mm, A3 transverse width 4. 4-5. 6mm; pale green; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.3mm long, slightly constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 105A and 116H) Other Descriptions Kendall 1961a (larva), Pyle 1981 (egg, larva, pupa), Scott 1986 (egg, larva, pupa), Minno and Emmel 1993 (egg, larva, pupa). Host Plants AMARANTHACEAE : Amaranthus species (Draudt 1924) [erroneous]. FABACEAE: Baptisia lactea (MCM collection), Baptisia species (Macy and Shepard 1941), Centrosema virginianum (Burns 1964a, probably; Scott 1986), Clitoria mariana (Burns 1964a, probably; Scott 1986), Desmodium species (Shapiro 1966, probably) [erroneous],

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175 Galactia elliottii (Minno 1992), Galactia regularis (Minno 1992), Galactia volubilis (MCM collection), Galactia wrightii (Bailowitz and Brock 1991), Glycine max (DHH collection), Indigofera caroliniana (Minno 1992), Indigofera suffruticosa (MCM collection), Lespedeza hirta (Burns 1964a, probably) Lysiloma bahamensis (USNM collection) [erroneous], Robinia pseudoacacia (Kendall 1961a), Sesbania grandiflora (Bates 1935), Sesbania longifolia (Opler and Krizek 1984), Sesbania macrocarpa (MacNeill 1975), Sesbania punicea (DHH collection, MCM collection, USNM collection), Sesbania vescicaria (Burns 1964a, probably; Scott 1986), Vicia floridana (DHH collection), Wisteria frutescens (Burns 1964a, probably; Scott 1986). Specimens Examined EGGS: FLORIDA: PUTNAM CO.: Lot 1534, 2 eggs (MCM); Lot 1538, 1 egg (MCM). LARVAE: FLORIDA: Lot 1256, 4 larva ( instar 5) (DHH). ALACHUA CO.: Lot 197, 6 larvae (instar 5) (MCM); Lot 718, 3 larvae (instar 5) (MCM); Lot 780, 2 larvae (instar 5) (USNM); Lot 1209, 1 larva (instar 5) (DHH); Lot 1243, 1 larva (instar 5) (DHH); Lot 1244, 2 larvae (instar 5) (DHH); Lot 1247, 1 larva (instar 5) (DHH); Lot 1248, 2 larvae (instar 5) (DHH); Lot 1249, 2 larvae (instar 5) (DHH); Lot 1250, 1 larva (instar 5) (DHH); Lot 1251, 1 larva (instar 5) (DHH); Lot 1253, 1 larva (instar 5) (DHH); Lot 1254; 1 larva (instar 5) (DHH); Lot 1257, 1 larva (instar 5) (DHH); Lot 1258, 1 larva (instar 5) (DHH); Lot 1259, 1 larva (instar 5) (DHH); Lot 1260, 1 larva

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176 (instar 5) (DHH); Lot 1261, 3 larvae (instar 5) (DHH); Lot 1263, 8 larvae ( instars 3, 5) (DHH); Lot 1264, 1 larva (instar 5) (DHH). BAKER CO.: Lot 783, 2 larvae (instar 5) (USNM). DUVAL CO.: Lot 717, 1 larva (instar 5) (MCM) ; Lot 1262, 2 larvae (instar 5) (DHH). BREVARD CO.: Lot 781, 1 larva (instar 5) (USNM). HIGHLANDS CO.: Lot 189, 1 larva (instar 5) (MCM); Lot 190, 1 larva (instar 5) (MCM); Lot 191; 1 larva (instar 5) (MCM); Lot 192, 1 larva (instar 3) (MCM); Lot 193, 1 larva (instar 5) (MCM); Lot 198, 3 larvae (instar 5) (MCM). LAKE CO.: Lot 1252, 1 larva (instar 5) (DHH). LEON CO.: Lot 1245, 1 larva (instar 5) (DHH). LEVY CO.: Lot 196, 2 larvae (instar 5) (MCM); Lot 203, 1 larva (instar 5) (MCM). LIBERTY CO.: Lot 194, 2 larvae (instars 3, 5) (MCM). MARION CO.: Lot 1265, 2 larvae (instar 5) (DHH). MONROE CO.: Lot 205, 5 larvae (instar 5) (MCM); Lot 206, 5 larvae (instar 5) (MCM); Lot 208, 2 larvae (instar 5) (MCM); Lot 719, 1 larva (instar 5) (MCM). OSCEOLA CO.: Lot 200, 1 larva (instar 5) (MCM). PUTNAM CO.: Lot 202, 1 larva (instar 5) (MCM); Lot 716, 1 larva (instar 5) (MCM); Lot 1246, 1 larva (instar 5) (DHH). VOLUSIA CO.: Lot 195, 1 larva (instar 5) (MCM). GEORGIA: HARRIS CO.: Lot 188, 2 larvae (instar 5) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 199, 2 pupae (MCM); Lot 1243, 2 pupae (DHH). BAKER CO.: Lot 783, 1 pupa (USNM); Lot 1256, 3 pupae (DHH). LIBERTY CO.: Lot 194, 1 pupa (MCM). MARION CO.: Lot 201, 1 pupa (MCM). MONROE CO.: Lot 205, 1 pupa (MCM); Lot 206, 1 pupa

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177 (MCM); Lot 207, 1 pupa (MCM) ; Lot 208, 2 pupae (MCM) PUTNAM CO.: Lot 720, 1 pupa (MCM). 94. Erynnis funeralis (Scudder and Burgess) Diagnosis EGG: height 0.7-0. 8mm, width 0.6-0. 7mm, 12-16 ribs. LAST INSTAR LARVA: BODY: length 2 0-3 4mm, A4 transverse width 3.8-6mm; preserved specimens pale, subdorsal line narrow, pale yellow; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 27E, black with orange patches at apex and lateral margin or light brown with a dark W-shape on face, orange eye patches present; transverse width 3.8-6mm; sculpturing rough, small knobs at apex; mandibles with teeth; setae feathery, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 40N); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle or near circle. PUPA: Figures 67B and 87H, length 17-20mm, A3 transverse width 4. 8-6. 2mm; preserved specimens pale; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.4mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 105B and 1161)

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178 Other Descriptions Coquillett 1899 (larva), Coolidge 1924 (egg, larva, pupa), Comstock 1927b (egg, larva, pupa), Comstock 1930b (egg, larva, pupa), Kendall 1961a (larva), Emmel and Emmel 1973 (egg, larva, pupa), Hogue 1974 (larva). Host Plants FABACEAE: Astragalus species (Brown et al 1992), Cracca edwardsi (Bailowitz and Brock 1991), Geoffroea decorticans (Hayward 1941), Indigofera leptosepala (Kendall 1960), Lotus crassifolius (GRB collection), Lotus grandiflorus (Tietz 1972), Lotus scoparius (Coquillett 1899), Medicago hispida (Field 1938), Medicago sativa (Coolidge 1924), Olneya tesota (Burns 1964a), Robinia neomexicana (Burns 1964a), Sesbania drummondii (Scott 1986), Sesbania macrocarpa (Emmel and Emmel 1973), Vicia leavenworthii (ROK collection), Vicia texana (Kendall 1960). HYDROPHYLLACEAE : Nemophila membranacea (Coolidge 1924) [erroneous]. MALPIGHIACEAE : Malpighia glabra (ROK collection) Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 940, 6 eggs (ROK). LARVAE: USA: NO DATA: Lot 551, 1 larva (instar 5) (NM). CALIFORNIA: SAN BERNARDINO CO.: Lot 591, 1 larva (instar 5). TEXAS: BEXAR CO.: Lot 939, 1 larva (instar 5) (ROK); Lot 940, 2 larvae (instar 5) (ROK). CAMERON CO.: Lot 936, 24 larvae (instars 2, 3, 4, 5) (ROK). PUPAE: TEXAS: BEXAR CO.: Lot 939, 2 pupae (ROK); Lot 940, 3 pupae (ROK). CAMERON CO.: Lot 936, 3 pupae (ROK).

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179 95. Erynnis lucilius (Scudder and Burgess) I could not find specimens of this northeastern species for study. Other Descriptions Lintner 1878 (egg, larva, pupa), Scudder 1889a (egg, larva, pupa), Lindsey 1927 (larva). Host Plants CHENOPODIACEAE : Chenopodium album (Scudder 1889a, b) [erroneous]. RANUNCULACEAE : Aquilegia brevistyla (Klassen et al 1989, possibly), Aquilegia canadensis (Lintner 1872), Aquilegia vulgaris (Shapiro 1974a). SALICACEAE: Populus species (Mead 1875) [erroneous], Salix species (Mead 1875) [erroneous]. 96. Erynnis baptisiae (Forbes) Diagnosis LAST INSTAR LARVA: BODY: length 172 6mm, A4 transverse width 4. 3-5. lmm; green, heart line dark, subdorsal line narrow, pale yellow; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 27F, dark brown or black with pale orange patches on apex and laterally, orange eye patches present, occasionally uniformly black or light brown with a dark W-shape on the face; transverse width 3. 1-3. 7mm; sculpturing rough, small knobs at apex; mandibles with teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.2mm; stemmata subegual (Figure 40O); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 70 crochets, multiordinal ca. 2-3 ranks, arranged in a circle or near circle. PUPA: Figures 67C and 88A, length

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180 13.5-17mm, A3 transverse width 4-5. 2mm; pale green; thoracic spiracle guard a small black knob; setae simple, to 0.1mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.2mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 105C and 116J) Other Descriptions Pyle 1981 (egg, larva, pupa). Host Plants FABACEAE: Amorpha canescens (Ebner 1970), Astragalus canadensis (Scott 1986), Baptisia australis (Opler and Krizek 1984), Baptisia leucantha (Burns 1964a), Baptisia leucophaea var. laevicaulis (Kendall 1965), Baptisia tinctoria (Forbes 1936, associated with; Comstock 1940), Coronilla varia (Wheeler 1974), Crotalaria sagittalis (Opler and Krizek 1984, possibly; Scott 1986), Crotalaria species (Shapiro 1966, possibly), Lupinus perennis (Shapiro 1974a), Sesbania vescicaria (ROK collection), Thermopsis mollis (Scott 1986, in lab), Thermopsis villosa (Opler and Krizek 1984). RANUNCULACEAE : Aquilegia species (Burns 1964a, in lab) Specimens Examined LARVAE: INDIANA: TIPPECANOE CO.: Lot 151, 3 larvae ( instar 5) (MCM) ; Lot 153, 1 larva ( instar 5) (MCM). KENTUCKY: WHITLEY CO.: Lot 993, 2 larvae (instar 5) (MCM). PENNSYLVANIA: MONTGOMERY CO.: Lot 1000, 2 larvae (instar 5) (FSCA). TEXAS: WILSON CO.: Lot 934, 4 larvae (instar 5) (ROK). PUPAE: INDIANA: JASPER CO.: Lot 152, 1

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181 pupa (MCM). KENTUCKY: WHITLEY CO.: Lot 993, 1 pupa (MCM) TEXAS: BASTROP CO.: Lot 568, 1 pupa ( JRH) 97. Erynnis afranius (Lintner) Diagnosis LAST INSTAR LARVA: BODY: length 25mm, A4 transverse width 4.3mm; preserved specimen pale; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 27G, light brown with a dark W-shape on the face; transverse width 2.8mm; sculpturing rough, pebbly at apex; mandibles with teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 40P); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle. Other Descriptions Lindsey 1927 (larva), Comstock and Dammers 1932b (egg?, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa), McCabe and Post 1977 (larva). Host Plants FABACEAE: Astragalus species (Scott and Scott 1980), Lotus purshianus (Comstock and Dammers 1932b), Lotus scoparius (Tilden and Smith 1986), Lupinus argenteus (Scott and Scott 1980), Lupinus species (Lindsey 1927), Thermopsis rhombifolia (Royer 1988, possibly), Trifolium species (Comstock 1927b). RHAMNACEAE: Ceanothus oliganthus (Comstock and Dammers 1932b) [erroneous]. SALICACEAE: Populus species (Comstock 1927b) [erroneous], Salix species (Comstock 1927b) [erroneous].

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182 Specimens Examined LARVAE: CALIFORNIA: SAN DIEGO CO.: Lot 592, 1 larva (instar 5) (GRB) 98. Erynnis persius (Scudder) Diagnosis LAST INSTAR LARVA: BODY: length 16-22. 5mm, A4 transverse width 3. 3-4. 8mm; green with numerous tiny white spots, heart line dark, subdorsal line narrow, pale yellow; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 27H, light brown with a dark W-shape on the face, area around the mouth, stemmata, and posterior dark brown; transverse width 2. 7-3. 2mm; sculpturing rough, small knobs at apex; mandibles with teeth; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 40Q) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a circle or near circle. PUPA: Figures 67D and 88B, length 15-16mm, A3 transverse width 4. 5-4. 8mm; pale green; thoracic spiracle guard a small black knob; setae simple, to 0.2mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.1mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 105D and 116K)

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183 Other Descriptions Scudder 1889a (egg, larva, pupa), Folsum 1896 (egg), Comstock 1927b (egg, larva, pupa), Lindsey 1927 (larva), Emmel et al 1992 (larva). Host Plants FABACEAE: Astragalus flexuosus (Scott 1986), Astragalus bisulcatus (Scott and Scott 1980, probably), Baptisia tinctoria (Tietz 1952), Baptisia species (Wild 1939), Galactia species (Davenport and Dethier 1938), Lespedeza capitata (Scudder 1881), Lotus purshianus (Tietz 1952), Lotus scoparius (Scott 1986, probably), Lupinus argenteus (Scott 1986), Lupinus latifolius (Scott 1986), Lupinus perennis (Opler and Krizek 1984), Oxytropis splendens (Emmel et al. 1992), Thermopsis divaricarpa (Scott 1986), Thermopsis macrophylla (Burns 1964a), Thermopsis pinetorum (Burns 1964a), Trifolium ciliolatum (Tietz 1952). FAGACEAE: Quercus ilicifolia (Davenport and Dethier 1938) [erroneous]. RANUNCULACEAE : Aquilegia canadensis (Saunders 1869b, possibly) [erroneous]. RHAMNACEAE: Ceanothus oliganthus (Tietz 1952) [erroneous]. SALICACEAE: Populus balsamifera (Scudder 1881) [erroneous], Populus grandidentata (Scudder 1889a, b) [erroneous], Populus tremuloides (Scudder 1889a, b) [erroneous], Salix humilis (Scudder 1881) [erroneous], Salix sericea (Shapiro 1966, probably) [erroneous], Salix species (Edwards 1885a) [erroneous] Specimens Examined LARVAE: COLORADO: FREMONT CO. : Lot 159, 4 larvae (instar 5) (MCM); Lot 160, 5 larvae ( instar 5)

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184 (MCM); Lot 161, 5 larvae (instar 5) (MCM) ; Lot 162, 2 larvae (instar 5) (MCM). PUPAE: PENNSYLVANIA: CLINTON CO.: Lot 161, 2 pupae (MCM) 99. Pyrgus centaureae (Rambur) Diagnosis EGG: height 0.8-0. 9mm, width 0.5-0. 8mm, 16-20 ribs, pale green to white. LAST INSTAR LARVA: BODY: length 12-22. 5mm, A4 transverse width 2. 8-3. 8mm; green with numerous tiny white spots, heart line dark, subdorsal and lateral lines narrow, white; some setae with slightly expanded tips, to 0.5mm long on A4 dorsum; spiracles tan. HEAD: Figure 28A, black; transverse width 2. 6-3. lmm; sculpturing rough; mandibles with teeth; setae simple, to 0.4mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 40R) ; postocciput moderately wide. THORAX: legs dark brown; shield broad, tan. ABDOMEN: prolegs each with about 30 crochets, multiordinal, ca. 2-3 ranks, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 48F). Other Descriptions Emmel et aJ 1992 (larva). Host Plants LAMIACEAE: Mentha species (Scudder 1889a, b) [erroneous]. ROSEACEAE: Fragaria virginiana (Opler and Krizek 1984), Fragaria species (Scott and Scott 1980), Potentilla canadensis (Clark and Clark 1951, associated with), Potentilla diversifolia (Scott and Scott 1980).

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185 Specimens Examined EGGS: COLORADO: PARK CO.: Lot 451, 2 eggs (MCM). LARVAE: COLORADO: PARK CO.: Lot 451, 13 larvae (instars 1, 2, 5) (MCM). 100. Pyrgus ruralis (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 24.5mm, A4 transverse width 3.6mm; preserved specimen pale; some setae with slightly expanded tips, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: dark brown; transverse width 2.7mm; sculpturing rough; mandibles with teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.7mm; stemmata subegual (Figure 40S); postocciput moderately wide. THORAX: Tl legs brown, T2-3 legs pale; shield broad, brown. ABDOMEN: prolegs each with about 40 crochets, multiordinal ca. 2-3 ranks, arranged in a mesal penellipse. Other Descriptions Coolidge 1909 (egg). Host Plants MALVACEAE: Sidalcea malvae flora (Garth and Tilden 1986, questionable) [erroneous]. ROSEACEAE: Horkelia bolanderi var. clevelandii (Emmel and Emmel 1973, probably), Horkelia fusca (Lembert 1894), Horkelia tenuiloba (Comstock 1927b), Potentilla ambigens (Stanford 1981), Potentilla drummondii (Emmel and Emmel 1962, probably). Specimens Examined LARVAE: CALIFORNIA: SAN DIEGO CO.: Lot 587, 1 larva ( instar 5) (GRB) 101. Pyrgus xanthus Edwards The immature stages of this species are undescribed, but the larvae feed on roseaceous plants such as Potentilla

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186 ambigens (Scott 1975b, associated with), Potentilla hippiana (Scott and Scott 1980), and Potentilla pulcherrima (Scott 1986) 102. Pyrgus scriptura (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 12. 5-22. 5mm, A4 transverse width 2. 6-4. 2mm; green, heart line dark; some setae with expanded tips, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2-2. 3mm; sculpturing rough; mandibles with teeth; setae slightly feathery, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 40T) ; postocciput moderately wide. THORAX: Tl legs light brown, T2-3 legs tan to pale; shield indistinct or anterior tan, posterior brown. ABDOMEN: prolegs each with about 40 crochets, multiordinal, ca. 2-3 ranks, arranged in a circle. PUPA: Figures 67E and 88C, length ll-14mm, A3 transverse width 3. 9-4. 3mm; dark brown; thoracic spiracle guard distinct; some setae bifurcate, to 0.6mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 0.9mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 105E and 116L) Other Descriptions Emmel et al 1992 (larva, pupa). Host Plants MALVACEAE: Sida hederacea (Davenport and Dethier 1938), Sphaeralcea ambigua (Scott 1986), Sphaeralcea coccinea (Scott 1975b).

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187 S pecimens Examined LARVAE: CALIFORNIA: RIVERSIDE CO. : Lot 586, 2 larvae ( instar 5) (GRB). YOLO CO.: Lot 465, 11 larvae (instar 5) (MCM) PUPAE: CALIFORNIA: YOLO CO.: Lot 465, 7 pupae (MCM). 103. Pyrgus communis (Grote) Diagnosis EGG: height 0.7mm, width 0.6mm, 20 ribs, pale green to white. LAST INSTAR LARVA: BODY: length 12-2 8mm, A4 transverse width 2. 6-4. 8mm; green, heart line dark; some setae with slightly expanded tips, to 0.3mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2.13.1mm; sculpturing rough; mandibles with teeth; setae branching, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 40U) ; postocciput moderately wide. THORAX: Tl-2 legs dark brown, T3 legs tan to pale; shield broad, anterior light brown, posterior dark brown. ABDOMEN: prolegs each with about 50 crochets, multiordinal, ca. 2-3 ranks, arranged in a circle. PUPA: Figures 58H, 67F, and 88D, length 13-17mm, A3 transverse width 3. 9-5. 2mm; green with dark markings; thoracic spiracle guard distinct; some setae bifurcate, to 0.5mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.3mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 105F and 117A)

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188 Other Descriptions Scudder 1889a (egg, larva, pupa), Comstock 1927b (egg, larva, pupa), Dethier 1944b (egg, larva, pupa), Emmel et al 1992 (egg, larva, pupa). Host Plants CHENOPODIACEAE : Chenopodium album (Shapiro 1968; oviposition, but larva refused it). MALVACEAE: Abutilon abutiloides (Comstock 1940), Abutilon theophrasti (Scudder 1889a, b), Althaea officinalis (Shapiro and Shapiro 1973, possibly), Althaea rosea (French 1897), Althaea species (Scudder 1889a, b), Anoda lavaterioides (Hayward 1947), Callirhoe leiocarpa (Kendall 1965), Hibiscus trionum (Comstock 1940), Iliamna species (Field 1938), Malva moschata (Shapiro 1968), Malva neglecta (French 1897), Malva nicaeensis (Coolidge 1908), Malva parviflora (Shapiro 1974b), Malva sylvestris (Shapiro 1974b), Malva verticillata (Klassen et al. 1989), Malva species (Scudder 1889a, b), Malvastrum americanum (ROK collection), Malvastrum coromandelianum (Scott 1986), Malvastrum rotundifolium (Scott 1986), Modiola caroliniana (Scott 1986), Sida filicaulis (Kendall 1965), Sida hederacea (Shapiro 1974b, c), Sida lindheimeri (Kendall 1965), Sida rhombifolia (Kendall 1965), Sida spinosa (Comstock 1940), Sidalcea glaucescens (Emmel and Emmel 1962), Sidalcea neomexicana (Scott and Scott 1980), Sidalcea oregana ssp. spicata (Shapiro et al. 1981), Sphaeralcea ambigua (Austin and Austin 1981), Sphaeralcea angustifolia (Kendall 1965), Sphaeralcea angustifolia ssp. cuspidata (Kendall 1965), Sphaeralcea

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189 coccinea (Scott and Scott 1980), Sphaeralcea digitata (Hayward 1947), Sphaeralcea lindheimeri (Hayward 1947), Sphaeralcea munroana (Scott 1986), Sphaeralcea parvifolia (Scott 1986) Specimens Examined EGGS: OREGON: BENTON CO.: Lot 634, 1 egg (TCE) LARVAE: CALIFORNIA: SACRAMENTO CO.: Lot 15, 3 larvae ( instar 5) (MCM) TOULUMNE CO.: Lot 455, 3 larvae (instars 4, 5) (MCM). YOLO CO.: Lot 453, 7 larvae (instar 5) (MCM); Lot 454, 14 larvae (instars 3, 4, 5) (MCM); Lot 456, 3 larvae (instar 3) (MCM). FLORIDA: ALACHUA CO.: Lot 452, 2 larvae (instar 5) (MCM); Lot 737, 1 larva (instar 5) (MCM); Lot 1283, 1 larva (instar 5) (DHH); Lot 1290, 3 larvae (instars 2, 3) (DHH); Lot 1291, 1 larva (instar 5) (DHH). MISSOURI: BOONE CO.: Lot 1286, 1 larva (instar 5) (DHH). NORTH DAKOTA: CASS CO.: Lot 541, 1 larva (instar 5) (TLM). OREGON: BENTON CO.: Lot 634, 24 larvae (instars 1, 2, 3, 4, 5) (TCE). PUPAE: CALIFORNIA: SACRAMENTO CO.: Lot 15, 2 pupae (MCM). TUOLUMNE CO.: Lot 455, 1 pupa (MCM). YOLO CO.: Lot 453, 6 pupae (MCM). FLORIDA: ALACHUA CO.: Lot 452, 2 pupae (MCM). NORTH DAKOTA: CASS CO.: Lot 541, 2 pupae (TLM). OREGON: BENTON CO.: Lot 634, 3 pupae (TCE) 104. Pyrgus albescens Plotz Diagnosis LAST INSTAR LARVA: BODY: length 27mm, A4 transverse width 4.2mm; preserved specimen pale; some setae with slightly expanded tips, to 0.3mm long on A4 dorsum;

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190 spiracles tan. HEAD: black; transverse width 2. 1-3. lmm; sculpturing rough; mandibles with teeth; setae branching, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 40V); postocciput moderately wide. THORAX: Tl legs brown, T2 legs tan, T3 legs pale; shield broad, anterior light brown, posterior dark brown. ABDOMEN: prolegs each with about 60 crochets, multiordinal ca. 2-3 ranks, arranged in a circle. Other Descriptions See P. communis. Identification of these two species is difficult and their distinction in the literature is not clear. Host Plants MALVACEAE: Abutilon species (Brown et al 1992), Hibiscus species (Brown et al 1992), Malva parviflora (GRB collection), Sida hederacea (Orsak 1978), Sidalcea species (Garth 1950), Sphaeralcea fendleri (Bailowitz and Brock 1991), Sphaeralcea laxa (Bailowitz and Brock 1991) Specimens Examined LARVAE: CALIFORNIA: RIVERSIDE CO.: Lot 588, 1 larva (instar 5) (GRB). 105. Pyrgus oileus (Linnaeus) Diagnosis LAST INSTAR LARVA: BODY: Figure 20C, length 19-24mm, A4 transverse width 3-4. 4mm; green with numerous tiny white spots, heart line dark, subdorsal faint, dark, outlined by pale yellow; some setae with slightly expanded tips, to 0.5mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2. 1-3. lmm; sculpturing rough;

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191 mandibles with teeth; setae feathery, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 40W); postocciput moderately wide. THORAX: Tl legs brown, T2 legs tan, T3 legs pale; shield broad, brown with pale mesodorsal and later spots. ABDOMEN: prolegs each with about 65 crochets, multiordinal ca. 2-3 ranks, arranged in a circle. PUPA: Figures 67G and 88E, length 14-17mm, A3 transverse width 3. 9-4. 5mm; green with dark markings; thoracic spiracle guard distinct; some setae slightly feathery, to 0.8mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.3mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 105G and 117B) Other Descriptions Panton 1897 (egg, larva, pupa), Swainson 1901 (larva, pupa), Dethier 1940a (egg, young larva), Dethier 1942b (larva, pupa), Moss 1949 (larva), Wolcott 1951 (larva, pupa), Minno and Emmel 1993 (egg, larva, pupa) Host Plants MALVACEAE: Abutilon species (Tietz 1952), Althaea rosea (Tietz 1952), Hibiscus trionum (Tietz 1952), Malva neglecta (Tietz 1952), Malvastrum americanum (Tietz 1952), Malvastrum corchorifolium (Minno and Emmel 1993, probably), Malvastrum coromandelianum (Tietz 1972), Sida acuta (Minno and Emmel 1993, probably), Sida antillensis (Wolcott 1923), Sida carpinifolia (Wolcott 1923), Sida

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192 retusa (Grossbeck 1917), Sida rhombifolia (Bottimer 1926), Sida salviaefolia (Kendall 1976), Sida spinosa (Tietz 1952), Sidalcea humilis (Tietz 1952), Sidalcea malvaeflora (Tietz 1972) Specimens Examined DOMINICAN REPUBLIC: LA VEGA PROV. : Lot 461, 1 larva ( instar 5) (MCM) PUERTO PLATA PROV.: Lot 462, 1 larva (instar 5) (MCM). FLORIDA: ALACHUA CO.: Lot 736, 1 larva (instar 5) (MCM); Lot 1284, 1 larva (instar 5) (DHH); Lot 1285, 1 larva (instar 3) (DHH); Lot 1495, 1 larva (instar 5) (DHH). BROWARD CO.: Lot 459, 2 larvae (instar 5) (MCM). PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 461, 1 pupa (MCM). FLORIDA: BROWARD CO.: Lot 459, 4 pupae (MCM) 106. Pyrgus philetas Edwards The immature stages of this species have not been described, but the larvae feed on malvaceous plants such as Sida procumbens and probably Sida spinosa (Bailowitz and Brock 1991) 107. Heliopetes domicella (Erichson) The immature stages of this species have not been described, but the larvae feed on malvaceous plants such as Abutilon incanum and Herissantia crispa (Bailowitz and Brock 1991). 108. Heliopetes ericetorum (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 22-31mm, A4 transverse width 3. 9-6. 4mm; green, heart line dark; some

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193 setae with blunt or slightly expanded tips, to 0.4mm long on A4 dorsum; spiracles tan. HEAD: Figure 28B, black; transverse width 2. 3-3. 4mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.3mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 40X) ; postocciput moderately wide. THORAX: Tl legs dark brown, T2-3 legs tan to pale; shield indistinct. ABDOMEN: prolegs each with about 35 crochets, irregularly biordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 49A) PUPA: Figures 67H and 88F, length 15-20mm, A3 transverse width 4. 4-5. 2mm; light brown with dark spots; thoracic spiracle guard distinct; setae simple, to 0.8mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.5mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 105H and 117C) Other Descriptions Coolidge 1923 (egg, larva, pupa), Comstock 1927b (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants AMARANTHACEAE : Amaranthus graecizans (Coolidge 1923) [erroneous], Amaranthus species (Comstock 1927b) [erroneous]. MALVACEAE: Althaea rosea (Tietz 1972), Althaea species (Coolidge 1923), Eremalche rotundifolia (Garth and Tilden 1986), Hibiscus denudatus (Scott 1986), Iliamna rivularis (Newcomer 1964), Malacothamnus davidsonii

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194 (Coolidge 1923), Malacothamnus fasciculatus (Wright 1905), Malacothamnus fremontii ssp. cercophorum (Scott 1986), Malacothamnus orbiculatus (Coolidge 1923), Malva nicaeensis (Coolidge 1923), Malvastrum exile (Coolidge 1923), Malvastrum rotundifolium (Coolidge 1923), Sphaeralcea ambigua (Coolidge 1923), Sphaeralcea angustifolia ssp. cuspidata (Coolidge 1923), Sphaeralcea grossulariaefolia var. pedata (Austin and Austin 1981), Sphaeralcea munroana (Scott 1986). STERCULIACEAE : (Bailowitz and Brock 1991), Fremontia californica (TCE collection) Specimens Examined LARVAE: CALIFORNIA: CONTRA COSTA CO.: Lot 246, 7 larvae (instar 5) (MCM) LOS ANGELES CO.: Lot 635, 6 larvae ( instars 3, 4, 5) (TCE). RIVERSIDE CO.: Lot 585, 2 larvae (instar 5) (GRB) PUPAE: CALIFORNIA: CONTRA COSTA CO.: Lot 246, 1 pupa (MCM). LOS ANGELES CO.: Lot 635, 2 pupae (TCE) 109. Heliopetes lavianus (Hewitson) Diagnosis EGG: height 1mm, width 0.7mm, with about 35 vertical rows of short spines. LAST INSTAR LARVA: BODY: length 21-25mm, A4 transverse width 3. 8-5. 3mm; preserved specimens pale; some setae with slightly expanded tips, to 0.5mm long on A4 dorsum; spiracles pale. HEAD: black; transverse width 2. 3-3. 4mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.3mm long at apex, a few ventral setae to 0.8mm; stemmata subegual (Figure 40Y); postocciput moderately wide. THORAX: Tl legs brown, T2

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195 legs tan, T3 legs tan to pale; shield broad, light brown with a dark brown border. ABDOMEN: prolegs each with about 50 crochets, irregularly biordinal, arranged in a circle or near circle. PUPA: Figures 68A and 88G, length 16-19mm, A3 transverse width 5. 2-5. 4mm; light brown with dark spots; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen moderately long; cremaster to 1.8mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 1051 and 117D) Host Plants CONVOLVULACEAE : Ipomoea species (da Costa Lima 1936) [erroneous]. MALVACEAE: Abutilon abutiloides (Kendall 1965), Abutilon hypoleucum (Hayward 1947), Abutilon incanum (Kendall 1965, in lab), Abutilon lignosum (Scott 1986), Malvastrum americanum (Kendall 1965), Pseudabutilon lozani (Hayward 1947), Sida filipes (Kendall 1965), Sida hederacea (Garth and Tilden 1986), Sphaeralcea species (Garth and Tilden 1986), Wissadula holosericea (Kendall 1965, in lab) Specimens Examined EGGS: TEXAS: SAN PATRICIO CO.: Lot 943, 6 eggs (ROK) LARVAE: TEXAS: SAN PATRICIO CO.: Lot 560, 3 larvae (instars 4, 5) ( JRH) ; Lot 943, 6 larvae (instars 3, 5) (ROK). PUPAE: TEXAS: SAN PATRICIO CO.: Lot 560, 1 pupa (JRH); Lot 943, 2 pupae (ROK).

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196 110. Heliopetes macaira (Reakirt) Diagnosis EGG: height 0.8mm, width 0.6mm, with about 35 vertical rows of short spines. LAST INSTAR LARVA: BODY: length 9-17. 5mm, A4 transverse width 2. 8-3. 6mm; preserved specimens pale; some setae with slightly expanded tips, to 0.7mm long on A4 dorsum; spiracles pale. HEAD: brown, posterior dark brown; transverse width 2. 8-3. 2mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.4mm long at apex, a few ventral setae to 0.8mm; stemmata subequal (Figure 40Z); postocciput moderately wide. THORAX: legs tan; shield indistinct or broad, tan. ABDOMEN: prolegs each with about 30 crochets, irregularly biordinal, arranged in a circle or near circle. Host Plants MALVACEAE: Malvaviscus drummondii (Kendall 1965) Specimens Examined EGGS: TEXAS: SAN PATRICIO CO.: Lot 944, 6 eggs (ROK) LARVAE: TEXAS: SAN PATRICIO CO.: Lot 944, 10 larvae ( instars 1, 2, 5) (ROK). 111. Heliopetes arsalte (Linnaeus) Diagnosis LAST INSTAR LARVA: BODY: length 30-30. 5mm, A4 transverse width 4. 3-4. 4mm; green, heart line dark; some setae with slightly expanded tips, to 0.9mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2.93mm; sculpturing rough; mandibles with teeth; setae slightly feathery, to 0.3mm long at apex, a few ventral setae to 0.8mm; stemmata subequal (Figure 40AA); postocciput

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197 moderately wide. THORAX: legs tan; shield indistinct or broad, tan. ABDOMEN: prolegs each with about 50 crochets, irregularly biordinal, arranged in a circle or near circle. Host Plants MALVACEAE: Sida species (MCM collection). Specimens Examined LARVAE: COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 244, 1 larva (instar 5) (MCM); Lot 245, 2 larvae (instars 4, 5) (MCM) 112. Celotes nessus (Edwards) Diagnosis EGG: height 0.8mm, width 0.5-0. 6mm, with 36-38 vertical rows of short spines. LAST INSTAR LARVA: BODY: length 13. 5-2 3. 5mm, A4 transverse width 3. 5-4. 4mm; preserved specimens pale; some setae with slightly expanded tips, to 0.4mm long on A4 dorsum; spiracles pale. HEAD: Figure 28C, black; transverse width 2. 6-3. lmm; sculpturing rough; mandibles with some small, rounded teeth; setae feathery, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 40BB); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 30 crochets, irregularly biordinal, arranged in mesal penellipse; suranal plate rounded, unmarked (Figure 49B). PUPA: Figures 68B and 88H, length 12-15mm, A3 transverse width 3. 8-4. 6mm; light brown with dark spots; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.1mm long, blunt with numerous

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198 hooked setae in a cluster at the tip (Figures 105J and 117E) Host Plants MALVACEAE: Abutilon incanum (Kendall 1960), Althaea rosea (Kendall 1965), Sida filipes (Kendall 1965), Sphaeralcea angustifolia var. lobata (Kendall 1965), Wissadula amplissima (Kendall 1965), Wissadula holosericea (Burns 1964b). STERCULIACEAE : Ayenia compacta (Bailowitz and Brock 1991) Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 920, 5 eggs (ROK) LARVAE: TEXAS: BEXAR CO.: Lot 920, 8 larvae (instar 5) (ROK). PUPAE: TEXAS: BEXAR CO.: Lot 920, 6 pupae (ROK). 113. Celotes limpia Burns Diagnosis EGG: height 1.1mm, width 0.6-0. 7mm, with about 37-45 vertical rows of short spines. LAST INSTAR LARVA: BODY: length 17-18mm, A4 transverse width 3.8-4mm; preserved specimens pale; some setae with slightly expanded tips, to 0.4mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2. 7-2. 8mm; sculpturing rough; mandibles with some small, rounded teeth; setae feathery, to 0.2mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 40CC); postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 30 crochets, irregularly biordinal, arranged in mesal penellipse. PUPA: length 12-15mm, A3 transverse width 3. 8-4. 6mm; light brown with dark spots; thoracic

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199 spiracle guard distinct; setae simple, to 0.4mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 1.1mm long, blunt with numerous hooked setae in a cluster at the tip. Host Plants MALVACEAE: Abutilon incanum (Burns 1964b), Abutilon malacum (Burns 1964b), Althaea rosea (Burns 1964b, in lab), Malvastrum americanum (Burns 1964b, in lab), Sphaeralcea angustifolia var. lobata (Burns 1964b), Wissadula holosericea (Burns 1964b). Specimens Examined EGGS: TEXAS: JEFF DAVIS CO.: Lot 919, 5 eggs (ROK) LARVAE: TEXAS: JEFF DAVIS CO.: Lot 919, 3 larvae (instar 5) (ROK). 114. Pholisora catullus (Fabricius) Diagnosis LAST INSTAR LARVA: BODY: Figure 2 OB, length 12-21. 5mm, A4 transverse width 2. 4-4. 9mm; green with numerous tiny white spots, heart line dark, subdorsal narrow, dark, outlined with pale yellow; some setae with slightly expanded tips, to 0.3mm long on A4 dorsum; spiracles tan. HEAD: Figure 28D, black; transverse width 2. 4-3. lmm; sculpturing rough; mandibles with teeth; setae feathery, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 40DD); postocciput moderately wide. THORAX: legs pale to tan; shield moderately wide, a dark brown band between the annuli. ABDOMEN: prolegs each with about 45 crochets, irregularly

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200 biordinal, arranged in mesal penellipse; suranal plate rounded, unmarked (Figure 49C) PUPA: Figures 68C and 89A, length 12-14mm, A3 transverse width 3. 8-4. 7mm; light brown; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; head rounded; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 0.7mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 105K and 117F). Other Descriptions Scudder 1889a (egg, larva, pupa), Edwards 1885b (egg, larva, pupa), Comstock 1927b (egg, larva, pupa), Comstock and Dammers 1935 (egg), Emmel and Emmel 1973 (egg, larva, pupa), Capman 1990 (egg, larva). Host Plants AMARANTHACEAE : Amaranthus albus (Scudder 1889a, b) Amaranthus caudatus (Kendall 1965), Amaranthus graecizans (Klots 1951), Amaranthus hybridus (Kendall 1965), Amarahthus retroflexus (Kendall 1965), Amaranthus spinosus (Kendall 1965), Amaranthus species (Scudder 1872), Celosia argentea var. cristata (Shapiro 1968). ASTERACEAE: Ambrosia species (Scudder 1881) [erroneous], CAPRIFOLIACEAE: Symphoricarpos rivularis (Tietz 1952) [erroneous], Symphoricarpos species (Comstock 1927b) [erroneous]. CHENOPODIACEAE : Atriplex rosea (Shapiro 1974b), Chenopodium album (Scudder 1872), Chenopodium ambrosioides (Kendall 1965), Chenopodium berlandieri (Kendall 1965), Chenopodium foliosum (Scott 1986),

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201 Chenopodium murale (Shapiro 1974b), Chenopodium paganum (Shapiro 1968). LAMIACEAE: Marrubium species (Shapiro 1966) [erroneous], Mentha species (Shapiro 1966, possibly) [erroneous], Monarda punctata ( Scudder 1872) [erroneous], Origanum vulgare (Edwards 1885b) [erroneous]. MALVACEAE: Malva neglecta (Newcomer 1964) [erroneous]. POACEAE: Saccharum officinarum (Bodkin 1913) [erroneous], Zea mays (Mather and Mather 1958) [erroneous]. Specimens Examined LARVAE: CALIFORNIA: SAN BERNARDINO CO.: Lot 581, 1 larva (instar 5) (GRB). YOLO CO.: Lot 368, 3 larvae (instars 3, 5) (MCM); Lot 370, 3 larvae (instar 5) (MCM); Lot 371, 2 larvae (instar 5) (MCM). FLORIDA: ALACHUA CO.: Lot 1266, 1 larva (instar 5) (DHH) ILLINOIS: Lot 616, 7 larvae (instars 4, 5) (SP). INDIANA: TIPPECANOE CO.: Lot 367, 2 larvae (instars 3, 5) (MCM). KANSAS: DOUGLAS CO.: Lot 637, 1 larva (instar 5) (TCE). MARYLAND: PRINCE GEORGE'S CO.: Lot 1, 4 larvae (instars 3, 4, 5) (MCM); Lot 369, 1 larva (instar 5) (MCM). MISSOURI: BOONE CO.: Lot 1267, 1 larva (instar 5) (DHH); Lot 1269, 1 larva (instar 5) (DHH). Lot 1270, 3 larvae (instar 5) (DHH). NORTH DAKOTA: CASS CO.: Lot 539, 2 larvae (instar 5) (TLM) SOUTH CAROLINA: NEWBERRY CO.: Lot 1271, 3 larvae (instar 1) (DHH). TEXAS: WICHITA CO.: Lot 366, 5 larvae (instar 5) (MCM). PUPAE: CALIFORNIA: YOLO CO.: Lot 368, 8 pupa (MCM). ILLINOIS: Lot 616, 1 pupa (SP) TEXAS: WICHITA CO.: Lot 366, 1 pupa (MCM).

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202 115. Pholisora mejicana (Reakirt) I could not locate immatures of this southwestern species for study, but Scott (1986) has briefly described the egg, larva, and pupa. Host Plants AMARANTHACEAE : Amaranthus graecizans (Stanford 1981), Amaranthus retroflexus (Scott and Scott 1980). CHENOPODIACEAE: Chenopodium album (Scott and Scott 1980, in lab), Chenopodium species (Scott and Scott 1980). 116. Pholisora libya (Scudder) I could not locate immatures of this southwestern species for study, but Comstock and Dammers (1932b) and Emmel and Emmel (1973) have described the immature stages. Host Plants CHENOPODIACEAE: Atriplex canescens (Comstock and Dammers 1932b; McCabe and Post 1977), Atriplex conferti folia (Scott 1986, associated with). 117. Pholisora alpheus (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 12-28. 5mm, A4 transverse width 2 .6-4 .9mm; preserved specimens pale; some setae with slightly expanded tips, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2.23mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.3mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 41A) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 40 crochets, irregularly biordinal, arranged in mesal penellipse. PUPA: Figures 68D

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203 and 89B, length 12mm, A3 transverse width 3.8mm; light brown with dark spots; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the wing tips; abdomen short; cremaster to 0.7mm long, constricted, blunt with numerous hooked setae in a cluster at the tip (Figures 105L and 117G). Host Plants CHENOPODIACEAE: Atriplex argentea var. expansa (Davenport and Dethier 1938), Atriplex canescens (MacNeill 1970). Specimens Examined EGGS: TEXAS: BREWSTER CO.: Lot 963, 1 egg (ROK) LARVAE: TEXAS: BREWSTER CO.: Lot 963, 1 larva (instar 5) (ROK). CAMERON CO.: Lot 558, 6 larvae (instars 3, 4, 5) ( JRH) CALIFORNIA: KERN CO.: Lot 593, 2 larvae (instar 5) (GRB) 118. Pholisora gracielae MacNeill Diagnosis LAST INSTAR LARVA: BODY: length 21mm, A4 transverse width 3.9mm; preserved specimens pale; some setae with slightly expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: black; transverse width 2.3mm; sculpturing rough; mandibles with teeth; setae feathery, to 0.3mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41B) ; postocciput moderately wide. THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 35 crochets, irregularly triordinal, arranged in mesal penellipse.

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204 Other Descriptions Comstock 1929 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants CHENOPODIACEAE : Atriplex argentea var. expansa (Comstock 1929), Atriplex canescens (Scott 1986, associated with), Atriplex lentiformis (MacNeill 1970), Chenopodium species (Scott 1986). Specimens Examined LARVAE: CALIFORNIA: SAN BERNARDINO CO.: Lot 1498, 1 larva ( instar 5) (GRB). Subfamily Heteropterinae Too few specimens were available to provide an adeguate diagnosis of this subfamily. 119. Carterocephalus palaemon (Pallas) No preserved eggs, mature larvae, or pupae of this boreal species were available for study. Other Descriptions Fletcher 1888, 1889 (egg, larva), Frohawk 1892 (egg, larva, pupa). Host Plants POACEAE: Agropyron repens (Fletcher 1889, in lab), Bromus asper (Frowawk 1892, in lab), Bromus ciliatus var. asper (Tietz 1972), Calamagrostis purpurascens (MacNeill 1975), Digitaria sanguinalis (Fletcher 1889, in lab), Echinochloa crusgalli (Fletcher 1889, in lab), Panicum species (Forbes I960), Poa pratensis (Fletcher 1888, in lab) Specimens Examined LARVAE: CALIFORNIA: SIERRA CO.: Lot 123, 6 larvae (instar 1) (MCM)

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205 120. Piruna pirus (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 12-25mm, A4 transverse width 3 .4-3 .8mm; preserved specimens pale; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 28E, pale with brown medial and lateral stripes, front with a few brown stripes, postocciput dark brown, transverse width 2. 3-2. 5mm; sculpturing rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata 5 and 6 relatively small (Figure 41C) ; paraclypeal hooks present. THORAX: legs pale; shield a narrow dark brown band between the annuli. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 49D) Host Plants POACEAE: Lolium species (ROK collection, in lab), Poa species (Scott and Scott 1980, probably). Specimens Examined LARVAE: MEXICO: NUEVO LEON: Lot 965, 4 larvae (instar 5) (ROK). 121. Piruna polingii (Barnes) Nothing is known of the biology of this southwestern species, except that the grass, Dactylis glomerata, may be a host (Bailowitz and Brock 1991). 122. Piruna microstictus (Godman) Nothing is known of the biology of this southwestern species.

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206 123. Piruna haferniki H. A. Freeman Nothing is known of the biology of this southwestern species. Subfamily Hesperiinae Diagnosis EGG: with polygonal sculpturing, usually pale green or white, occasionally pink, gray, or brownish, sometimes with a red ring, height 0.4-1. 2mm, width 0.61.6mm. LAST INSTAR LARVA: BODY: length 9.8-63mm, A4 transverse width 1.6-9. 5mm; setae usually simple, < 0.11.3mm long; spiracles pale to dark, Tl and A8 spiracles largest. HEAD: pale, dark, or patterned; transverse width 1.5-4. 4mm; sculpturing rough to pitted; mandibles without teeth, hesperiine type articulation; labial-submental complex relatively small; setae simple, < 0.1-1. 8mm long; stemmata usually subegual, 5 or 6 sometimes small; postocciput narrow. THORAX: prothorax about the same size as head or smaller; legs pale to black; shield narrow to broad; ventral prothoracic gland present. ABDOMEN: A4 proleg with 50-230 crochets, more or less triordinal, arranged in a circle, a near circle, or a mesal penellipse, posterior prolegs always with a mesal penellipse; suranal plate usually rounded, sometimes pointed, occasionally with dark markings; anal comb well developed; wax glands sometimes present in the form of longitudinal or transverse patches or spots on the ventral side of Al, A3-6, or A7-8. PUPA: length 12. 5-43. 5mm, A3 transverse width 2. 1-8. 5mm;

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207 green or brown; thoracic spiracle guard distinct; setae < 0.1-0. 8mm; pilifers separated; head usually rounded, occasionally with a long pointed process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5 to beyond the body; abdomen moderately long or long; cremaster bluntly pointed or rounded, hooked setae sometimes absent; lenticles present on the prothorax, dorsal abdomen, and proleg scars. 124. Synapte malitiosa (Herrich-Schaf f er) Diagnosis LAST INSTAR LARVA: BODY: length 20. 5-21. 5mm, A4 transverse width 3-3. 6mm; green frosted with tiny white spots, heart line, subdorsal, and lateral lines dark green (unfrosted); setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 28F, rounded, pale, medial line dark brown along adfrontals, light brown at vertex, lateral line dark brown, front with a few dark brown stripes, postocciput black; transverse width 2. 2-2. 3mm; sculpturing rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemmata subegual (Figure 41D) THORAX: legs pale or tan; shield a very narrow dark brown band between the annul i. ABDOMEN: pro legs with about 130 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 49E) ; wax glands broad, transverse patches posterior of prolegs on A3-A6 (Figure 15C). PUPA: Figures 59F, 68E, and 89C, length 1616.5mm, A3 transverse width 3-4. 8mm; cream-colored; setae

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208 simple, to 0.6mm long on head; thoracic spiracle guard distinct; pilifers separated; head with a short, mesal process on cap; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the cremaster; abdomen moderately long; cremaster to 1.8mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 105M and 117H); lenticles present on the proleg scars. Host Plants POACEAE: Panicum maximum (Kendall 1976), Paspalum species (Scott 1986). Specimens Examined LARVAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 470, 1 larva ( instar 5) (MCM) ; Lot 471, 2 larvae (instars 4, 5) (MCM). PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 471, 4 pupae (MCM). 125. Synapte salenus (Mabille) Nothing is known of the biology of this neotropical species. 126. Corticea corticea (Plotz) The immature stages of this common neotropical species are undescribed and I could not locate specimens for study. Host Plants POACEAE: Saccharum officinarum (Moss 1949). 127. Callimormus saturnus (Herrich-Schaf fer) Nothing is known of the biology of this species from tropical America. 128. Vidius perigenes (Godman) Diagnosis LAST INSTAR LARVA: BODY: length 26mm, A4 transverse width 3.2mm; preserved specimen pale, frosted

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209 with tiny white spots, heart, subdorsal, and lateral lines unfrosted; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 28G, pale, medial and epicranial lines convergent at midadfrontals brown, lateral line and postocciput dark brown, frontal sclerite with a few brown lines; transverse width 2.6mm; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemma 6 small (Figure 41E) THORAX: legs pale to tan; shield a narrow black line between the annuli. ABDOMEN: prolegs each with about 130 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked (Figure 49F) Host Plants POACAEAE: Spartina spartinae (ROK collection), Stenotaphrum secundatum (Kendall 1966a, in lab). Specimens Examined LARVAE: TEXAS: CAMERON CO.: Lot 989, 4 larvae ( instars 3, 4, 5) (ROK). 129. Monca tyrtaeus (Plotz) Nothing is known of the biology of this neotropical species. 130. Nastra lherminier (Latreille) Diagnosis LAST INSTAR LARVA: BODY: length 2 2-2 6mm, A4 transverse width 2. 8-3. 3mm; green with a yellowish cast, heart line dark green, subdorsal line narrow, faint, yellowish; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 28H, pale, medial and

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210 epicranial lines separate, reddish, lateral line reddish (These markings often fade on preserved specimens.); transverse width 1.8-2. lmm; sculpturing rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 6 small (Figure 41F) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked (Figure 49G) PUPA: Figures 60A, 68F, and 89D, length 20.5mm, A3 transverse width 3. 3-3. 4mm; pale green, heart line darker green, faintly outlined with white, subdorsal and lateral lines outlined with faint white bands; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers nearly touching; head with a long pointed process on cap; antennal tip lies cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.9mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 105N and 1171); lenticles present on the proleg scars. Other Descriptions Laurent 1908 (egg, young larva). Host Plants POACEAE: Andropogon scoparius (Shapiro 1966), Andropogon species (MCM collection). Specimens Examined LARVAE: GEORGIA: HARRIS CO.: Lot 330, 4 larvae ( instar 5) (MCM). MARYLAND: PRINCE GEORGE'S CO.: Lot 331, 4 larvae (instars 3, 4, 5) (MCM). PUPAE: GEORGIA: HARRIS CO.: Lot 330, 1 pupa (MCM). MARYLAND: PRINCE GEORGE'S CO.: Lot 331, 2 pupae (MCM).

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211 131. Nastra julia (H. A. Freeman) Diagnosis LAST INSTAR LARVA: BODY: length 2 62 7mm, A4 transverse width 3. 3-3. 5mm; preserved specimens pale; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: pale, medial and epicranial lines separate, reddish, lateral line reddish; transverse width 1.9-2. 3mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 6 small (Figure 41G) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs with about 80 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked. PUPA: Figures 68G and 89E, length 19mm, A3 transverse width 3.6mm; pale; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies cephalad of tip of middle leg; proboscis extending into A8; abdomen long; cremaster to 1.8mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 1050 and 117J); lenticles indistinct. Host Plants POACEAE: Cynodon dactylon (Scott 1986), Sorghum sudanense (USNM collection), Stenotaphrum secundatum (Kendall 1960, in lab). Specimens Examined LARVAE: TEXAS: BEXAR CO.: Lot 951, 1 larva (instar 5) (ROK) MAVERICK CO.: Lot 774, 1 larva (instar 5) (USNM). PUPAE: TEXAS: BEXAR CO.: Lot 951, 1 pupa ( ROK )

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212 132. Nastra neamathla (Skinner and R. C. Williams) Diagnosis LAST INSTAR LARVA: BODY: length 13-26. 5mm, A4 transverse width 2. 3-3. 5mm; green with a yellowish cast, heart line darker green, subdorsal line narrow, faint, yellowish; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: pale, medial and epicranial lines separate, reddish, lateral line reddish; transverse width 22.4mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.1mm; stemma 6 small (Figure 41H) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked. PUPA: Figures 68H and 89F, length 19-19. 5mm, A3 transverse width 3.2mm; pale green, heart line darker green, faintly outlined with white; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies cephalad of tip of middle leg; proboscis extending to the cremaster; abdomen long; cremaster to 1.8mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 105P and 117K) ; lenticles indistinct. Other Descriptions Minno and Emmel 1993 (larva, pupa). Host Plants POACEAE: Andropogon virginicus var. glaucus (MCM observation, probably for N. neamathla) Andropogon species (Minno and Emmel 1993), Sorghum halepense (ROK collection)

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213 Specimens Examined LARVAE: TEXAS: LEON CO.: Lot 952, 1 larva (instar 5) (ROK) Nastra species, probably N. neamathla: FLORIDA: ALACHUA CO.: Lot 1067, 1 larva (instar 4) (DHH). DUVAL CO.: Lot 334, 2 larvae (instar 3) (MCM); Lot 337, 2 larvae (instar 5) (MCM) ; Lot 701, 1 larva (instar 5) (MCM); Lot 703, 1 larva (instar 5) (MCM). PUTNAM CO.: Lot 332, 6 larvae (instar 5) (MCM); Lot 338, 1 larva (instar 4) (MCM). ST. JOHNS CO.: Lot 335, 1 larva (instar 5) (MCM). VOLUSIA CO.: Lot 336, 1 larva (instar 5) (MCM); Lot 702, 1 larva (instar 5) (MCM); Lot 1066, 1 larva (instar 5) (DHH). PUPAE: Nastra species, probably N. neamathla: FLORIDA: DUVAL CO.: Lot 333, 1 pupa (MCM). PUTNAM CO.: Lot 332, 2 pupae (MCM). 133. Cymaenes tripunctus (Herrich-Schaf fer) Diagnosis LAST INSTAR LARVA: BODY: length 192 9mm, A4 transverse width 2. 53. 8mm; green, lightly frosted with white, heart line darker green, lateral line narrow, green outlined by faint white lines; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 29A, pale, medial and epicranial lines broadly convergent along the adfrontals, dark brown, front mostly pale, lateral line and postocciput dark brown; transverse width 2-2. 4mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.3mm; stemma 6 very small (Figure 411). THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 100 crochets, irregularly triordinal, arranged in a near circle;

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214 suranal plate rounded, unmarked (Figure 50A) PUPA: Figures 59E, 69A, and 89G, length 15.5-21mm, A3 transverse width 2. 5-3. 6mm; pale green, heart line darker green outlined with white; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers nearly touching; head with a long pointed process on cap; antennal tip lies slightly cephalad of tip of middle leg; proboscis extending to the cremaster; abdomen long; cremaster to 1.8mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 106A and 117L) ; lenticles present on the prothorax and proleg scars. Other Descriptions Dethier 1939b, 1942d (egg, larva, pupa), Peterson 1962 (larva), Minno and Emmel 1993 (larva, pupa ) Host Plants POACEAE: Bambusa vulgaris (Bruner et al 1945), Brachiaria mutica (MCM collection), Digitaria sanguinalis (Minno and Emmel 1993), Panicum maximum (Comstock 1944), Paspalum setaceum (MCM collection), Saccharum officinarum (Dethier 1939b, in lab), Tripsacum dactyloides (MCM observation) Specimens Examined LARVAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 134, 3 larvae (instar 5) (MCM); Lot 136, 6 larvae (instars 2, 5) (MCM). PUERTO PLATA PROV.: Lot 135, 4 larvae (instars 4, 5) (MCM). SANTIAGO PROV.: Lot 133, 1 larva (instar 5) (MCM). FLORIDA: BROWARD CO.: Lot 129, 1 larva (instar 5) (MCM); Lot 131, 3 larvae (instar 5) (MCM);

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215 Lot 132, 1 larva ( instar 5). MONROE CO.: Lot 128, 8 larvae (instar 5) (MCM); Lot 130, 4 larvae (instar 5) (MCM); Lot 665, 1 larva (instar 5) (MCM). PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 136, 3 pupae (MCM). PUERTO PLATA PROV. : Lot 135, 1 pupa (MCM). USA: FLORIDA: BROWARD CO.: Lot 132, 1 pupa (MCM). MONROE CO.: Lot 128, 4 pupae (MCM); Lot 130, 5 pupae (MCM); Lot 666, 1 pupa (MCM). 134. Cymaenes odilia (Burmeister) The immature stages of this tropical species are undescribed, and I could not find specimens for study. Host Plants POACEAE: Panicum maximum (Kendall 1976), Paspalum species (Scott 1986). 135. Lerema accius (J. E. Smith) Diagnosis EGG: height 1.2-1. 3mm, width 0.7-0. 9mm, polygonal sculpturing, pale green. LAST INSTAR LARVA: BODY: Figure 21E, length 15.5-35mm, A4 transverse width 34.9mm; green, frosted with white, heart, subdorsal, and later lines darker green outlined with white; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 29B, pale, medial line dark brown becoming light brown at vertex, epicranial line brown, convergent with medial line along the adfrontals, front with a few dark brown lines, lateral line and postocciput dark brown; transverse width 2. 5-3. lmm; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41J) THORAX: legs pale; shield narrow, a dark brown band between

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216 the annul i. ABDOMEN: prolegs each with about 130 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked (Figure 50B) ; wax glands forming a ventrolateral patch on A7-8 (Figure 15B) PUPA: Figures 69B and 90A, length 21-28mm, A3 transverse width 3. 8-5. 2mm; pale green, heart and lateral lines darker green bounded by white; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies cephalad of tip of middle leg; proboscis extending nearly to distal tip of cremaster; abdomen long; cremaster to 2.5mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 106B and 118A) ; lenticles present on the prothorax and proleg scars. Other Descriptions Edwards and Chapman 1879 (larva, pupa), Scudder 1889a (egg, larva, pupa), Ainslie 1922 (egg, larva, pupa), Minno and Emmel 1993 (larva, pupa). Host Plants FABACEAE: Wisteria frutescens (Scudder 1889a, b) [erroneous]. POACEAE: Andropogon species (Pyle 1981), Arundinaria gigantea (MCM collection), Arundo donax (MCM collection), Bambusa species (DHH collection), Brachiaria mutica (MCM collection, DHH collection), Chasmanthium latifolium (MCM observation) Chasmanthium sessiliflorum (MCM collection) Cynodon dactylon (MCM collection, in lab), Echinochloa crusgalli (MCM collection), Echinochloa muriacata (MCM collection), Echinochloa

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217 poiretiana (Hayward 1941), Echinochloa species (Pyle 1981), Erianthus alopecuroides (Edwards and Chapman 1879), Erianthus giganteus (MCM collection), Erianthus species (Davenport and Dethier 1938), Heteropogon melanocarpus (MCM observation), Imperata cylindrica (MCM observation), Oryza sativa (Ainslie 1922), Oryzopsis species (Tietz 1952), Panicum capillare (Ainslie 1922, probably), Panicum clandestinum (MCM observation), Panicum dichotomiflorum (MCM observation), Panicum gymnocarpum (MCM collection), Panicum hallii var. filipes (ROK collection), Panicum hemitomon (MCM collection), Panicum laxiflorum (MCM observation), Panicum maximum (MCM collection) Panicum rigidulum (MCM collection), Paspalidium geminatum (MCM observation), Paspalum ciliatifolium (Scott 1986), Paspalum dilitatum (MCM collection), Paspalum floridanum (MCM collection), Paspalum langei (MCM observation) Paspalum pubiflorum (MCM collection), Paspalum repens (DHH collection), Paspalum setaceum (Lenczewski 1980), Pennisetum ciliare (Lenczewski 1980), Pennisetum purpureum (MCM observation), Phragmites australis (ROK collection) Phyllostachys aurea (MCM collection), Saccharum officinarum (MCM collection), Setaria grisebachii (MCM collection), Setaria macrosperma (Minno and Emmel 1993), Sorghum halepense (Ainslie 1922), Stenotaphrum secundatum (Kendall 1960), Tridens flavus (MCM collection), Tripsacum dactyloides (MCM collection), Zea mays (Scudder 1872), Zizaniopsis miliacea (MCM collection).

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218 S pecimens Examined EGGS: FLORIDA: ALACHUA CO. : Lot 1514, 8 eggs (MCM) LARVAE: FLORIDA: ALACHUA CO.: Lot 275, 1 larva ( instar 5) (MCM); Lot 279, 1 larva (instar 5) (MCM); Lot 284, 1 larva (instar 5) (MCM); Lot 285, 1 larva (instar 5) (MCM); Lot 289, 1 larva (instar 4) (MCM); Lot 290, 4 larvae (instar 1, 5) (MCM); Lot 293, 1 larva (instar 5) (MCM); Lot 294, 3 larvae ( instars 3, 5) (MCM); Lot 307, 2 larvae (instar 5) (MCM); Lot 608, 1 larva (instar 4) (SP); Lot 683, 1 larva (instar 5) (MCM); Lot 685, 1 larva (instar 2) (MCM); Lot 686, 1 larva (instar 5) (MCM); Lot 687, 1 larva (instar 2) (MCM); Lot 688, 1 larva (instar 5) (MCM); Lot 689, 1 larva (instar 2) (MCM); Lot 690, 1 larva (instar 5) (MCM); Lot 691, 1 larva (instar 3) (MCM); Lot 1050, 1 larva (instar 5) (DHH); Lot 1055, 1 larva (instar 5) (DHH); Lot 1057, 1 larva (instar 3) (DHH); Lot 1058, 1 larva (instar 3) (DHH); Lot 1059, 1 larva (instar 5) (DHH); Lot 1060, 3 larvae (instars 3, 5) (DHH); Lot 1514, 4 larvae (instar 1) (MCM). BAKER CO.: Lot 775, 1 larva (instar 5) (USNM). BREVARD CO.: Lot 1054, 1 larva (instar 5) (DHH). BROWARD CO.: Lot 272, 1 larva (instar 5) (MCM); Lot 274, 2 larvae (instar 3) (MCM); Lot 278, 1 larva (instar 5) (MCM); Lot 286, 1 larva (instar 5) (MCM). COLLIER CO.: Lot 12, 3 larvae (instar 5) (MCM); Lot 277, 2 larvae (instar 5) (MCM). DADE CO.: Lot 291, 3 larvae (instar 5) (MCM); Lot 776, 1 larva (instar 5) (USNM); Lot 1048, 1 larva (instar 4) (DHH). HIGHLANDS CO.: Lot 684, 1 larva (instar 5) (MCM). LEVY

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219 CO.: Lot 271, 1 larva (instar 5) (MCM) MADISON CO.: Lot 1047, 1 larva (instar 5) (DHH). ORANGE CO.: Lot 1049, 1 larva (instar 5) (DHH). PALM BEACH CO.: Lot 1051, 3 larvae (instar 5) (DHH); Lot 1052, 1 larva (instar 5) (DHH). PUTNAM CO.: Lot 1056, 1 larva (instar 3) (DHH). SARASOTA CO.: Lot 287, 1 larva (instar 5) (MCM). VOLUSIA CO.: Lot 288, 1 larva (instar 5) (MCM). MARYLAND: PRINCE GEORGE'S CO.: Lot 301, 4 larvae (instar 5) (MCM). NORTH CAROLINA: CRAVEN CO.: Lot 303, 1 larva (instar 3) (MCM). SOUTH CAROLINA: COLLETON CO.: Lot 298, 1 larva (instar 5) (MCM). GEORGETOWN CO.: Lot 299, 2 larvae (instar 5) (MCM). TEXAS: BEXAR CO.: Lot 295, 3 larvae (instar 5) (MCM); Lot 296, 4 larvae (instar 5) (MCM); Lot 297, 1 larva (instar 5) (MCM). SAN PATRICIO CO.: Lot 557, 6 larvae (instars 2, 5) ( JRH) PUPAE: FLORIDA: ALACHUA CO.: Lot 279, 1 pupa (MCM); Lot 284, 1 pupa (MCM); Lot 285, 1 pupa (MCM); Lot 292, 2 pupae (MCM); Lot 294, 1 pupa (MCM); Lot 302, 1 pupa (MCM). BROWARD CO.: Lot 273, 1 pupa (MCM). COLLIER CO.: Lot 12, 2 pupae (MCM). LAKE CO.: Lot 280, 1 pupa (MCM); Lot 281, 6 pupae (MCM). LEVY CO.: Lot 271, 1 pupae (MCM). PALM BEACH CO.: Lot 1053, 3 pupae (DHH). NORTH CAROLINA: COLUMBUS CO.: Lot 300, 2 pupae (MCM). SOUTH CAROLINA: COLLETON CO.: Lot 298, 1 pupa (MCM). GEORGETOWN CO.: Lot 299, 1 pupa (MCM). TEXAS: BEXAR CO.: Lot 296, 2 pupae (MCM).

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220 136. Lerema liris Evans Diagnosis EGG: height 1.2mm, width 0.7mm, polygonal sculpturing. LAST INSTAR LARVA: BODY: length 20-3 3mm, A4 transverse width 2. 8-4. 6mm; preserved specimens pale; setae simple, to 0.2mra long on A4 dorsum; spiracles tan. HEAD: pale, medial and epicranial lines convergent along the adfrontals, brown, front with a few dark brown lines, lateral line dark brown, posterior of head brown; transverse width 2. 5-2. 6mm; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 4 IK) THORAX: legs pale; shield narrow, a dark brown band between the annul i. ABDOMEN: prolegs each with about 130 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked. PUPA: Figures 69C and 90B, length 2024mm, A3 transverse width 3. 6-4. 4mm; preserved specimens pale; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching to nearly touching; head with a long pointed process on cap; antennal tip lies cephalad of tip of middle leg; proboscis extending to tip of the cremaster; abdomen moderately long; cremaster to 2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 106C and 118B) ; lenticles present on the prothorax and proleg scars. Host Plants POACEAE: Bambusa vulgaris (Kendall and McGuire 1975), Saccharum officinarum (Kendall and McGuire 1975)

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221 Specimens Examined LARVAE: MEXICO: TAMAULIPAS: Lot 948, 4 larvae ( instars 3, 4, 5) (ROK) ; Lot 949, 3 larvae ( instars 3, 5) (ROK). PUPAE: MEXICO: TAMAULIPAS: Lot 948, 2 pupae (ROK); Lot 949, 2 pupae (ROK). 137. Perichares philetes (Gmelen) Diagnosis LAST INSTAR LARVA: BODY: length 27.5-41mm, A4 transverse width 5. 1-7. 2mm; pale green, heart line darker green bounded by white; setae simple, to 0.9mm long on A4 dorsum; spiracles tan. HEAD: Figure 29C, pale green, unmarked; transverse width 3. 4-4. 3mm; setae simple, to 0.8mm long at apex, a few ventral setae to 1.1mm; stemmata subegual (Figure 41M) THORAX: legs dark brown; shield indistinct. ABDOMEN: prolegs each with about 210 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 50C); wax glands forming a ventral, transverse patch on Al (Figure 15G) PUPA: Figures 60B, 69D, and 90C, length 30-42. 5mm, A3 transverse width 5. 2-8. 2mm; pale green, heart line darker green bounded by pale yellow; thoracic spiracle guard indistinct; setae simple, to 0.4mm long on head; pilifers separated; head with a long, slightly downward pointing process on the cap; antennal and middle leg tips subegual; proboscis extending beyond the cremaster; abdomen long; cremaster to 4.2mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 106D and 118C); lenticles absent.

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222 Other Descriptions Panton 1897 (egg, larva, pupa), Wolcott 1922 (larva, pupa), Dethier 1942d (larva), Otero 1943 (larva), Moss 1949 (larva, pupa). Host Plants ARECACEAE: Desmonicus species (Moss 1949) [guestionable] Hyospathe elegans (Moss 1949) [questionable]. POACEAE: Arundo donax (Bruner et al 1945), Bambusa vulgaris (Wolcott 1941, Kendall and McGuire 1975), Brachiaria mutica (Bates 1935), Oryza sativa (Otero 1943), Panicum maximum (Kaye 1926), Saccharum officinarum (Wolcott 1921), Zea mays (Kaye 1926). Specimens Examined LARVAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 359, 1 larva ( instar 5) (MCM); Lot 361, 4 larvae (instars 2, 3, 5) (MCM). PUERTO PLATA PROV.: Lot 362, 6 (instar 2, 3, 5) (MCM). HONDURAS: Lot 614, 1 larva (instar 4) (SP). PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 359, 2 pupae (MCM). PUERTO PLATA PROV.: Lot 360, 1 pupa (MCM); Lot 362, 1 pupa (MCM). 138. Rhinthon osca (Plotz) Nothing is known of the biology of this neotropical species. 139. Decinea percosius (Godman) The immature stages of this tropical species are undescribed, and I could not find specimens for study. Host Plants POACEAE: Cynodon dactylon (Kendall and Rickard 1976, in lab), Lolium perenne (Kendall and Rickard 1976, in lab), Sorghum halepense (Kendall and Rickard 1976,

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223 in lab), Stenotaphrum secundatum (Kendall and Rickard 1976, in lab) 140. Conga chydaea (Butler) Nothing is known of the biology of this species from tropical America. 141. Ancyloxypha numitor (Fabricius) Diagnosis EGG: height 0.6-0. 7mm, width 0.4-0. 5mm, polygonal sculpturing, pale green with a red ring. LAST INSTAR LARVA: BODY: length 14.5-23mm, A4 transverse width 2-2. 8mm; green frosted with white, heart line darker green; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 29D, pale, medial and epicranial lines convergent along the adfrontals, brown, front with a few brown lines, lateral line and posterior of head brown; transverse width 1.5-1. 9mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 6 small (Figure 41L) THORAX: legs pale to tan; anterior of shield white, posterior black. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 50D) PUPA: Figures 69E and 90D, length 12.5-16mm, A3 transverse width 2. 1-2. 7mm; cream-colored, head and prothorax black; thoracic spiracle guard absent; setae simple, < 0.1 long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long with a transverse row of short

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224 spines on the dorsum of A5-8, two short ventrolateral rows on A5-7, short spines on proleg scars of A5-6; cremaster to 1.1mm long, broadly rounded with a small downward pointing spine at tip, with numerous hooked setae in a row along distal margin (Figures 106E and 118D); lenticles present on the prothorax, dorsum of abdomen, and proleg scars of A7-8. Other Descriptions Harris 1862 (pupa), Edwards 1884 (egg), Scudder 1889a (egg, larva, pupa), Dethier 1938b (larva, pupa ) Host Plants FABACEAE: Lespedeza capitata (Scudder 1869) [erroneous], POACEAE: Agrostis hiemalis (ROK collection), Leersia hexandra (MCM collection), Leersia oryzoides (Shapiro and Shapiro 1973), Oryza sativa (Ross and Lambremont 1963), Panicum species (Pyle 1981, associated with: Scott 1986), Phalaris arundinacea (Layberry et al 1982, possibly), Poa species (Shapiro 1966), Setaria species (Beutenmiiller 1889), Spartina species (Field 1938), Zea mays (Tietz 1952), Zizaniopsis miliacea (Kendall 1966b). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1522, 10 eggs (MCM). LARVAE: FLORIDA: ALACHUA CO.: Lot 1522, 1 larva ( instar 1) (MCM). BROWARD CO.: Lot 55, 1 larva (instar 5) (MCM); Lot 56, 1 larva (instar 5) (MCM); Lot 57, 7 larvae (instar 5) (MCM). TEXAS: SAN SABA CO.: Lot 901, 3 larvae ( instars 4, 5) (ROK). PUPA: FLORIDA: BROWARD CO.: Lot 56, 1 pupa (MCM); Lot 57, 5 pupae (MCM).

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225 142. Ancyloxypha arene (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 12-17mm, A4 transverse width 1.6-2. lmm; preserved specimens pale; setae simple, to 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 29E, pale, epicranial line wide and convergent with medial line along the adfrontals, black, front with a few black lines, lateral line black, posterior of head dark brown; transverse width 1.5-1. 7mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 6 small (Figure 41N) THORAX: legs pale to tan; anterior of shield white, posterior black. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with two black longitudinal lines (Figure 50E) PUPA: Figures 69F and 90E, length 1313.5mm, A3 transverse width 2. 3 -2. 6mm; preserved specimens pale, head and prothorax blackish; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long with a transverse row of short spines on the dorsum of A6-8, two short ventrolateral rows on A5-7, short spines on proleg scars of A5-6; cremaster to 1.1mm long, broadly rounded with a small downward pointing spine at tip, numerous hooked setae in a row along distal margin (Figures 106F and 118E); lenticles present on the prothorax and dorsum of abdomen.

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226 Host Plants POACEAE: Echinochloa crusgalli (Bailowitz and Brock 1991), Polypogon simiverticillata (Tilden and Smith 1986), Polypogon viridis (Bailowitz and Brock 1991). Specimens Examined LARVAE: TEXAS: KINNEY CO.: Lot 900, 5 larvae (instar 5) (ROK). PUPAE: TEXAS: KINNEY CO.: Lot 900, 2 pupae (ROK) 143. Oarisma poweshiek (Parker) I could not find immature specimens of this midwestern species for study. Willard (1892) and McAlpine (1973) described the egg, larva, and pupa. Host Plants CYPERACEAE: Carex species (McCabe and Post 1977), Eleocharis elliptica (Opler and Krizek 1984). POACEAE: Poa species (McCabe and Post 1977, in lab). 144. Oarisma garita (Reakirt) I could not find immature specimens of this western species for study, but Gibson (1910) gave a description of the egg and larva. Host Plants POACEAE: Blepharoneuron tricholepis (Scott and Scott 1980), Bouteloua gracilis (Scott 1986), Poa agassizensis (Scott 1986), Poa pratensis (Gibson 1910, in lab), Sitanion hystrix (Scott and Scott 1980), Stipa columbiana (Scott and Scott 1980). 145. Oarisma edwardsii (Barnes) Nothing is known of the biology of this western species.

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227 146. Copaeodes aurantiacus (Hewitson) Diagnosis EGG: height 0.9-lmm, width 0.5-0. 6mm, polygonal sculpturing. LAST INSTAR LARVA: BODY: length 152 3. 5mm, A4 transverse width 2-2. 6mm; preserved specimens pale, subdorsal and lateral lines outlined with white; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 29F, pale, unmarked, with two adjacent, forward pointing processes at apex; transverse width 1.7-1. 9mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.1mm; stemma 6 small (Figure 410). THORAX: legs pale to tan; shield indistinct. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate pointed, unmarked (Figure 50F) PUPA: Figures 69G and 90F, length 16-19mm, A3 transverse width 2.1-3mm; preserved specimens pale, heart, subdorsal, and lateral lines outlined with white; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.4mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 106G and 118F) ; lenticles absent. Other D escriptions Comstock 1929 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants POACEAE: Bouteloua curtipendula (Bailowitz and Brock 1991), Cynodon dactylon (Wright 1905), Digitaria

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228 sanguinalis (Tietz 1972), Distichlis spicata (Gunder 1930), Leptochloa dubia (Bailowitz and Brock 1991). Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 931, 5 eggs (ROK) LARVAE: ARIZONA: SANTA CRUZ CO.: Lot 788 (probably), 1 larva ( instar 5) (USNM) TEXAS: BEXAR CO.: Lot 931, 11 larvae (instar 5) (ROK). PUPAE: TEXAS: BEXAR CO.: Lot 931, 8 pupae (ROK). 147. Copaeodes minimus (Edwards) Diagnosis EGG: Figure 3H, height 0.7-0. 8mm, width 0.40.5mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: length 14-19mm, A4 transverse width 2. 1-3. lmm; green, heart line darker green outlined with white, lateral line yellow; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 29G, green, unmarked, rounded; transverse width 1.7-1. 8mm; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 1 and 6 small (Figure 41P) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 50 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate pointed, unmarked (Figure 50G) PUPA: Figures 70A and 90G, length 15-16mm, A3 transverse width 2. 3-2. 8mm; green, heart line and lateral lines faintly outlined with white; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen long; cremaster to 1.6mm long, blunt with numerous hooked

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229 setae in a cluster at the tip (Figures 106H and 118G) ; lenticles absent. Other Descriptions Minno and Emmel 1993 (larva, pupa). Host Plants POACEAE: Cynodon dactylon (Kendall 1960). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1517, 6 eggs (MCM) LARVAE: FLORIDA: ALACHUA CO.: Lot 1061, 1 larva (instar 5) (DHH); Lot 1063, 1 larva (instar 5) (DHH); Lot 1517, 11 larvae (instar 1) (MCM). BROWARD CO.: Lot 127, 10 larvae (instars 1, 5) (MCM). TEXAS: LASALLE CO.: Lot 796, 3 larvae (instars 4, 5) (USNM) ; Lot 797, 1 larva (instar 5) (USNM). PUPAE: FLORIDA: BROWARD CO.: Lot 127, 4 pupae (MCM). 148. Adopaeoides prittwitzi (Plotz) The immature stages of this southwestern species are undescribed, and I could not find specimens for study. Host Plants POACEAE: Paspalum disticum (Bailowitz and Brock 1991) 149. Thymelicus lineola (Ochsenheimer) Diagnosis LAST INSTAR LARVA: BODY: length 21mm, A4 transverse width 4.1mm; preserved specimens pale; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 29H, tan, unmarked; transverse width 2.2mm; setae simple, to 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 1 and 6 small (Figure 41Q) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 60

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230 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 51A) Other Descriptions Forbes 1960 (larva, pupa), Pengelly 1961 (egg, larva, pupa), Toliver 1987 (larva). Host Plants CYPERACEAE: Carex species (Pengelly 1961, larval feeding, but doubtful if sedge served as an oviposition substrate) POACEAE: Agropyron repens (Pengelly 1961), Agrostis alba (Pengelly 1965), Avena sativa (Pengelly 1961), Bromus inermis (Pengelly 1961, in lab), Dactylis glomerata (Bucher and Arthur 1961), Festuca elatior (Pengelly 1961), Lolium perenne (Pengelly 1961), Phleum pratense (Bucher and Arthur 1961), Poa compressa (Arthur and Smith 1974), Poa pratensis (Pengelly 1965). Specimens Examined LARVAE: NEW YORK: TOMPKINS CO.: Lot 545, 1 larva ( instar 4) (TLM) 150. Hylephila phyleus (Drury). Diagnosis EGG: Figure 3E, height 0.7-0. 8mm, width 0.50.6mm, polygonal sculpturing, pale green. LAST INSTAR LARVA: BODY: length 14-34. 5mm, A4 transverse width 3.25.3mm; brown, heart line darker brown, subdorsal line faint, dark; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 30A, black with two pale lines parallel to vertex, adfrontals mostly pale, front with two pale spots, a pale eye patch containing a central black spot present; transverse width 2. 7-3. 3mm; sculpturing rough to pitted; setae simple, < 0.1mm long at apex, a few ventral

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231 setae to 0.3mm; stemmata subequal (Figure 41R) THORAX: legs very dark brown; shield broad, very dark brown. ABDOMEN: prolegs each with about 120 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 51B) ; no wax glands. PUPA: Figures 70B and 91A, length 15-28mm, A3 transverse width 3. 8-5. 2mm; cream-colored with a dark brown dorsal line on the thorax and abdomen, other short dark brown lines on head and mesothorax, tiny dark brown spots on the head, thorax, and abdomen; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pi lifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen moderately long; cremaster to 1.3mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 1061 and 118H) ; lenticles present on the prothorax, dorsum of abdomen, and proleg scars. Other Descriptions Edwards and Chapman 1879 (larva, pupa), Panton 1897 (egg, larva, pupa), Coquillett 1899 (larva), Coolidge 1925 (egg, larva, pupa), Comstock 1927b (egg, larva, pupa), Comstock and Dammers 1933a (egg, larva, pupa), Dethier 1939b (larva), Moss 1949 (larva), Peterson 1962 (larva), Emmel and Emmel 1973 (egg, larva, pupa), Hogue 1974 (larva), Tashiro and Mitchell 1985 (egg, larva, pupa), Minno and Emmel 1993 (egg, larva, pupa).

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232 Host Plants CANNACEAE: Canna species (da Costa Lima 1936) [erroneous]. EUPHORBIACEAE: Euphorbia species (Orsak 1978, oviposition substrate only). POACEAE: Agrostis species (Bohart 1947), Axinopus compressus (Brown and Heineman 1972), Cynodon dactylon (Coolidge 1925), Digitaria sanguinalis (Scudder 1872), Eragrostis hypnoides (Scott 1986), Panicum species (Bates 1935), Paspalum conjugatum (Kaye 1926), Poa pratensis (Tietz 1952), Saccharum officinarum (Dethier 1942d, in lab), Stenotaphrum secundatum (Kendall 1960). ROSACEAE: Rosa species (Wright 1905) [ erroneous ] Specimens Examined EGGS: CALIFORNIA: SAN BERNARDINO CO.: Lot 631, 4 eggs (TCE) FLORIDA: PUTNAM CO.: Lot 1524, 12 eggs (MCM) LARVAE: CALIFORNIA: RIVERSIDE CO.: Lot 575, 2 larvae (instar 5) (GRB); Lot 1040, 1 larva (instar 5) (DHH). SAN BERNARDINO CO.: Lot 631, 9 larvae ( instars 1, 2, 3, 4, 5) (TCE). YOLO CO.: Lot 261, 14 larvae (instars (2, 3, 4, 5) (MCM); Lot 266, 21 larvae (instars 2, 3, 4, 5) (MCM). FLORIDA: ALACHUA CO.: Lot 9, 5 larvae (instar 5) (MCM); Lot 258, 5 larvae (instar 5) (MCM); Lot 260, 4 larvae (instar 5) (MCM); Lot 1039, 1 larva (instar 5) (DHH); Lot 1042 (possibly), 1 larva (instar 5) (DHH). BROWARD CO.: Lot 259, 4 larvae (instar 5) (MCM). PUTNAM CO.: Lot 1524, 14 larvae (instar 1) (MCM). NEW MEXICO: BERNALILLO CO.: Lot 678, 5 larvae (instar 5) (MCM); Lot 679, 1 larva (instar 5) (MCM); Lot 680, 4 larvae (instar 5) (MCM); Lot 682, 4

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233 larvae (instar 5) (MCM) PUPAE: USA: CALIFORNIA: SAN BERNARDINO CO.: Lot 631, 1 pupa (TCE) YOLO CO.: Lot 262, 26 pupae (MCM); Lot 263, 25 pupae (MCM); Lot 264, 25 pupae (MCM). FLORIDA: ALACHUA CO.: Lot 9, 2 pupae (MCM); Lot 258, 3 pupae (MCM); Lot 260, 2 pupae (MCM). BROWARD CO.: Lot 259, 5 pupae (MCM). NEW MEXICO: BERNALILLO CO.: Lot 678, 4 pupae (MCM); Lot 679, 1 pupa (MCM); Lot 680, 6 pupae (MCM); Lot 681, 7 pupae (MCM); Lot 682, 6 pupae (MCM). 151. Yvretta rhesus (Edwards) The immature stages of this western species are undescribed, and I could not find specimens for study. Little is known of the biology of this species, except that the grass, Bouteloua gracilis, is a host (Scott and Scott 1980). 152. Yvretta cams (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 28-28. 5mm, A4 transverse width 4. 8-5. 3mm; preserved specimens pale; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 30B, black with two pale lines parallel to vertex, adfrontals pale, front with two small pale spots; a pale eye patch present; transverse width 3.1mm; sculpturing rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41S). THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with three black

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234 longitudinal lines (Figure 51C). PUPA: Figures 70C and 91B, length 27.5mm, A3 transverse width 4.8mm; light brown; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen moderately long; cremaster to 1.9mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 106J and 1181); lenticles present on the prothorax and proleg scars. Host Plants POACEAE: Cynodon dactylon (MCM collection, in lab). Specimens Examined LARVAE: ARIZONA: SANTA CRUZ CO.: Lot 528, 2 larvae (instar 5) (MCM). PUPAE: ARIZONA: SANTA CRUZ CO.: Lot 528, 1 pupa (MCM). 153. Pseudocopaeodes eunus (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 22.5mm, A4 transverse width 4.8mm; preserved specimen pale; some setae with blunt tips, to 0.2mm long on A4 dorsum; spiracles black. HEAD: Figure 30C, black with two pale lines parallel to vertex extending onto the adfrontals, front without pale spots, a pale eye patch present; transverse width 2.9mm; sculpturing rough to pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 41T) THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: pro legs each with about 50 crochets, irregularly triordinal, arranged in a near circle;

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235 suranal plate rounded, unmarked (Figure 51D) PUPA: Figures 70D and 91C, length 16mm, A3 transverse width 4.3mm; light brown; thoracic spiracle guard distinct; setae simple, to 0.2mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen moderately long; cremaster to 1 3mm long blunt with numerous hooked setae in a cluster at the tip (Figures 106K and 118J) ; lenticles present on the prothorax and proleg scars. Other Descriptions Comstock 1932a (egg). Host Plants POACEAE: Distichlis spicata (Emmel and Emmel 1973, probably), Distichlis spicata var. stricta (MacNeill 1975) Specimens Examined LARVAE: CALIFORNIA: INYO CO. : Lot 434, 1 larva ( instar 5) (MCM) PUPAE: CALIFORNIA: INYO CO.: Lot 434, 1 pupa (MCM). 154. Stinga morrisoni (Edwards) The immature stages of this western species are mostly undescribed, and I could not find specimens for study. Emmel et al. (1992) noted that the eggs are relatively large and white. Nothing is known of the biology of this species. Stanford (1981) thought that the grasses, Andropogon scoparius and Bouteloua gracilis may be hosts. 155. Hesperia uncas Edwards Diagnosis LAST INSTAR LARVA: BODY: length 21-29mm, A4 transverse width 4. 6-5. 4mm; brown; setae simple, < 0.1mm

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236 long on A4 dorsum; spiracles dark brown. HEAD: Figure 30D, black with two pale lines parallel to vertex, adfrontals pale, front with two small pale spots; transverse width 3.33.5mm; sculpturing rough to pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41U) THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 51E) PUPA: Figures 70E and 91D, length 18-22mm, A3 transverse width 3. 7-4. lmm; pale green, abdomen cream-colored with small dark spots; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.9mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 106L and 118K); lenticles present on the prothorax, abdomen, and proleg scars. Other Descriptions MacNeill 1964 (egg), Scott 1975a (larva, pupa), Emmel et al. 1992 (egg, larva, pupa). Host Plants POACEAE: Bouteloua gracilis (Shields et ai. 1970), Bouteloua uniflora (Scott 1986), Erioneuron pilosum (McGuire 1982), Poa pratensis (McCabe and Post 1977, in lab), Stipa nevadensis (MacNeill 1975), Stipa pinetorum (McGuire 1982)

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237 Specimens Examined LARVAE: COLORADO: EL PASO CO.: Lot 256, 6 larvae (instars 3, 4, 5) (MCM) ; Lot 257, 12 larvae (instars 2, 3, 4) (MCM). NORTH DAKOTA: SLOPE CO.: Lot 550, 1 larva ( instar 5) (TLM). PUPAE: COLORADO: EL PASO CO.: Lot 256, 3 pupae (MCM). 156. Hesperia juba (Scudder) Diagnosis LAST INSTAR LARVA: BODY: length 22.5-39mm, A4 transverse width 4.4-6mm; preserved specimen brownish; some setae with expanded tips, to 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: black with two pale lines parallel to the vertex, adfrontals pale, front with two small pale spots; transverse width 3. 6-3. 8mm; sculpturing rough to pitted; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41V). THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked; wax glands in transverse patches on the ventral side of A7 and A8 (Figure 15E). Other Descriptions Lindsey 1923 (egg, larva), MacNeill 1964 (egg, larva, pupa). Host Plants POACEAE: Bromus rubens (McGuire 1982), Deschampsia elongata (McGuire 1982), Poa agassizensis (Scott 1986), Poa pratensis (Scott 1986), Stipa species (McGuire 1982)

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238 Specimens Examined LARVAE: CALIFORNIA: SAN BERNARDINO CO.: Lot 578, 2 larvae (instar 5) (GRB). 157. Hesperia comma (Linnaeus) Diagnosis EGG: height 1.1mm, width 0.7mm, polygonal sculpturing. LAST INSTAR LARVA: BODY: length 15-28. 5mm, A4 transverse width 4.3-6mm; brownish; some setae with blunt or slightly expanded tips, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: black with two pale lines parallel to the vertex, adfrontals mostly pale, front with two small pale spots; transverse width 3-3. 4mm; sculpturing rough to pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 41W) THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 75 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked. PUPA: Figures 70F and 91E, length 17. 5-24. 5mm, A3 transverse width 5-5. 6mm; cream-colored; thoracic spiracle guard distinct; setae simple, to 0.3mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 2mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107A and 119A); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions. Fletcher 1888 (egg, larva), Fyles 1895 (egg, larva, pupa), Cockayne 1952 (larva), MacNeill 1964

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239 (egg, larva, pupa), Scott 1975a (egg, larva, pupa), McCabe and Post 1977 (larva), Emmel et al. 1992 (egg, larva). Host Plants POACEAE: Andropogon saccharoides (Scott 1986), Bouteloua gracilis (Scott 1986), Browns species (Hardy 1954), Festuca rubra (MacNeill 1964), Koeleria cristata (Klassen et al. 1989), Lolium species (Hardy 1954), Muhlenbergia species (McGuire 1982), Phleum pratense (Fyles 1895, in lab), Poa arctica (Scott 1986), Poa glauca (Scott 1986), Poa pratensis (Tietz 1972), Poa scabrella (MacNeill 1964), Stipa thurberiana (MacNeill 1975), Vulpia octaflora (McGuire 1982, associated with). S pecimens Examined EGGS: CANADA: NEW BRUNSWICK: VICTORIA CO.: Lot 999, 7 eggs (FSCA). LARVAE: CANADA: NEW BRUNSWICK: VICTORIA CO.: Lot 999, 13 larvae (instars 1, 5) (FSCA). CALIFORNIA: SAN DIEGO CO.: Lot 254, 2 larvae (instar 5) (MCM) COLORADO: TELLER CO.: Lot 255, 8 larvae (instars 1, 2, 5) (MCM). NORTH DAKOTA: SLOPE CO.: Lot 548, 1 larva (instar 5) (TLM). PUPAE: CANADA: NEW BRUNSWICK: VICTORIA CO.: Lot 999, 1 pupa (FSCA). CALIFORNIA: SAN DIEGO CO.: Lot 254, 2 pupae (MCM). 158. Hesperia woodgatei (R. C. Williams) Diagnosis EGG: height 1.1-1. 3mm, width 0.8-lmm, polygonal sculpturing. LAST INSTAR LARVA: BODY: length 27mra, A4 transverse width 4.8mm; preserved specimens brownish; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: black with two pale lines

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240 parallel to the vertex, adfrontals mostly pale, front with two small pale spots; transverse width 3.7mm; sculpturing pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41X) THORAX: legs black; shield broad, black. ABDOMEN: prolegs each with about 80 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked. PUPA: length 14.5mm, A3 transverse width 4.83mm; cream-colored with dark markings on the head and thorax; thoracic spiracle guard distinct; setae simple, to 0.2mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen short; cremaster to 1.6mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107B and 119B); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions MacNeill 1964 (egg), Pyle 1981 (egg, larva, pupa). Host Plants POACEAE: Bouteloua uni flora var. coahuilensis (ROK collection). Specimen s Examined EGGS: MEXICO: COAHUILA: Lot 947, 2 eggs (ROK), probably this species. LARVAE: MEXICO: COAHUILA: Lot 947, 6 larvae (instars 1, 4, 5) (ROK), probably this species. PUPAE: MEXICO: COAHUILA: Lot 947, 1 pupa (ROK), probably this species.

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241 159. Hesperia ottoe Edwards The egg and larva of this species was described by Nielsen (1958, 1960), but I could not find specimens for study Host Plants ASTERACEAE: Echinacea pallida (Dana 1981) [oviposition substrate only]. POACEAE: Andropogon gerardii (Stanford 1981, probably; Scott 1986), Andropogon scoparius (McGuire 1982), Bouteloua curtipendula (Scott 1986), Bouteloua gracilis (Scott 1986), Bouteloua hirsuta (Scott 1986), Bromus species (Scott 1986), Leptoloma cognatum (Nielsen 1958), Panicum (wilcoxianum?) (Scott 1986), Sporobolus neglecta (Scott 1986). Specimens Examined LARVAE: NORTH DAKOTA: BILLINGS CO.: Lot 542, 1 larva ( instar 1) (TLM) 160. Hesperia leonardus Harris Diagnosis LAST INSTAR LARVA: BODY: length 13 -15mm, A4 transverse width 3. 5-3. 6mm; preserved specimens brownish; some setae with expanded tips, < 0.1mm long on A4 dorsum; dark brown. HEAD: Figure 30E, black with two pale lines parallel to vertex, adfrontals mostly pale, front with two pale spots, a pale eye patch present; transverse width 3.5mm; sculpturing pitted; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 41Y). THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: pro legs each with about 60 crochets,

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242 irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked. Other Descriptions Scudder 1889a (egg, young larva), Dethier 1939a (young larva), Dethier 1948 (larva, pupa), Scott 1975a (egg, larva, pupa). Host Plants ASTERACEAE: Taraxacum officinale (Tietz 1972) [oviposition substrate only]. POACEAE: Agrostis species (Scudder 1893), Andropogon species (Opler and Krizek 1984, associated with), Danthonia spicata (McGuire 1982), Eragrostis alba (Shapiro 1966), Panicum virgatum (Shapiro 1966), Sporobolus heterlepis (Scott 1986). S pecimens Examined LARVAE: PENNSYLVANIA: HUNTINGDON CO. : Lot 1008, 2 larvae (instar 5) (FSCA). 161. Hesperia pawnee Dodge Scott (1975a) has described the egg, larva, and pupa of this upper Great Plains species, but I could not find specimens for study. Host Plants POACEAE: Andropogon scoparius (Scott 1986), Bouteloua curtipendula (Scott 1986), Bouteloua gracilis (Stanford 1981, probably; Scott and Stanford 1982), Bouteloua hirsuta (Scott 1986), Cynodon dactylon (Scott and Stanford 1982, in lab), Poa pratensis (Scott and Stanford 1982, in lab), Stipa comata (McCabe and Post 1977), Tridens species (Pyle 1981).

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243 162. Hesperia pahaska (Leussler) Diagnosis LAST INSTAR LARVA: BODY: length 25. 52 9mm, A4 transverse width 4. 8-5. 5mm; preserved specimens pale; some setae with slightly expanded tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 30F, black with pale patches on upper face, two pale lines parallel to the vertex, adfrontals mostly pale, front with two small pale spots, a pale eye patch present; transverse width 3. 53.7mm ; sculpturing pitted; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 41Z). THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded, unmarked. Other Descriptions MacNeill 1964 (egg, larva, pupa), Scott 1975a (larva, pupa). Host Plants POACEAE: Bouteloua gracilis (MacNeill 1975), Erioneuron pulchellum (Emmel and Emmel 1973), Tridens pulchellus (MacNeill 1964). Specimens Examined LARVAE: CALIFORNIA: SAN BERNARDINO CO.: Lot 1497, 4 larvae ( instar 5) (MCM) 163. Hesperia Columbia (Scudder) MacNeill (1964) described the egg, larva, and pupa of this western species, but I could not find specimens for study

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244 Host Plants POACEAE: Danthonia californica (Emmel and Emmel 1973, probably), Koeleria cristata (MacNeill 1964), Koeleria macrantha (Garth and Tilden 1986). 164. Hesperia metea Scudder I could not locate eggs, mature larvae, or pupae of this species for study. Figure 41AA shows the stemmatal pattern of a fourth instar larva. Other Descriptions Scudder 1881 (larva), Scudder 1889a (egg, young larva), Laurent 1908 (larva, pupa), Heitzman and Heitzitian 1970b (egg, larva). Host Plants POACEAE: Andropogon gerardii (Heitzman and Heitzman 1970b), Andropogon glomeratus (Shapiro 1965, associated with; Shapiro 1966), Andropogon scoparius (Shapiro 1965, associated with; Shapiro 1966), Andropogon virginicus var. abbreviatus (Shapiro 1966, possibly; Scott 1986), Andropogon species (Macy and Shepard 1941), Panicum species (Scudder 1869, probably). Specimens Examined LARVAE: GEORGIA: DEKALB CO.: Lot 945, 1 larva (instar 4) (ROK) 165. Hesperia viridis (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 14mm, A4 transverse width 4.3mm; preserved specimens brown; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: black with two pale lines parallel to the vertex, adfrontals pale, front with two pale spots; transverse width 3.7mm; sculpturing pitted; setae

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245 simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 4 IBB) THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 65 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked. PUPAL EXUVIUM: cremaster blunt with numerous hooked setae in a cluster at the tip (Figures 107C and 119C) Other Descriptions MacNeill 1964 (egg), Scott 1975a (larva, pupa) Host Plants POACEAE: Bouteloua curtipendula (Scott 1986), Bouteloua gracilis (Kendall 1965, possibly; MacNeill 1975), Buchloe dactyloides (McGuire 1982), Cynodon dactylon (Kendall 1965, in lab), Erioneuron pilosum (McGuire 1982), Lolium perenne (Kendall 1965, in lab), Tridens muticus (McGuire 1982) Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 946, 2 egg shells (ROK) LARVAE: TEXAS: BEXAR CO.: Lot 946, 12 larvae (instars 1, 3, 5) (ROK). 166. Hesperia attalus (Edwards) Diagnosis EGG: Figure 4D, height 1.2-1. 5mm, width 0.91.2mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: Figure 20E, length 20.5-28mm, A4 transverse width 3. 6-5. 6mm; olive green or brownish; some setae with blunt or slightly expanded tips, < 0.1mm long on A4 dorsum; spiracles black. HEAD: black with two pale lines parallel to the vertex, adfrontals mostly pale, front with two pale spots;

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246 transverse width 3. 3-5. 6mm; sculpturing pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemmata subegual (Figure 42A) THORAX: Tl-2 legs dark brown, T3 legs brown; shield broad, dark brown. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked; wax glands forming two ventral transverse patches on A7-8. PUPA: Figures 59A, 70G, and 91F, length 19-23mm, A3 transverse width 4. 5-4. 8mm; pale green, abdomen creamcolored with small light brown spots; thoracic spiracle guard distinct; setae simple, to 0.2mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A8; abdomen long; cremaster to 1.8mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107D and 119D) ; lenticles present on the prothorax and lenticle scars. Host Plants POACEAE: Aristida stricta (MCM collection, in lab), Aristida virgata (McGuire 1982), Bothriochloa barbinodis (McGuire 1982, possibly), Bouteloua curtipendula var. caespitosa (McGuire 1982), Cynodon dactylon (MCM collection, in lab), Leptoloma cognatum (McGuire 1982), Panicum virgatum (Shapiro and Shapiro 1973, associated with) Specimens Examined EGGS: FLORIDA: LEVY CO.: Lot 248, 1 egg (MCM). ST. JOHNS CO.: Lot 1045, 4 eggs (DHH). LARVAE:

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247 FLORIDA: ALACHUA CO.: Lot 247, 1 larva (instar 1) (MCM); Lot 251, 9 larvae (instars 1, 2) (MCM). LEVY CO.: Lot 248, 11 larvae (instars 1, 2) (MCM); Lot 252, 5 larvae (instars 4, 5) (MCM). PUTNAM CO.: Lot 249, 1 larva (instar 5) (MCM); Lot 250, 1 larva (instar 4) (MCM); Lot 675, 1 larva (instar 5) (MCM); Lot 676, 6 larvae (instar 5) (MCM); Lot 677, 7 larvae (instars 3, 4) (MCM); Lot 997, 3 larvae (instars 4, 5) (MCM); Lot 1525, 3 larvae (instar 1) (MCM). ST. JOHNS CO.: Lot 1044, 1 larva (instar 1) (DHH); Lot 1046, 3 larvae (instar 1) (DHH). PUPAE: FLORIDA: LEVY CO.: Lot 252, 1 pupa (MCM). PUTNAM CO.: Lot 676, 2 pupae (MCM); Lot 997, 2 pupae (MCM). 167. Hesperia meskei (Edwards) Diagnosis EGG: height 1.4mm, width 0.9mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: length 1620mm, A4 transverse width 4. 4-4. 6mm; brown; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: black with two pale lines parallel to the vertex; transverse width 3. 6-3. 8mm; sculpturing pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 42B) THORAX: legs black; shield broad, black. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked.

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248 Host Plants POACEAE: Andropogon scoparius (McGuire 1982), Aristida purpurascens (McGuire 1982, associated with; Scott 1986), Cynodon dactylon (MCM collection, in lab). Specimens Examined EGGS: FLORIDA: DUVAL CO.: Lot 1043, 2 eggs (DHH). LARVAE: FLORIDA: PUTNAM CO.: Lot 994, 3 larvae (instars 3, 5) (MCM). 168. Hesperia dacotae (Skinner) McCabe and Post (1977) and McCabe (1981) described the larva of this upper Great Plains species, but the only specimen that I could find for examination was a single first instar larva. Host Plants CYPERACEAE: Carex species (McCabe 1981, in lab). POACEAE: Andropogon gerardii (McCabe 1981, in lab), Andropogon scoparius (Scott 1986), Aristida species (Pyle 1981), Bouteloua species (Pyle 1981), Koeleria cristata (McCabe 1981, in lab), Panicum species (Scott 1986), Phleum pratense (McCabe 1981, in lab), Poa pratensis (McCabe 1981, in lab), Stipa spartea (McCabe 1981, in lab). Specimens Examined LARVAE: MINNESOTA: FELTON CLAY CO.: Lot 543, 1 larva (instar 1) (TLM) 169. Hesperia lindseyi (Holland) MacNeill (1964) described the egg, larva, and pupa of this western species, but I could not find specimens for study

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249 Host Plants POACEAE: Danthonia californica (MacNeill 1964), Festuca idahoensis (MacNeill 1964), Vulpia megalura (Scott 1986) 170. Hesperia sassacus Harris Scudder (1889a) described the egg and larva of this northeastern species, but I could not find specimens for study. Host Plants CYPERACEAE: Car ex varia (Fletcher 1888, in lab) POACEAE: Agrostis vulgaris (Fletcher 1888, in lab), Andropogon scoparius (McGuire 1982), Danthonia spicata (Fletcher 1888, in lab), Digitaria sanguinalis (Scudder 1889a, b), Festuca obtusa (Scott 1986), Festuca paradoxa (Shapiro 1974a), Festuca rubra (Opler and Krizek 1984), Panicum species (Scudder 1893), Poa annua (Shapiro 1966, possibly) 171. Hesperia miriamae MacNeill Little is known of the biology of this high-altitude, western species. MacNeill (1975) briefly described the egg and listed the grass, Andropogon scoparius, as a host. McGuire (1982) corrected the host identification to Festuca br achy phy 11a 172. Hesperia nevada (Scudder) I could not locate eggs, last instar larvae, or pupae of this western, montane species for study. Figure 42C shows the stemmatal pattern of a fourth instar larva.

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250 Other Descriptions MacNeill 1964 (egg, larva), Emmel et al. 1992 (egg, larva). Host Plants POACEAE: Festuca ovina (Pyle 1981), Koeleria species (Scott and Scott 1980), Poa pratensis (MCM collection, in lab), Sitanion hystrix (Emmel et al 1971), Stipa occidentalis (MacNeill 1975). Specimens Examined LARVAE : COLORADO : TELLER CO : Lot 253, 14 larvae ( instars 1, 3, 4) (MCM). 173. Polites cor as (Cramer) Diagnosis LAST INSTAR LARVA: BODY: length 18.5-25mm, A4 transverse width 3. 8-4. 6mm; brown with tiny dark spots, heart line darker brown, subdorsal line faint, brown; setae simple, to 0.4mm long on A4 dorsum; spiracles black. HEAD: Figure 30G, black with tan patches on upper face, two pale lines parallel to vertex, adfrontals mostly pale, a pale eye patch present; transverse width 2.6-3mm; sculpturing pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.3mm; stemmata subegual (Figure 42D) THORAX: legs very dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with dark markings (Figure 51F). PUPA: Figures 70H and 91G, length 15-18mm, A3 transverse width 3. 8-4. 3mm; blackish brown, abdomen cream-colored; thoracic spiracle guard distinct; setae simple, to 0.8mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle

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251 leg; proboscis extending into A8; abdomen long; cremaster to 1.1mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107E and 119E) ; lenticles present on the prothorax and proleg scars Other Descriptions Saunders 1869a (egg, young larva), Dethier 1940b (larva, pupa). Host Plants POACEAE: Cynodon dactylon (MCM collection, in lab), Leersia oryzoides (Shapiro 1974a). Specimens Examined LARVAE: MARYLAND: PRINCE GEORGE'S CO.: Lot 14, 6 larvae ( instar 5); Lot 386, 2 larvae (instar 5) (MCM); Lot 388, 4 larvae (instar 5) (MCM). NEW YORK: Lot 650?, 1 larva (instar 4) (USNM) PUPAE: MARYLAND: PRINCE GEORGE'S CO.: Lot 14, 4 pupae (MCM); Lot 385, 6 pupae (MCM); Lot 386, 2 pupae (MCM); Lot 387, 5 pupae (MCM); Lot 388, 2 pupae (MCM). 174. Polites sabuleti (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 15-25mm, A4 transverse width 2. 9-4. 5mm; green or brown, heart line darker, subdorsal line faint, dark, lateral line bounded by dark bands; setae simple, to 0.4mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 30H, black, upper face tan with two pale lines parallel to vertex, adfrontals pale, a small pale eye patch present; transverse width 2. 3-2. 9mm; sculpturing pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 42E). THORAX: legs very dark brown; shield broad, dark brown.

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252 ABDOMEN: prolegs each with about 65 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with dark markings (Figures 51G and 52A) PUPA: Figures 71A and 91H, length 13-16mm, A3 transverse width 3.5-4mm; brown, abdomen cream-colored; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennae tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to lmm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107F and 119F); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Corns tock 1929 (egg, young larva), Dethier 1944a (egg, larva, pupa), Newcomer 1966 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants CYPERACEAE: Carex filifolia (Lembert 1894) [oviposition substrate only]. FABACEAE: Tri folium monanthum (Lembert 1894) [oviposition substrate only]. POACEAE: Agrostis scabra (Shapiro 1977), Cynodon dactylon (Wright 1905), Digitaria sanguinalis (Tietz 1972), Distichlis spicata (Wright 1905, probably; Shapiro 1974b, c) Distichlis spicata var. stricta (Scott 1986), Eragrostis trichodes (Scott and Scott 1980), Festuca brachyphylla (McGuire 1982), Festuca idahoensis (Shapiro et al 1981), Poa pratensis (Newcomer 1966, associated with; Scott 1986). Specimens Examined LARVAE: CALIFORNIA: NEVADA CO.: Lot 403, 14 larvae (instars 3, 4, 5) (MCM) ORANGE CO.: Lot

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253 577, 1 larva (instar 5) (GRB). SACRAMENTO CO.: Lot 399, 1 larva (instar 3) (MCM); Lot 400, 6 larvae (instar 5) (MCM); Lot 401, 5 larvae (instar 5) (MCM); Lot 402, 4 larvae (instar 5) (MCM). PUPAE: CALIFORNIA: NEVADA CO.: Lot 403, 2 pupae (MCM). SACRAMENTO CO.: Lot 400, 2 pupae (MCM); Lot 401, 2 pupae (MCM); Lot 402, 4 pupae (MCM). 175. Polites mar don (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 19 5-21 5mm, A4 transverse width 3. 7-4. 2mm; preserved specimens brown; setae simple, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 31A, black with pale patches on upper face, two pale lines parallel to the vertex, adfrontal mostly pale, a pale eye patch present; transverse width 2.62.8mm; sculpturing pitted; setae simple, to 0.1mm long at apex, a few ventral setae to 0.2mm; stemmata subegual (Figure 42F) THORAX: legs very dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 65 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with dark markings (Figure 52B) PUPA: Figures 71B and 92A, length 17mm, A3 transverse width 4-4. lmm; brown; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers touching; head rounded; antennae tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to lmm long, bluntly pointed with numerous hooked setae in a cluster at the tip

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254 (Figures 107G and 119G) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Newcomer 1966 (egg, larva, pupa). Host Plants POACEAE: Bromus carinatus (Newcomer 1966, probably), Festuca idahoensis (Pyle 1981, suspected), Festuca ovina (Newcomer 1966, probably), Poa pratensis (MCM collection, in lab). Specimens Examined LARVAE: CALIFORNIA: DEL NORTE CO.: Lot 391, 2 larvae ( instar 5) (MCM), Lot 392, 4 larvae (instar 4) (MCM); Lot 576, 1 larva (instar 5) (GRB). PUPAE: CALIFORNIA: DEL NORTE CO.: Lot 391, 2 pupae (MCM). 176. Polites draco (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 17-22mm, A4 transverse width 3. 3-4. 8mm; brown, heart line darker brown, subdorsal line faint, dark, lateral line bounded by dark bands; setae simple, to 0.4mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 3 IB, black with two pale lines parallel to vertex, adfrontals mostly pale; transverse width 2. 1-3. lmm; sculpturing pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.3mm; stemmata subegual (Figure 42G) THORAX: legs very dark brown; shield broad, black. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with dark markings (Figure 52C) PUPA: Figures 71C and 92B, length 14-26. 5mm, A3 transverse width 3. 6-4. 8mm; dark brown, abdomen cream-colored; thoracic spiracle guard

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255 distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.5mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107H and 119H); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Emmel et al 1992 (larva, pupa). Host Plants POACEAE: Poa pratensis (MCM collection, in lab), Poa species (Emmel et al 1992). Specimens Examined LARVAE: COLORADO: TELLER CO.: Lot 389, 7 larvae ( instars 1, 2, 3, 5) (MCM); Lot 390, 12 larvae (instars 4, 5) (MCM). PUPAE: COLORADO: TELLER CO.: Lot 389, 2 pupae (MCM); Lot 390, 11 pupae (MCM). 177. Polites baracoa (Lucas) Diagnosis EGG: Figure 4E, height 0.9mm, width 0.6-0. 7mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: length 19mm, A4 transverse width 4.1mm; brown, heart line darker brown, subdorsal line dark brown, lateral line orange brown bounded by dark bands; some setae with expanded tips, < 0.1mm long on A4 dorsum; spiracles brown. HEAD: Figure 31C, black, upper face tan with dark pits, two pale lines parallel to vertex, adfrontals partly pale, front with two pale spots, a pale eye patch present; transverse width 2.1mm; sculpturing pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemmata subegual

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256 (Figure 42H) THORAX: legs dark brown; shield broad, black. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with three broad black longitudinal stripes convergent along the distal margin (Figure 52D). PUPA: Figures 71D and 92C, length 14-15. 5mm, A3 transverse width 3. 3-4. 0mm; cream-colored with brown markings on the head and thorax; thoracic spiracle guard distinct; setae simple, to 0.3mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 1071 and 1191); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Dethier 1939b (egg, larva). Host Plants POACEAE: Cynodon dactylon (MCM collection, in lab), Saccharum officinarum (Dethier 1939b, in lab). Specimens Examined EGGS: FLORIDA: PUTNAM CO.: Lot 1527, 7 eggs (MCM). LARVAE: FLORIDA: BROWARD CO.: Lot 384, 9 larvae (instars 1, 5) (MCM). PUTNAM CO.: Lot 998, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: BROWARD CO.: Lot 384, 11 pupae (MCM) 178. Polites themistocles (Latreille) Diagnosis EGG: height 1-1. lmm, width 0.6-0. 8mm, polygonal sculpturing, pinkish. LAST INSTAR LARVA: BODY: length 1930mm, A4 transverse width 3.9-5mm; green or brown, heart

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257 line dark, subdorsal line faint, dark, lateral line bounded by dark bands; setae simple, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 31D, black with two pale lines parallel to vertex, adfrontals mostly pale, or occasionally head uniformly black; transverse width 2.5-3mm; sculpturing rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 421). THORAX: legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with dark markings (Figure 52E) PUPA: Figures 71E and 92D, length 14.5-24mm, A3 transverse width 3. 8-5. lmm; green, abdomen cream-colored; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen long; cremaster to 1.5mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107J and 119J) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Fletcher 1888 (egg, larva, pupa), Scudder 1889a (egg, larva, pupa), Fyles 1896 (larva), Laurent 1908 (egg, larva, pupa), Dethier 1938a (larva), Dethier 1942a (larva, pupa), Emmel et al 1992 (egg, larva, pupa ) Host Plants POACEAE: Agropyron repens (Fletcher 1888, in lab), Cynodon dactylon (MCM collection, in lab), Digitaria

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258 filiformis (Opler and Krizek 1984), Echinochloa crusgalli (Fletcher 1888, in lab), Eremochloa ophiuroides (MCM collection), Panicum aciculare (Minno 1992), Panicum clandestinum (Shapiro 1966), Phleum pratense (Fletcher 1888, in lab), Poa pratensis (Scott 1986), Poa species (Stanford 1981) Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1529, 11 eggs (MCM). LARVAE: COLORADO: FREMONT CO.: Lot 409, 7 larvae (instar 5) (MCM); Lot 410, 3 larvae (instars 1, 5) (MCM); Lot 411, 3 larvae (instar 5) (MCM). FLORIDA: ALACHUA CO.: Lot 413, 8 larvae (instar 5) (MCM); Lot 1529, 3 larvae (instar 1) (MCM). FLAGLER CO.: Lot 408, 2 larvae (instar 5) (MCM). HIGHLANDS CO.: Lot 412, 2 larvae (instar 2) (MCM). PUPAE: COLORADO: FREMONT CO.: Lot 409, 7 pupa (MCM); Lot 410, 1 pupa (MCM). FLORIDA: ALACHUA CO.: Lot 413. 4 pupae (MCM). FLAGLER CO.: Lot 408, 1 pupa (MCM). HIGHLANDS CO.: Lot 411, 2 pupae (MCM); Lot 412, 2 pupae (MCM) 179. Polites origenes (Fabricius) Diagnosis LAST INSTAR LARVA: BODY: length 2 33 0mm, A4 transverse width 4-5. 4mm; brown, heart line darker brown, subdorsal line faint, dark; setae simple, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 31E, uniformly black; transverse width 3-3. 5mm; sculpturing rough to pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 42J) THORAX:

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259 legs dark brown; shield broad, dark brown. ABDOMEN: prolegs each with about 75 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with a few small dark spots (Figure 52F); wax glands forming a ventral transverse patch on A7-8. PUPA: Figures 71F and 92E, length 19-22. 5mm, A3 transverse width 4. 3-5. 6mm; green, abdomen cream-colored; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.7mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107K and 119K) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Scudder 1889a (egg, larva), Laurent 1892 (egg, larva, pupa), Laurent 1908 (egg), Dethier 1938a ( e g9/ young larva), Dethier 1941 (larva). Host Plants POACEAE: Andropogon scoparius (Scott 1986), Cynodon dactylon (MCM collection, in lab), Poa pratensis (MCM collection, in lab), Tridens flavus (Shapiro 1966). Specimens Examined LARVAE: COLORADO: EL PASO CO.: Lot 394, 2 larvae ( instars 3, 4) (MCM); Lot 395, 5 larvae (instar 5) (MCM); Lot 396, 5 larvae (instar 5) (MCM). MARYLAND: PRINCE GEORGE'S CO.: Lot 397, 2 larvae (instar 5) (MCM). PENNSYLVANIA: CAMBRIA CO.: Lot 398, 4 larvae (instar 3) (MCM). PUPAE: COLORADO: EL PASO CO.: Lot 395,

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260 4 pupae (MCM); Lot 396, 2 pupae (MCM) MARYLAND: PRINCE GEORGE'S CO.: Lot 397, 4 pupae (MCM). 180. Polites mystic (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 14-2 7. 5mm, A4 transverse width 3. 1-5. lmm; brown, heart line darker brown; setae simple, to 0.1mm long on A4 dorsum; spiracles black. HEAD: Figure 3 IF, uniformly black; transverse width 2.23mm; sculpturing rough to pitted; setae simple, to 0.1mm long at apex, a few ventral setae to 0.2mm; stemmata subegual (Figure 42K) THORAX: legs dark brown; shield broad, black. ABDOMEN: prolegs each with about 65 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with a few small dark spots (Figure 53A). PUPA: Figures 71G and 92F, length 20mm, A3 transverse width 4.8mm; dark brown; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.8mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107L and 120A); lenticles present on the prothorax and proleg scars. Other Descriptions Saunders 1869a (egg, larva), Fyles 1896 (larva), Laurent 1908 (egg, larva), Dethier 1938a (egg, young larva), Dethier 1940c (larva, pupa). Host Plants CYPERACEAE: Carex species (Tietz 1972) [oviposition substrate only]. POACEAE: Agropyron repens

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261 (Tietz 1952), Bromus inermis (TLM collection), Cynodon dactylon (MCM collection, in lab), Echinochloa crusgalli (Tietz 1952), Festuca idahoensis (MacNeill 1975, possibly), Phleum pratense (Tietz 1952), Poa pratensis (MCM collection, in lab), Poa species (Shapiro 1966). Specimens Examined LARVAE : NEW YORK : BROOME CO : Lot 393, 3 larvae (instars 3, 5) (MCM). NORTH DAKOTA: CASS CO.: Lot 549, 1 larva (instar 4) (TLM). PUPAE: USA: NEW YORK: BROOME CO.: Lot 393, 1 pupa (MCM). 181. Polites sonora (Scudder) Diagnosis LAST INSTAR LARVA: BODY: length 17-2 3. 5mm, A4 transverse width 3. 8 -4. 8mm; brown, heart line darker brown; setae simple, < 0.1mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 31G, uniformly dark brown; transverse width 2. 6-2. 8mm; sculpturing rough to pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 42L) THORAX: legs dark brown; shield dark brown. ABDOMEN: prolegs each with about 65 crochets, irregularly triordinal, arranged in a circle or near circle; suranal plate rounded with a few small dark spots (Figure 53B). PUPA: Figures 71H and 92G, length 15.5-17mm, A3 transverse width 3. 6-4. lmm; blackish, abdomen brown; thoracic spiracle guard distinct; setae simple, to 0.3mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.3mm long, bluntly

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262 pointed with numerous hooked setae in a cluster at the tip (Figures 107M and 120B) ; lenticles present on the prothorax and dorsal abdomen. Other Descriptions Newcomer 1966 (egg, young larva), Emmel et al. 1992 (egg, larva, pupa). Host Plants POACEAE: Festuca idahoensis (Newcomer 1966, associated with); Poa pratensis (MCM collection, in lab). Specimens Examined LARVAE: CALIFORNIA: SAN BERNARDINO CO.: Lot 407, 1 larva (instar 5) (MCM). COLORADO: TELLER CO.: Lot 404, 7 larvae (instar 5) (MCM); Lot 405, 7 larvae (instar 5) (MCM); Lot 406, 7 larvae (instars 1, 4, 5) (MCM). PUPAE: COLORADO: TELLER CO.: Lot 404, 3 pupae (MCM); Lot 405, 5 pupae (MCM) 182. Polites vibex (Geyer) Diagnosis EGG: Figures 3D and 5B, height 0.9-lmm, width 0.6-0. 7mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: length 15-27. 5mm, A4 transverse width 4-5mm; green, heart line dark; some setae with blunt to slightly expanded tips, < 0.1mm long on A4 dorsum; spiracles black. HEAD: Figure 31H, mostly light brown with dark pits, two pale lines parallel to vertex, adfrontals pale, front with two pale spots, a pale eye patch present; transverse width 2.63mm; sculpturing rough to pitted; setae simple, < 0.1mm long at apex, a few ventral setae to 0.3mm; stemmata subegual (Figure 42M) THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about

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263 105 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with a few black spots (Figure 53C) ; wax glands forming ventral transverse patches on A7-8. PUPA: Figures 72A and 92H, length 18-21mm, A3 transverse width 4. 2-5. 3mm; green; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pi lifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending to the cremaster; abdomen moderately long; cremaster to 1.2mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 107N and 120C) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Edwards and Chapman 1879 (egg, larva, pupa), Minno and Emmel 1993 (larva, pupa). Host Plants POACEAE: Cynodon dactylon (Kendall 1965), Digitaria villosa (MCM collection), Paspalum ciliatifolium (Scudder 1889a, b), Paspalum setaceum (Edwards and Chapman 1879), Stenotaphrum secundatum (Kendall 1965). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1528, 7 eggs (MCM). NASSAU CO.: Lot 1537, 1 egg (MCM). FLORIDA: ALACHUA CO.: Lot 415, 12 larvae ( instar 2) (MCM); Lot 417, 1 larva (instar 5) (MCM); Lot 418, 7 larvae (instar 5) (MCM); Lot 1038, 1 larva (instar 5) (DHH); Lot 1080, 1 larva (instar 4) (DHH); Lot 1528, 7 larvae (instar 1) (MCM). BROWARD CO.: Lot 419, 1 larva (instar 5) (MCM); Lot 421, 6 larvae (instar 5) (MCM); Lot 422, 1 larva (instar 5) (MCM).

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264 LEVY CO.: Lot 1079, 1 larva (instar 4) (DHH). PUPAE: FLORIDA: ALACHUA CO.: Lot 418, 4 pupae (MCM) ; Lot 422, 5 pupae (MCM). BROWARD CO.: Lot 421, 6 pupae (MCM). 183. Wallengrenia otho (J. E. Smith) Diagnosis LAST INSTAR LARVA: BODY: length 162 5mm, A4 transverse width 3. 8-4. 8mm; pinkish brown, heart line dark, lateral line pale orange, well-developed on thorax, fading out on abdomen; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 32A, uniformly black; transverse width 2. 7-2. 9mm; sculpturing rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.3mm; stemmata subegual (Figure 42N) THORAX: legs dark brown; shield broad, black. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 53D). PUPA: Figures 72B and 93A, length 17-21mm, A3 transverse width 4. 1-4. 8mm; cream-colored; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.3mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 1070 and 120D) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Minno and Emmel 1993 (larva). Host Plants GENTIANACEAE : Sabatia brevi folia (Scudder 1881) [erroneous], Sabatia campanulata (Tietz 1952)

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265 [erroneous]. POACEAE: Digitaria sanguinalis (Scudder 1889a, b), Eleusine indica (MCM collection), Oryza sativa (Riley 1975), Panicum species (Lenczewski 1980), Paspalum caespitosum (Minno and Emmel 1993), Paspalum species (Lenczewski 1980), Saccharum officinarum (Riley 1975), Stenotaphrum secundatum (Kendall 1960, in lab). Specimens Examined EGGS: FLORIDA: LAKE CO.: Lot 1530, 1 egg (MCM). LARVAE: FLORIDA: COLLIER CO.: Lot 525, 1 larva (instar 4) (MCM). MONROE CO.: Lot 526, 1 larva (instar 5) (MCM); Lot 527, 8 larvae (instars 3, 5) (MCM). TEXAS: COMAL CO.: Lot 990, 3 larvae (instars 4, 5) (ROK) PUPAE: FLORIDA: MONROE CO.: Lot 527, 3 pupae (MCM); Lot 990, 1 pupa (ROK) 184. Wallengrenia egeremet (Scudder) Diagnosis LAST INSTAR LARVA: BODY: Figure 20G, length 24-26. 5mm, A4 transverse width 4. 5-4. 8mm; pinkish brown, heart line dark, lateral line very pale orange, welldeveloped on thorax, fading out on abdomen; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: uniformly black; transverse width 2. 8-2. 9mm; sculpturing rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.3mm; stemmata subegual (Figure 420). THORAX: legs dark brown; shield broad, black. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked. PUPA: Figures 72C and 93B, length 18-20mm, A3 transverse width 4.6-5mm; cream-

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266 colored; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A8; abdomen long; cremaster to 1.3mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 107P and 120E) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Scudder 1889a (egg, larva, pupa), Laurent 1908 (egg, larva), Dethier 1938a (egg, larva). Until relatively recently, this species and W. otho were considered to be conspecific, thus some of these descriptions may actually refer to the latter. Host Plants GENTIANACEAE: Sabatia gracilis (Hayward 1947) [erroneous]. IRIDACEAE: Sisyrinchium species (Draudt 1924) [erroneous]. POACEAE: Digitaria sanguinalis (Duffy and Garland 1978), Panicum clandestinum (Shapiro 1966), Panicum dichotomum (Shapiro 1974a), Paspalum setaceum (MCM collection, in lab). Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 523, 6 larvae ( instars 4, 5) (MCM); Lot 524, 3 larvae (instars 4, 5) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 523, 2 pupae (MCM); Lot 524, 2 pupae (MCM). 185. Pompeius verna (Edwards) Diagnosis LAST INSTAR LARVA: BODY: Figure 20F, length 22-24mm, A4 transverse width 4. 3-4. 7mm; green, heart line dark; setae simple, to 0.1mm long on A4 dorsum; spiracles

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267 dark brown. HEAD: Figure 32B, uniformly black, shiny; transverse width 3mm; sculpturing pitted to rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 42P) THORAX: Tl legs black, T23 legs pale; shield broad, black. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 53E). PUPA: Figures 59B, 72D, and 93C, length 20.5-21mm, A3 transverse width 4. 3-4. 5mm; cream-colored, head and dorsum of thorax black, wing veins outlined with dark brown, cremaster black; thoracic spiracle guard distinct; setae simple, to 0.8mm long on head; pi lifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.7mm long, sguarish with small irregularly sized teeth along distal margin, numerous stiff setae in a row along distal margin (Figures 108A and 120F); lenticles present on prothorax, dorsal abdomen, and proleg scars. Other Descriptions Scudder 1889a (egg, larva), Laurent 1908 (egg, larva), Dethier 1939a (egg, young larva). Host Plants POACEAE: Cynodon dactylon (MCM collection, in lab), Erianthus alopecuroides (Tietz 1952), Panicum species (Stanford 1981), Tridens flavus (Shapiro 1966). Specimens Examined LARVAE: PENNSYLVANIA: CAMBRIA CO.: Lot 432, 24 larvae ( instars 3, 4) (MCM); Lot 433, 2 larvae

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268 (instar 5) (MCM) PUPAE: PENNSYLVANIA: CAMBRIA CO.: Lot 43 3, 2 pupae (MCM) 186. Atalopedes campestris (Boisduval) Diagnosis EGG: Figure 5A, height 0.9-lmm, width 0.60.7mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: length 18-40mm, A4 transverse width 3.5-6mm; brown, heart line dark; some setae with blunt or expanded tips, < 0.1mm long on A4 dorsum; spiracles black. HEAD: Figure 32C, black with two pale lines parallel to vertex, adfrontals mostly pale, front black; transverse width 2.73.7mm; sculpturing pitted to rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subequal (Figure 42Q) THORAX: Tl-2 legs dark brown, T3 legs brown; shield broad, dark brown. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded with a few small dark spots (Figure 53F); wax glands forming ventral transverse patches on A7-8. PUPA: Figures 72E and 93D, length 18-22. 5mm, A3 transverse width 4. 3-5. 83mm; brown, abdomen cream-colored with brown spots; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.6mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 108B

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269 and 120G) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Scudder 1889a (egg, larva, pupa), Comstock 1929 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa), Emmel et al 1992 (larva, pupa), Minno and Emmel 1993 (egg, larva, pupa). Host Plants POACEAE: Cynodon dactylon (Scudder 1889a, b), Digitaria sanguinalis (Tietz 1972), Distichlis spicata (Tietz 1952), Eleusine indica (Shapiro 1966), Festuca rubra (Scott 1986), Imperata cylindrica (Bryson 1985), Poa pratensis (Garth and Tilden 1986), Stenotaphrum secundatum (Kendall 1960). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1526, 13 eggs (MCM) LARVAE: CALIFORNIA: SAN BERNARDINO CO.: Lot 574, 1 larva (instar 5) (GRB) YOLO CO.: Lot 74, 10 larvae (instars 2, 5) (MCM); Lot 81, 5 larvae (instar 5) (MCM); Lot 82, 4 larvae (instar 5) (MCM). FLORIDA: ALACHUA CO.: Lot 75, 4 larvae (instar 5) (MCM); Lot 77, 3 larvae (instar 5) (MCM); Lot 78, 3 larvae (instar 5) (MCM); Lot 1526, 8 larvae (instar 1) (MCM). BROWARD CO.: Lot 76, 3 larvae (instar 5) (MCM); Lot 79, 3 larvae (instar 5) (MCM). DADE CO.: Lot 651, 3 larvae (instar 5) (MCM). LEVY CO.: Lot 65, 1 larva (instar 2) (MCM); Lot 66, 9 larvae (instar 5) (MCM); Lot 67, 7 larvae (instar 5) (MCM); Lot 68, 6 larvae (instar 5) (MCM); Lot 69, 8 larvae (instar 5) (MCM); Lot 71, 1 larva (instar 5) (MCM); Lot 1029, 1 larva (instar

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270 5) (DHH). NEW MEXICO: HIDALGO CO.: Lot 786, 2 larvae (instar 5) (USNM) TEXAS: EDWARDS CO.: Lot 785, 5 larvae (instars 4, 5) (USNM). PUPAE: CALIFORNIA: YOLO CO.: Lot 81, 2 pupae (MCM) ; Lot 82, 2 pupae (MCM) FLORIDA: ALACHUA CO.: Lot 75, 2 pupae (MCM); Lot 77, 2 pupae (MCM); Lot 78, 2 pupae (MCM). BROWARD CO.: Lot 76, 3 pupae (MCM); Lot 79, 2 pupae (MCM). LEVY CO.: Lot 70, 6 pupae (MCM); Lot 71, 6 pupae (MCM); Lot 72, 6 pupae (MCM); Lot 73, 8 pupae (MCM). TEXAS: EDWARDS CO.: Lot 785, 1 pupa (USNM). 187. Atrytone arogos (Boisduval and Leconte) Diagnosis LAST INSTAR LARVA: BODY: length 20-29mm, A4 transverse width 2. 8-3. 7mm; green; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 3 2D, pale, medial, epicranial, and lateral lines light brown, epicranial line interrupted midway, separate from medial line; transverse width 2. 3-2. 8mm; sculpturing slightly rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.3mm; stemma 6 small (Figure 42R) THORAX: legs pale; shield narrow, a dark line between the annuli. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 53G) PUPA: Figures 72G and 94A, length 15mm, A3 transverse width 3.1mm; cream-colored, head and prothorax black; thoracic spiracle guard distinct; setae simple, to 0.2mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle

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271 leg; proboscis extending to A5; abdomen moderately long; cremaster to 0.8mm long, rounded without hooked setae (Figures 108C and 120H) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Heitzman 1966 (egg, larva, pupa). Host Plants POACEAE: Andropogon gerardii (Heitzman 1966), Andropogon glomeratus (Shapiro and Shapiro 1973, in lab), Andropogon scoparius (Shapiro and Shapiro 1973), Panicum species (Scudder 1872), Sorghastrum secundum (MCM collection) Specimens Examined LARVAE: FLORIDA: DUVAL CO.: Lot 95, 1 larva (instar 4) (MCM); Lot 652, 4 larvae (instars 3, 5) (MCM); Lot 653, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: DUVAL CO.: Lot 94, 1 pupa (MCM). 188. Atrytone del aware (Edwards) Diagnosis EGG: Figures 3F and 4F, height 1-1. 2mm, width 0.5-0. 8mm, polygonal sculpturing, green with a red ring. LAST INSTAR LARVA: BODY: Figure 21A, length 21-38mm / A4 transverse width 3. 7-5. 4mm; bluish green, heart line slightly dark; setae simple, to 0.3mm long on A4 dorsum; spiracles tan. HEAD: Figure 32E, pale, medial, epicranial, and lateral lines broad, black; transverse width 2. 7-3. 2mm; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 42S). THORAX: Tl legs dark brown, T2-3 legs pale; shield broad, black. ABDOMEN: prolegs each with about 115 crochets, irregularly

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272 triordinal, arranged in a circle or near circle; suranal plate rounded, distal margin black and a black transverse crescent (Figure 54A) PUPA: Figures 72F and 94B, length 18-25mm, A3 transverse width 3.7-5mm; cream-colored, head and prothorax black; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending beyond the cremaster; abdomen long; cremaster to 2mm long with a downward pointing spine at the tip, hooked setae absent (Figures 108D and 1201); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Edwards and Chapman 1879 (larva, pupa). Host Plants POACEAE: Andropogon gerardii (McCabe and Post 1977), Andropogon species (Shapiro 1966), Erianthus alopecuroides (Edwards and Chapman 1879), Erianthus giganteus (MCM collection), Panicum hemitomon (Minno 1992), Panicum rigidulum (MCM observation) Panicum virgatum (Shapiro 1966), Sorghastrum nutans (MCM observation), Sorghastrum secundum (MCM collection) Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1518, 25 eggs (MCM). LARVAE: FLORIDA: ALACHUA CO.: Lot 102, 1 larva (instar 4) (MCM); Lot 658, 1 larva (instar 4) (MCM); Lot 1030, 1 larva (instar 3) (DHH); Lot 1031, 1 larva (instar 5) (DHH); Lot 1032, 1 larva (instar 5) (DHH); Lot 1518, 8 larvae (instar 1) (MCM). BROWARD CO.: Lot 107, 1 larva (instar 4) (MCM); Lot 108, 1 larva (instar 4) (MCM).

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273 DUVAL CO.: Lot 656, 1 larva ( instar 5) (MCM) HIGHLANDS CO.: Lot 10, 1 larva (instar 5) (MCM); Lot 98, 1 larva (instar 5) (MCM); Lot 99, 2 larvae (instars 3, 5) (MCM); Lot 100, 5 larvae (instar 1) (MCM); Lot 655, 1 larva (instar 5) (MCM). LEVY CO.: Lot 113, 2 larvae (instar 4) (MCM); Lot 114, 1 larva (instar 5) (MCM). MARION CO.: Lot 103, 2 larvae (instar 5) (MCM); Lot 110, 2 larvae (instar 5) (MCM). PUTNAM CO.: Lot 101, 2 larvae (instar 5) (MCM); Lot 111, 1 larva (instar 4) (MCM); Lot 654, 1 larva (instar 5) (MCM). SARASOTA CO.: Lot 106, 2 larvae (instar 5) (MCM). SOUTH CAROLINA: BEAUFORT CO.: Lot 112, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 109, 1 pupa (MCM). DUVAL CO.: Lot 97, 1 pupa (MCM); Lot 115, 1 pupa (MCM). HIGHLANDS CO.: Lot 99, 4 pupae (MCM); Lot 117, 1 pupa (MCM). LEVY CO.: Lot 105, 1 pupa (MCM); Lot 116, 1 pupa (MCM) 189. Problema byssus (Edwards) Diagnosis EGG: Figure 4G, height 1.4-1. 6mm, width 1mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: Figure 21B, length 24-43. 5mm, A4 transverse width 4. 5-6. 2mm; bluish green with a yellowish cast to the thorax and posterior segments, heart line dark; setae simple, to 0.3mm long on A4 dorsum; spiracles tan. HEAD: Figure 32F, pale, medial, epicranial, and lateral lines broad, black, front with a few black stripes, epicranial line largely separate from medial line, sometimes with some small lateral

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274 connections, posterior of head black; transverse width 2.83.8mm; sculpturing rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 42T) THORAX: Tl legs tan to brown, T2-3 legs pale; shield moderately broad, black. ABDOMEN: prolegs each with about 165 crochets, irregularly triordinal arranged in a near circle; suranal plate rounded, unmarked (The suranal plate of earlier instars is mostly black or bears a black crescent.) (Figures 54B and C); wax glands forming ventral transverse patches on A7-8 (Figure 15A) PUPA: Figures 72H and 94C, length 26-28. 5mm, A3 transverse width 5. 5-6. 3mm; cream-colored with brown markings on the head and thorax; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A5; abdomen long; cremaster to 2.1mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 108E and 120J) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Heitzman 1965b (egg, larva, pupa). Host Plants POACEAE: Andropogon gerardii (Scott 1986), Chasmanthium sessiliflorum (MCM collection), Erianthus alopecuroides (MCM collection), Erianthus giganteus (MCM collection), Tripsacum dactyloides (Heitzman 1965a). Specimens Examined EGGS: FLORIDA: PUTNAM CO.: Lot 1521, 2 eggs (MCM). LARVAE: GEORGIA: HARRIS CO.: Lot 7, 2

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275 larvae ( instar 5) (MCM) FLORIDA: ALACHUA CO.: Lot 435, 1 larva (instar 5) (MCM); Lot 437, 1 larva (instar 5) (MCM); Lot 442, 2 larvae ( instars 3, 4) (MCM); Lot 443, 3 larvae (instars 3, 5) (MCM); Lot 1069, 1 larva (instar 4) (DHH). DUVAL CO.: Lot 1068, 1 larva (instar 4) (DHH). FLAGLER CO.: Lot 440, 1 larva (instar 3) (MCM). LAKE CO.: Lot 441, 4 larvae (instars 2, 3) (MCM); Lot 444, 1 larva (instar 5) (MCM); Lot 445, 3 larvae (instar 5) (MCM); Lot 446, 5 larvae (instars 3, 5) (MCM); Lot 447, 3 larvae (instar 5) (MCM). LEVY CO.: Lot 438, 2 larvae (instars 3, 4) (MCM). MISSOURI: JACKSON CO.: Lot 566, 3 larvae (instar 5) ( JRH) SOUTH CAROLINA: GEORGETOWN CO.: Lot 436, 2 larvae (instar 5) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 435, 1 pupa (MCM); Lot 443, 2 pupae (MCM). LAKE CO.: Lot 444, 1 pupa (MCM). GEORGIA: HARRIS CO.: Lot 7, 1 pupa (MCM). 190. Problema Jbulenta (Boisduval and Leconte) Cromartie and Schweitzer (1993) described the egg and larva of this species, but I could not find specimens for study Host Plants POACEAE: Phragmites australis Spartina alterniflora (Cromartie and Schweitzer 1993, possibly), Spartina cynosuroides (Cromartie and Schweitzer 1993), Zizaniopsis miliacea (Opler and Krizek 1984, associated with).

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276 191. Ochlodes sylvanoides (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 20-24mm, A4 transverse width 4. 4-5. lmm; preserved specimens pale; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 32G, pale, medial and lateral lines black, front pale, posterior of head black; transverse width 2. 4-2. 8mm; sculpturing slightly rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 42U) THORAX: legs pale; shield narrow, a dark line between the annuli. ABDOMEN: prolegs each with about 75 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with a few small black spots. PUPA: Figures 73A and 94D, length 16-28mm, A3 transverse width 3. 5-4. 3mm; cream-colored; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 1.5mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 108F and 121A); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Corns tock 1929 (larva, pupa), Corns tock 1930a (egg), Emmel and Emmel 1973 (larva, pupa). Host Plants POACEAE: Agropyron caninum ssp. ma jus (Scott 1986), Cynodon dactylon (Scott 1986), Elymus cinereus (Scott

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277 1986), Elymus glaucus (MCM collection), Phalaris californica (Shapiro 1974b), Phalaris lemmoni (Shapiro 1974b). Specimens Examined LARVAE: CALIFORNIA: RIVERSIDE CO.: Lot 344, 1 larva (instar 5) (MCM); Lot 704, 3 larvae ( instar 5) (MCM). PUPAE: CALIFORNIA: RIVERSIDE CO.: Lot 704, 2 pupae (MCM) 192. Ochlodes agricola (Boisduval) Diagnosis LAST INSTAR LARVA: BODY: length 19.5mm, A4 transverse width 4.1mm; preserved specimen pale; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 32H, pale, medial and lateral lines black, posterior of head black; transverse width 2.6mm; sculpturing rough to pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemmata subequal (Figure 42V). THORAX: legs pale; shield narrow, a dark line between the annuli. ABDOMEN: prolegs each with about 75 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked; wax glands forming two ventrolateral spots on A7-8 (Figure 15F) Other Descriptions Comstock 1927a (egg, young larva), Comstock 1930a (egg), Emmel and Emmel 1973 (egg, larva). Specime ns Examined LARVAE: CALIFORNIA: SOLANO CO.: Lot 343, 1 larva (instar 5) (MCM). 193. Ochlodes snowi (Edwards) The immature stages of this western species are undescribed, and I could not find specimens for study.

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278 Host Plants POACEAE: Blepharoneuron tricholepis (Scott and Scott 1980) 194. Ochlodes yuma (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 32-35mm, A4 transverse width 4 .7-5. 7mm; preserved specimens pale; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 33A, pale, medial and lateral lines black, posterior of head black; transverse width 3-3. 3mm; sculpturing slightly rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 42W) THORAX: legs pale; shield narrow, a dark line between the annuli. ABDOMEN: prolegs each with about 135 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with some tiny dark spots (Figure 54D) Other Descriptions Emmel and Emmel 1973 (egg, pupa), MacNeill 1975 (egg, larva, pupa). Host Plants POACEAE: Phragmites australis (Tilden 1965). Specimens Examined LARVAE: CALIFORNIA: INYO CO.: Lot 345, 2 larvae (instar 5) (MCM) 195. Poanes massasoit (Scudder) Laurent (1908) described the egg and young larva of this northeastern species, but I could not find specimens for study. Host Plants CYPERACEAE: Carex aquatilis (Layberry et al 1982, probably), Carex stricta (Shapiro and Shapiro 1973,

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279 probably; Shapiro 1974a). POACEAE: (Pyle 1981) [erroneous] 196. Poanes hobomok (Harris) Diagnosis LAST INSTAR LARVA: BODY: length 29mm, A4 transverse width 5.5mm; preserved specimen pale; setae simple, to 0.3mm long on A4 dorsum; spiracles brown. HEAD: Figure 33B, brown, darker brown at vertex, scattered pits on upper face dark brown; transverse width 3.4mm; sculpturing rough to pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 42X) THORAX: legs tan; shield narrow, a brown line between the annuli. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, with tiny brown spots (Figure 54E); wax glands forming ventral transverse bands on A7-8. PUPA: Figures 73B and 95A, length 24mm, A3 transverse width 6mm; preserved specimen pale; thoracic spiracle guard distinct; setae simple, to 0.7mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.8mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 108G and 121B) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Saunders 1869a (egg, young larva), Scudder 1881 (egg, larva), French 1886 (egg, larva), Scudder 1889a (egg, larva, pupa). In the past, this species and P.

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280 zabulon were considered to be conspecific, thus some of these descriptions may actually refer to the latter. Host Plants POACEAE: Panicum species (Shapiro 1966), Poa species (Shapiro 1966). Specimens Examined LARVAE: WISCONSIN: WAUKESHA CO.: Lot 372, 1 larva ( instar 5) (MCM) PUPAE: WISCONSIN: WAUKESHA CO.: Lot 372, 1 pupa (MCM). 197. Poanes zabulon (Boisduval and Leconte) Diagnosis LAST INSTAR LARVA: BODY: length 12-23mm, A4 transverse width 3. 2-4. 7mm; preserved specimens brown; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 3 3C, brown; transverse width 2. 5-2. 3mm; sculpturing rough to pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.3mm; stemmata subequal (Figure 42Y). THORAX: legs tan; shield narrow, a brown line between the annuli. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with some tiny brown spots. Other Descriptions Laurent 1908 (egg, larva). Host Plants POACEAE: Eragrostis species (Shapiro 1966), Tridens species (Shapiro 1966). Specimens Examined LARVAE: PENNSYLVANIA: MONTGOMERY CO.: Lot 1001, 6 larvae (instar 5) (FSCA). 198. Poanes taxiles (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 21-27mm, A4 transverse width 4. 7-5. 3mm; pinkish brown, heart line dark,

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281 subdorsal and lateral lines faint, outlined by dark bands; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 3 3D, light brown with darker brown pits on upper face; transverse width 3. 1-3. 3mm; sculpturing rough to pitted; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 43A) THORAX: legs pale; shield narrow, a brown line between the annuli. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with some tiny dark spots. PUPA: Figures 7 3C and 95B, length 18-21. 5mm, A3 transverse width 4. 1-4. 9mm; creamcolored, abdomen with dark spots, head and thorax with dark markings; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A8; abdomen long; cremaster to 1.7mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 108H and 121C); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Scott 1986 (larva, pupa). Host Plants POACEAE: Agropyron repens (Scott 1986), Agropyron smithii (Scott and Scott 1980), Agropyron trachycaulum (Scott 1986), Agrostis gigantea (Scott 1986), Bromus anomalus (Bailowitz and Brock 1991), Dactylis glomerata (Scott 1986), Dactylis species (Stanford 1981), Elymus arizonicus (Bailowitz and Brock 1991), Elymus

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282 canadensis (Scott 1986), Poa pratensis (Scott 1986), Puccinellia airoides (Scott and Scott 1980). Specimens Examined LARVAE : COLORADO : EL PASO CO : Lot 374, 5 larvae ( instar 5) (MCM) ; Lot 375, 5 larvae ( instar 5) (MCM); Lot 376, 1 larva (instar 5) (MCM); Lot 377, 7 larvae (instars 1, 3, 5) (MCM); Lot 378, 5 larvae (instar 5) (MCM). PUPAE: COLORADO: EL PASO CO.: Lot 373, 10 pupae (MCM); Lot 374, 4 pupae (MCM); Lot 375, 2 pupae (MCM); Lot 378, 4 pupae (MCM) 199. Poanes aaroni (Skinner) Diagnosis LAST INSTAR LARVA: BODY: length 3 3mm, A4 transverse width 4.8mm; pinkish brown, heart line dark, subdorsal and lateral lines bounded by dark bands; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 33E, light brown, medial and epicranial lines black, convergent at midadfrontals, area around stemmata black, posterior of head black; transverse width 2.9mm; sculpturing rough; setae simple, to 0.3mm long at apex, a few ventral setae to 1mm; stemmata subegual (Figure 43B) THORAX: legs pale; shield narrow, a dark brown line between the annul i. ABDOMEN: prolegs each with about 95 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with tiny dark spots. PUPA: Figures 73D and 95C, length 22mm, A3 transverse width 4.3mm; cream-colored, head and prothorax blackish, faint brown spots and markings on thorax and abdomen; thoracic spiracle guard distinct; setae simple,

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283 to 0.5mm long on head; pi lifers touching; head rounded, cap somewhat bulbous; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen moderately long; cremaster to 1.2mm long, sharply angled laterally, distal tip with a mesal indentation, hooked setae absent (Figures 1081 and 121D) ; lenticles present on the prothorax, dorsum of abdomen, and proleg scars. Other Descriptions Laurent 1908 (egg, young larva). Host Plants POACEAE : Panicum hemitomon (MCM collection, associated with, in lab) Spartina alternifolia (Clark and Clark 1951, associated with). Specimens Examined LARVAE: FLORIDA: LAKE CO.: Lot 995, 5 larvae (instars 3, 4, 5) (MCM). PUPAE: FLORIDA: LAKE CO.: Lot 995, 1 pupa (MCM). 200. Poanes yehl (Skinner) Diagnosis EGG: Figure 4H, height 1-1. 2mm, width 0.60.8mm, polygonal sculpturing, brownish white. LAST INSTAR LARVA: BODY: length 3 0-3 4mm, A4 transverse width 5.25.5mm; pinkish brown, heart line dark, subdorsal line bounded by faint dark bands; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 33F, black, orangebrown eye patches present; transverse width 3. 3-3. 5mm; sculpturing rough; setae simple, to 0.5mm long at apex, a few ventral setae to 1mm; stemmata subegual (Figure 43C). THORAX: legs pale; shield narrow, a dark band between the annuli. ABDOMEN: prolegs each with about 160 crochets,

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284 irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, tan with tiny dark spots; no wax glands. PUPA: Figures 59C, 73E, and 95D, length 20.5mm, A3 transverse width 4.3mm; cream-colored, head and dorsum of thorax blackish, a few dark spots on abdomen; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers touching; head rounded, cap somewhat bulbous; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 1mm long, sharply angled laterally, distal tip with a mesal indentation, hooked setae absent (Figures 108J and 121E); lenticles present on the prothorax, dorsum of abdomen, and proleg scars. Host Plants POACEAE: Arundinaria gigantea (Scott 1986; MCM collection, in lab). Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1523, 2 eggs (MCM). LARVAE: FLORIDA: ALACHUA CO.: Lot 383, 3 larvae (instar 1) (MCM); Lot 1509, 3 larvae (instar 5) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 1509, 1 pupa (MCM). 201. Poanes viator (Edwards) Diagnosis EGG: height 1.1-1. 2mm, width 0.7-0. 8mm, polygonal sculpturing, brownish white. LAST INSTAR LARVA: BODY: length 20-4 3mm, A4 transverse width 3. 7-5. 5mm; pinkish brown, heart line dark, subdorsal line bounded by dark bands; setae simple, to 0.3mm long on A4 dorsum;

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285 spiracles tan. HEAD: Figure 33G, light brown, medial line black, scattered small dark spots on face, area around the stemmata black; transverse width 3. 1-3. 9mm; sculpturing rough; setae simple, to 0.3mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 43D) THORAX: legs pale; shield narrow, a dark brown line between the annuli. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, tan with small dark spots. PUPA: Figures 73F and 96A, length 22-32mm, A3 transverse width 4. 2-5. 6mm; cream-colored, head and dorsum of thorax blackish, abdomen with dark spots; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers touching; head rounded, cap bulbous, lobed along the margin; antennal tip lies far cephalad of tip of middle leg; proboscis extending slightly beyond the wings in A4; abdomen moderately long; cremaster to 1.8mm long, sharply angled laterally, distal tip with a mesal indentation, hooked setae absent (Figures 109A and 121F); lenticles present on the prothorax, dorsum of abdomen, and proleg scars. Other Descriptions Laurent 1908 (egg, young larva), Pyle 1981 (egg, larva). Host Plants CYPERACEAE: Carex lacustris (Shapiro 1971), Carex rostrata (Layberry et al 1982), Cyperus esculentus (MCM collection, in lab). POACEAE: Cynodon dactylon (MCM collection, in lab), Panicum species (Pyle 1981), Phragmites

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286 australis (Shapiro 1966, suspected: Shapiro and Shapiro 1973), Zizania aquatica (Forbes 1960, associated with; Shapiro 1966), Zizaniopsis miliacea (Kendall 1966b). Specimens Examined EGGS: TEXAS: GUADALUPE CO.: Lot 966, 8 eggs (MCM) LARVAE: FLORIDA: Lot 1289, 1 larva (instar 4) (DHH). ALACHUA CO.: Lot 379, 2 larvae (instars 2, 3); Lot 380, 2 larvae (instar 5) (MCM); Lot 707, 5 larvae (instar 5) (MCM); Lot 382, 9 larvae (instars 1, 3, 4) (MCM); Lot 1070, 1 larva (instar 5) (DHH); Lot 1071, 1 larva (instar 1) (DHH); Lot 1072, 1 larva (instar 5) (DHH). NORTH DAKOTA: RICHLAND CO.: Lot 544, 3 larvae (instar 1). SOUTH CAROLINA: BEAUFORT CO.: Lot 381, 1 larva (instar 5) (MCM). TEXAS: GUADALUPE CO.: Lot 966, 1 larva (instar 5) (ROK) PUPAE: FLORIDA: ALACHUA CO.: Lot 380, 1 pupa (MCM). TEXAS: GUADALUPE CO.: Lot 966, 3 pupae (ROK). 202. Paratrytone melane (Edwards) Diagnosis EGG: height 0.7-0. 8mm, width 1.1-1. 2mm, polygonal sculpturing. LAST INSTAR LARVA: BODY: length 12-29. 5mm, A4 transverse width 3. 9-5. 3mm; preserved specimens pale; setae simple, to 0.1mm long on A4 dorsum, spiracles tan. HEAD: Figure 3 3H, brown; transverse width 3-3. 8mm; sculpturing rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 43E) THORAX: legs pale; shield narrow, a brown line between the annuli. ABDOMEN: prolegs each with about 110 crochets, irregularly triordinal, arranged in a circle;

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287 suranal plate rounded, tan with some small dark spots (Figure 54F) PUPA: Figures 73G and 96B, length 21mm, A3 transverse width 4.3mm; preserved specimens pale, brown markings on the head and thorax, abdomen with brown spots; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.8mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures 109B and 121G) ; lenticles on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Comstock and Dammers 1931 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants CYPERACEAE: Carex spissa (Brown 1984). POACEAE: Bromus carinatus (Scott 1986), Bromus inermis (Tietz 1972), Cynodon dactylon (Wright 1905), Deschampsia caespitosa (Emmel and Emmel 1973), Digitaria ischaemum (Tietz 1972), Digitaria sanguinalis (Tietz 1972), Lamarckia aurea (Brown 1984), Stenotaphrum secundatum (Brown 1984). Specimens Examined EGGS: MEXICO: NUEVO LEON: Lot 960, 4 eggs (ROK). LARVAE: MEXICO: NUEVO LEON: Lot 960, 13 larvae (instars 1, 3, 5) (ROK). USA: CALIFORNIA: Lot 639, 1 larva (instar 5) (TCE). PUPAE: CALIFORNIA: LOS ANGELES CO.: Lot 639, 1 pupa (TCE).

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288 203. Choranthus radians (Lucas) Dethier (1939b) described the egg and larva of this Caribbean species, but I could not find specimens for study. Host Plants POACEAE: Saccharum officinarum (Dethier 1939b) 204. Choranthus haitensis (Skinner) Diagnosis LAST INSTAR LARVA: BODY: length 28.5mm, A4 transverse width 3.5mm; light green, heart line dark; setae simple, to 0.2mra long on A4 dorsum; spiracles tan. HEAD: Figure 34A, light brown, epicranial and lateral lines black, posterior of head black; transverse width 2.3mm; sculpturing rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 43F) THORAX: legs pale; shield broad, black. ABDOMEN: prolegs each with about 145 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded with a large black patch (Figure 55A); wax glands forming ventral spots posterior of the prolegs on A3-6. PUPA: Figures 7 3H and 96C, length 1923mm, A3 transverse width 3. 7-4. 2mm; cream-colored; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 1.3mm long, rounded with a few short dorsolateral spines, numerous hooked setae in a row along distal margin (Figures 109C and 121H) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars.

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289 Other Descriptions Wolcott 1921 (egg, larva, pupa). Host Plants POACEAE: Panicum maximum (Minno 1990), Saccharum officinarum (Wolcott 1921). Specimens Examined LARVAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 125, 7 larvae (instars 2, 4) (MCM) PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 125, 2 pupae (MCM). 205. Mellana eulogius (Plotz) This nonresident species from tropical America has been reported as feeding on sugar cane {Saccharum officinarum) (Dyar 1914), but the immature stages are undescribed. 206. Mellana mexicana (Bell) Nothing is known of the biology of this neotropical species. 207. Euphyes arpa (Boisduval and Leconte) Diagnosis LAST INSTAR LARVA: BODY: Figure 2 ID, length 22.5-42mm, A4 transverse width 4. 3-6. 5mm; green, reticulated with pale yellow, bases of setae black; setae simple, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 34B, pale, white on upper face, medial line modified into an elongate spot, epicranial line brown, becoming black near the apex, lateral line and posterior of head brown; transverse width 3. 5-4. 2mm; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemma 5 and 6 small (Figure 43G) THORAX: legs pale; anterior shield white, posterior black. ABDOMEN: prolegs each with about 140 crochets, irregularly

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290 triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 55B) ; wax glands forming ventral transverse patches on A7-8. PUPA: Figures 74A and 96D, length 23-28mm, A3 transverse width 5. 6-5. 8mm; light brown; thoracic spiracle guard distinct; setae simple, to 0.5mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 1.3mm long, rounded with a pair of dorsolateral spines, hooked setae absent (Figures 109D and 1211); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Edwards and Chapman 1879 (larva, pupa), Minno and Emmel 1993 (egg, larva). Host Plants ARECACEAE: Sabal etonia (Minno 1992), Sabal palmetto (Hayward 1947) [erroneous], Sabal species (Forbes 1960) [erroneous], Serenoa repens (Edwards and Chapman 1879). CYPERACEAE: Cladium jamaicense (Tietz 1972) [erroneous], Cyperus esculentus (MCM collection, in lab). Specimens Examined LARVAE: FLORIDA: Lot 1034, 1 larva (instar 5) (DHH). LEVY CO.: Lot 210, 9 larvae (instars 1, 3, 4) (MCM); Lot 211, 1 larva (instar 4) (MCM); Lot 1033, 1 larva (instar 5) (DHH). HIGHLANDS CO.: Lot 212, 4 larvae (instars 3, 5) (MCM); Lot 214, 3 larvae (instar 5) (MCM), Lot 215, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: HIGHLANDS CO.: Lot 213, 2 pupae (MCM).

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291 208. Euphyes pilatka (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 3 6-4 6mm, A4 transverse width 4. 2-5. 5mm; green, reticulated with pale yellow, bases of setae black; setae simple, to 0.2mm long on A4 dorsum; spiracles dark brown. HEAD: Figure 34C, pale, white on upper face, medial line modified into an elongate spot, epicranial line brown, becoming black near the apex, lateral line and posterior of head brown; transverse width 3.1-4mm; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemma 5 and 6 small (Figure 43H) THORAX: legs pale; anterior shield white, posterior black. ABDOMEN: prolegs each with about 120 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked; wax glands forming ventral transverse patches on A7-8. PUPA: Figures 74B and 97A, length 28mm, A3 transverse width 5.3mm; brown; thoracic spiracle guard distinct; setae simple, to 0.3mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A4; abdomen long; cremaster to 1.3mm long, rounded with a pair of dorsolateral spines, hooked setae absent (Figures 109E and 121J) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other D escriptions Edwards and Chapman 1879 (larva), Minno and Emmel 1993 (egg, larva).

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292 Host Plants CYPERACEAE: Cladium jamaicense (Edwards and Chapman 1879; Minno and Emmel 1993). Specimens Examined LARVAE: FLORIDA: BROWARD CO.: Lot 221, 16 larvae (instar 1) (MCM) ; Lot 222, 1 larva (instar 5) (MCM) MARION CO.: Lot 220, 1 larva (instar 5) (MCM). MONROE CO.: Lot 225, 3 larvae (instar 5) (MCM); Lot 673, 1 larva (instar 5) (MCM). PUTNAM CO.: Lot 674, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: MONROE CO.: Lot 223, 1 pupa (MCM). 209. Euphyes dion (Edwards) I could not find specimens of the immature stages of this eastern species for study. Other Descriptions Laurent 1908 (egg, young larva), Scott 1986 (egg, larva, pupa). Host Plants CYPERACEAE: Car ex emoryi (Shuey 1986), Car ex hyalinolepis (Opler and Krizek 1984), Carex lacustris (Shapiro 1971), Carex walteriana (Clark and Clark 1951, associated with; Opler and Krizek 1984), Scirpus species (Shapiro 1966) 210. Euphyes alabanae Diagnosis LAST INSTAR LARVA: BODY: length 38-45mm, A4 transverse width 4. 8-5. 8mm; green, reticulated with pale white; setae simple, to 0.2mm long on A4 dorsum; spiracles black. HEAD: Figure 34D, pale, medial line modified into an elongate spot, epicranial and lateral lines brown, posterior of head brown; transverse width 3.5-4mm;

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293 sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.6mm; stemma 5 and 6 small (Figure 431). THORAX: legs pale; anterior shield white, posterior black. ABDOMEN: prolegs each with about 125 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked; wax glands forming ventral transverse patches on A7-8. PUPA: Figures 74C and 97B, length 23.5mm, A3 transverse width 5.3mm; brown; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 1.5mm long, rounded with a pair of dorsolateral spines, hooked setae absent (Figures 109F and 121K) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Host Plants CYPERACEAE: Carex emoryi (ROK collection), Carex lacustris (Harris 1972), Cyperus esculentus (MCM collection, in lab), Scirpus cyperinus (Harris 1972). Specimens Examined LARVAE: FLORIDA: LEVY CO.: Lot 218, 1 larva (instar 3) (MCM). SOUTH CAROLINA: BEAUFORT CO.: Lot 219, 2 larvae (instar 5) (MCM). TEXAS: TARRENT CO.: Lot 941, 1 larva (instar 5) (ROK). PUPAE: SOUTH CAROLINA: BEAUFORT CO.: Lot 219, 1 pupa (MCM). 211. Euphyes dukesi (Lindsey) Diagnosis LAST INSTAR LARVA: BODY: length 4 0-4 7mm, A4 transverse width 5. 1-5. 8mm; green, reticulated with pale

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294 white, bases of setae black; setae simple, to 0.3mm long on A4 dorsum; spiracles black. HEAD: Figure 34E, pale, white on upper face, medial line modified into an elongate spot, epicranial and lateral lines brown, posterior of head brown; transverse width 3.5-4mm; sculpturing slightly rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.6mm; stemma 6 small (Figure 43J) THORAX: legs pale; anterior shield white, posterior black. ABDOMEN: prolegs each with about 110 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked; wax glands forming ventral transverse patches on A7-8. PUPA: Figures 74D and 97C, length 24-25mm, A3 transverse width 5. 3-5. 6mm; brown; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 1.3mm long, rounded with a pair of dorsolateral spines, hooked setae absent (Figures 109G and 121L); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Host Plants CYPERACEAE: Carex hyalinolepis (Irwin and Downey 1973, probably; Opler and Krizek 1984), Carex lacustris (Forbes 1960, perhaps; Opler and Krizek 1984), Carex lupulina (MCM collection, in lab), Carex walteriana (Scott 1986), Rhynchosia inundata (MCM collection).

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295 Specimens Examined LARVAE: FLORIDA: PASCO CO.: Lot 672, 4 larvae (instar 5) (MCM) PUPAE: FLORIDA: PASCO CO.: Lot 669, 1 pupa (MCM); Lot 670, 4 pupae (MCM); Lot 671, 3 pupae (MCM) 212. Euphyes conspicuus (Edwards) The immature stages of this midwestern species are undescribed, and I could not locate specimens for study. Host Plants CYPERACEAE: Carex stricta (Clark and Clark 1951) 213. Euphyes berry i (Bell) Diagnosis PUPA: Figures 74E and 97D, length 23.5mm, A3 transverse width 4.3mm; brown; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen long; cremaster to 1.5mm long, rounded with a pair of dorsolateral spines, hooked setae absent (Figures 109H and 121M); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Host Plants CYPERACEAE: Carex species (Shuey 1986, probably), Cyperus esculentus (MCM collection, in lab). Specimens Examined EGGS: FLORIDA: VOLUSIA CO.: Lot 1036, 1 egg (DHH) LARVAE: FLORIDA: VOLUSIA CO.: Lot 1037, 2 larvae (instar 1) (DHH); Lot 1539, 9 larvae (instar 1) (MCM). PUPAE: FLORIDA: VOLUSIA CO.: Lot 217, 1 pupa (MCM)

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296 214. Euphyes macguirei H. A. Freeman Diagnosis LARVAL EXUVIUM: stemmata subequal (Figure 43K) PUPAL EXUVIUM: cremaster to 1.1mm long, rounded with a pair of dorsolateral spines, hooked setae absent with numerous hooked setae at tip (Figures 1091 and 122A) Host Plants CYPERACEAE: Carex emoryi (ROK collection). 215. Euphyes bimacula (Grote and Robinson) Diagnosis LAST INSTAR LARVA: BODY: length 18-2 5mm, A4 transverse width 3.9-4mm; preserved specimens brownish; setae simple, to 0.1mm long on A4 dorsum; spiracles black. HEAD: Figure 34F, pale, white on upper face, medial line modified into an elongate spot, epicranial and lateral lines brown, posterior of head brown; transverse width 2. 93.2mm ; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subequal (Figure 43L) THORAX: legs pale; anterior shield white, posterior black. ABDOMEN: crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked. PUPA: Figures 74F and 97E, length 17.5mm, A3 transverse width 3.8mm; brown; thoracic spiracle guard distinct; setae simple, to 0.3mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen moderately long; cremaster to 0.7mm long, rounded with a pair of dorsolateral spines, hooked setae absent

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297 (Figures 109J and 122B) ; lenticles present on the prothorax and proleg scars. Other Descriptions Laurent 1908 (egg), Scott 1986 (egg, larva) Host Plants CYPERACEAE: Carex trichocarpa (Shapiro and Shapiro 1973). POACEAE: (Tietz 1952) [erroneous]. Specimens Examined LARVAE: PENNSYLVANIA: TIOGA CO.: Lot 1007, 4 larvae ( instars 4, 5) (FSCA). PUPAE: PENNSYLVANIA: TIOGA CO.: Lot 1007, 1 pupa (FSCA). 216. Euphyes ruricola (Boisduval) Diagnosis EGG: Figures 3G, 41 and 5E, height 1.1-1. 3mm, width 0.7-0. 8mm, polygonal sculpturing, green with a red ring. LAST INSTAR LARVA: BODY: length 19-34mm, A4 transverse width 3. 5-5. 2mm; green, reticulated with pale white, heart line dark, bases of setae black; setae simple, to 0.2mm long on A4 dorsum; spiracles black. HEAD: Figure 34G, pale, white on upper face, medial line modified into an elongate spot, epicranial and lateral lines brown, posterior of head black; transverse width 2. 7-3. 4mm; sculpturing slightly rough; setae simple, to < 0.1mm long at apex, a few ventral setae to 0.4mm; stemma 6 small (Figure 43M) THORAX: legs pale; anterior shield white, posterior black. ABDOMEN: prolegs each with about 120 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked or sometimes distal margin edged with black and bearing a black crescent (Figure 55C); wax glands

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298 forming ventral transverse patches on A7-8. PUPA: Figures 75A and 97F, length 17.5-22mm, A3 transverse width 3.94.3mm; head and thorax blackish, abdomen cream-colored; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen long; cremaster to 1.2mm long, rounded with a pair of dorsolateral spines, hooked setae absent (Figures 109K and 122C); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Moffat 1894 (larva, pupa), Heitzman 1964a (egg, larva, pupa), Brown and McGuire 1983 (egg, larva, pupa), Emmel et al 1992 (egg, larva). Host Plants CYPERACEAE: Carex geophila (Scott 1986), Car ex heliophila (Scott and Scott 1980), Carex lacustris (Scott 1986), Carex lupulina (MCM collection), Carex spissa (Brown and McGuire 1983), Carex stricta (Shuey 1986), Cyperus esculentus (Heitzman 1964a, in lab; MCM collection, in lab), Cyperus species (Emmel and Emmel 1973). POACEAE: (Moffat 1894, in lab), Tridens flavus (Shapiro 1966) [guestionable] Specimens Examined EGGS: FLORIDA: DUVAL CO.: Lot 1519, 10 eggs (MCM). LARVAE: CALIFORNIA: SAN DIEGO CO.: Lot 1011, 1 larva (instar 5) (SDNHM); Lot 1012, 1 larva (instar 5) (SDNHM); Lot 1013, 1 larva (instar 5) (SDNHM); Lot 1014, 1 larva (instar 5) (SDNHM); Lot 1015, 1 larva (instar 3)

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299 (SDNHM); Lot 1016, 1 larva ( instar 2) (SDNHM). COLORADO: EL PASO CO.: Lot 237, 6 larvae (instars 4, 5); Lot 239, 1 larva (instar 4) (MCM) FLORIDA: ALACHUA CO.: Lot 236, 2 larvae (instar 5) (MCM). DUVAL CO.: Lot 1519, 7 larvae (instar 1) (MCM). PUPAE: COLORADO: EL PASO CO.: Lot 237, 1 pupa (MCM). FLORIDA: ALACHUA CO.: Lot 235, 1 pupa (MCM); Lot 236, 3 pupae (MCM). 217. Asbolis capucinus (Lucas) Diagnosis LAST INSTAR LARVA: BODY: Figure 21C, length 25-57. 5mm, A4 transverse width 4. 8-8. 2mm; yellowish green, heart line dark; some setae with blunt tips, to 0.3mm long on A4 dorsum; spiracles black. HEAD: Figure 34H, pale orange; transverse width 3. 2-4. 4mm; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 43N) THORAX: legs pale; shield narrow, a black line between the annuli. ABDOMEN: prolegs each with about 235 crochets, irregularly triordinal, arranged in a near circle; suranal plate broad, rounded, edged with black (Figure 55D) ; wax glands forming small ventral spots posterior of the prolegs on A3-6 (Figure 15D) PUPA: Figures 75B and 98A, length 28-35. 5mm, A3 transverse width 5-7mm; cream-colored to pale green, head and thorax blackish; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers separated; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to

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300 2.8mm long, broadly rounded with numerous hooked setae in a row along distal margin (Figures 109L and 122D) ; lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Minno and Emmel 1993 (larva, pupa). Host Plants ARECACEAE: Acoelorrhaphe wrightii (Kimball 1965), Chrysalidocarpus lutescens (Minno and Emmel 1993), Coccothrinax argentata (Minno and Emmel 1993), Cocos nucifera (Bruner et al 1945), Dictyosperma album (DHH collection), Dictyosperma rubrum (Bruner et al. 1945), Phoenix dactylifera (Opler and Krizek 1984), Phoenix species (Kimball 1965), Roystonea regia (Bruner et al 1945), Sabal etonia (Minno 1992), Sabal palmetto (MacNeill 1975), Sabal species (Bruner et al. 1945), Serenoa repens (Minno and Emmel 1993), Thrinax radiata (Minno and Emmel 1993), Veitchia merrillii (Minno and Emmel 1993). Specimens Examined LARVAE: FLORIDA: BROWARD CO.: Lot 86, 1 larva ( instar 5) (MCM) ; Lot 87, 1 larva ( instar 5) (MCM). COLLIER CO.: Lot 1026, 1 larva (instar 5) (DHH). DADE CO.: Lot 92, 1 larva (instar 5) (MCM); Lot 660, 1 larva (instar 5) (MCM); Lot 789, 1 larva (instar 5) (USNM); Lot 791, 1 larva (instar 5) (USNM); Lot 1024, 1 larva (instar 5) (DHH); Lot 1025, 4 larvae ( instars 3, 5) (DHH). HIGHLANDS CO.: Lot 84, 1 larva (instar 5) (MCM). MONROE CO.: Lot 85, 1 larva (instar 5) (MCM); Lot 90, 2 larvae (instar 5) (MCM); Lot 91, 2 larvae (instar 5) (MCM); Lot 790, 2 larvae (instar 5) (USNM); Lot 1022, 3 larvae (instar

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301 5) (DHH); Lot 1028, 2 larvae ( instar 5) (DHH) PALM BEACH CO.: Lot 877, 2 larvae (instar 5) (USNM) ; Lot 1019, 1 larva (instar 4) (DHH); Lot 1020, 3 larvae (instars 4, 5) (DHH); Lot 1021, 1 larva (instar 4) (DHH); Lot 1023, 2 larvae (instar 5) (DHH). PUPAE: FLORIDA: BROWARD CO.: Lot 86, 1 pupa (MCM). DADE CO.: Lot 661, 1 pupa (MCM) ; Lot 1018, 1 pupa (DHH). MONROE CO.: Lot 85, 1 pupa (MCM); Lot 89, 4 pupae (MCM); Lot 90, 1 pupa (MCM); Lot 790, 1 pupa (USNM); Lot 1022, 1 pupa (DHH); Lot 1028, 1 pupa (DHH). PALM BEACH CO.: Lot 877, 1 pupa (USNM); Lot 1019, 1 pupa (DHH); Lot 1027, 3 pupae (DHH) 218. Atrytonopsis hianna (Scudder) Diagnosis LAST INSTAR LARVA: BODY: Figure 20H, length 30mm, A4 transverse width 5.2mm; pinkish green or brown; setae simple, to 1.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 35A, brown, paler around the mouth; transverse width 3.8mm; sculpturing rough, knobby at apex; setae simple, to 1.3mm long at apex, a few ventral setae to 1.3mm; stemmata subequal (Figure 430). THORAX: legs pale; shield narrow, a black line between the annuli. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, tan (Figure 55E); wax glands forming small one or two ventral transverse patches on A7-8. Other Descriptions Harris 1972 (larva), Heitzman and Heitzman 1974 (egg, larva, pupa).

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302 Host Plants FABACEAE: Glycine species (Scudder 1869) [erroneous]. POACEAE: Andropogon gerardii (Heitzman and Heitzman 1970b), Andropogon scoparius (Shapiro and Shapiro 1973), Andropogon species (Shapiro 1966, associated with), Calamovilfa brevipilis (MCM collection) Specimens Examined LARVAE: MISSOURI: BENTON CO.: Lot 555, 4 larvae ( instars 1, 4) ( JRH) NEW JERSEY: BURLINGTON CO.: Lot 659, 1 larva ( instar 5) (MCM). NORTH DAKOTA: RANSOM CO.: Lot 538, 3 larvae (instars 1, 3) (TLM) 219. Atrytonopsis deva (Edwards) Nothing is known of the biology of this southwestern species. 220. Atrytonopsis lunus (Edwards) Nothing is known of the biology of this southwestern species. Bailowitz and Brock (1991) listed the grass, Muhlenbergia emersleyi as a probable larval host. 221. Atrytonopsis vierecki (Skinner) Nothing is known of the biology of this western species. 222. Atrytonopsis loammi (Whitney) Diagnosis LAST INSTAR LARVA: BODY: length 34mm, A4 transverse width 5. 5-5. 7mm; pinkish green; setae simple, to 1.3mm long on A4 dorsum; spiracles tan. HEAD: light brown; transverse width 3. 7-3. 8mm; sculpturing rough, knobby at apex; setae simple, to 1.1mm long at apex, a few ventral setae to 1.8mm; stemmata subegual (Figure 43P) THORAX:

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303 legs pale; shield narrow, a brown line between the annuli. ABDOMEN: prolegs each with about 80 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, tan; wax glands forming small one or two ventral transverse patches on A7-8. PUPA: Figures 75C and 98B, length 19mm, A3 transverse width 3.8mm; cream-colored, abdomen reddish dorsal ly; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A7; abdomen long; cremaster to 2.3mm long, bluntly pointed with a few hooked setae in a cluster at the tip (Figures 110A and 122E); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Host Plants POACEAE: Andropogon species (MCM collection, in lab) Specimens Examined LARVAE: FLORIDA: ST. JOHNS CO.: Lot 93, 2 larvae ( instar 5) (MCM). PUPAE: FLORIDA: ST. JOHNS CO.: Lot 93, 1 pupa (MCM). 223. Atrytonopsis pittacus (Edwards) Nothing is known of the biology of this southwestern species. 224. Atrytonopsis python (Edwards) MacNeill (1975) described the egg and larva of this southwestern species, but I could not find specimens for study

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304 225. Atrytonopsis cestus (Edwards) Nothing is known of the biology of this southwestern species. 226. Atrytonopsis ovinia (Hewitson) The immature stages of this western species are undescribed and I could not find specimens for study. Host Plants POACEAE: Bouteloua curtipendula (Bailowitz and Brock 1991), Leptochloa dubia (Bailowitz and Brock 1991, suspected) 227. Amblyscirtes simius Edwards The immature stages of this western species are undescribed, and I could not find specimens for study. MacNeill (1975) listed the grass, Bouteloua gracilis as a larval host. 228. AmJblyscirtes exoteria (Herrich-Schaf fer) Nothing is known of the immature stages of this southwestern species. Bailowitz and Brock (1991) listed the grass, Muhlenberg ia emersleyi as a probable larval host. 229. Amblyscirtes cassus Edwards The immature stages of this southwestern species are undescribed, and I could not find specimens for study. The grass, Panicum bulbosum, is a larval host (Bailowitz and Brock 1991)

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305 230. Amblyscirtes aenus Edwards Scott (1986) described the egg, larva, and pupa of this southwestern species, but I could not find specimens for study Host Plants POACEAE: Bouteloua curtipendula (Bailowitz and Brock 1991), Bromus anomalus (Bailowitz and Brock 1991), Chasmanthium latifolium (Scott 1986), Digitaria sanguinalis Eragrostis curvula, Setaria leucopila, Sorghum halepense (Bailowitz and Brock 1991, probably), Stenotaphrum secundatum (ROK collection, in lab). 231. Amblyscirtes linda H. A. Freeman Diagnosis LAST INSTAR LARVA: BODY: length 19-2 lmm, A4 transverse width 3. 3-3. 8mm; preserved specimens pale; setae simple, to 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 35B, pale, medial and epicranial lines broadly convergent along the adfrontals, black, front with a few dark lines, lateral line and posterior black; transverse width 2.3mm; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.3mm; stemma 6 small (Figure 43Q) ; paracylpeal hooks present. THORAX: legs pale; shield narrow, a black line between the annuli. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 55F). Other Descriptions Heitzman and Heitzman 1970a (egg, larva, pupa)

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306 Host Plants POACEAE: Chasmanthium latifolium (Heitzman and Heitzman 1970a). Specimens Examined LARVAE : ARKANSAS : CARROLL CO : Lot 571, 2 larvae ( instar 5) ( JRH) 232. Amblyscirtes oslari (Skinner) The immature stages of this western species are undescribed, and I could not find specimens for study. Host Plants POACEAE: Bouteloua gracilis (Stanford 1981, probably), Cynodon dactylon (Bailowitz and Brock 1991, in lab) 233. Amblyscirtes erna H. A. Freeman Nothing is known of the biology of this southwestern species. 234. Amblyscirtes hegon (Scudder) Scudder (1889a) and Klots (1966) described the larva and pupa of this eastern species, but I could not find specimens for study. Host Plants POACEAE: Andropogon species (French 1886), Chasmanthium latifolium (Heitzman and Heitzman 1970a), Poa pratensis (Klots 1966), Sorghastrum nutans (MacNeill 1975), Sorghastrum secundum (Scudder 1889a), Sorghum vulgare var. bicolor (Tietz 1952). 235. Amblyscirtes texanae Bell Nothing is known of the immature stages of this southwestern species. Bailowitz and Brock (1991) listed the grass, Panicum bulbosum, as a possible larval host.

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307 236. Amblyscirtes prenda Evans Nothing is known of the biology of this southwestern species. 237. Amblyscirtes aesculapius (Fabricius) Diagnosis LAST INSTAR LARVA: BODY: length 9. 8-1 2ram, A4 transverse width 2-3. 3mm; yellowish green, heart line dark; setae simple, to 0.2mm long on A4 dorsum; spiracles tan. HEAD: Figure 35C, pale, medial line black, front with a few dark lines, lateral line and posterior black; transverse width 1.9-2. 3mm; sculpturing slightly rough; setae simple, to 0.2mm long at apex, a few ventral setae to 0.4mm; stemma 6 small (Figure 43R); paracylpeal hooks present. THORAX: legs pale; shield moderately broad, black. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked; wax glands forming two ventrolateral patches on A7-8. PUPA: Figures 75D and 98C, length 16mm, A3 transverse width 3.33.7mm; cream-colored; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal and middle leg tips nearly subegual; proboscis extending into A8; abdomen long; cremaster to 1.2mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures HOB and 122F); lenticles present on the prothorax and proleg scars. Host Plants POACEAE: Arundinaria gigantea (Opler and Krizek 1984)

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308 Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 59, 1 larva ( instar 5) (MCM); Lot 61, 1 larva (instar 5) (MCM). PUPAE: FLORIDA: ALACHUA CO.: Lot 60, 2 pupae (MCM); Lot 61, 1 pupa (MCM). 238. Amblyscirtes Carolina (Skinner) Nothing is known of the immature stages of this eastern species. Clark and Clark (1951) noted that the adults are associated with the grass, Arundinaria gigantea the probably larval host. 239. Amblyscirtes reversa (F. M. Jones) Nothing is known of the immature stages of this eastern species. Opler and Krizek (1984) listed the grass, Arundinaria gigantea, as the larval host. 240. Amblyscirtes nereus (Edwards) Nothing is known of the biology of this southwestern species. 241. Amblyscirtes nysa Edwards Heitzman (1964b) described the egg, larva, and pupa of this raidwestern species, but I could not find specimens for study. Host Plants POACEAE: Digitaria sanguinalis (Heitzman 1964b), Echinochloa crusgalli (Heitzman 1964b), Echinochloa muriacata (Scott 1986), Leptochloa dubia (Bailowitz and Brock 1991, in lab), Paspalum species (Scott 1986), Setaria glauca (Heitzman 1964b), Stenotaphrum secundatum (Kendall 1960)

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309 242. Amblyscirtes eos (Edwards) Nothing is known of the immature stages of this southwestern species. Bailowitz and Brock (1991) listed the grass, Panicum obtusum, as a probable larval host. 243. Amblyscirtes vialis (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 11-23. 5mm, A4 transverse width 3. 5-4. 5mm; green, heart line dark; setae simple, to 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 35D, pale, medial and epicranial lines broadly convergent along the adfrontals, black, front with a few dark lines, lateral line and posterior black; transverse width 2. 1-2. 5mm; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.3mm; stemma 6 small (Figure 43S); paracylpeal hooks present. THORAX: legs pale; shield narrow, a black line between the annuli. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked; wax glands forming two ventrolateral patches on A7-8. PUPA: length 13-15mm, A3 transverse width 3. 4-3. 6mm; cream-colored; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers touching; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending to the cremaster; abdomen long; cremaster to 1.2mm long, bluntly pointed with numerous hooked setae in a cluster at the tip (Figures HOC and 122G); lenticles indistinct or absent.

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310 Other Descriptions Fletcher 1888 (young larva), Scudder 1889a (egg, larva, pupa). Host Plants POACEAE: Agrostis species (Shapiro 1966), Avena striata (Scudder 1889a, in lab), Chasmanthium latifolium (Heitzman and Heitzman 1970a), Cynodon dactylon (Kendall 1965, in lab), Erianthus alopecuroides (MCM collection), Poa pratensis (Fletcher 1888, in lab). Specimens Examined LARVAE: NORTH CAROLINA: IREDELL CO.: Lot 62, 1 larva (instar 4) (MCM); Lot 63, 3 larvae ( instar 5) (MCM). TEXAS: BROWN CO.: Lot 907, 20 larvae ( instars 1, 3, 5) (ROK). PUPAE: TEXAS: BROWN CO.: Lot 907, 2 pupae (ROK) 244. Amblyscirtes celia Skinner Diagnosis EGG: height 0.8-lmm, width 0.6-0. 7mm, polygonal sculpturing. LAST INSTAR LARVA: BODY: length 16-21mm, A4 transverse width 2. 9-3. 7mm; preserved specimens pale; setae simple, to 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 35E, pale, medial line brown, front with a few brown lines, lateral line and posterior brown; transverse width 22.1mm; sculpturing slightly rough; setae simple, to 0.1mm long at apex, a few ventral setae to 0.3mm; stemmata subequal (Figure 43T); paracylpeal hooks present. THORAX: legs pale; shield broad, black. ABDOMEN: prolegs each with about 90 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked. PUPA: Figures 75E and 98D, length 15.5-17mm, A3 transverse width

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311 3. 5-3 .7mm; preserved specimens pale; thoracic spiracle guard distinct; setae simple, to 0.4mm long on head; pilifers narrowly separated; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis extending into A8 ; abdomen moderately long; cremaster to 1mm long, bluntly with a few hooked setae in a cluster at the tip (Figures HOD and 122H) ; lenticles present on the prothorax and proleg scars. Host Plants POACEAE: Paspalum species (Scott 1986), Stenotaphrum secundatum (Kendall 1960). Specimens Examined EGGS: TEXAS: BEXAR CO.: Lot 906, 8 eggs (ROK) LARVAE: TEXAS: BEXAR CO.: Lot 906, 3 larvae (instar 5) (ROK). PUPAE: TEXAS: BEXAR CO.: Lot 906, 3 pupae (ROK) 245. Amblyscirtes belli H. A. Freeman Heitzman (1965b) described the egg, larva, and pupa of this eastern species, but I could not find specimens for study Host Plants POACEAE: Chasmanthium latifolium (Heitzman 1965b) 246. Amblyscirtes alternata (Grote and Robinson) Nothing is known of the biology of this southeastern species. 247. Amblyscirtes phylace (Edwards) Nothing is known of the biology of this southwestern species.

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312 248. Amblyscirtes fimbriata (Plotz) Coolidge (1911) described the egg and young larva of this species, but I could not find specimens for study. Host Plants POACEAE: Bromus anomalus (Bailowitz and Brock 1991), Bromus inermis (Bailowitz and Brock 1991, suspected), Dactylis glomerata (Bailowitz and Brock 1991, suspected), Elymus arizonicus (Bailowitz and Brock 1991). 249. Lerodea eufala (Edwards) Diagnosis EGG: height 1-1. 2mm, 0.6-0. 8mm, polygonal sculpturing, pale green to white. LAST INSTAR LARVA: BODY: Figure 21G, length 14-31mm, A4 transverse width 2. 3-4. 2mm; green with tiny white spots, heart, subdorsal, and lateral lines dark, narrow; setae simple, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 35F, pale, medial line brown at vertex, broadly convergent with the brown epicranial lines, front brown, lateral line and posterior brown; transverse width 1.7-2. 4mm; sculpturing slightly rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 6 small (Figure 43U) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 56A) ; wax glands forming two ventrolateral patches on A7-8. PUPA: Figures 75F and 98E, length 16-20mm, A3 transverse width 2. 1-3. 8mm; pale green, heart line dark, faintly bounded by white; thoracic spiracle guard distinct; setae simple, < 0.1mm long

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313 on head; pilifers narrowly separated; head with a long pointed process on cap; antennal tip lies cephalad of tip of middle leg; proboscis extending into A8 ; abdomen long; cremaster to 1.8mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 110E and 1221); lenticles present on the prothorax and proleg scars. Other Descriptions Coolidge 1922 (egg, larva, pupa), Hayward 1926 (egg, young larva), Comstock 1929 (larva, pupa), Comstock 1930a (egg), Dethier 1939b (larva), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants CYPERACEAE: Cyperus species (Hayward 1926, oviposition, but no larval feeding). FABACEAE: Medicago sativa (Field 1938) [erroneous]. POACEAE: Bothriochloa barbinodis var. perforata (MCM collection), Brachiaria subquadripara (MCM collection), Cynodon dactylon (Tilden 1965, associated with; Shapiro 1974b, c), Digitaria species (MCM collection, in lab), Echinochloa crusgalli (Shapiro 1974b, c), Eustachys petraea (MCM observation), Oryza sativa (Bruner et al 1945), Paspalum urvillei (MCM collection), Poa pratensis (Tietz 1972), Saccharum officinarum (Dethier 1942d, in lab), Setaria verticillata (Shapiro 1974b), Sorghum halepense (Shapiro 1974b, c), Sorghum sudanense (MCM collection), Sorghum vulgare var. bicolor (Shapiro 1974b), Stenotaphrum secundatum (Kendall 1960, in lab; MCM observation), Zea mays (Scott 1986).

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314 Specimens Examined EGGS: FLORIDA: ALACHUA CO.: Lot 1513, 15 eggs (MCM) LARVAE: ARIZONA: YUMA CO.: Lot 820, 4 larvae (instar 5) (USNM) CALIFORNIA: RIVERSIDE CO.: Lot 573, 2 larvae (instar 5) (GRB) YOLO CO.: Lot 311, 1 larva (instar 5) (MCM); Lot 312, 7 larvae ( instars 3, 4, 5) (MC). FLORIDA: ALACHUA CO.: Lot 1513, 6 larvae (instar 1) (MCM). BROWARD CO.: Lot 313, 1 larva (instar 5) (MCM); Lot 314, 1 larva (instar 5) (MCM). DADE CO.: Lot 771, 1 larva (instar 5) (USNM). TEXAS: BEXAR CO.: Lot 309, 1 larva (instar 5) (MCM). HIDALGO CO.: Lot 772, 2 larvae (instars 3, 5) (USNM). JIM WELLS CO.: Lot 773, 1 larva (instar 3) (USNM). PUPAE: CALIFORNIA: YOLO CO.: Lot 312, 2 pupae (MCM). FLORIDA: BROWARD CO.: Lot 315, 1 pupa (MCM). TEXAS: BEXAR CO.: Lot 309, 2 pupae (MCM). 250. Lerodea arabus (Edwards) The immature stages of this southwestern species are undescribed, and I could not find specimens for study. Host Plants POACEAE: Cynodon dactylon (Bailowitz and Brock 1991), Leptochloa dubia (Bailowitz and Brock 1991). 251. Lerodea dysaules Godman The immature stages of this southwestern species are undescribed, and I could not find specimens for study. Host Plants POACEAE: Cynodon dactylon (Kendall 1976, in lab)

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315 252. Oligoria maculata (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 24-30mm, A4 transverse width 4. 3-4 .8mm; pinkish green; setae simple, to 0.2mm long on A4 dorsum; spiracles tan to brown. HEAD: Figure 35G, brown; transverse width 3. 5-3. 6mm; sculpturing rough, pebbly at apex; setae simple, 0.2mm long at apex, a few ventral setae to 0.4mm; stemma 6 slightly small (Figure 44A) THORAX: legs pale; shield broad, dark brown. ABDOMEN: prolegs each with about 180 crochets, irregularly triordinal, arranged in a mesal penellipse; suranal plate rounded, unmarked (Figure 56B) PUPA: Figures 59D, 75G, and 98F, length 25mm, A3 transverse width 4.5mm; creamcolored; thoracic spiracle guard distinct; setae simple, to 0.6mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A8; abdomen moderately long; cremaster to 1.8mm long, blunt with numerous hooked setae in a row along the distal margin (Figures 110F and 122J); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Edwards and Chapman 1879 (larva, pupa), Minno and Emmel 1993 (egg, larva, pupa). Host Plants POACEAE: Andropogon species (MCM collection), Digitaria species (MCM collection, in lab). Specimens Examined LARVAE: FLORIDA: ALACHUA CO.: Lot 347, 1 larva (instar 2) (MCM). BROWARD CO.: Lot 346, 2

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316 larvae (instar 5) (MCM) DUVAL CO.: Lot 95, 1 larva (instar 4) (MCM). PUPAE: FLORIDA: BROWARD CO.: Lot 346, 1 pupa (MCM) 253. Calpodes ethlius (Stoll) Diagnosis EGG: height 1.2mm, width 0.7-0. 8mm, polygonal sculpturing, gray. LAST INSTAR LARVA: BODY: length 2663mm, A4 transverse width 4. 3-9. 5mm; cuticle transparent, appearing dark green due to gut contents; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 35H, tan with a black spot in center of face, stemmatal area black; transverse width 3. 5-4. 3mm; sculpturing rough; setae simple to blunt, 0.1mm long at apex, a few ventral setae to 0.5mm; stemmata subegual (Figure 44B) THORAX: Tl-2 legs brown and tan, T3 legs pale; shield pale dorsally, brown laterally. ABDOMEN: prolegs each with about 195 crochets, multiordinal ca. 2-3 ranks, arranged in a circle; suranal plate rounded, unmarked (Figure 57B); wax glands forming two ventrolateral patches on A7-8. PUPA: Figures 60C, 76A, and 99A, length 3743.5mm, A3 transverse width 7-8. 5mm; pale green, lenticles black, process on cap and free portion of proboscis tan; thoracic spiracle guard indistinct; setae simple, to 0.2mm long on head; pilifers separated; head with a slender, upturned process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending beyond the cremaster; abdomen long; cremaster to 3.6mm long, bluntly pointed with

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317 numerous hooked setae in a cluster at the tip (Figures HOG and 122K); lenticles present on the prothorax, dorsal abdomen, and proleg scars. Other Descriptions Dodge 1872, 1876 (larva, pupa), King 1882 (egg, larva, pupa), Scudder 1889a (egg, larva, pupa), Cockerell 1892, 1894 (egg, larva, pupa), Dyar 1898 (egg, larva, pupa), Chittenden 1905, 1912 (egg, larva, pupa), Mosher 1916 (pupa), Koehler 1927 (egg, larva, pupa), Weigel and Middleton 1926 (larva), Wolcott 1936 (larva), Dethier 1939b (larva), Moss 1949 (larva, pupa), Kirkpatrick 1957 (larva), Peterson 1962 (larva), Young 1982 (larva, pupa), Minno and Emmel 1993 (egg, larva, pupa). Host Plants APIACEAE: Apium graveolens (Kimball 1965) [erroneous]. ARACEAE: Colocasia esculentum (Chittenden 1905) [erroneous]. BRASSICACEAE: (Lenczewski 1980) [erroneous]. CANNACEAE: Canna aurantiacea (Hayward 1941), Canna coccinea (Wolcott 1923), Canna edulis (Wolcott 1923), Canna flaccida (Scudder 1889a, b), Canna X generalis (DHH collection), Canna indica (Dodge 1872). CUCURBITACEAE: Lagenaria species (da Costa Lima 1936) [erroneous]. EUPHORBIACEAE: Phyllanthus species (Kimball 1965) [erroneous]. MARANTACEAE: Maranta arundinacea (Ogilvie 1928), Maranta species (Bates 1935), Thalia geniculata (Opler and Krizek 1984). MUSACEAE: Heliconia species (USNM collection). ZINGIBERACEAE: (MCM collection)

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318 S pecimens Examined EGGS: FLORIDA: ALACHUA CO. : Lot 1516, 4 eggs (MCM) LARVAE: NO DATA: Lot 1128, 1 larva (instar 5) (DHH). COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 119, 2 larvae (instar 5) (MCM); Lot 120, 1 larva (instar 5) (MCM). COSTA RICA: GUANACASTE PROV.: Lot 1108, 2 larvae (instar 5) (DHH). DOMINICAN REPUBLIC: LA VEGA PROV.: Lot 118, 1 larva (instar 5) (MCM). JAMAICA: Lot 805, 5 larvae (instars 3, 4) (USNM). MEXICO: Lot 803, 1 larva (instar 2) (USNM). VIRGIN ISLANDS: ST. THOMAS: Lot 798, 2 larvae (instars 4, 5) (USNM). FLORIDA: ALACHUA CO.: Lot 663, 1 larva (instar 5) (MCM); Lot 1078, 2 larvae (instars 2, 3) (DHH); Lot 1091, 1 larva (instar 5) (DHH); Lot 1092, 2 larvae (instar 5) (DHH); Lot 1093, 2 larvae (instar 5) (DHH); Lot 1094, 1 larva (instar 5) (DHH); Lot 1095, 2 larvae (instar 5) (DHH); Lot 1096, 2 larvae (instars 3, 4) (DHH); Lot 1097, 4 larvae (instar 5) (DHH); Lot 1098, 1 larva (instar 5) (DHH); Lot 1099, 1 larva (instar 5) (DHH); Lot 1100, 1 larva (instar 5) (DHH); Lot 1102, 1 larva (instar 5) (DHH); Lot 1103, 1 larva (instar 5) (DHH); Lot 1104, 1 larva (instar 5) (DHH); Lot 1105, 3 larvae (instars 3, 5) (DHH); Lot 1106, 1 larva (instar 5) (DHH); Lot 1107, 2 larvae (instar 5) (DHH); Lot 1111, 3 larvae (instar 5) (DHH); Lot 1113, 2 larvae (instar 5) (DHH); Lot 1114, 3 larvae (instars 4, 5) (DHH); Lot 1116, 1 larva (instar 5) (DHH); Lot 1117, 1 larva (instar 5) (DHH); Lot 1118, 1 larva (instar 5) (DHH); Lot 1119, 1 larva (instar 3) (DHH); Lot

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319 1120, 1 larva (instar 4) (DHH); Lot 1121, 1 larva (instar 5) (DHH); Lot 1122, 1 larva (instar 5) (DHH); Lot 1123, 1 larva (instar 3) (DHH); Lot 1124, 1 larva (instar 5) (DHH); Lot 1125, 1 larva (instar 5) (DHH); Lot 1126, 3 larvae (instar 5) (DHH); Lot 1129, 1 larva (instar 5) (DHH); Lot 1131, 1 larva (instar 5) (DHH); Lot 1132, 1 larva (instar 5) (DHH); Lot 1133, 1 larva (instar 5) (DHH); Lot 1135, 1 larva (instar 5) (DHH); Lot 1136, 1 larva (instar 3) (DHH); Lot 1138, 1 larva (instar 5) (DHH); Lot 1141, 1 larva (instar 5) (DHH); Lot 1142, 4 larvae ( instars 3, 5) (DHH); Lot 1143, 1 larva (instar 5) (DHH); Lot 1145, 1 larva (instar 3) (DHH); Lot 1146, 1 larva (instar 5) (DHH); Lot 1147, 2 larvae (instar 5) (DHH); Lot 1148, 1 larva (instar 4) (DHH); Lot 1149, 1 larva (instar 3) (DHH); Lot 1150, 1 larva (instar 5) (DHH); Lot 1152, 3 larvae (instar 2) (DHH); Lot 1153, 1 larva (instar 5) (DHH); Lot 1154, 1 larva (instar 5) (DHH); Lot 1155, 3 larvae (instars 3, 5) (DHH); Lot 1156, 1 larva (instar 3) (DHH); Lot 1157, 1 larva (instar 5) (DHH); Lot 1158, 1 larva (instar 5) (DHH); Lot 1159, 2 larvae (instar 5) (DHH); Lot 1160, 1 larva (instar 5) (DHH); Lot 1161, 1 larva (instar 5) (DHH); Lot 1162, 3 larvae (instars 2, 3, 5) (DHH); Lot 1164, 2 larvae (instar 5) (DHH); Lot 1166, 1 larva (instar 2) (DHH); Lot 1168, 1 larva (instar 5) (DHH). BROWARD CO.: Lot 11, 1 larva (instar 5) (MCM) ; Lot 121, 4 larvae (instar 3) (MCM); Lot 122, 1 larva (instar 5) (MCM). COLLIER CO.: Lot 1127, 2 larvae (instar 5) (DHH). DUVAL

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320 CO.: Lot 1110, 2 larvae ( instars 3, 5) (DHH). HARDEE CO.: Lot 1165, 1 larva ( instar 3) (DHH). HIGHLANDS CO.: Lot 1101, 3 larvae (instars 3, 4) (DHH); Lot 1163, 1 larva (instar 5) (DHH). INDIAN RIVER CO.: Lot 662, 1 larva (instar 5) (MCM) ; Lot 664, 4 larvae (instars 4, 5) (MCM) MARION CO.: Lot 1134, 1 larva (instar 5) (DHH). POLK CO.: Lot 1140, 1 larva (instar 5) (DHH). SARASOTA CO.: Lot 1137, 1 larva (instar 5) (DHH). TEXAS: CAMERON CO.: Lot 787, 2 larvae (instar 5) (USNM) PUPAE: DOMINICAN REPUBLIC: LA VEGA PROV. : Lot 118, 1 pupa (MCM). USA: ARIZONA: COCHISE CO.: Lot 540, 1 pupa (TLM) FLORIDA: Lot 1139, 1 pupa (DHH). ALACHUA CO.: Lot 1096, 3 pupae (DHH); Lot 1109, 4 pupae (DHH); Lot 1112, 4 pupae (DHH); Lot 1115, 3 pupae (DHH); Lot 1116, 1 pupa (DHH); Lot 1130, 1 pupa (DHH); Lot 1151, 2 pupa (DHH); Lot 1156, 1 pupa (DHH); Lot 1160, 1 pupa (DHH); Lot 1164, 1 pupa (DHH). BROWARD CO.: Lot 121, 1 pupa (MCM). INDIAN RIVER CO.: Lot 662, 1 pupa (MCM); Lot 664, 1 pupa (MCM). 254. Panoquina panoquin (Scudder) Diagnosis EGG: height 0.9mm, width 0.6-0. 8mm, polygonal sculpturing, pale green. LAST INSTAR LARVA: BODY: Figure 21F, length 28. 5-36. 5mm, A4 transverse width 3. 1-4. 3mm; green, heart line dark, subdorsal line narrow, pale yellow; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: Figure 3 6A, green; transverse width 2. 4-2. 8mm; sculpturing slightly rough; setae simple to

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321 blunt, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 1, 5, and 6 small (Figure 44C) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 70 crochets, irregularly triordinal, arranged in a circle; suranal plate bluntly pointed, unmarked (Figure 56C) PUPA: Figures 76B and 99B, length 24-25mm, A3 transverse width 3.8mm; pale green, heart and subdorsal lines dark, outlined by pale yellow; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending slightly beyond the wings in A4; abdomen long; cremaster to 2.2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 110H and 123A) ; lenticles indistinct or absent. Other Descriptions Skinner 1890 (egg, young larva), Laurent 1908 (egg, young larva), Pyle 1981 (egg, larva). Host Plants CYPERACEAE: Scirpus species (Clark and Clark 1951, associated with; Shapiro 1966) [erroneous]. POACEAE: Distichlis spicata (Shapiro and Shapiro 1973, associated with; MCM collection), Spartina alternifolia (MCM collection, DHH collection). Specimens Examined EGGS: FLORIDA: BREVARD CO.: Lot 1515, 16 eggs (MCM). LARVAE: FLORIDA: BREVARD CO.: Lot 705, 4 larvae ( instar 5) (MCM). FLAGLER CO.: Lot 1511, 1 larva (instar 5) (MCM). LEVY CO.; Lot 353, 3 larvae (instar 5) (MCM). VOLUSIA CO.: Lot 706, 2 larvae (instars 2, 5)

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322 (MCM); Lot 1510, 1 larva (instar 4) (MCM) PUPAE: FLORIDA: LEVY CO.: Lot 353, 1 pupa (MCM). VOLUSIA CO.: Lot 706, 1 pupa (MCM) 255. Panoquina panoquinoides (Skinner) Diagnosis LAST INSTAR LARVA: BODY: length 14.5-28mm, A4 transverse width 2-3. 7mm; green, heart line dark, subdorsal line narrow, pale yellow; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: green, sometimes with faint reddish stripes; transverse width 2.12.4mra; sculpturing slightly rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.2mm; stemma 1, 5, and 6 small (Figure 44D) THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 60 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 56D) PUPA: Figures 76C and 99C, length 19-20. 5mm, A3 transverse width 3. 1-3. 4mm; pale green, heart and subdorsal lines dark, outlined by pale yellow; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending slightly beyond the wings in A4; abdomen moderately long; cremaster to 2.3mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 1101 and 123B); lenticles present on dorsal abdomen and proleg scars.

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323 Other Descriptions Brown and Turner 1966 (egg, larva, pupa), Minno and Emmel 1993 (larva, pupa). Host Plants FABACEAE: Mimosa pudica (Brown and Heineman 1972) [erroneous]. POACEAE: Cynodon dactylon (Brown and Heineman 1972, in lab), Distichlis spicata (Pyle 1981, possibly; Minno and Emmel 1993), Saccharum officinarum (Riley 1975), Sporobolus virginicus (Clench and Bjorndal 1980, probably; Minno and Emmel 1993). Specimens Examined LARVAE: FLORIDA: DADE CO.: Lot 356, 2 larvae (instar 5) (MCM) MONROE CO.: Lot 354, 2 larvae (instar 5) (MCM); Lot 355, 2 larvae (instar 5) (MCM). PUPAE: FLORIDA: MONROE CO.: Lot 354, 2 pupae (MCM). 256. Panoquina errans (Skinner) I could not locate specimens of this western species for study. Other Descriptions Dyar 1892 (egg, larva, pupa), Comstock 1931 (egg, larva, pupa), Brown and Turner 1966 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants POACEAE: Cynodon dactylon (Comstock 1931, in lab), Distichlis spicata (Dyar 1892). 257. Panoquina ocola (Edwards) Diagnosis LAST INSTAR LARVA: BODY: length 2 1-3 5mm, A4 transverse width 3. 3-4. 6mm; green, heart line dark, subdorsal and lateral lines narrow, pale yellow; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: green; transverse width 2. 4 -3. lmm; sculpturing

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324 slightly rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.3mm; stemma 1, 5, and 6 small (Figure 44E). THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 145 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 56E) PUPA: Figures 76D and 99D, length 23-24. 5mm, A3 transverse width 4-4. lmm; pale green, heart and subdorsal lines dark, outlined by pale yellow; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers touching; head with a long pointed process on cap; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A5; abdomen long; cremaster to 2.9mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 110J and 123C); lenticles absent or indistinct. Other Descriptions Pyle 1981 (egg, larva, pupa), Minno and Emmel 1993 (larva, pupa). Host Plants POACEAE: Cynodon dactylon (MCM collection, in lab), Hymenachne amplexicaulis (Wolcott 1923), Leersia hexandra (MCM collection), Oryza sativa (Bell 1940), Panicum repens (Minno 1992), Saccharum officinarum (Wolcott 1923). Specimens Examined LARVAE: HONDURAS: DEPT. FRANCISCO MORAZAN: Lot 611, 2 larvae ( instar 5) (SP) FLORIDA: ALACHUA CO.: Lot 13, 5 larvae (instar 5) (MCM); Lot 348, 3 larvae (instar 5) (MCM); Lot 1041, 1 larva (instar 3) (DHH); Lot 1063, 1 larva (instar 5) (DHH); Lot 1064, 1 larva

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325 (instar 5) (DHH); Lot 1065, 1 larva (instar 5) (DHH). BROWARD CO.: Lot 350, 1 larva (instar 5) (MCM) Lot 352, 1 larva (instar 5) (MCM). HIGHLANDS CO.: Lot 351, 5 larvae (instars 3, 5) (MCM). PUPAE: HONDURAS: DEPT. FRANCISCO MORAZAN: Lot 611, 1 pupa (SP) FLORIDA: ALACHUA CO.: Lot 13, 1 pupa (MCM); Lot 348, 2 pupae (MCM). BROWARD CO.: Lot 349, 1 pupa (MCM) 258. Panoquina hecebola (Scudder) Nothing is known of the biology of this neotropical species. 259. Panoquina sylvicola (Herrich-Schaf fer) Diagnosis LAST INSTAR LARVA: BODY: length 20-29mm, A4 transverse width 2.8-4mm; green, heart line dark, subdorsal and lateral lines narrow, pale yellow; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles tan. HEAD: green; transverse width 2. 8-3. 2mm; sculpturing slightly rough; setae simple, < 0.1mm long at apex, a few ventral setae to 0.4mm; stemmata subegual (Figure 44F). THORAX: legs pale; shield indistinct. ABDOMEN: prolegs each with about 120 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 56F). PUPA: Figures 76E and 99E, length 27.5mm, A3 transverse width 4.5mm; pale green, heart and subdorsal lines dark, outlined by pale yellow; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers separated; head with a long pointed process on cap; antennal tip lies far

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326 cephalad of tip of middle leg; proboscis extending into A5; abdomen moderately long; cremaster to 2.2mm long, blunt with numerous hooked setae in a cluster at the tip (Figures 111A and 123D); lenticles present on dorsal abdomen and proleg scars. Other Descriptions Wolcott 1921 (larva, pupa), Jones and Wolcott 1922 (egg, larva, pupa), Dethier 1939b (egg, larva), Dethier 1942d (egg, larva, pupa). Host Plants POACEAE: Axinopus compressus (Brown and Heineman 1972), Bambusa vulgaris (Jones and Wolcott 1922), Brachiaria mutica (Wolcott 1923), Eriochloa polystachya (Jones and Wolcott 1922), Oryza sativa (Jones and Wolcott 1922), Saccharum officinarum (Van Dine 1913), Sorghum halepense (Jones and Wolcott 1922). Specimens Examined LARVAE: COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 357, 1 larva (instar 5) (MCM); Lot 358, 5 larvae (instar 5) (MCM). PUPAE: COLOMBIA: DEPT. VALLE DEL CAUCA: Lot 357, 1 pupa (MCM). 260. Panoquina evansi (H. A. Freeman) Nothing is known of the biology of this neotropical species. 261. Nyctelius nyctelius (Latreille) Diagnosis LAST INSTAR LARVA: BODY: length 26-39mm, A4 transverse width 4. 3-6. 6mm; bluish green, heart line dark; some setae with blunt tips, < 0.1mm long on A4 dorsum; spiracles pale. HEAD: Figure 36B, pale with broad, black

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327 medial, epicranial, and lateral lines, occasionally black with two pale lines from near vertex to base of mandibles and pale eye patches, Caribbean populations: tan with light brown medial and epicranial stripes, becoming blackish toward middle of face and stemmatal area black; transverse width 3. 3-3. 5mm; sculpturing rough to pitted; setae simple, to 0.1mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 44G) THORAX: Tl-2 legs black, T3 legs dark brown; shield broad, black. ABDOMEN: prolegs each with about 85 crochets, irregularly triordinal, arranged in a circle; suranal plate rounded, unmarked (Figure 57A) PUPA: Figures 77A and 100A, length 27.5mm, A3 transverse width 6.5mm; green shaded with black; thoracic spiracle guard distinct; setae simple, to 0.9mm long on head; pilifers touching; head rounded; antennal tip lies far cephalad of tip of middle leg; proboscis extending into A6; abdomen long; cremaster to 1.8mm long, squarish with numerous hooked setae in a row along the distal margin (Figures 111B and 123E) ; lenticles present on the prothorax, dorsal abdomen and proleg scars. Other Descriptions Wolcott 1921 (larva, pupa), Jones and Wolcott 1922 (egg, larva, pupa), Dethier 1939b (larva, pupa), Dethier 1942d (egg, larva, pupa). Host Plants POACEAE: Brachiaria mutica (Bates 1935), Oryza sativa (Wolcott 1922), Panicum maximum (MCM

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328 collection), Saccharum officinarum (Smyth 1919), Sorghum species (SP collection), Zea mays (Comstock 1944). Specimens Examined LARVAE: COSTA RICA: Lot 1087, 1 larva (instar 3) (DHH), probably this species. DOMINICAN REPUBLIC: LA VEGA PROV. : Lot 340, 2 larvae ( instars 3, 5) (MCM). SANTIAGO PROV.: Lot 342, 1 larva (instar 5) (MCM) HONDURAS: Lot 607, 1 larva (instar 5) (SP); Lot 621, 2 larvae (instars 3, 4) (SP); Lot 622, 1 larva (instar 5) (SP); Lot 624, 1 larva (instar 5) (SP). DEPT. FRANCISCO MORAZAN: Lot 623, 1 larva (instar 5) (SP). MEXICO: Lot 817, 1 larva (instar 5) (USNM). NICARAGUA: Lot 816, 2 larvae (instar 5) (USNM); Lot 818, (instar 5) (USNM). PUERTO RICO: Lot 625, 1 larva (instar 5) (SP) PUPAE: DOMINICAN REPUBLIC: PUERTO PLATA PROV.: Lot 341, 1 pupa (MCM). NICARAGUA: Lot 818, 1 pupa (USNM). 262. Thespieus macareus (Herrich-Schaf fer) Nothing is known of the biology of this neotropical species. Subfamily Megathyminae Diagnosis EGG: height 1.3-1. 8mm, width 2.5-3mm, polygonal sculpturing, green, red, brown, or white. LAST INSTAR LARVA: BODY: length 19-86mm, A4 transverse width 5.7511.67mm; cream-colored or bluish; setae; spiracles tan to dark, Tl and A8 spiracles largest. HEAD: uniformly colored; transverse width 3. 3-5. 8mm; sculpturing rough to pebbly; mandibles with or without teeth, pyrgine type

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329 articulation; labial-submental complex relatively large; setae simple; stemmata usually subequal; postocciput wide or moderately wide. THORAX: prothorax larger than head; legs brown to black; shield broad; ventral prothoracic gland absent. ABDOMEN: A4 proleg with 65 to 130 crochets, irregularly triordinal, arranged in a circle, a near circle, or transverse bands, posterior prolegs always with a mesal peneiiipse; suranai Diate rounded. unmarKed: anai co-.n absent or poorly developed; wax glands present in the form of transverse patches on the ventral side of A7-8. PUPA: length 34-52mm, A3 transverse width 8-10. 2mm; brown; thoracic spiracle guard distinct; pilifers separated; head rounded; antennal and middle leg tips subequal; proboscis short; abdomen lonq to verv lonq; cremaster blunt lv pointed without setae or broad and rounded with numerous setae. nnnx&n aezraez an"!flnr 263. Agathymus neumoegeni (Edwards) Freeman (1951a) described the egg, larva, and pupa of this southwestern species, but I could not find specimens for study. Host Plants AGAVACEAE: Agave chrysantha (Freeman 1951a), Agave deserti (Tietz 1972), Agave lechuguilla (Scott 1986), Agave palmeri (Freeman 1951a), Agave parryi (Tinkham 1954). 264. Agathymus carlsbadensis (D. Stallings and Turner) The immature stages of this species are undescribed, Snu x CGuju PiOt f luu SpcClmSnS TOT StUuV

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330 Host Plants AGAVACEAE: Agave parry i (Freeman 1969). 265. Agathymus florenceae (D. Stallings and Turner) The immature stages of this species are undescribed, and I could not find specimens for study. Host Plants AGAVACEAE: Agave parryi (Freeman 1969). 266. Agathymus judithae (D. Stallings and Turner) Stallings and Turner (1957) illustrated the pupal terminalia of this species, but I could not find specimens for study. Host Plants AGAVACEAE: Agave parryi (Freeman 1960). 267. Agathymus diabloensis H. A. Freeman Freeman (1962) described the larva and pupa of this species, but I could not find specimens for study. Host Plants AGAVACEAE: Agave parryi (Freeman 1962). 268. Agathymus mcalpinei (H. A. Freeman) Freeman (1955) commented on the larva and pupa of this species, but I could not find specimens for study. Host Plants AGAVACEAE: Agave scabra (Freeman 1955). 269. Agathymus chisosensis (H. A. Freeman) Freeman (1952b) briefly described the pupa of this species, but I could not find specimens for study. Host Plants AGAVACEAE: Agave scabra (Freeman 1952b). 270. Agathymus aryxna (Dyar) Roever (1975) briefly described the egg of this species.

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331 Host Plants AGAVACEAE: Agave chrysantha (Roever 1975), Agave deserti (Bailowitz and Brock 1991), Agave palmeri (Stallings and Turner 1957). 271. Agathymus baueri (D. Stallings and Turner) Roever (1964) described the egg of this species. Host Plants AGAVACEAE: Agave chrysantha (Roever 1975), Agave parry i (Stallings and Turner 1954). 272. Agathymus freemani D. Stallings and Turner The immature stages of this species are undescribed. Host Plants AGAVACEAE: Agave deserti (Freeman 1969). 273. Agathymus evansi (H. A. Freeman) Comstock (1956a) described the larva and pupa of this species, but I could not find specimens for study. Host Plants AGAVACEAE: Agave palmeri (Freeman 1969), Agrave parry i (Comstock 1956a). 274. Agathymus mariae (Barnes and Benjamin) Bonniwell (1931), Freeman (1951a), Stallings and Turner (1957), Freeman (1964a, b), and Roever (1975) have described immature stages of this species, but I could not find specimens for study. Host Plants. AGAVACEAE: Agave lechuguilla (Tinkhara 1944). 275. Agathymus chinatiensis H. A. Freeman Freeman (1964b) briefly described the larva and pupa of this species. Host Plants. AGAVACEAE: Agave lechuguilla (Freeman 1964b).

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332 276. Agathymus lajitaensis H. A. Freeman Freeman (1964b) briefly described the larva and pupa of this species. Host Plants AGAVACEAE: Agave lechuguilla (Freeman 1964b). 277. Agathymus rindgei H. A. Freeman Freeman (1964b) described the larva and pupa of this species. Host Plants AGAVACEAE: Agave lechuguilla (Freeman 1964b). 278. Agathymus gilberti H. A. Freeman Host Plants Agave lechuguilla (Freeman 1964b). 279. Agathymus valverdiensis Freeman Diagnosis LAST INSTAR LARVA: BODY: length 2 3 -2 4mm, A4 transverse width 6. 5-7. 5mm; preserved specimens pale; setae simple, to 0.4mm long on A4 dorsum; spiracles tan. HEAD: Figure 36C, tan; transverse width 3. 4-3. 6mm; sculpturing slightly rough; setae simple, to 0.4mm long at apex, a few ventral setae to 0.6mm; stemmata subegual (Figure 44H) THORAX: legs tan; shield broad, black dorsally, tan laterally. ABDOMEN: prolegs each with about 130 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked (Figure 57C). PUPAL EXUVIA: Figure 100B, light brown; thoracic spiracle guard distinct; setae simple, < 0.1mm long on head; pilifers separated; head rounded; antennal tip extending nearly to tip of middle leg; proboscis extending midway between tip of middle leg and tip of wings; abdomen moderately long; cremaster to 1.7mm long,

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333 bluntly pointed, setae absent (Figure 111C); lenticles absent Host Plants Agave lechuguilla (Freeman 1966). Specimens Examined LARVAE: USA: TEXAS: VAL VERDE CO.: Lot 888, 9 larvae (instars 2, 3, 4, 5). 280. Agathymus stephensi (Skinner) Diagnosis LAST INSTAR LARVA: BODY: length 26mm, A4 transverse width 7.6mm; preserved specimen brownish; setae simple, to 0.8mm long on A4 dorsum; spiracles tan to brown. HEAD: tan; transverse width 3.3mm; sculpturing slightly rough; setae simple, to 0.6mm long at apex, a few ventral setae to 0.8mm; mandibles with a shallow tooth; stemma 5 slightly small (Figure 441). THORAX: Tl-2 legs brown, T3 legs tan; shield broad, black dorsally, tan laterally. ABDOMEN: prolegs each with about 105 crochets, irregularly triordinal, arranged in a near circle; suranal plate rounded, unmarked. Other Descriptions Corns tock and Dammers 1934 (egg, larva, pupa), Comstock 1957 (larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa). Host Plants AGAVACEAE: Agave deserti (Skinner 1912). Specimens Examined LARVAE: CALIFORNIA: SAN DIEGO CO.: Lot 1006, 1 larva (instar 5) (FSCA). 281. Agathymus polingi (Skinner)

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334 Stallings and Turner (1957) and Roever (1964) described some aspects of the immature stages, but I could not find specimens for study. Host Plants AGAVACEAE: Agave palmeri (Freeman 1951a), Agave schottii (Barnes and McDunnough 1912), Agave toumeyana (Roever 1975) 282. Agathymus alliae (D. Stallings and Turner) Stallings and Turner (1957), Emmel and Emmel (1973), and Roever (1975) described the immature stages, but I could not find specimens for study. Host Plants AGAVACEAE: Agave utahensis (Freeman 1969). 283. Megathymus yuccae (Boisduval and Leconte) Diagnosis EGG: height 2.9-3mm, width 1.3-1. 8mm, polygonal sculpturing, pinkish brown. LAST INSTAR LARVA: BODY: Figure 21H, length 27-64mm, A4 transverse width 7-10. 5mm; cream-colored; setae simple, to 0.6mm long on A4 dorsum; spiracles brown. HEAD: Figure 36D, reddish brown; transverse width 4. 3-5. 8mm; sculpturing roucjh to pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.7mm; stemma 5 and 6 slightly small (Figure 44J) THORAX: legs brown; shield broad, black and brown dorsal ly. ABDOMEN: prolegs each with about 90 crochets, multiordinal ca. 2-3 ranks, arranged in transverse bands; suranal plate rounded, tan to brown (Figure 57D); wax glands forming ventral transverse patches on A7-8. PUPAL EXUVIUM: Figure 100C, brown; thoracic

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335 spiracle guard distinct; setae simple, < 0.1mm long on head; pilifers separated; head rounded; antennal tip extending nearly to tip of middle leg; proboscis slightly longer than tip of middle leg; abdomen very long; cremaster to 3.8mm long, broadly rounded with numerous simple setae, hooked setae absent (Figures HID and 123F); lenticles absent. Other Descriptions Riley 1876a, b (egg, larva, pupa), Skinner 1917 (pupa), Harris 1972 (pupa), Matthews 1986 (egg, larva) Host Plants AGAVACEAE: Agave species (Remington 1959, feeding, but no survival), yucca aloifolia (Riley 1876a, b) Yucca filamentosa (Riley 1876a, b), Yucca flaccida (Freeman 1952a), Yucca glauca (Tietz 1972), Yucca gloriosa (Riley 1876a, b), Yucca smalliana (Freeman 1952a). Specimens Examined EGGS: FLORIDA: MARION CO.: Lot 699, 1 egg (MCM) SUWANNEE CO.: Lot 700, 4 eggs (MCM) LARVAE: FLORIDA: HIGHLANDS CO.: Lot 4, 1 larva (instar 5) (MCM); Lot 318, 1 larva (instar 5) (MCM); Lot 320, 1 larva (instar 5) (MCM); Lot 321, 1 larva (instar 5) (MCM); Lot 328, 1 larva (instar 5) (MCM). LIBERTY CO.: Lot 324, 1 larva (instar 5) (MCM); Lot 325, 1 larva (instar 5) (MCM); Lot 326, 1 larva (instar 5) (MCM). MARTIN CO.: Lot 316, 1 larva (instar 5) (MCM); Lot 317, 1 larva (instar 5) (MCM). ORANGE CO.: Lot 1005, 7 larvae (instar 4) (FSCA). PUTNAM CO.: Lot 322, 1 larva (instar 5) (MCM); Lot 323, 1 larva (instar 3) (MCM); Lot 698, 1 larva (instar 5) (MCM).

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336 LOUISIANA: SABINE PARISH: Lot 893, 7 larvae (instars 3, 5) (ROK). TEXAS: SMITH CO.: Lot 894, 1 larva ( instar 5) (ROK). 284. Megathymus coloradensis Riley Diagnosis LAST INSTAR LARVA: BODY: length 3 4-8 6mm, A4 transverse width 7. 5-11. 7mm; cream-colored; setae simple, to 0.7mm long on A4 dorsum; spiracles brown. HEAD: dark reddish brown; transverse width 4. 8-5. 8mm; sculpturing rough to pebbly; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.8mm; stemma 5 and 6 slightly small (Figure 44K) THORAX: legs dark brown; shield broad, dark brown and tan dorsally. ABDOMEN: prolegs each with about 90 crochets, multiordinal ca. 2-3 ranks, arranged in transverse bands; suranal plate rounded, tan to brown; wax glands forming ventral transverse patches on A7-8. PUPA: Figures 77B and 100D, length 41.5-44mm, A3 transverse width 8.5mm; brown; thoracic spiracle guard indistinct; setae simple, < 0.1mm long on head; pilifers separated; head rounded; antennal tip lies cephalad of tip of middle leg; proboscis shorter than tip of middle leg; abdomen very long; cremaster to 3.3mm long, broadly rounded with numerous simple setae, hooked setae absent (Figures 111E and 123G); lenticles absent. Other Descriptions Corns tock and Dammers 1934 (egg, larva, pupa), Tinkham 1954 (egg, larva, pupa), Emmel and Emmel 1973 (egg, larva, pupa), Toliver 1987 (larva).

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337 Host Plants AGAVACEAE: Yucca angustifolia (Skinner 1911), Yucca arizonica (Bailowitz and Brock 1991), Yucca arkansana (Freeman 1951b), Yucca baccata (Brown et al. 1957), Yucca bailey i (Brown et al. 1957), Yucca brevifolia (Comstock and Dammers 1934), Yucca campestris (Freeman 1963), Yucca carnerosana (Stallings and Turner 1957), Yucca constricta (Freeman 1965), Yucca elata (Tinkham 1954), Yucca freemanii (Freeman 1969), Yucca glauca (Freeman 1963), Yucca harrimaniae (Brown et al 1957), Yucca louisianensis (Freeman 1963), Yucca navajoa (Scott 1986), Yucca necopina (Freeman 1965), Yucca pallida (Freeman 1963), Yucca reverchoni (Freeman 1963), Yucca rupicola (Freeman 1963), Yucca schidigera (Comstock and Dammers 1934), Yucca schottii (Stallings and Turner 1956), Yucca tenuistyla (Scott 1986), Yucca thompsoniana (Freeman 1963), Yucca thornberyi (Tinkham 1954), Yucca torreyi (Freeman 1963), Yucca treculeana (Stallings and Turner 1957), Yucca verdiensis (Wielgus et al. 1971). Specimens Examined LARVAE: ARIZONA: COCONINO CO.: Lot 793, 1 larva (instar 5) (USNM). MOHAVE CO.: Lot 792, 1 larva (instar 5) (USNM). YAVAPAI CO.: Lot 794, 1 larva (instar 5) (USNM). CALIFORNIA: LOS ANGELES CO.: Lot 641, 2 larvae (instar 5) (TCE); Lot 642, 1 larva (instar 5) (TCE). SAN BERNARDINO CO.: Lot 1496, 1 larva (instar 5) (GRB). TEXAS: BEXAR CO.: Lot 889, 1 larva (instar 5) (ROK); Lot 891, 3 larvae (instar 5) (ROK) BAILEY CO.: Lot

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338 890, 1 larva (instar 5) (ROK). PUPAE: CALIFORNIA: LOS ANGELES CO.: Lot 641, 2 pupae (TCE). 285. Megathymus cofaqui (Strecker) Diagnosis EGG: Figures 31 and 5F, height 2.8-3mm, width 1.5-1. 7mm, polygonal sculpturing, white. LAST INSTAR LARVA: BODY: length 19-22. 5mm, A4 transverse width 5.8-7mm; creamcolored; setae simple, to 0.9mm long on A4 dorsum; spiracles brown. HEAD: dark reddish brown; transverse width 3.64.2mm; sculpturing rough to pebbly; mandibles without teeth; setae simple, to 0.7mm long at apex, a few ventral setae to 0.8mm; stemma 5 and 6 slightly small (Figure 44L) THORAX: legs tan; shield broad, tan dorsal ly. ABDOMEN: prolegs each with about 65 crochets, irregularly triordinal, arranged in transverse bands; suranal plate rounded, tan. PUPA: Figures 77C and 100E, length 34mm, A3 transverse width 8mm; brown; thoracic spiracle guard distinct; setae simple, 0.5mm long on head; pilifers separated; head rounded; antennal and middle leg tips nearly subequal; proboscis extending to tip of middle leg; abdomen long; cremaster to 2.7mm long, broadly rounded with numerous simple setae, hooked setae absent (Figures 111F and 123H); a few lenticles present on the dorsal abdomen. Other Descriptions Skinner 1917 (pupa), Matthews 1986 ( egg 1 arva pupa )

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339 Host Plants AGAVACEAE: Yucca aloifolia (Bonniwell 1916), Yucca filamentosa (Roever 1975), Yucca gloriosa (Roever 1975, perhaps), Yucca smalliana (Roever 1975). Specimens Examined EGGS: FLORIDA: PUTNAM CO.: Lot 327, 2 eggs (MCM) ; Lot 693, 1 egg (MCM) ; Lot 694, 1 egg (MCM); Lot 1512, 2 eggs (MCM). MARION CO.: Lot 695, 2 eggs (MCM). LARVAE: FLORIDA: PUTNAM CO.: Lot 329, 1 larva ( instar 5) (MCM); Lot 694, 1 larva (instar 1) (MCM); Lot 696, 1 larva (instar 4) (MCM). PUPAE: FLORIDA: HILLSBOROUGH CO.: Lot 1004, 1 pupa (FSCA) 286. Megathymus harrisi H. A. Freeman Harris (1954, 1972) described the egg, larva, and pupa of this species. Host Plants AGAVACEAE: Yucca filamentosa (Harris 1954), Yucca smalliana (Remington 1959). 287. Megathymus streckeri (Skinner) Leussler (1930) and Wielgus and Stallings (1974) described the immature stages of this southwestern species, but I could not find specimens for study. Host Plants AGAVACEAE: Yucca angustissima (Roever 1975), Yucca baileyi (Brown et al. 1957), Yucca glauca (Holland 1898), Yucca navajoa (Scott 1986).

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340 288. Megathymus texanus Barnes and McDunnough Wielgus and Stallings (1974) and McCabe and Post (1977) described the immature stages of this Great Plains species. Host Plants AGAVACEAE: Yucca constricta (Roever 1975), Yucca glauca (Freeman 1951b). 289. Megathymus ursus Poling Diagnosis LAST INSTAR LARVA: BODY: length 34-58mm, A4 transverse width 6. 3-11. 2mm; preserved specimens pale; setae simple, to 0.6mm long on A4 dorsum; spiracles tan. HEAD: dark reddish brown; transverse width 3. 9-5. 4mm; sculpturing rough; mandibles without teeth; setae simple, to 0.2mm long at apex, a few ventral setae to 0.8mm; stemmata subequal (Figure 44M) THORAX: legs brown; shield broad, black and brown dorsally. ABDOMEN: prolegs each with about 85 crochets, multiordinal ca. 2-3 ranks, arranged in transverse bands; suranal plate rounded, dark brown. PUPA: Figures 77D and 100F, length 52mm, A3 transverse width 10.2mm; brown; thoracic spiracle guard indistinct; setae simple, to 0.3mm long on head; pilifers separated; head rounded; antennal and middle leg tips nearly subequal; proboscis shorter than tip of middle leg; abdomen long; cremaster to 4mm long, broadly rounded with numerous simple setae, hooked setae absent (Figures 111G and 1231); lenticles absent. Other Descriptions. Wielgus et al. 1972 (larva, pupa), Roever 1975 (egg, larva).

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341 Host Plants AGAVACEAE: Yucca arizonica (Roever 1975), Yucca baccata (Stallings and Turner 1956; Wielgus et al. 1972; Roever 1975), Yucca brevifolia (Bailowitz and Brock 1991), Yucca faxoniana (Roever 1975), Yucca schottii (Stallings and Turner 1956), Yucca thornberyi (Wielgus et al. 1973), Yucca torreyi (Stallings and Turner 1956), Yucca treculeana (Roever 1975). Specimens Examined LARVAE: ARIZONA: PINAL CO.: Lot 795, 1 larva ( instar 4) (USNM) TEXAS: BREWSTER CO.: Lot 892, 1 larva (instar 5) (ROK). PUPAE: TEXAS: BREWSTER CO.: Lot 892, 2 pupae (ROK). 290. Stallingsia maculosa (H. A. Freeman) Freeman (1955) and Roever (1975) described the immature stages of this southwestern species, but I could not find specimens for study. Host Plants AGAVACEAE: Polianthes maculosa (= Manfreda maculosa) (Freeman 1951a).

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342 Figure 19. Last instar larva of Autochton cellus (A), Epargyreus clarus (B), Thorybes py lades (C), Urbanus proteus (D), Urbanus dorantes (E), Polygonus leo (F), Gesta gesta (G) and Erynnis horatius (H).

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343

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344 Figure 20. Last instar larva of Achlyodes thraso (A), Pholisora catullus (B) Pyrgus oileus (C) Staphylus hayhurstii (D), Hesperia attalus (E), Pompeius verna (F) Wallengrenia egeremet (G) and Atrytonopsis hianna (H)

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345

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346 Figure 21. Last instar larva of Atrytone delaware (A), Problema byssus (B) Asbolis capucinus (C), Euphyes arpa (D), Lerema accius (E), Panoquina panoquin (F), Lerodea eufaia (G) and Megathymus yuccae (H)

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347

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348 EYE PATCH Figure 22. Frontal view of the larval head of Phocides palemon (A), Epargyreus clarus (B), Polygonus leo (C) Chioides catillus (D) Typhedanus undulatus (E), and Polythrix mexicana (F). All drawings are enlarged 12 times

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349 Figure 23. Frontal view of the larval head of Polythrix procera (A), Codatractus alcieus (B), c. arizonensis (C) Urbanus proteus (D) Urbanur, dorantes (E) and Astraptes fulgerator (F). All drawinq;: are enlarged 12 times.

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350 Figure 24. Frontal view of the larval head of Astraptes gilberti (A), Autochton cellus (B), Achalarus lyciades (C) Thorybes bathyllus (D), Cabares potrillo (E), and Cogia hippalus (F). All drawings are enlarged 12 times.

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351 Figure 25. Frontal view of the larval head of Nisoniades rubescens (A), Staphylus ceos (B), Carrhenes canescens (C), Xenophanes trixus (D), Systasea pulverulenta (E) Achlyodes thraso (F), Timochares ruptifasciatus (G) and Gesta gesta (H) All drawings are enlarged 12 times.

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352 Figure 26. Frontal view of the larval head of Ephyriades brunneus (A), Erynnis icelus (B), E. brizo (C), E. juvenalis (D), E. propertius (E) and E. horatius (F). All drawings are enlarged 12 times.

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353 Figure 27. Frontal view of the larval head of Erynnis tristis (A), E. martialis (B), E. pacuvius (C), E. zarucco (D), E. funeralis (E) E. baptisiae (F), E. afranius (G) and E. persius (H) All drawings are enlarged 12 times.

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354 MEDIAL LINE EPICRANIAL LINE LATERAL LINE Figure 28. Frontal view of the larval head of Pyrgus centaureae (A), Heliopetes ericetorum (B), Celotes nessus (C), Pholisora catullus (D), Piruna pirus (E), Synapte malitiosa (F) Vidius perigenes (G) and Nastra lherminier (H). All drawings are enlarged 12 times.

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355 Figure 29. tripunctus Ancyloxypha numitor (D), (F), C. minimus (G), and A-C and H are enlarged 12 Frontal view of the larval head of Cymaenes (A), Lerema accius (B), Perichares philetes (C) A. arene (E) Copaeodes aurantiacus Thymelicus lineola (H) Drawings enlarged 25 times. times; D-G are

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356 Figure 30. phyleus (A) Hesperia uncas Polites cor as (G) enlarged 12 times. Frontal view of the larval head of Hylephila retta cams (B) Pseudocopaeodes eunus (C) (E), H. pahaska (F) (D), H. leonardus and P. sabuleti (H) All drawings are

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357 Figure 31. Frontal view of the larval head of Polites mardon (A), P. draco (B), P. baracoa (C) P. themistocles (D), P. ongenes (E) P. mystic (F) P. sonora (G) and P. vibex (H). All drawings are enlarged 12 times.

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358 Figure 32. Frontal view of the larval head of Wallengrenia otho (A), Pompeius verna (B), Atalopedes campestris (C), At ry tone arogos (D) A. delaware (E), Problema byssus (F) Ochlodes sylvanoides (G), and Ochlodes agricola (H) All' drawings are enlarged 12 times.

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359 Figure 33. (A) Poanes aaroni (E) melane (H) Frontal view of the larval head of Ochlodes yuma hobomok (B), P. zabulon (C) P. taxiles (D), P. P. yehl (F), P. viator (G) and Paratrytone All drawings are enlarged 12 times.

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360 Figure 34. Frontal view of the larval head of Choranthus haitensis (A), Euphyes arpa (B) E. pilatka (C) E. alabamae (D), E. dukesi (E), E. bimacula (F), E. ruricola (G) and Asbolis capucinus (H). All drawings are enlarged 12 times

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361 Figure 35. Frontal view of the larval head of Atrytonopsis hianna (A), Amblyscirtes linda (B), A. aesculapias (C) A. vialis (D), A. celia (E), Lerodea eufala (F), Oligoria maculata (G) and Calpodes ethlius (H) All drawings are enlarged 12 times.

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362 Figure 36 Frontal view of the larval head of Panoquina panoquin (A), Nyctelius nyctelius (B) Agathymus valverdiensis (C), and Megathymus yuccae (D) All drawings are enlarged 12 times. gs

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363 o O o o 1 o o o o £? A B c o o o o o o o o o o o o D E F o o o o o o o o o o o O o o G o H I o o o o o o o o v^ o o o o J K L Figure 37. Stemmatal pattern of Pyrrhopyge araxes (A) Phoczdes pigmalion (B) P. palemon (c), Epargyreus zestos (D),E. clarus (E) E. exadeus (F), Polygons leo (G), Chioides catillus (H) Typnedanus undulatus (I), Polythrix mexicana (J), p. procera (K) and Codatractus aJcaeus (L). All drawings are enlarged 25 times.

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364 o o o o c o o o
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365 A o o o o o B o o o c o o o D o o o o o o o o o E o o o o I o o o o J o F o o o o o o o K o G o o o o o o L H o M o o o o M o o O o o o o P o o o Q O o o o o o R Figure 39. Stemmatal pattern of Thorybes bathyllus (A), T. pylades (B), r. diversus (C) T. mexicanus (D), T. confusis (E), I*. drusius (F), Cajbares potrillo (G) Cogia hippalus (H), C. outis (I), C. caicus (J), Nisoniades rubescens (K) Stapnylus mazans (L) S. hayhurstii (M) Carrhenes canescens (N), Xenophanes trixus (0) Systasea pulverulenta (P), S. zampa (Q), and Achlyodes thraso (R) All drawings are enlarged 25 times.

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366 o o o o ^ o o •*\ o o V— o o o O o 1 n o o o o O ^ o A o B o C D o E o o F o c £? o o O o n o o o v^ o o O o o 1 o & o G H I J K o o o O o o O o o o o c o 1 o o o L M N p Q o o o o o R C O 'o s O V o T o o o o O o o u o o V o o o o w o c o X o o o o o O o o o o o o o o o o o o o o o o o o o Y z AA L BB cc DD Figure 40. Stemmatal pattern of Timochares ruptifasciatus (A), Chiomara asychis (B), Gesta gesta (C) Ephyriades brunneus (D), Erynnis icelus (E), E. jbrizo (F), £. juvenalis (G), £. propertius (H), £. horatius (I), £. tristis (J), £. martialis (K) £. pacuvius (L), £. zarucco (M) E. funeralis (N), £. baptisiae (0), £. afranius (P), £. persius (Q), Pyrgus centaureae (R) P. ruralis (S), P. scriptura (T) P. communis (U) P. albescens (V), P. oileus (W) Heliopetes encetorum (X) tf. lavianus (Y), tf. macaira (Z), tf. arsaite (AA), Ceiotes Jiessus (BB) C. limpa (CC) and Pholisora catullus (DD). All drawings are enlarged 25 times.

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367 0^0 o o o o O o o o A B C d E F o o o O o OoOo oO 0 O O o o O o o o o G oo o rt o HI J o K O C? „ o o u o n • • o O o O o o O o O N P Q o O R M o o o o O o O a O q 0. O^ O o v w o o X O o o o o o u o O o o O Y Z AA BB Figure 41. Stemmatal pattern of Pholisora alpheus (A), P. gracielae (B), Piruna pirus (C), Synapte malitiosa (D), Vidius perigenes (E), Nastra lherminier (F), N. julia (G) W. neamathla (H) Cymaemes tripunctus (I), Lerema accius (J), L. liris (K), Aiicyloxypha numitor (L) Perichares philetes (M) A. arejie (N) Copaeodes aurantiacus (0), C. minimus (P), rhymelicus lineola (Q) Hylephila phyleus (R), Yvretta carus (S), Pseudocopaeodes eunus (T), Hesperia uncas (U), tf. ju&a (V), if. comma (W) if. woodgratei (X), if. leonardus (Y) //. pahaska (Z), //. metea (AA) and if. viridis (BB). All drawings are enlarged 25 times.

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368 o o O o A o o o o o o o o o o o o A o B c D E o o c 1 o o o o o o o o O O o c o 1 F G H I J o o ^"\ o o o o o K o o L o o o o M o o o N (9 o o o o o o o o o o o o_ o O o o o p o o o Q o o o R o o o o s o o o o i T o o o o o o o o o o o o o O o o u V o w o X o Y Figure 42. Stemmatal pattern of Hesperia attalus (A) H meskei (B) If. .evada (C) Polices coras (D) P. saouieti (E), p mardon (F), P. draco (G), P. ftaracoa (H) P. themzstocles (I) P. origenes (J), P. cystic (K) P. SM ora (L), P. vitex (M), Wallengrenia otho (N) W. egeremet (o) Pompeius verna (P) Atalopedes campestris (Q) Atrytone arogos R) a. del aware (S), Probieraa oyssus (T) Ochlodes sylvanoxdes (U) O. agricola (V) O. yuL (W) Poanes 25 ?Ses! ^ ^ P Zai?UiOJ2 (Y) A11 Swings are enlarged

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369 O o c 7 o o o A o B O o o 1 O o o o o o c D E O o o o o o O o o o F o o o G o 1 o o o H o o o I o o O o J o o o o O o O o o o o o o O o ^3 O o K L M
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370 o o o O o o o o o o A o o B C o o D O o o o E o o o O O o o o o F o O o G o o H o i o o o < o o o o J o o K o o o L o o H Figure 44. Stemmatal pattern of Oligoria maculata (A) Calpodes ethlius (B), Panoquina panoquin (C) P panoquinoides (D), P. ocola (E), P. sylvicol'a (F) Nyctelius nyctelius (G), Agathymus valverdiensis (H) A. Stephens! (I), Megathyinus yuccae ( J) coloradensis (K), M. cofaqui (L), and w. ursus (M) All drawings are enlarged 25 times

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371 Dorsal Figure 45. Pyrrhopyge zestos (C), and Polygonus enlarged six times. view of the larval terminalia of Phocides pigmalion (B), Epargyreus leo (D). All drawings are

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372 Figure 46 Dorsal view of the larval terminalia of Chioides catillus (A), Typhedanus undulatus (B), Polythrix (C), Codatractus alcaeus (D), Urbanus Astraptes fulgerator (F). All times. mexicana proteus (E), and drawings are enlarged six

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373 Figure 47. Dorsal view of the larval terminalia of Autochton cellus (A), Achalarus lyciades (B), Thorybes bathyllus (C) Cabares potrillo (D), Cogia hippalus (E), Nisomades rubescens (F), Staphylus ceos (G), Carrhenes canescens (H) and Xenophanes trixus (I). All drawings are enlarged six times.

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374 Figure 48. Dorsal view of the larval terminalia of Achlyodes thraso (A), Timochares ruptifasciatus (B) Gesta gesta (C) Ephyriades brunneus (D), Erynnis icelus E) and Pyrgus centaureae (F). All drawings are enlarged six times

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375 Figure 49. Dorsal view of the larval terminalia of Heliopetes ericetorum (A), Celotes nessus (B), Pholisora catullus (C), Piruna pirus (D), Synapte malitiosa (E) Vidius pengenes (F), and Nastra lherminier (G). All drawings are enlarged six times.

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376 Dorsal Figure 50. tripunctus Ancyloxypha numitor view of the larval terminalia of Cymaenes (A), Lerema accius (B) Perichares philetes (c) (D), A. arene (E), Copaeodes aurantiacus (F), and C. minimus (G) Drawings D and F-G times; E is enlarged 12 times. are enlarged 6

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377 ?£££?• 7 D rs f 1 Vlew of the larval terminalia of Thymelicus lmeola (A), Hylephila phyleus (B), Yvretta cams cnk/ltf OCO Z ae r> deS GUnUS (D) Besperia uncas (E), Follies tZls. ( V 'Tc r;/ a ^ ieti (G) D ^ wi ^s A-E are enlarged 6 times, F and G are enlarged 12 times.

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378 Sni^ o V i SW f the larval te ™inalia of Polites sabuleti (A) P mardon (B), P. draco (C), P. baracoa (D), P.thermstocles (E) and P. origei.es (F) Drawings A-E are enlarged 12 times; F is enlarged 6 times.

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379 Figure 53 Dorsal view of the larval terminalia of Polites (D) Pompeius verna (E), Atalopedes campestris (F) and r lll ne i ar 9 T { V-' Drawin 9 s A-C are enlarged 12 times; G are enlarged 6 times. D-

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380 Figure 54 Dorsal view of the larval terminalia of Atrytone Delaware (A), third instar Problema byssus (B), last insta? and Paratrytone melane (F). Drawings A and c-F are enlarged 6 times; B is enlarged 12 times. enlarged

PAGE 409

381 Figure 55. Dorsal view of the larval terminalia of Choranthus haitensis (A), Euphyes arpa (B) E. ruricola Asbolis capucinus (D), Atrytonopsis hianna (E), and Amblyscirtes linda (F). All drawings are enlarged six times. ^ (C)

PAGE 410

382 Ufalt ll\ ? rsal view of the larval terminalia of Lerodea eufala (A ) Oligona maculata (B) Panoquina panoquin (C) P. panoquinozdes (D), P. ocola (E) and P. sylviclla (F) times" 95 are enlarged 6 time s; E-F are enlarged 12

PAGE 411

383 Figure 57. Dorsal view of the larval terminalia of enlarged six > times. S,aW!y US YUCCae fD) Al1 d "9S are

PAGE 412

384 Figure 58. Pupa of Epargyreus clarus (A), Thorybes bathyllus (B), Urbanus proteus (C) Phocides pigmalion (D) Polygonus leo (E), Anastrus sempiternus (F), Achlyodes thraso (G) and Pyrgus communis (H)

PAGE 413

385

PAGE 414

386 Figure 59. Pupa of Hesperia attalus (A), Pompeius verna (B), Poanes yehl (C) Oligoria maculata (D), Cymaenes tripunctus (E), and Synapte malitiosa (F).

PAGE 415

387

PAGE 416

388 Figure 60. Pupa of Nastra lherminier (A), Perichares philetes (B), and Calpodes ethlius (C)

PAGE 417

389 /* WJh

PAGE 418

390 EPICRANIAL SUTURE PROTHORAX -MESOTHORAX METATHORAX SPIRACLE GUARD n^ r ?61 \,? rSal VieW of ant erior pupa of Phocides citrus* m, () L^' A 9 1 1 (B) E P a ^yreus zestos (C) and E clarus (D). All drawings are enlarged six times.

PAGE 419

391 fi? Ur ? h 62 -w Dors ^ 7 view f anterior pupa of Polygonus leo (A), Chioides catillus (B), Polythrix mexicanus (C), proteus (F). All drawings are enlarged six times.

PAGE 420

392 Figure 63. Dorsal esmeraldus (A) U. (D), and Astraptes fulgerator enlarged six times. view of anterior pupa of Urbanus dorantes (B) U. teleus (C), U (E) All drawings are procne

PAGE 421

393 Figure 64. Dorsal view of anterior pupa of Astraptes gilberti (A), Autochton cellus (B), Achalarus lyciades Thorybes bathyllus (D) T. pylades (E), and T. confusis All drawings are enlarged six times. (F)

PAGE 422

394 Figure 65. Dorsal view of anterior pupa of Cajbares potrillo (A), Cogia outis (B) Staphylus ceos (C) S. hayhurstii (D) Carrhenes canescens (E), Systasea pulverulenta (F), Achlyodes thraso (G) and Chiomara asychis (H). All drawings are enlarged six times.

PAGE 423

395 m Figure 66. Dorsal view of anterior pupa of Gesta gesta (A), Ephyriades brunneus (B), Erynnis icelus (C), E. brizo (D) E. juvenalis (E), E. horatius (F), E. tristis (G) and E. martialis (H) All drawings are enlarged six times.

PAGE 424

396 m Figure 67. Dorsal view of anterior pupa (A), E. funeralis (B), E. baptisiae (C) Pyrgus scriptura (E), P. communis (F), P. Heliopetes ericetorum (H) All drawings times. of Erynnis zarucco E. persius (D) oileus (G), and are enlarged six

PAGE 425

397 Figure 68. Dorsal view of anterior pupa of Heliopetes lavianus (A), Celotes nessus (B), Pholisora catullus (C), P. alpheus (D), Synapte malitiosa (E), Nastra lherminier (F) N. julia (G), and N. neamathla (H) All drawings are enlarged six times.

PAGE 426

398 Figure 69. Dorsal view of anterior pupa of Cymaenes tripunctus (A), Lerema accius (B) L. liris (C) Perichares philetes (D) Ancyloxypha numitor (E), A. arene (F), and Copaeodes aurantiacus (G) All drawings are enlarged six times

PAGE 427

399 Figure 70. Dorsal view of anterior pupa of Copaeodes minimus (A), Hylephila phyleus (B) Yvretta carus (C), Pseudocopaeodes eunus (D) Hesperia uncas (E) H. comma (F) H. attalus (G), and Polites coras (H) All drawings are enlarged six times.

PAGE 428

400 Figure 71. Dorsal view of anterior pupa of Polites sabuleti (A), P. mardon (B), P. draco (C) P. baracoa (D) P. themistocles (E), P. origenes (F), P. mystic (G), and P. sonora (H) All drawings are enlarged six times.

PAGE 429

401 Figure 72. Dorsal view of anterior pupa of Polites vibex (A), Wallengrenia otho (B), W. egeremet (C), Pompeius verna (D), Atalopedes campestris (E), At ry tone del aware (F) A arogos (G), and Problema byssus (H). All drawings enlarged six times. are

PAGE 430

402 Figure 73. Dorsal view of anterior pupa of Ochlodes sylvanoides (A), Poanes hobomok (B), P. taxiles (c) aaroni (D), P. yehl (E), P. viator (F), (G), and Choranthus haitensis (H). All enlarged six times. (C), Paratrytone melane drawings are

PAGE 431

403 V Figure 74. Dorsal view of anterior pupa of Euphyes arpa (A), E. pilatka (B) E. alabamae (C) E. dukesi (D) E. berryi (E), and E. bimacula (F). All drawings are enlarged six times.

PAGE 432

404 Figure 75. Dorsal view of anterior pupa of Euphyes ruricola (A), Asbolis capucinus (B), Atrytonopsis loammi (C), Amblyscirtes aesculapias (D), A. celia (E), Lerodea eufala (F), and Oligoria maculata (G). All drawings are enlarged six times.

PAGE 433

405 Figure 76. Dorsal view of anterior pupa of Calpodes ethlius (A), Panoquina panoquin (B), P. panoquinoides (C), P. ocola (D), and P. sylvicola (E). All drawings are enlarged six times.

PAGE 434

406 Figure 77. Dorsal view of anterior pupa of Nyctelius nyctelius (A) Megathymus coloradensis (B), M. cofaqui (C) and M. ursus (D) Drawing A is enlarged six times ;Vd are enlarged three times.

PAGE 435

407 PILIFER FORE WINK EYE FORE LEG MIDDLE LEG ANTENNA TIP OF HIND LEG Figure 78, pigmalion six times. Ventral view of anterior pupa of Phocides (A) and P. palemon (B). All drawings are enlarged

PAGE 436

408 Figure 79. Ventral view of anterior pupa of Epargyreus zestos (A) and E. clams (B). All drawings are enlarged six L. IMG S •

PAGE 437

409 Figure 80. Ventral view of anterior pupa of Polygonus leo (A) and Chioides catillus (B). All drawings are enlarged six times.

PAGE 438

410 Figure 81. Ventral view of anterior pupa of Polythrix mexicana (A), Codatractus alcaeus (B) and C. arizonensis (C). All drawings are enlarged six times.

PAGE 439

411 Figure (A), U are enlarged 82. Ventral view of anterior pupa of Urbanus proteus esmeraldus (B), and U. dorantes (C). All drawings six times.

PAGE 440

412 Figure 83. Ventral view of anterior pupa of Urbanus teleus (A) and U. procne (B). All drawings are enlarged six times

PAGE 441

413 Figure 84. Ventral view of anterior pupa of Astraptes fulgerator (A), A. gilberti (B), and Autochton cellus (C) All drawings are enlarged six times.

PAGE 442

414 Figure 85. Ventral view of anterior pupa of Thorybes bathyllus (A), T. pylades (B), T. confusis (C) Cabares potrillo (D), and Cogia outis (E). All drawings are enlarged six times.

PAGE 443

415 Figure 86. Ventral view of anterior pupa of Staphylus ceos (A), S. hayhurstii (B) Carrhenes canescens (C), Systasea pulverulenta (D), Achlyodes thraso (E), Chiomara asychis (F), Gesta gesta (G), and Ephyriades brunneus (H) All drawings are enlarged six times.

PAGE 444

416 Figure 87. Ventral view of anterior pupa of Erynnis icelus (A), E. brizo (B), E. juvenalis (C) E. horatius (D) E. tristis (E), E. martialis (F), E. zarucco (G) and E. funeralis (H) All drawings are enlarged six times.

PAGE 445

417 Figure 88. Ventral view of anterior pupa of Erynnis baptisiae (A), E. persius (B), Pyrgus scriptura (C), P. communis (D), P. oileus (E), Heliopetes ericetorum (F), lavianus (G), and Celotes nessus (H) All drawings are enlarged six times. H.

PAGE 446

418 Figure 89. Ventral view of anterior pupa of Pholisora catullus (A), P. alpheus (B) Synapte malitiosa (C), Nastra Ihermimer (D), N. julia (E) N. neamathla (F) and Cymaenes tripunctus (G). All drawings are enlarged six times.

PAGE 447

419 Figure 90. Ventral view of anterior pupa of Lerema accius (A), L. liris (B), Perichares philetes (C) Ancyloxypha numitor (D) A. arene (E), Copaeodes aurantiacus (F), and C. minimus (G) Drawings A-B and D-G are enlarged six times; C is enlarged three times.

PAGE 448

420 Figure 91. Ventral view of anterior pupa of Hylephila phyleus (A), Yvretta carus (B), Pseudocopaeodes eunus (C) Hesperia uncas (D), H. comma (E), H. attalus (F), Polites coras (G), and P. sabuleti (H) All drawings are enlarged six times.

PAGE 449

421 Figure 92. Ventral view of anterior pupa of Polites mardon (A), P. draco (B), P. baracoa (C), P. themistocles (D) P ongenes (E), P. mystic (F), P. sonora (G), and P. viipex (H). All drawings are enlarged six times.

PAGE 450

422 Figure 93 otho (A), Atalopedes times Ventral view of anterior pupa of Wallengrenia W. egeremet (B) Pompeius verna (C), and campestris (D). All drawings are enlarged six

PAGE 451

423 Figure 94. Ventral view of anterior pupa of Atrytone arogos (A), A. del aware (B) Problema byssus (C), and Ochlodes sylvanoides (D). All drawings are enlarged six times.

PAGE 452

424 Figure 95. Ventral view of anterior pupa of Poanes hobomok (A), P. taxiles (B), P. aaroni (C), and P. yehl (D) All drawings are enlarged six times.

PAGE 453

4 25 Figure 96. Ventral view of anterior pupa of Poanes viator (A), Paratrytone melane (B) Choranthus haitensis (C), Euphyes arpa (D). All drawings are enlarged six times, and

PAGE 454

426 Figure (A), E bimacula enlarged 97. Ventral view of anterior pupa of Euphyes pilatka alabamae (B), E. dukesi (C) E. berryi (D), E. ruricola (F). All drawings are (E) six and E. times.

PAGE 455

427 Figure 98. Ventral view of anterior pupa of Asbolis capucmus (A), Atrytonopsis loammi (B), Amblyscirtes aesculapias (C), A. celia (D), Lerodea eufala (E), and Oligona maculata (F). All drawings are enlarged six times

PAGE 456

428 Figure 99. Ventral view of anterior ethlius (A), Panoquina panoquin (B) P. ocola (D), and P. sylvicola (E). enlarged six times. pupa of Calpodes P. panoquinoides (C) All drawings are

PAGE 457

429 Figure 100. Ventral view of anterior pupa of Nyctelius nyctelius (A), Agathymus valverdiensis (B), Megathymus yuccae (C), M. coloradensis (D), M. cofaqui (E), and M. ursus (F). All drawings are enlarged three times.

PAGE 458

430 Figure 101. Lateral view of the cremaster of Phocides pigmalion (A), P. palemon (B), Epargyreus zestos (C), E. clarus (D), Polygonus leo (E), Chioides catillus (F), Polythrix mexicana (G), Codatractus alcaeus (H) and C. arizonensis (I). All drawings are enlarged 12 times.

PAGE 459

431 Figure 102. Lateral view of the cremaster of Urbanus proteus (A), U. esmeraldus (B) U. dorantes (C) U. teleus (D), U. procne (E), Astraptes fulgerator (F), A. gilbert i (G), Autochton cellus (H) and Achalarus lyciades (I). All drawings are enlarged 12 times.

PAGE 460

432 y> Figure 103. Lateral view of the cremaster of Thorybes bathyllus (A), T. pylades (B), T. confusis potrillo (D), Cogia hippalus (E), C. outis (G) Nisoniades rubescens (H) and drawings are enlarged 12 times. (C), Cabares (F) C. caicus Staphylus ceos (I). All

PAGE 461

433 Figure 104 mazans (A) Xenophanes thraso (F) (H), Gesta icelus (K) E. tristis enlarged 1 Lateral view of the cremaster of Staphylus S. hayhurstii (B), Carrhenes canescens (C) trixus (D) Systasea pulverulenta (E), Achlyodes Timochares ruptifasciatus (G), Chiomara asychis gesta (I), Ephyriades brunneus (J), Erynnis E. brizo (L), E. juvenalis (M) £. horatius (N) (0), and £. martialis (P). All drawings are 2 times.

PAGE 462

434 Figure 105. Lateral view zarucco (A), E. funeralis ( D ) Pyrgus scriptura ( E ) Heliopetes ericetorum (H) (J), Pholisora catullus (K) P. alpheus (L) malitiosa (M) Nastra lherminier (N) N. julia (0) neamathla (P). All drawings are enlarged 12 times of the cremaster of (B) E. baptisiae P. communis (F) H. lavianus (I) Erynnis (C) E. persius P. oileus (G) Celotes nessus Synapte and N.

PAGE 463

435 H Figure 106. Lateral view of the cremaster of Cymaenes tripunctus (A), Lerema accius (B), L. liris (C), Perichares philetes (D) Ancyloxypha numitor (E) A. arene (F), Copaeodes aurantiacus (G), C. minimus (H) Hylephila phyleus (I), Yvretta carus (J), Pseudocopaeodes eunus (K) and Hesperia uncas (L) Drawings A-C and E-L are enlarged 12 times; D is enlarged 6 times.

PAGE 464

436 Figure (A), H. Polites (H), P. P. mystic otho (0) times. 107. Lateral view of the cremaster of Hesperia comma woodgatei (B), H. viridis (C) H. attains (D), coras (E), P. sabuleti (F), P. mardon (G) P. draco baracoa (I), P. themistocles (J), P. origenes (K) (L), P. so/iora (M) and W. egeremet (P) P. vibex (N), Wallengrenia All drawings are enlarged 12

PAGE 465

437 Figure 108. Lateral view of the cremaster of Pompeius verna (A), Atalopedes campestris (B), Atrytone arogos (C), A. delaware (D), Problema byssus (E) Ochlodes sylvanoides (F), Poanes hobomok (G) P. taxiles (H), P. aaroni (I), and P. yehl (J). All drawings are enlarged 12 times.

PAGE 466

438 Figure 109. Lateral view of the cremaster of Poanes viator (A), Paratrytone melane (B), Choranthus haitensis (C), Euphyes arpa (D), E. pilatka (E), E. alabamae (F), E. dukesi (G), E. berryi (H) E. macguirei (I), E. ruricola (K) and Asbolis capucinus (L) H-K are enlarged 12 times; D-G and L are bimacula ( J) E. Drawings A-C and enlarged 6 times.

PAGE 467

439 Figure 110. Lateral view of the cremaster of Atrytonopsis loammi (A), Amblyscirtes aesculapias (B), A. vialis (C), A. celia (D), Lerodea eufala (E), Oligoria maculata (F), Calpodes ethlius (G) Panoquina panoquin (H) P. panoquinoides (I), and P. are enlarged 12 times; G ocola ( J) is enlarged Drawings 6 times. A-F and H-J

PAGE 468

440 Figure 111. Lateral view of the cremaster of Panoquina sylvicola (A), Nyctelius nyctelius (B), Agathymus valverdiensis (C), Megathymus yuccae (D), M. coloradensis (E), M. cofaqui (F), and M. ursus (G) Drawings A-B are enlarged 12 times; C-G are enlarged 6 times.

PAGE 469

441 Figure 112. Dorsal view of the cremaster of Phocides pigmalion (A), P. palemon (B), Epargyreus zestos (C), E. clarus (D), Polygonus leo (E), and Chioides catillus (F) Drawings A and B are enlarged 6 times; C-F are enlarged 12 times

PAGE 470

442 Figure 113. Dorsal view of the cremaster of Polythrix mexicana (A), Codatractus alcaeus (B), C. arizonensis (C) Urbanus proteus (D) U. esmeraldus (E), U. dor antes (F), U, teleus (G), U. procne (H) and Astraptes fulgerator (I). All drawings are enlarged 12 times.

PAGE 471

443 Figure 114. gilberti (A) Thorybes bathyllus (D) T. Cabares potrillo (G), Cogia hippalus All drawings are enlarged 12 times. Dorsal view of the cremaster of Astraptes Autochton cellus (B) Achalarus lyciades (C) (D), T. pylades (E), T. confusis (F), (H) and C. outis (I)

PAGE 472

444 Figure 115. Dorsal view of the cremaster of Cogia caicus (A), Nisoniades rubescens (B), Staphylus ceos (C), S. mazans (D), S. hayhurstii (E), Carrhenes canescens (F), Xenophanes trixus (G), Systasea pulverulenta (H) Achlyodes thraso Timochares ruptifasciatus (J), Chiomara asychis (K) and Gesta gesta (L) All drawings are enlarged 12 times. (I)

PAGE 473

445 Figure 116. Dorsal view of the cremaster of Ephyriades brunneus (A), Erynnis icelus (B) E. brizo (C) E. juvenalis (D), E. horatius (E), E. tristis (F), E. martialis (G) E. zarucco (H) E. funeralis (I), E. baptisiae (J), E. persius (K), and Pyrgus scriptura (L). All drawings are enlarged 12 times.

PAGE 474

446 Figure 117. Dorsal view of the cremaster of Pyrgus communis (A), P. oileus (B), Heliopetes ericetorum (C), H. lavianus (D), Celotes nessus (E), Pholisora catullus (F), P. alpheus (G), Synapte malitiosa (H) Nastra lherminier (I), N. julia (J), N. neamathla (K) and Cymaenes tripunctus (L). All drawings are enlarged 12 times.

PAGE 475

447 Figure 118. Dorsal view of the cremaster of Lerema accius (A), L. liris (B) Perichares philetes (C), Ancyloxypha numitor (D) A. arene (E), Copaeodes aurantiacus (F), C. minimus (G), Hylephila phyleus (H), Yvretta carus (I), Pseudocopaeodes eunus (J), and Hesperia uncas (K) Drawings A-B and D-K are enlarged 12 times; C is enlarged 6 times.

PAGE 476

448 Figure 119. Dorsal view of the cremaster of Hesperia comma (A), H. woodgatei (B) H. viridis (C), H. attalus Polites coras (E), P. sabuleti (F) P. mardon (G) (H), P. baracoa (I), p. themistocles (J), and P. (K) All drawings are enlarged 12 times. (D) P. draco origenes

PAGE 477

449 Figure 120. Dorsal view of the cremaster of Polites mystic (A), P. sonora (B), P. vibex (C) Wallengrenia otho (D), W. egeremet (E), Pompeius verna (F), Atalopedes campestris (G) ( Atrytone arogos (H), A. del aware (I), and Projbiema byssus (J). Drawings A-I are enlarged 12 times; J is enlarged 6 times.

PAGE 478

450 w I — / w. w. ** Figure 121. sylvanoides aaroni (D), P. yehi (G), Choranthus (J) Dorsal view of the (A) Poanes hobomok (E) P. viator (F) cremaster of Ochlodes (B) taxiles (C) P. Par atry tone melane haitensis (H) Euphyes arpa (I), £. pilatka (K), £. dukesi (L) and £. jberryi (M) are enlarged 12 times; B, F, and Drawings A, C-E, I-L are enlarged G-H, and 6 times. M

PAGE 479

451 ^mTrnnrn^ Figure 122. Dorsal view of the cremaster of Euphyes macguirei (A), E. bimacula (B) E. ruricola (C), Asbolis capucinus (D), Atrytonopsis loammi (E), Amblyscirtes aesculapias (F) A. vialis (G) A. celia (H) Lerodea eufala (I) Oligoria maculata (J), and Calpodes ethlius (K) Drawings A-C and E-J are enlarged 12 times; D and K are enlarged 6 times.

PAGE 480

452 Figure 123. Dorsal view of the cremaster of Panoquina panoquin (A), P. panoquinoides (B), P. ocola (C), P. sylvicola (D), Nyctelius nyctelius (E), Meqathymus yuccae (F), M. coloradensis (G) M. cofaqui (H), and M. ursus (I) Drawings A-E are enlarged 12 times; F-I are enlarged 6 times.

PAGE 481

CHAPTER 6 TAXONOMIC KEYS TO THE IMMATURE STAGES Preliminary Key to Eggs of U. S. Hesperiidae (Subfamily and Genus ) la. Egg diameter > 2mm Megathyminae (5) lb Egg diameter < 2mm ( 2 ) 2a. Egg with 30 to 50 vertical rows of short spines, on Malvaceae Celotes Heliopetes Systasea 2b. Egg without rows of spines ( 3 ) 3a. Egg with 10 to 30 rows of vertical ribs, usually on dicotyledonous plants Pyrginae (6) 3b. Egg with polygonal sculpturing or smooth, not ribbed (4) 4a. Egg on dicotyledonous plants (Fabaceae) Cogia ( Pyrginae ) 4b. Egg on monocotyledonous plants Heteropterinae Hesperiinae ( 11 ) 5a. Egg green with red blotches or entirely red, on Agave species Agathymus 5b. Egg brownish or white, on Yucca species Megathymus 6a. Egg red Phocides 6b Not as above / y \ 453

PAGE 482

454 7a. Egg yellow or orange Achlyodes Autochton, Chioides, Gesta, Erynnis Timochares Urbanus Zestusa 7b. Not as above / 8 x 8a. Egg green Cabares Epargyreus Erynnis, Polygonus Pyrgus Thorybes Urbanus 8b Not as above / g v 9a Egg white Achalarus, Ephyriades Pholisora, Pyrgus, Thorybes 9b. Not as above 1Q 10a. Egg gray Typhedanus 10b. Egg brown Pholisora, Staphylus 11a. Egg green, yellow, or white with a red ring Amblyscirtes Ancyloxypha, Atrytone, Choranthus Euphyes lib Egg lacking a red ring 2) 12a. Egg pink Polites 12b. Not as above ,. 13a. Egg pale green or whitish Amblyscirtes, Atalopedes Carterocephalus Copaeodes Cymaenes, Hesperia, Hylephila, Lerema, Lerodea, Nastra, Nyctelius, Oarisma, Ochlodes Panoquina, P ar atrytone Perichares, Poanes Polites, Pompeius Problema, Pseudocopaeodes Thymelicus Wallengrenia 13b. Not as above ,. M % (14) 14a. Egg gray Calpodes 14b. Egg brown Atrytonopsis Oligoria

PAGE 483

455 Preliminary Key to Last Instar Larvae of U. S. Hesperiidae (Subfamily and Genus) la. Lenticles and usually an anal comb and a ventral prothoracic gland present; larva living in a leaf shelter or boring in Agavaceae Hesperiidae (2) lb. Not as above other Lepidoptera 2a. More than 3 supraspiracular lenticles present on each side of abdominal segment 4; anal comb weakly developed or absent; larva boring in stems, roots, or leaves of Agavaceae Megathyminae ( 5 ) 2b. Fewer than 3 supraspiracular lenticles present on each side of abdominal segment 4; anal comb well developed; larva usually living in a leaf shelter (3) 3a. Mandibles of pyrgine type; proleg lenticles one or two; bright yellow or red pigments present; larva usually feeding on dicotyledonous plants ( 4 ) 3b. Mandibles of hesperiine type; three or more proleg lenticles present; bright yellow or red pigments absent; larva feeding on monocotyledonous plants Heteropterinae Hesperiinae ( 35 ) 4a. Head with low vertical ridges; setae > 5mm long; ground color dull red with transverse yellow stripes; on Quercus species Pyrrhopyginae (Pyrrhopyge araxes) 4b. Head lacking ridges; setae < 1mm long; ground color green, yellowish, or occasionally red Pyrginae (6)

PAGE 484

456 5a. Crochets arranged in transverse bands; thoracic segments enlarged; boring in Yucca species .Megathymus 5b. Crochets arranged in a circle; abdominal segments 3-5 largest; boring into Agave species Agathymus 6a. Prothoracic legs brown, dark brown, or black (7) 6b. Prothoracic legs pale, red, or lightly sclerotized (tan) (14) 7a. Head with yellow, orange, or reddish eye patches ...(8) 7b. Head without colorful eye patches absent (io) 8a. Body setae with expanded tips, on Combretaceae Polythrix 8b Body setae simple on Fabaceae ( 9 ) 9a. Head reddish brown with large yellow eye patches, body with a broad, yellow lateral stripe Autochton 9b. Head brown, lower face blackish, with orange or reddish eye patches, lacking a broad, yellow lateral stripe ... Urbanus 10a. Setae on head feathery or branching, on Malvaceae Heliopetes Pyrgus 10b. Setae on head simple (H) 11a. Head light brown with 6 black spots on face, on Fabaceae Gesta lib. Head black / 12 ) 12a. Crochets of each proleg of uniform size (irregularly triordinal ) on Fabaceae Urbanus

PAGE 485

457 12b. Posterior crochets of each proleg much larger than anterior crochets (irregularly triordinal) (13) 13a. Prothoracic shield tan dorsally, brown laterally, on Lamiaceae and Verbenaceae Cabares 13b. Prothoracic shield darkly sclerotized, on Fabaceae Achalarus Thorybes 14a. Prothoracic shield distinctly present, tan, brown, or black in color ( 15 ) 14b. Prothoracic shield absent or indistinct (23) 15a. Setae on head feathery or branching (16) 15b. Setae on head simple (18) 16a. Head with yellow or orange eye patches present, on Fabaceae Typhedanus 16b. Head uniformly dark, without pale eye patches, on Amaranthaceae and Chenopodiaceae ( 17 ) 17a. Prothoracic shield darkly sclerotized Pholisora 17b. Prothoracic shield lightly sclerotized (tan) or indistinct Staphylus 18a. Head uniformly black, without eye patches, on Convolvulaceae Nisoniades 18b. Head not black, colorful yellow or orange eye patches usually present ( 19 ) 19a. Body white, on Myrtaceae or Rhizophoraceae Phocides 19b. Body green or red with fine or broad yellow stripes (20)

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458 20a. Head uniformly brown or reddish brown (except for eye patches ) ( 21 ) 20b. Head brown with black patterns ( 22 ) 21a. Body with patches of black microspines and numerous narrow, pale yellow transverse stripes per segment, on Fabaceae Epargyreus 21b. Body lacking patches of black microspines, yellow stripes longitudinal or body red with 1 transverse yellow stripe per segment, usually on Fabaceae, occasionally on Aguifoliaceae, Asteraceae, Rhamnaceae, Trigoniaceae, or Verbenaceae Astraptes 22a. Head with two black eye spots on lower face, on Fabaceae Chioides 22b. Head without black eyespots, lower part of face black, on Fabaceae Codatractus 23a. Crochets arranged in a mesal penellipse (24) 23b. Crochets arranged in a complete or nearly complete circle ( 2 g) 24a. Head color light brown or with a contrasting pattern (25) 24b. Head uniformly dark brown or black (27) 25a. Crochets uniordinal, on Malvaceae Systacea 25b. Crochets multiordinal ( 26 ) 26a. Head setae branching, small yellow eye patches usually present, on Fabaceae Cogia

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459 26b. Head setae simple, eye patches absent, on Malpighiaceae Timochares 27a. Larva on Malvaceae Celotes 27b. Larva on Chenopodiaceae (28) 28a. Larva on Chenopodium species Staphylus 28b. Larva on Atriplex species Pholisora 29a. Head pale with 2 black eye spots on upper face, on Fabaceae (30) 29b. Head brown or dark brown, without black eyespots ..(31) 30a. Head with a yellow and black lateral stripe .Polythrix 30b. Head with a black lateral stripe Polygonus 31a. Upper most eye (stemmata 1) at least twice as large as the other eyes, usually on Rutaceae Achlyodes 31b. Eyes subequal in size (32) 32a. Head uniformly dark brown or black, without eye patches on Malvaceae Carrhenes 32b. Head brown or light brown, usually with orange spots or eye patches ( 33 ) 33a. Body setae > 0.4mm in length, on Malvaceae Xenophanes 33b. Body setae < 0.2mm in length, not on Malvaceae (34) 34a Larva on Malpighiaceae Ephyriades 34b. Larva usually on Fabaceae or Fagaceae (sometimes on Betulaceae, Rhamnaceae, or Salicaceae) Erynnis 35a. Prothoracic legs dark brown or black (36)

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460 35b. Prothoracic legs pale, concolorous with body or lightly sclerotized (tan) (49) 36a. Head pale with or without dark stripes (37) 36b. Head black with or without pale stripes or spots ..(39) 37a. Body setae > 0.5mm long, head pale Perichares 37b. Body setae < 0.5mm long, head with dark stripes ...(38) 38a. Suranal plate with a black crescent Atrytone 38b. Suranal plate without dark markings Nyctelius 39a Head black unmarked ( 40 ) 39b. Head black with pale stripes or spots (42) 40a. Crochets arranged in a mesal penellipse, mesoand metathoracic legs pale Pompeius 40b. Crochets arranged in a circle or near circle, all thoracic legs dark (41) 41a. Thorax with a pale (yellowish or reddish in life) lateral stripe, larva living in a leaf case clipped from the host Wallengrenia 41b. Thorax lacking a pale lateral stripe, larva living in shelters of folded or tied leaves Polites 42a Frons bearing 2 pale spots ( 43 ) 42b. Frons without pale spots (47) 43a. Pale eye patch with a central black spot Hylephila 43b. Eye patch absent or lacking a black spot (44) 44a. Suranal plate without black markings (45) 44b. Suranal plate with black markings (46) 45a. Body setae simple Polites

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461 45b. Body setae with expanded tips Hesperia 46a. Suranal plate with 3 sharply delineated black lines Yvretta 46b. Suranal plate markings irregular or diffuse .Polites 47a. Body setae with expanded tips Atalopedes 47b. Body setae simple (48) 48a. Suranal plate with dark markings Polites 48b. Suranal plate without dark markings . Pseudocopaeodes 49a. Prothoracic shield pale, indistinct (50) 49b. Prothoracic shield brown or black (at least the groove between the major annuli dark) (55) 50a. Head pale, unmarked (occasionally faint stripes) ..(51) 50b. Head pale with bold reddish, brown, or black markings (53) 51a. Crochets arranged in a mesal penellipse, suranal plate pointed, body length < 20mm Copaeodes 51b. Crochets arranged in a circle or near circle, suranal plate rounded, body length usually > 20mm (52) 52a. Body setae with blunt tips Panoquina 52b. Body setae simple Thymelicus 53a. Head brown with paler stripes, medial stripe dark brown along the vertex Lerodea 53b. Head pale with dark stripes (54) 54a. Head with reddish stripes, the epicranial stripe separate from the medial stripe Nastra

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462 54b. Head with dark brown or black stripes, epicranial stripe broadly convergent with the medial stripe along the adfrontal sclerite Cymaenes 55a Paraclypeal hooks present .Amblyscirtes (Hesperiinae) Piruna (Heteropterinae) 55b Paraclypeal hooks absent ( 56 ) 56a. Head color uniform brown or pale orange (57) 56b. Head with dark stripes ( 60 ) 57a Body setae > 1mm long Atrytonopsis 57b Body setae < 1mm long ( 58 ) 58a. Head light orange, suranal plate broadly rounded and edged with black, on Arecaceae Asbolis 58b. Head brown, suranal plate not edged with black, on Poaceae ( 59 ) 59a. Posterior half of prothoracic dorsum dark, on Poaceae Oligoria 59b. Only the groove between the major annuli of the dorsal prothorax dark, on Poaceae and Cyperaceae Poanes Paratrytone 60a. Head light brown with a central black spot on face and on area around the eyes, cuticle transparent in life, usually on Cannaceae and Marantaceae Calpodes 60b. Head with dark stripes, cuticle opague (61) 61a. Only the groove between the major annuli of dorsal prothorax dark ( 62 ) 61b. Posterior half of prothoracic dorsum dark (65)

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463 62a. Epicranial stripe present on face (63) 62b. Epicranial stripe absent (64) 63a. Epicranial stripe convergent with the medial stripe along the adfrontal sclerite Lerema, Vidius 63b. Epicranial stripe separate from the medial stripe Atrytone 64a. Frontal sclerite with dark stripes Synapte 64b. Frontal sclerite pale, without dark stripes Ochlodes 65a. Epicranial stripe broadly convergent with the medial stripe along the adfrontal sclerite (66) 65b. Epicranial stripe separate from the medial stripe or with narrow lateral connections (not broadly joined) (67) 66a. Medial stripe with a black elongate spot along the vertex (larva with a cyclopian appearance), body length usually > 25mm, on Arecaceae and Cyperaceae .Euphyes 66b. Medial stripe not formed into an elongate spot, body length < 25mm, on Poaceae Ancyloxypha 67a. Frontal sclerite with dark stripes Problema 67b. Frontal sclerite pale, without dark stripes Choranthus Prelimin ary Key to Pupae of U. s. Hesperiidae (Subfamily and Genus 1 la. Lenticles present on prothorax and/or abdomen, proboscis extending beyond the wing tips, pilifers usually touching Hesperiinae (11)

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464 lb. Lenticles absent, proboscis extending to the wing tips or shorter, pilifers separated by the labial sclerite (2) 2a Setae > 2mm in length Pyrrhopyginae (Pyrrhopyge araxes) 2b Setae < 2mm in length ( 3 ) 3a. Proboscis extending to the wing tips Pyrginae (5) 3b. Proboscis much shorter than the wing tips, only reaching about to the tip of the middle legs Megathyminae ( 4 ) 4a. Cremaster broadly rounded with abundant stiff setae Megathymus 4b. Cremaster pointed, setae absent Agathymus 5a. Posterior margin of the prothorax crenulated, irregular (6) 5b. Posterior margin of the prothorax straight (7) 6a. Pupal cap with a short pointed process Polythrix 6b. Pupal cap rounded Achalarus Astraptes Autochton, Cabares, Chioides Codatractus Thorybes Urbanus 7a. Thoracic spiracle guard absent or indistinct (8) 7b. Thoracic spiracle guard well developed (9) 8a. Pupal cap and anterior margin of eyes each with a short, pointed process Polygonus 8b. Pupal cap with a mesal bulge bearing a shallow indentation Phocides

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465 9a. Longest setae > 0.5mm Carrhenes Celotes Heliopetes, Pholisora, Pyrgus Staphylus Systasea 9b Longest setae < 5mm m o ) 10a. Pupal cap with a short pointed process Achlyodes 10b Pupal cap rounded Cogia, Chiomara, Ephyriades Erynnis Gesta 11a. Pupal cap with a pointed process (12) lib. Pupal cap rounded (17) 12a. Antennal tip extending to near tip of middle leg ..(13) 12b. Antennal tip far cephalad of tip of middle leg (14) 13a. Body length < 25mm, process on pupal cap short Synapte 13b. Body length > 25mm, process on cap relatively long Perichares 14a. Proboscis extending beyond abdominal segment 7 (15) 14b. Proboscis not extending beyond A 7 (16) 15a. Proboscis extending well beyond the tip of the cremaster, body length > 40mm Calpodes 15b. Proboscis not extending beyond the tip of the cremaster, body length < 40mm Cymaenes Lerema Lerodea Nastra 16a. Body length < 25mm Copaeodes 16b. Body length > 25mm Panoquina 17a. Cremaster with a short, medial, ventral spine (18) 17b. Cremaster without a medial spine ( 19 \ 18a. Proboscis extending beyond the cremaster Atrytone

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466 18b. Proboscis not extending beyond abdominal segment 6 .... Ancyloxypha 19a. Cremaster tapering to a blunt point (27) 19b. Cremaster broadly rounded or sguarish, not tapering to a point ( 20 ) 20a Cremaster with hooked setae ( 21 ) 20b. Cremaster without hooked setae (25) 21a. Proboscis extending into abdominal segment 7 (22) 21b. Proboscis extending only into A 6 Pompeius 22a Cremaster rounded ( 23 ) 22b. Cremaster sguarish Nyctelius 23a. Cremaster with a few short dorsolateral spines Choranthus 23b. Cremaster without spines ( 24 ) 24a. Cremaster with hooked setae in a medial tuft Wallengrenia 24b. Cremaster with hooked setae in a row along the distal margin Asbolis 25a. Cremaster with a few short dorsolateral spines Euphyes 25b. Cremaster without spines ( 26 ) 26a. Distal tip of cremaster with a mesal indentation Poanes 26b. Distal tip of cremaster rounded, not indented Atrytone

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467 aei. Distal tip of cremaster bearing a tuft of hooked setae Amblyscirtes Atalopedes Atrytonopsis Hesperia, Hylephila, Oc hi odes Paratrytone Poanes Polites, Problema, Pseudocopaeodes Yvretta 27b. Cremaster with hooked setae in a row along the distal mar, ? in Oligoria

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CHAPTER 7 SUMMARY AND CONCLUSIONS Although a large literature exists on the immature stages of U. S. skippers, previous authors have largely described superficial characters, without reference to other species. This project furthers our understanding of the biology and morphology of immature stages of U. S. hesperiids by determining the extent of their biological and morphological variation, comparing their biology and morphology in a standardized way, and providing keys for their identification. Biology Most hesperiids are leaf feeders that hide in shelters of folded or tied leaves. The Megathyminae have diverged greatly from other skipper groups in that these larvae are borers in the leaves, stems, and roots of plants in the Agavaceae. The genus Agathymus (Megathyminae) is unique in largely feeding on the host's sap during the later instars. Skipper butterflies tend to be specialized in their choice of host plants. Some 92% of the U. S. hesperiids feed on only one family of plants. At the other extreme, one species, Astraptes fulgerator (Pyrginae), has been recorded from plants of six different families. Species 468

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469 having relatively broad host ranges, but mostly within one plant family, include Lerema accius (48 hosts), Urbanus proteus (38 host spp.)/ Pyrgus communis (35 spp. ) Epargyreus clarus (31 spp.), Thorybes py lades (27 spp.), and Megathymus yuccae (27 spp.). Overall, 31 families of plants are eaten by hesperiids recorded from the U. S., including 8 families of monocots and 23 families of dicots. Hesperiinae, Heteropterinae and rarely Pyrginae feed on monocots, whereas Pyrrhopyginae and most Pyrginae eat dicots. At least 138 new host records are reported for 55 species of U. S. Hesperiidae. Host plants remain to be discovered for 22% of the 290 skipper butterflies recorded from the U. S. The eggs of skippers may be brownish, pinkish, white, yellow, green, or red. Adult females usually deposit the eggs singly on the host leaves, but some species oviposit on other substrates. A few pyrgines such as Autochton cellus and Urbanus proteus lay short strings of eggs. The head of the first instar larva is usually black. Later instars may develop stripes, spots, or colorful eye patches on the head. Frass is usually propelled away from the larva by the anal comb. Megathymines, however, have degenerate anal combs and simply drop the frass pellet from the anus. Hesperiid larvae are attacked by parasitic wasps, such as braconids, ichneumonids, and eulophids, as well as tachinid flies. Secretions from the ventral prothoracic

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470 gland may deter invertebrate predators. Defenses used to protect against vertebrate predation include dropping to the ground and remaining motionless, cryptic as well as aposematic coloration, aggressive displays, regurgitation of fluids from the mouth, and defecation. Skipper larvae usually have five larval instars, and occasionally more. Most U. S. species overwinter as partly or fully grown larvae and sometimes as fully formed larvae within eggs, first instars, or pupae. The last instar of hesperiines and megathymines may produce particles of white wax from specialized glands on the ventral side of the abdomen. The wax is used by the larva to coat the inside of the cocoon or plug the pupal chamber. The pupa of most skipper butterflies is supported in the cocoon by a silk thread about the middle, and the cremaster is anchored by hooked setae to a small pad of silk. Megathyminae and some hesperiines do not have hooked setae on the cremaster and do not anchor the pupa to the cocoon. Megathyminae pupate within the larval burrows. Future research on the biology of U. S. hesperiids should be directed towards investigating species whose life history is currently not known. In addition, errors in the use of particular plants by skipper larvae have been incorporated into the literature due to mistaken plant and butterfly identification, misinterpretation of old illustrations and common names, ovipositional mistakes or

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471 deliberate oviposition on nonhosts, and laboratory rearings. Biologists should attempt to discover new host plant associations, as well as verify published records. Morphology The eggs of U. S. hesperiids range in height from 0.4 to 1.8mm and in diameter from 0.6 to 3mm. Megathyminae Hesperiinae, and rarely Pyrginae have polygonal patterns of sculpturing on the eggs. Pyrgines usually have 10 to 50 vertical ribs or rows of short spines and vary from 0.5 to 1.3mm in height and 0.6 to 1.8mm in width. Similarly, hesperiine eggs range in height from 0.4 to 1.2mm and width of 0.6 to 1.6mm. In comparison, megathymines have very large eggs (height = 1.3-1. 8mm, width 2.5-3mm). No specimens of Pyrrhopyginae or Heteropterinae eggs were located for study. Those of the Heteropterinae have been described as having vertical grooves. Last instar larvae of U. S. hesperiids vary in length from 8 to 86mm. The width at A4 ranges from 1.6 to 11.7mm and the transverse width of the head may be 1.5 to 7.2mm. Pyrgines tend to be somewhat shorter and thicker bodied than hesperiines. Pyrgine lengths range from 8 to 49mm, while hesperiines may be 9.8 to 63mm long. Body widths vary 1.8 to 9.8 for pyrgines and 1.6 to 9.5mm for hesperiines. Pyrgines also have wider heads than hesperiines, 2 to 6.4mm and 1.5 to 4.4mm respectively. Megathymines are usually much larger and thicker than other skippers. Body lengths

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472 of megathymines range from 19 to 86mm. Body width varies from 5.8 to 11.7mm, and head width may be 3.3 to 5.8mm. Interesting morphological characteristics of hesperiid larvae that may be useful in phylogenetic analyses include the type of mandible articulation, setal types, sculpturing of the head, lenticle distribution, wax gland patterns, and presence of red or yellow pigments. Pyrrhopygines, pyrgines, and megathymines have mandibles typical of higher lepidopterans. Heteropterines and hesperiines have a modified type of articulation. Pyrgines freguently have modified setae with expanded tips on the body or branching setae on the head. Some hesperiines also have setae with expanded tips. Pyrrhopygines have ridges on the head, a sculpturing pattern not found in other groups. Lenticle patterns also vary between subfamilies. Pyrgines sometimes have numerous lenticles on the ventral side of the prothorax, a feature shared by some Old World coeliadines. Only one or two lenticles on the prolegs are characteristic of pyrrhopygines and pyrgines. Hesperiinae have 4 to 15 lenticles at this position. Subdorsal lenticles are not usually present in pyrrhopygines or pyrgines, but freguently occur on hesperiines. Megathymines also have lenticles, but differ from other subfamilies in their distribution. In megathymines, the lenticles on the abdomen are often in a supraspiracular cluster, ranging from 3 to 15 per side of

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473 segment. Wax glands occur only in the Megathyminae Hesperiinae, and probably Heteropterinae. Patches of wax glands may occur on the ventral side of abdominal segments one, three to six, or seven and eight. Bright red, orange and yellow pigments freguently occur in Pyrginae and Pyrrhopyginae larvae. Hesperiid pupae range in length from 10.5 to 52mm and in width from 2.1 to 10.3mm. Megathymines have the largest pupae (length = 34-52mm, width = 8-10. 2mm). Pyrgines tend to be shorter and wider than hesperiines. Lengths range from 10.5 to 32mm for pyrgines and 12.5 to 43.5mm for hesperiines. Widths vary from 3.2 to 10.3mm for pyrgines and 2.1 to 8.5mm for hesperiines. Promising comparative characteristics of hesperiid pupae include the shape of the pupal cap (usually rounded, sometimes pointed), the length of the antennae and proboscis, cremaster shape, type of thoracic spiracle guard, and presence of crenulations on the posterior margin of the prothorax. Hesperiines and some pyrgines have pointed processes on the head of the pupa. Pyrrhopygines many pyrgines, megathymines, and rarely hesperiines have antennae that extend to the tip of the middle leg. in some pyrgines and most hesperiines, the distal tip of the antenna lies far cephalad of the distal tip of the middle leg. The proboscis of megathymines is greatly reduced (much shorter than the wing tips), whereas most hesperiines have long or very long

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474 probosci (extending beyond the wing tips or sometimes the beyond the cremaster) In pyrrhopygines and pyrgines, the proboscis is subequal to the wing tips. The cremaster of hesperiids usually tapers to a blunt point with a cluster of hooked setae at the tip, but hesperiines display several other variations. The cremaster of Megathymus species is broadly rounded and has many stiff, unhooked setae. Agathymus species usually have a pointed cremaster that lacks setae. The spiracle guard of pyrgines is often greatly enlarged and positioned on a small cuticular rise. Hesperiines simply have a small mound of microspines protecting the spiracle. A few species lack the spiracle guard. Lastly, many pyrgines have small crenulations on the posterior margin of the prothorax. No other subfamilies have this character. Descriptions are presented for the first time for the eggs of 16 species, the larvae of 41 species, and the pupae of 35 species of Hesperiidae found in the U. S. Overall, diagnoses are given for the eggs of 49 species, larvae of 156 species, and pupae of 120 species. Immature stages of 26% of the 290 U. S. species remain to be described. About half of the undescribed species are true residents that maintain breeding populations. The other half are tropical species that rarely enter the U. S. and do not reproduce within its limits. Very little is known of the immature stages of the diverse neotropical fauna. Moss's (1949)

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475 brief descriptions, drawings, and biological notes remain the most comprehensive treatment. Preliminary taxomonic keys are provided for the known eggs, larvae, and pupae of the Hesperiidae of the U. S. Taxonomic Insights and Problems The immature stages of butterflies are rarely considered in classification, but may provide an alternate set of data guite useful in determining phylogenetic relationships. Of the species described above, some clearly show close relationship, but may be widely separated in the Miller and Brown (1984) check list. Among other problems are species with very different immatures included presently within a single genus. The following observations are intended to point out hesperiid taxa in need of closer scrutiny, and perhaps, revisionary studies. At the subfamily level, the immature stages of Pyrrhopyginae and Pyrginae are similar. Likewise, Hesperiinae and Megathyminae are related. The preceding subfamily diagnoses list shared derived characteristics of each of these groups. Very few specimens of Heteropterinae were available for study. Published descriptions of Carterocephalus palaemon indicate a close affinity with Hesperiinae. The mature larvae of Piruna pints (Heteropterinae) have paraclypeal hooks, a derived characteristic observed only in one other genus, Amblyscirtes (Hesperiinae). I can find no evidence

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476 among the immature stages to support subfamily rank for Heteropterinae, based on the larva of this one species. Some related genera within Pyrginae are Epargyreus Chioides, Codatractus Urbanus Astraptes Autochton, Achalarus, Thorybes and Cabares. The pupae of these genera have crenulations on the posterior margin of the prothorax. In addition, their larvae are often very colorful with large yellow or orange eye patches and usually feed on legumes. The larvae of Phocides and Cogia also have larvae with colorful eye patches, but their pupae lack the crenulations. Staphylus Carrhenes Xenophanes Systasea, Pyrgus Heliopetes Celotes and Pholisora also appear to be related. The larvae of these genera usually have black or dark brown heads with branching setae, and feed on Malvaceae, Chenopodiaceae, or Amaranthaceae A third group of pyrgines includes T imoc hares Chiomara, Gesta, Ephyriades and Erynnis The larvae of these genera have an indistinct prothoracic shield, short setae, and feed on a variety of plants, particularly Malpighiaceae, Fabaceae, and Fagaceae. Urbanus, Astraptes, and Polythrix are polyphyletic taxa. Urbanus species with iridescent blue adults have colorful larvae. Species with brown adults have brown larvae that are similar to Achalarus and Thorybes. The larvae of some Astraptes species resemble Epargyreus while others have a Pyrrhopygelike appearance. The larvae of

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477 Polythrix mexicanus and Polythrix procera are so wildly different, that they must represent different genera. It is interesting that the larvae of Polygonus leo and Polythrix mexicanus are similar, but their pupae are not. The remaining genera, Nisoniades and Achlyodes do not seem to have any close relatives in the U. S. fauna, but Nisoniades probably belongs more with Timochares-Erynnis rather than Staphylus-Pholisora Within the hesperiines, Synapte, Vidius Lerema, Nastra, Cymaenes Copaeodes Thymelicus Lerodea, Calpodes and Panoquina seem to be related. These larvae are green or whitish and usually have striped heads. Some develop longitudinal wax glands on the ventral side of A7-8. The pupae all have a pointed process on the cap. Vidius and Lerema are very similar. The larvae of Ancyloxypha, Atrytone, Problema, Ochlodes Euphyes, and Nyctelius are greenish with striped heads and live on the leaves of grasses. Hylephila, Yvretta, Pseudocopaeodes Hesperia, Atalopedes and Polites have brownish larvae with dark heads and live at the base of the host plant. Wallengrenia and Pompeius are closest to this group of hesperiines, but have a number of differences. All of these skippers have pupae with rounded heads. Other associations include Atrytonopsis and Oligoria whose larvae are similar in general appearance, and feed on Andropogon species. Asbolis and Choranthus have similar wax

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478 glands in the form of small spots posterior of the proleg on A3-6. Some Choranthus species feed on palms as does Asbolis capucinus. The immatures of Poanes and Paratrytone are very similar in general appearance. Perichares philetes is curious in having the antennae extending to the tip of the middle leg of the pupa (a pyrgine characteristic) and transverse wax glands on the ventral side of Al. The pupa of P. philetes has a long pointed process. Much remains to be discovered about the biology and immature stages of the Hesperiidae of the U. S. There is also much potential for using characteristics of the immature stages to develop hypotheses on the evolution of this family.

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REFERENCES Ainslie, G. G. 1922. The corn leaf -tier, Lerema accius S. & A. Florida Entomol., 6:1-4, 10-14. Ancona H. L. 1934. Los gusanitos del maguey. Aegiale (Acentrocneme) hesperiaris Kirby. An. Inst. Biol. Univ. Mex., 5:193-200. Arthur, A. P. 1966. The present status of the introduced skipper, Thymelicus lineola (Ochs.) (Lepidoptera: Hesperiidae) in North America and possible methods of control. Can. Entomol., 98:622-626. Arthur, A. P. and B. C. Smith 1974. Thymelicus lineola: Effects of mowing and silica sprays on populations. J. Econ. Entomol., 66:907-908. Austin, G. T. and A. T. Austin. 1981. Butterflies of Clark County, Nevada. J. Res. Lep. 19:1-63. Bailowitz, R. A. and J. P. Brock. 1991. Butterflies of Southeastern Arizona Sonoran Arthropod Studies, Inc., Tucson, Arizona, ix + 342 pp. Baker, W. L. 1972. Eastern forest insects. USDA Misc. Publ. 1175. 642 pp. Barnes, W. and J. McDunnough. 1912. A review of the Megathymidae Contr. Nat. Hist. Lep. N. Amer. 1(3) :143, 6 pis. Basinger, A. J. 1926. Thanaos clitus (Lepid: Hesperiidae) in California. Entomol. News, 37:301. Bates, M. 1935. The butterflies of Cuba. Bull. Mus. Comp. Zool., 78:65-258. Bell, E. L. 1940. New records and new species of Hesperiidae from Mexico (Lep. Hesp.). An. Escuela Nac. Cienc. Biol. Mexico, 2:455-468. Belwood, J. J. 1990. Anti-predator defences [sic] and ecology of neotropical forest katydids, especially the Pseudophyllinae. Pp. 8-14, pis. 1-20. In: W. J. Bailey and D. C. F. Rentz (eds.). The Tettiaoniidae: 479

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480 Biology, Systematics and Evolution Crawford House Press, Bathurst, New South Wales, Australia, ix + 395 pp. Beutelspacher, C. R. 1980. Mariposas Diurnas del Valle de Mexico Talleres Grdficos Victoria, Mexico. 134 + 33 pp. Beutenmiiller, W. 1889. On early stages of some Lepidoptera. Can. Entomol., 21:160. Beutenmuller, W. 1893. Descriptive catalogue of the butterflies found within fifty miles of New York City, together with a brief account of their life histories and habits. Bull. Amer. Mus. Nat. Hist., 5:241-310, pis. II-VI. Blasguez, D. I. 1870. Insectos del maguey. La Naturaleza, 1:282-284, 1 pi. Bodkin, G. E. 1913. Insects injurious to sugar cane in British Guiana, and their natural enemies. J. Board Agric. British Guiana, 7:29-32. Bohart, R. M. 1947. Sod webworms and other lawn pests in California. Hilgardia, 17:267-308. Bonniwell, J. C. 1931. Notes on Megathymus mariae Barnes & Benjamin. Ann. Carnegie Mus., 20:264-265. Bonniwell, J. G. 1916. Location of pupae of Megathymus cofaqui (Lep.). Entomol. News, 27:372. Borror, D. J., D. M. DeLong, and C. A. Triplehorn. 1954. An Introduction to the Study of Insects Fourth edition. Holt, Rinehart, & Winston, New York, xii + 852 pp. Bottimer, L. J. 1926. Notes on some Lepidoptera from eastern Texas. J. Agric. Res., 33:797-819. Box, H. E. 1928. Note upon the larva and pupa of Bungalotis astylos (Lepidoptera, Hesperiidae) Proc. Entomol. Soc. Lond. 1928:83-84. Bridges, C. A. 1983. Lepidoptera: Hesperiidae. Notes on species-group names. Published by the author, Urbana, Illinois, ii, 129, 41, 62, 30, and 13 pp. Brooks, M. and C. Knight. 1982. A Complete Guide to British Butterflies. Jonathan Cape Ltd., London, viii + 159 pp.

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506 from Arizona with observations and notes on its distribution and life history. Bull. Allyn Mus. 9. 11 pp. Wild, W. 1939. The butterflies of the Niagara frontier region and beginner's guide for collecting, rearing and preserving them. Bull. Buffalo Soc. Nat. Sci., 19:355, 8 pis. Willard, H. G. 1892. Thy. poweshiek. Entomol. News, 3:232. Wolcott, G. N. 1921. The minor sugar-cane insects of Porto Rico. J. Dept. Agric. Porto Rico, 5:1-46. Wolcott, G. N. 1923. "Jnsectae Portoricensis" A preliminary annotated check-list of the insects of Porto Rico, with descriptions of some new species. J. Dept. Agric. Puerto Rico, 7:1-313. Wolcott, G. N. 1933. An economic entomology of the West Indies. Entomol. Soc. Puerto Rico, San Juan, xviii + 688 pp. Wolcott, G. N. 1936. "Jnsectae Borinquenses" A revision of Insectae Portoricensis' A preliminary annotated check-list of the insects of Port Rico, with descriptions of some new species". J. Agric. Univ. Puerto Rico, 20:1-600. Wolcott, G. N. 1941. A supplement to "Insectae Boringuenses" J. Agric. Univ. Puerto Rico, 25:33-158. Wolcott, G. N. 1951. The insects of Puerto Rico. J. Agric. Univ. Puerto Rico, 32:417-749. Wright, W. G. 1905. The Butterflies of the West Coast of the United States The Whi taker & Ray Company, Publishers, San Francisco, California. 257 + vii pp., 32 pis. Young, A. M. 1982. Notes on the interaction of the skipper butterfly Calpodes ethlius (Lepidoptera: Hesperiidae) with its larval host plant Canna edulis (Cannaceae) in Mazatlan, State of Sinaloa, Mexico. J. New York Entomol. Soc, 90:99-114. Young, A. M. 1985. Natural history notes on Astraptes and Urbanus (Hesperiidae) in Costa Rica. J. Lep. Soc, 39:215-223.

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507 Young, A. M. 1991. Notes on the natural history of Quadrus (Pythonides) contubernalis (Hesperiidae) in Costa Rica. J. Lep. SOC, 45:366-371.

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BIOGRAPHICAL SKETCH Marc Claude Minno was born 2 March 1956 in Johnstown, Pennsylvania. While a young boy, many summer days were spent searching the fields and forests about his home in western Pennsylvania for wood nymphs, swallowtails, fritillaries, question marks, and other species. The metamorphosis of monarchs, swallowtails, and checkerspots held great interest and wonderment. Marc attended Vinco Elementary School grades 1-8. In 1974 he graduated from Central Cambria High School and went on to study entomology at Purdue University. While attaining his bachelor's degree from Purdue, Marc worked as a technician in the university entomological collection. Much time was devoted to collecting, preparing, and identifying aquatic insects, especially mayflies. It was also during this period that Marc was able to curate a large collection of Lepidoptera, renewing his interest in butterflies. After graduating from Purdue in May 1978, Marc entered the University of California at Davis. Working with Dr. Arthur M. Shapiro, he studied skippers in the Cauca Valley, Colombia, as well as in California. After graduating with a Master of Science degree in entomology in May 1981, Marc was employed briefly as a biologist for the California Department of Fish and 508

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509 Game. In March 1982, he began working for the University of Florida at the Institute of Food and Agricultural Sciences in Fort Lauderdale on the biological control of hydrilla. Marc married Maria Frances Rosiello Lain on 10 October 1982. Angela Katharine Minno was born on 25 April 1984. During the fall of 1984, Marc entered the University of Florida and with Dr. Thomas C. Emmel, continued to study butterflies in Florida, Colorado, and Hispaniola. Marc and Maria's second child, Ivan Alain Minno, was born on 17 September 1990. Marc hopes to continue his interest in the early stages and natural history of butterflies and moths.

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I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. \\jcy+44 Thomas C. Emmel, Chairman Professor of Zoology I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. Lincoln P. Brower Professor of Zoology I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. Jonathan Reiskind Associate Professor of Zoology I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. 'MSS*y/^ Dale H. Ha beck Professor of Entomology and Nematology I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. *£L Gary R. Huckinghai Assistant: Professor of Entomology and Nematology

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This dissertation was submitted to the Graduate Faculty of the Department of Zoology in the College of Liberal Arts and Sciences and to the Graduate School and was accepted as partial fulfillment of the reguirements for the degree of Doctor of Philosophy. April 1994 Dean, Graduate School

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mo mi UNIVERSITY OF FLORIDA 3 1262 08555 0795


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