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Title: Florida Entomologist
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Creator: Florida Entomological Society
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Place of Publication: Winter Haven, Fla.
Publication Date: 1987
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FLORIDA ENTOMOLOGIST

(An International Journal for the Americas)

Volume 70, No. 3 September, 1987

TABLE OF CONTENTS

CHOATE, P. M.-Biology of Ceratocanthus aeneus (Coleoptera: Scarabaeidae:
C eratocanthinae) .................................................. ........................ 301
BARANOWSKI, R. M.-A New Species of Ozophorafrom Costa Rica (Hemiptera:
Lygaeidae) ................................... ............................................. 305
SCHIFFHAUER, D. E., AND R. F. MIZELL III-Bionomics ofGlyphidocerajuni-
perella (Lepidoptera: Blastobasidae), A Newly Discovered Pest of Con-
tainer-Grown Juniper ............................................... ..................... 310
MIZELL, R. F., III, AND D. E. SCHIFFHAUER-Evaluation of Insecticides fbr
Control of Glyphidocera juniperella (Lepidoptera: Blastobasidae) in Con-
tainer-Grown Juniper .................................................................... 316
ALUJA, M., M. CABRERA, E. RIos, J. GUILLEN, H. CELEDONIO, J. HEND-
RICHS, AND P. LIEDO-A Survey of the Economically Important Fruit
Flies (Diptera: Tephritidae) Present in Chiapas and a Few Other Fruit
Growing Regions in Mexico ........................................ .................. 320
ALUJA, M., J. GUILLEN, G. DELA ROSA, M. CABRERA, H. CELEDONIO, P.
LIEDO, AND J. HENDRICHS-Natural Host Plant Survey of the Econom-
ically Important Fruit Flies (Diptera: Tephritidae) of Chiapas, Mexico .. 329
EGER, J. E., JR.-A Review of the Genus Tridates Stdl (Heteroptera: Penta-
tomidae: Scutelleridae) .......................................................... 339
SCHUSTER, D. J., AND J. L. TAYLOR-Residual Activity of Abamectin Against
Liriomyza trifolii (Diptera: Agromyzidae) ........................................ 351
SHAW, K. C., AND P. GALLIART-Acoustic and Mating Behavior of a Mexican
Katydid, Pterophylla beltrani (Orthoptera: Tettigoniidae) ..................... 354
PASSOA, S., AND D. H. HABECK-A Description of the Larva and Pupa of
Rupela albinella, A Pest of Rice in Latin America (Lepidoptera: Pyralidae
Schoenobiinae) .......................................................... ................... 368
ALEXANDER, J. B.-A New Species of Nemapalpus (Diptera; Psychodidae;
Bruchomyiinae) From Northeastern Colombia .................................... 376
BARANOWSKI, R. M., AND J. A. SLATER-New Genera and Species of Antil-
locorini From Trinidad and Brazil Hemiptera: Lygaeidae) ................... 381
LANDOLT, P. J., AND R. HEATH-Seasonal and Diel Patterns of Sex Attraction
of Male Harrisina americana and Acoloithus falsarius (Lepidoptera:
Zygaenidae) ...................... .................................... 392


Scientific Notes
CAIN, M. L.-Prey Capture and Diel Movement of Brachynemurus
(Neuroptera: Myrmeleontidae) Antlion Larvae in South Central
F lorida ...... .................................................................. 397
GLANCEY, B. M., J. C. E. NICKERSON, D. WOJCIK, J. TRAGER,
W. A. BANKS, AND C. T. ADAMS-The Increasing Incidence of the
Polyqynous Form of the Red Imported Fire Ant, Solenopsis invicta
(Hymenotera: Formicidae), in Florida ..................................... 400
HUDSON, W. G.-Variability in Development of Scapteriscus acletus
(Orthoptera: Gryllotalpidae) ............................ ................... 403
Continued on Back Cover

Published by The Florida Entomological Society














FLORIDA ENTOMOLOGICAL SOCIETY




OFFICERS FOR 1987-88
President ... ......... ............................. ........... J. L. Taylor
President-Elect .................................................................. R. S. Patterson
Vice-President .............................................. ..... ............... J. E. Eger
Secretary ...................................................... E. R. Mitchell
Treasurer .......... ............................ ............... A. C. Knapp


M. L. Wright, Jr.
C. O. Calkins
Other Members of the Executive Committee ............... LS. Osborne
G. Mathurin
A. Gettman
C. G. Witherington
J. R. McLaughlin

PUBLICATIONS COMMITTEE


E ditor ................................. ............................................ J. R M cL aughlin
A associate E ditors .............................. .............................................. A A li
C. S. Barfield
J. B. Heppner
M. D. Hubbard
L. S. Osborne
O. Sosa, Jr.
H. V. Weems, Jr.
W. W. Wirth
Business M manager .......................... .... .................................... A C. Knapp

FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30.00 per year in advance, $7.50 per
copy. Membership in the Florida Entomological Society, including subscription to
Florida Entomologist, is $25 per year for regular membership and $10 per year for
students. Inquires regarding membership, subscriptions, and page charges should be
addressed to the Business Manager, P. O. Box 7326, Winter Haven, FL 33883-7326.
Florida Entomologist is entered as second class matter at the Post Office in DeLeon
Springs and Winter Haven, FL.
Authors should consult "Instructions to Authors" on the inside cover of this issue
while preparing manuscripts or notes. When submitting a paper or note to the Editor,
please send the original manuscript, original figures and tables, and 3 copies of the
entire paper. Include an abstract in Spanish, if possible. Upon receipt, manuscripts and
notes are acknowledged by the Editor and assigned to an appropriate Associate Editor
who will make every effort to recruit peer reviewers not employed by the same agency
or institution as the authorss. Reviews from individuals working out-of-state or in
nearby countries (e.g. Canada, Mexico, and others) will be obtained where possible.
Page charges are assessed for printed articles.
Manuscripts and other editorial matter should be sent to the Editor, JOHN R.
McLAUGHLIN, 4628 NW 40th Street, Gainesville, FL 32606.


This issue mailed September 25, 1987











Choate: Biology of Uncommon Scarab 301

BIOLOGY OF CERATOCANTHUS AENEUS
(COLEOPTERA : SCARABAEIDAE: CERATOCANTHINAE)

PAUL M. CHOATE
Department of Entomology and Nematology
University of Florida
Gainesville, Florida 32611
and
Research Associate, Florida State Collection of Arthropods
Division of Plant Industry
Florida Department of Agriculture and Consumer Services
Gainesville, Florida 32602

ABSTRACT

Larval habitat of Ceratocanthus aeneus (MacLeay) is illustrated and discussed.
Adults and larvae occur together and may be subsocial in their relationship. Stridulation
was heard between adults. Pupae have unusual structures that also may permit sound
production.

RESUME

Se dispute e ilustra la habitaci6n de las larvas de Ceratocanthus aeneus (MacLeay).
Adultos y larvas ocuren al mismo tiempo y pueden que sean subsociales en sus re-
laciones. Se cyeron chirridos entire adults. Las pupas contienen estructuras inusuales
que pueden tambien permitir la production de sonidos.



Woodruff (1973) stated that perhaps less than 15 specimens of Ceratocanthus aeneus
(MacLeay) were to be found in all museums of the world. He summarized collection
data for this species and listed pertinent literature. The reader is referred to his paper
for complete literature citations.
The world fauna of Ceratocanthini is currently being revised by Renaud Paulian of
France. His papers (1968, 1973, 1974, 1975, 1977a, 1977b, 1977c, 1978, 1979a, 1979b,
1980, 1981a, 1981b, 1982a, 1982b, 1983, Paulian & Howden 1982) are listed to update
the literature for this tribe. Paulian (personal communication) wrote ... "I have person-
ally collected Ceratocanthids in Africa and Madagascar in termite nests (fungus growing
species) where they are occasionally very common, and in sifting forest litter, under
bark, or on low shrubs. Larvae have been found associated with termites, but few
larvae have been described ... no detailed study of a species has ever been attempted."
Paulian also stated that Ceratocanthus aeneus (MacLeay) is only represented in the
Paris museum by some very old specimens with no other data than "N. Carolina".
One further collecting note has been reported (Westcott 1980). He found 7
Ceratoeanthus (n. sp?) adults as night on the surface of recently(?) felled logs, on Plant-
ation Key, Florida, 5-XII-1970.
Nine adult C. aeneus were collected 13-III-1983 by P. M., T. L., and A. L. Choate.
Specimens were found in a tree hole (Fig. 1) at the edge of the flood plain of the
Apalachicola River, Torreya State Park, Liberty Co., Florida. A single adult was ob-
served on the surface of the moist contents of the tree hole. The shiny reflection of this
individual is what attracted us to the particular site.
The tree hole was approximately 1.5 ft. above the ground, 1 ft deep, and elliptical











Florida Entomologist 70(3)


in shape. The contents were very moist silt and decaying wood, red-brown in color.
Unlike many wet tree holes, no fly larvae of the families Syrphidae and Tabanidae were
found. This was not a fermenting foetid tree hole. Other Coleoptera collected included
Carabidae (Clivina pallida Say, Paratachys sp.); Histeridae (Epierus sp.);
Scaphidiidae, Pselaphidae, and Staphylinidae. Further digging produced 8 additional
adults and many scarab larvae that have been reared to adults. These larvae will be
described in a separate paper.
So seldom had this species been collected, and so restricted the apparent habitat,
that we chose to leave half of the tree hole contents intact, with many larvae, to insure
the existence of this population.
Larvae are typical scarabaeiform, although somewhat more elongate and less
scarabaeiform than those of some other genera. Larvae also appeared gregarious in
rearing containers. Two weeks were spent in pupal cells prior to pupation. During this
time the pre-pupae were quite active, almost constantly moving in a rocking fashion.
Pupae were also quite active, producing a knocking sound in their pupal cells. Approx-
imately 2 more weeks were required (21C) to emerge as adults. Teneral adults re-
mained in pupal cells at least 1 week prior to emergence. Pupation occurred in mid-May.
New adults probably are present by early June.
Pupae (Figs. 2 & 3) exhibit 6 peculiar knob-like structures around the head and
thorax (2 on front of head, 4 on thorax). The function of these remains unknown, but
(in rearing containers) appeared to serve as sound producing structures.
Adults stridulated and appeared gregarious. Stridulation occurred by a movement
of the abdomen against the elytra. Adults found with larvae were presumed to be
parents of some of the larvae. "Old" adults died at the approximate time of emergence
of the new adults. Newly emerged adults were maintained in the same container in
which the larvae had developed. After two weeks, all adults attempted to escape. Less
than a week later all new adults were dead. Apparently the food needed for adult
survival is not found in the larval habitat. Perhaps adults fly to flowers (canopy?) for
food (Edwards 1949).
Ceratocanthus adults quickly tuck into a compact ball when startled, but they also
were observed spending considerable time in this position when not threatened. This
behavior has been described as a death feign but may also serve another purpose.
Adults that were periodically active would return to a compact ball. Also, individuals
that were allowed to run freely would attempt to fly rather than tuck into a ball. The
compact nature of the contracted insect might enable individuals to survive long periods
of quiescence. The incised grooves on the tibial surfaces (Woodruff 1973, Fig. 392) may
permit air storage and possibly gas exchange for survival in a semi-liquid medium. The
paddle-shaped legs also may permit travel through a wet substrate. Flattened tibiae
fold up sheath-like, one on top of the other, as the adult retracts into a ball. This enables
a more complete folding up than seen in the related genus Cloeotus.
Ceratocanthus adults are capable of rapid running and appear quite ready to fly.
Occasional specimens have been taken by beating dead vines (Kissinger 1955, Schwarz
1878). Flower records (Edwards 1949) may refer to canopy flowers, an area almost
totally neglected by North American collectors. Ceratocanthus may possibly be tree
specific. The Carabidae, Clivina pallida Say, that occurred in the tree hole are also
quite abundant under pine bark during the winter months.
That this is indeed a site of continued use by Ceratocanthus was indicated by the
following sequence of events. In June 1983, I returned to the tree hole but failed to find
any sign of adults. During the remainder of the summer no evidence of Ceratocanthus
was found. However, in February, 1984 and 1985, the same tree hole was found to
contain numerous larvae and 3 newly emerged adults in pupal cells. Therefore this tree
is assumed to be a continuous site of larval development. No additional trees of this


302


September, 1987











Choate: Biology of Uncommon Scarab


Fig. 1. Larval habitat of Ceratocanthus aeneus. Liberty Co., Fla.; Torreya St. Park,
flood plain of Apalachicola River.
Fig. 2 & 3. Pupa of Ceratocanthus aeneus.

type have been found in the vicinity. Hopefully, collectors will now find that Ceratocan-
thus is not as rare as formally believed. Until such time, however, I believe that such
sites should be carefully preserved with only limited collecting.


303











Florida Entomologist 70(3)


ACKNOWLEDGEMENTS

I wish to express my appreciation to Angela and Teresa Choate for their assistance
in collecting, in rearing, and in studying these insects, to Francisco Alvarez for provid-
ing the Spanish abstract, and to Dr. R. E. Woodruff for continued assistance during
my studies of Florida Coleoptera.
Contribution No. 655, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, FL 32602.


REFERENCES CITED

EDWARDS, J. G. 1949. Coleoptera or beetles east of the Great Plains. Edwards Bros.,
Inc., Ann Arbor, Michigan. 181 p., 23 pl.
KISSINGER, D. G. 1955. New distribution and habitat records of North American Col-
eoptera. Coleopterists Bull. 9(1): 13-15.
LODING, H. P. 1945. Catalogue of the beetles of Alabama. Alabama Geol. Surv.
Monogr. 11: 1-172. (published posthumously).
PAULIAN, R. 1968. The scientific results of the Hungarian soil zoological expedition
to the Brazzaville-Congo. 33. Esp6ces de la famille Acanthoceridae (Coleoptera:
Scarabaeoidea). Opusc. zool. Budapest 8(1): 87-98, 8 figs.
--. 1973. Sur quelques Acanthoceridae (Coleoptera: Scarabaeoidea) de Nouvelle
Guinee. Opusc. zool. Budapest 12 (1-2): 87-89, 2 figs.
-- 1974. Six nouvelle esp6ces d'Acanthoceridae (Coleoptera) recoltses au Ghana.
Annales hist.-nat. Muz. nat. hung. 66: 205-209.
S1975. Sur quelques Acanthoceridae (Coleoptera) indo-australiens. Annales
hist.-nat. Muz. nat. hung. tom. 67: 151-154, 3 figs.
- 1977a. Un nouveau Besuchtetostes R. Paulian (Col. Ceratocanthidae) de
Malaisie. Revue Suisse Zool. 84(2): 441-442. (illus.)
- 1977b. The Australian Ceratocanthidae (Coleoptera: Scarabaeoidea). J. Aust.
Ent. Soc. 16(3): 261-265. (illus.)
- 1977c. Revision des Ceratocanthidae. Les esp6ces africaines. Revue Zool.
afric. 91(2): 253-316, 62 fig.
--. 1978. Revision des Ceratocanthidae II. Les esp6ces Orientales et Australien-
nes. Ann. Soc. ent. France (N.S.) 14(3): 479-514. (illus.)
- 1979a. A propos de quelques Ceratocanthidae africains (Coleoptera:
Scarabaeoidea). Revue Zool. afr. 93(1): 241-244.
1979b. Sur quelques ceratocanthides de Malaisie (Coleoptera: Scarabaeoidea).
Bull. Soc. ent. France 84(7-8): 171-174, illus.
- 1980. A propos de quelques ceratocanthides indo-africains (Coleoptera:
Scarabaeoidea). Bull. Soc. ent. Fr. 85(3-4): 57-59, 3 figs.
1981a. Un nouveau Madrosotes des iles Bismarck (Coleoptera: Ceratocan-
thidae). Rev. suisse Zool. 88(2): 343-344.
1981b. Trois nouveaux Ceratocanthidae (coleopteres). Bulletin mens. Soc. linn.
Lyon 50(10): 328-329.
1982a. Revision des ceratocanthides (Coleoptera: Scarabaeoidea) d'Amerique
de sud. Mem. Mus. nat. Hist. nat. (N.S., Ser A, Zool.) 124: 1-110, 18 pl.
- 1982b. Observations sur quelques ceratocanthides de Sumatra. Bull. Soc. ent.
France 87(5-6): 196-197.
- 1983. Un nouvel Aneilobolus Hesse (Col. Ceratocanthidae) Bull. Soc. Ent.
France 88: X-XI.
AND H. F. HOWDEN. 1982. Un nouveau genre de ceratocanthides des Antilles.
Bull. Soc. ent. France 87(7-8): 78-85.
SCHWARZ, E. A. 1878. The Coleoptera of Florida. Proc. American Phil. Soc. 17: 353-
469.
WESTCOTT, R. 1980. Note on collecting Ceratocanthus in Florida. In.
SCARABAEUS: A newsletter for those interested in Scarabaeidae. No. 3, p. 2.


304


September, 1987











Baranowski: New Species of Ozophora


WOODRUFF, R. E. 1973. The scarab beetles of Florida (Coleoptera: Scarabaeidae) Part
1. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae,
Ochodaeinae, Geotrupinae, Acanthocerinae). Arthropods of Florida and neigh-
boring land areas (Gainesville), 8: 1-218.





A NEW SPECIES OF OZOPHORAFROM COSTA RICA
(HEMIPTERA:LYGAEIDAE)

RICHARD M. BARANOWSKI
University of Florida, IFAS
Tropical Research and Education Center
Homestead, Florida 33031

ABSTRACT

Ozophora slateri, n. sp., is described and figured. Its relationship to members of the
0. laticephala complex is discussed, and the immature stages are described.

RESUME

La especie nueva Ozophora slateri es descrita e ilustrada. Se discuten sus relaciones
con miembros del complejo 0. laticephala. Se described sus estadios inmaduros.


Among the lygaeids collected during a recent trip to Costa Rica was a series of
specimens representing a new species in the Ozophora laticephala complex. Since this
group has recently been studied (Slater & O'Donnell 1979) this species is herein de-
scribed.
The genus Ozophora has, in recent years, been the subject of considerable study
(Slater & Baranowski 1978, 1979, 1984 (1983), Slater & O'Donnell 1979, Slater 1981,
Slater & Hassey 1981, Ashlock & Slater 1982 and Baranowski and Slater 1984 (1983).
Prior to these studies, 22 species were recognized in the genus. Including Ozophora
slater NEW SPECIES, the genus now contains 45 species. A considerable number
from the Neotropics remain to be described.
All measurements are given in mm.

Ozophora slater Baranowski, NEW SPECIES
Fig. 1

General coloration brownish, head, anterior pronotal lobe, thorax (laterally and ven-
trally) and abdomen brown. Pronotal color, except for dark brown spot at midline,
posterior pronotal lobe except for brown postero-lateral corners and brown spot on
midline at posterior margin and anterior border, and explanate lateral margins yel-
lowish. Scutellum brownish with apex white and a pair of yellow vittae, one on either
side of the midline, converging toward the apex. Antennal segments I, II, light brown,
III darker with distal 1/3 dark brown, IV dark brown with proximal 1/3 white. Legs
yellow brown except for a faint annulus near apex of hind femur, apex of anterior tibia
and apical tarsal segment darker. Labium yellow brown. Clavus with 5 rows of dark


305










Florida Entomologist 70(3)


.1


.7

/2


306


Fig. 1. Ozophora slater n. sp. Dorsal view.


punctures and a darkened area near claval commissure. Corium with a dark spot near
level of apex of scutellum, a broad dark band at level of distal end of claval commissure
with a lighter area at edge near commissure, apical tip dark brown. Membrane with an
infuscated area near apex.
Head broad, moderately declivent, clothed with short gold decumbent hairs; eyes
large. Head length 0.66, width 0.80, interocular space 0.38. Pronotum with brown punc-
tation, less dense on anterior calli, transverse impression complete. Lateral pronotal
margins strongly sinuate, distinctly but not prominently explanate. Pronotum length
0.78, width 1.20. Disk of scutellum slightly elevated. Scutellum length 0.70, width 0.70.


September, 1987


























i /











Baranowski: New Species of Ozophora


Hemelytra with lateral corial margins very slightly sinuate, explanate. Membrane well
exceeding end of abdomen. Claval commissure length 0.50. Midline distance apex clavus-
apex corium 0.86. Midline distance apex corium-apex membrane 0.76. Metathoracic
scent gland auricle slender, similar to that of 0. baranowskii Slater & O'Donnell.
Labium extending between mesocoxae, first segment reaching base of head. Labial
segments length I 0.54, II 0.50, III 0.40, IV 0.32. Antennal segments length I 0.36, II
0.96, III 0.80, IV 0.82. Total body length 3.76.
Clasper similar to that of 0. baranowskii.
HOLOTYPE: S COSTA RICA: Heredia Prov. Heredia 15-II-83 (F. Gilstrap, R. M.
Baranowski), U.S.N.M. no. 100255.
PARATYPES: 8 6, 12 9 same data as holotype; 16, 1 9 Prov. San Jose, San Jose,
Univ. Costa Rica, 13-II-83 (R. M. Baranowski, F. Gilstrap); 1 9 San Jose, Univ. Costa
Rica. 18-III-83 (R. Clayton). In U.S.N.M., Florida State Collection of Arthropods, R.
M. Baranowski and J. A. Slater collections.
Ozophora slateri is a member of the 0. laticephala complex. It will go to couplet 5
in the Slater and O'Donnell (1979) key, but will not pass thru either choice as it does
not have a pale median or broad dark median stripe on the posterior pronotal lobe.
However, in some specimens, the dark markings on the midline might be considered as
an incomplete stripe. With this interpretation, 0. slater will go to couplet 8 thru 7 and
key out to 0. baranowskii. It can be separated from 0. baranowskii by the distinct
yellow posterior pronotal lobe, the yellow scutellar markings and by the clasper shape
(Fig. 2).
All specimens were collected under Ficus sp. (Moraceae).
It is with great pleasure that I dedicate this species to Dr. James A. Slater, Univer-
sity of Connecticut, a friend and colleague for many years, in recognition of his many
outstanding contributions to our knowledge of the Lygaeidae.


Description of slateri nymphs

5th instar (in alcohol)
General color pale yellow, marked with brown as follows: a narrow pale stripe along
suture between tylus and jugae, joining to form a slightly broader stripe extending
posteriorly, a broad stripe along lateral margin of head from anteniferous tubercle to
posterior margin, meeting narrowly at margin; a broad median stripe along midline of
prothorax abruptly doubling in width in posterior half, narrowly along lateral and an-
terior margin except next to median stripe, a broad irregular stripe between lateral
and midline stripe broadly joined anteriorly and narrowly posteriorly to an irregular
spot on posterior margin; a narrow stripe along either side of midline of scutellar area
extending laterally along anterior margin, antero-laterally midway then sharply revers-
ing to posterior margin forming a hairpin loop; wing pads narrowly outlined laterally,
broadly posteriorly and mesally with a narrow triangular loop extending from postero-
lateral margin almost to base and a short wide stripe extending anteriorly from postero-
mesal margin; much of pro, meso and metapleuron except for 2 elongate areas on each
side. Antennal segments I, II yellow brown, III, IV dark brown except for proximal
tips and distal tip of the IV white. Acetabula and coxae white, legs dusky yellow except
for distal ends of femora, proximal tip and distal 1/3 of tibiae and 1st tarsal segment
yellow.
Dorsal surface of abdomen with a large brown spot between wing pads, another
smaller spot between scent glands 3-4 and 4-5 and a 3rd between scent glands 4-5 and
5-6, paired red dashes present on either side of scent glands and near midline of seg-
ments 4, 5 and 6.











Florida Entomologist 70(3)


Fig. 2. Ozophora slater n. sp. clasper.


Head moderately declivent, tylus extending to middle of 1st antennal segment. Head
length 0.68, width 0.80, interocular space 0.46. Pronotum quadrate, lateral margins
slightly sinuate, distinctly explanate, posterior margin almost straight, pronotal collar
distinct only at lateral margins. Pronotum length 0.58, width 1.12. Wing pad length
1.18. Abdomen length 1.96. Labium extending between mesocoxae; labial segments
length I 0.52, II 0.50, III 0.38, IV 0.30. Antennal segments length I 0.30, II 0.72, III
0.68, IV 0.74. Total body length 3.68.

4th instar (in alcohol)
General form and color as in 5th instar, except head brown with a pair of lighter
spots near tylus, pronotum brown except for a narrow mesally slanting stripe near
midline extending 1/2 the length of the pronotum from the anterior edge, 2 spots on
each side of midline on posterior margin and a narrow stripe just inside of lateral margin


308


September, 1987











Baranowski: New Species of Ozophora


yellow. Head length 0.44, width 0.60, interocular space 0.36. Pronotum length 0.38,
width 0.76. Wing pad length 0.48. Abdomen length 1.24. Labial segments length I 0.44,
II 0.36, III 0.26, IV 0.24. Antennal segments length I 0.20, II 0.40, III 0.40, IV 0.48.
Total body length 2.60.

3rd instar (in alcohol)
Similar in shape to 4th instar. Head and pronotum uniformly light brown, wing pads
with 5 irregular spots on each side of midline. Antennae and legs dusky straw color,
segments III, IV darker. Abdominal terga with irregular reddish markings only. Head
length 0.42, width 0.48, interocular space 0.30. Pronotum length 0.28, width 0.58, wing
pads 0.18. Abdomen length 1.16. Labial segments length I 0.32, II 0.28, III 0.22, IV
0.18. Antennal segments length I 0.16, II 0.30, III 0.28, IV 0.40. Total body length 2.10.

2nd instar (in alcohol)
Similar in shape and color to 3rd instar except no light markings on thorax. Head
length 0.32, width 0.40, interocular space 0.28. Pronotum length 0.20, width 0.42. Ab-
domen length 0.90. Labial segments length I 0.26, II 0.14, III 0.22, IV 0.18. Antennal
segments length I 0.14, II 0.24, III 0.20, IV 0.34. Total body length 1.64.

1st instar (in alcohol)
Similar in shape and color to 2nd instar. Head length 0.30, width 0.30, interocular
space 0.22. Pronotum length 0.14, width 0.30. Abdomen length 0.56. Labial segments
length I 0.26, II 0.14, III 0.22, IV 0.18. Antennal segments length I 0.14, II 0.24, III
0.20, IV 0.34. Total body length 1.64.

Egg (in alcohol)
Cream colored, elongate, tapering to a narrow rounded posterior end and a less
tapered truncate anterior end with typically 5 short club shaped micropylar processes.
Length 0.80, width across widest area 0.30.


ACKNOWLEDGEMENTS

This is No. 5481 of the Florida Agriculture Experiment Stations Journal Series. The
author is a Research Associate of the Florida State Collection of Arthropods, Florida
Department of Agriculture and Consumer Services, Gainesville.
I thank Ms. Mary Jane Spring, University of Connecticut, for preparation of the
illustrations, Drs. T. Henry, United States National Museum of Natural History and
J. Nation, University of Florida, for their helpful reviews.


REFERENCES CITED

ASHLOCK, P. D., AND J. A. SLATER. 1982. A review of the genera of Western Hemis-
phere Ozophorini with two new genera from Central America (Hemiptera-
Heteroptera:Lygaeidae). J. Kansas Ent. Soc. 55: 737-750.
BARANOWSKI, R. M., AND J. A. SLATER. 1984 (1983). The Ozophora pallescens com-
plex in the West Indies with the description of 4 new species. Florida Ent.
66: 463.
SLATER, J. A. 1981. A new species of Ozophora from Cocos Island (Hemipt-
era:Lygaeidae). J. Kansas Ent. Soc. 54: 22-26.
SLATER, J. A., AND R. M. BARANOWSKI. 1978. A new species of bromeliad lygaeid
from Jamaica. Florida Ent. 61: 83-88.
- 1979. New Species of Ozophora from the lesser Antilles with notes on the
biology and immature stages (Hemiptera:Lygaeidae). Florida Ent. 62: 244-259.


309











Florida Entomologist 70(3)


- 1984 (1983). The genus Ozophora in Florida. Florida Ent. 66: 416-440.
SLATER, J. A., AND J. E. O'DONNELL. 1979. An Analysis of the Ozophora
laticephala-complex with the description of eight new species (Hemiptera:
Lygaeidae). J. Kansas Ent. Soc. 52: 154-179.
SLATER, J. A., AND M. HASSEY. 1981. The distribution and systematics of Ozophora
atropicta Barber with the description of a new species from the Neotropics.
Florida Ent. 64: 246-259.





BIONOMICS OF GLYPHIDOCERA JUNIPERELLA
(LEPIDOPTERA: BLASTOBASIDAE), A NEWLY
DISCOVERED PEST OF CONTAINER-GROWN JUNIPER

D. E. SCHIFFHAUER AND R. F. MIZELL, III
Agricultural Research and Education Center, Monticello
Rt. 4 Box 63, Monticello, FL 32344

ABSTRACT

The life history of Glyphidocera juniperella Adamski (Lepidoptera: Blastobasidae)
is presented. Larvae of this newly described blastobasid moth are serious pests of
nursery, container-grown junipers. Larval feeding causes twig dieback and reduces
plant value. Head capsule measurements indicate 6 instars in the 2 summer generations
and 7 instars for the partial 3rd or overwintering generation.

RESUME

Se present la historic de la vida de Glyphidocerajuniperella Adamski (Lepidoptera:
Blastobasidae). Las nuevas descritas larvas de las alevillas blastocidas, son plagas series
en viveros de plants de junipero creciendo en macetas. Dafio por las larvas al comer,
causan la muerte de ramas pequefias y reduce el valor de las plants. Medidas de la
cabeza indican que hay 6 estadios en las 2 generaciones del verano y 7 estadios por la
parcial tercera o generaci6n hibernante.


The wholesale nursery industry in Florida is valued at approximately $412 million
per year. Container-grown junipers are one of the more important plants produced by
the Florida nursery industry. As with all nursery products the aesthetic appearance of
the plants must be maintained.
We recently discovered a new pest of container-grown juniper which was described
as Glyphidocera juniperella Adamski (Adamski & Brown 1987). The larvae are serious
pests of horizontal-growing ornamental juniper. This paper presents the bionomics of
the moth.

MATERIALS AND METHODS

Field collections of G. juniperella were made weekly from March, 1984 until March,
1985 at a nursery in n. Florida. The webbing masses produced by larvae in infested
branches were removed and placed in plastic bags for laboratory dissection. Addition-


310


September, 1987











Schiffhauer & Mizell: Bionomics of a New Juniper Pest 311

ally, 3 infested plants were brought into the laboratory each week and dissected for
eggs, larvae, and pupae. Larvae were collected from the webbing in the laboratory,
fixed for 30 sec in boiling water, and preserved in a 70% alcohol for later measurement
of head capsule width. Larval head capsule widths were measured to the nearest 0.01
mm for 1687 larvae by means of an ocular micrometer.
Pupae collected from the field were held individually in 1 oz diet cups until eclosion.
Emerging adults were sexed and 25 pairs were placed in 8.5 cm x 4.5 cm plastic
oviposition chambers along with fresh juniper twigs. Pairs were held until the female
died, males were replaced as needed. The pre-oviposition period, fecundity and longev-
ity of females and the time required for egg hatch were determined.
First instars were gathered from the oviposition chambers and placed on one of 6
artificial diets in an attempt to rear this species in the laboratory. The 6 diets used
were: modified pinto bean diet (Shorey & Hale 1965, Schroeder 1970), Douglas-fir tus-
sock moth diet (Lyon & Flake 1966), universal diet (Singh 1983), and each diet plus
dried juniper needles at 1% dry weight. All rearing and diet tests were conducted in a
rearing room with a photoperiod of 15:9 (L:D), temp 28 + 20C, and RH 50-80%.
For determination of pupal development time large larvae were collected from the
field and placed on pinto bean diet. Larvae were checked daily for onset of pupation
and pupal development time was recorded for 108 pupae. Larvae or pupae suspected
of being parasitized were placed in gelatin capsules and held for parasite emergence.
Pupal parasites were also collected from pupae placed in diet cups.


RESULTS AND DISCUSSION

Fig. 1 shows the distribution of eggs, larvae, and pupae collected from the field in
1984-85. Large larvae and pupae of the overwintering cohort were collected in March
of 1984. The 1st generation began in early May and the 2nd in late June-early July.
Eggs for the 3rd generation, which overwintered as late instars or pupae, were collected
in early October. Based on the field collections of early instars, development of each
summer generation required ca. 70 days.
Female G. juniperella reared from field collected pupae laid an average of (x+SD)
82.8 +39 eggs. Eggs collected in the field were found on detritus and needles near the
media surface. Eggs are laid singly or in groups up to 10. They are transparent at
oviposition, turn a salmon color ca. 2 days later, and finally are bright pinkish-red prior
to eclosion. Eggs hatched in 7.8 + 1.2 days at 28C in the laboratory. The preoviposition
period for newly emerged females in the laboratory was 1.3 0.6 days at 280C. Female
longevity (1 -SD n=21) in the laboratory was 3-14 days with a mean of 7.1 2.9 days.
Two hundred first instar larvae were placed individually on each of the 6 diets
tested. Survivorship on all artificial diets was very low. The highest percentage sur-
vivorship, 2.5%, was on pinto bean diet without juniper. The few G. juniperella which
did complete development on the diets were small and total development time was far
greater than in the field under similar temperatures. No difference in survivorship was
noted between diets with or without juniper.
Although survivorship and development of 1st instars was very low on the diets
tested, last instars collected in the field were able to develop to pupae on pinto bean
diet. The pupal stadium of field collected larvae placed on pinto bean diet lasted 9.3 +
1.7 days (n=108). The average weight of 17 male and 12 female pupae was 9.5 1.1
mg and 12.4 + 1.9 mg, respectively.
The frequency distribution of larval head capsule widths (Fig. 2) has 5 apparent
peaks centered at 0.22, 0.27, 0.38, 0.52, and 0.72 mm. Above the 0.72 mm peak the
distribution is less clear, with 1 or 2 peaks possible. The head capsules of the last instar













Florida Entomologist 70(3)


L: Larvae
L1 P= Pupae
LI
SE:Eggs
IP1
....... El

AA

S00
C..L2

P EP 2 15o

'3 L3

P3
............. E3




L4

Mar Apr. May June July Aug. Sept. Oct. Nov. Dec. Jn. Feb. Mar. Apr.

1984 1985

Fig. 1. Date of occurrence of eggs, larvae and pupae for 3 generations of Glyphidoc-
erajuniperella Adamski collected in the field from container grown juniper, 1984-1985.


of the 2 summer generations were smaller than those of the last instar in the overwin-
tering generation (Fig. 3). This explains the ambiguity of the frequency distribution
(Fig. 2) as to the number of instars above .71 mm. Various combinations of instar
sequences were fitted to the data and compared using analysis of variance. Table 1 lists
a 6 instar and a 7 instar model which best fit the data according to Dyar's rule (Dyar
1890). The 6-instar model was the best fit for head capsule widths from the twoesummer
generations while the 7-instar model best fit the head capsule data of the overwintering
generation. Data from the overwintering generation was comprised of pooled head
capsule measurements from March-May 1984 and Nov. 1984-March 1985.



60

50

40
z
w
D 30

(r 20
l.


02 03


04 05 06 07
HEAD CAPSULE WIDTH Imml


08 09 10 11 12


Fig. 2. Frequency distribution of head capsule widths of 1687 Glyphidocera
juniperella Adamski larvae collected from container grown juniper.


312


September, 1987















Schiffhauer & Mizell: Bionomics of a New Juniper Pest


313


.. *. 0




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Mor. Apr. May June July Aug. Sept. Oct Nov. Dec. Jan. Fb. Mar. Apr.

1984 1985


Fig. 3. Distribution of larval head capsule widths of Glyphidocera juniperella
Adamski vs collection date, 1984-1985.

Developmental polymorphism has been described for many lepidopteran species.
Schmidt & Lauer (1977) found varying instar numbers in each of 3 different Choris-
toneura spp., Watson and Johnson (1974) described 4 and 5 instar larvae for Pec-
tinophora gossypiella (Saunders), and Solomon (1973) found instar number varied in
Prionoxystus robiniae (Peck). Postulated reasons for variable instar numbers have
included nutrition, temperature, humidity, and photoperiod. G. juniperella overwinters
mainly as late instar larvae and these larvae are active on warm days during the winter.
The occurrence of an extra instar in the overwintering generation may accommodate
the long period of time spent in the last instar larvae.
Typical damage of G. juniperella on junipers includes removal of the outer bark and
phloem (Fig. 4). The twig is often girdled causing "flagging"-red colored needles.
Girdling behavior is most prevalent in late fall and winter. Damage is not evident during
the winter due to plant dormancy and cool temperatures, but with the onset of warm
weather the needles on the girdled limbs of infested plants turn red.
While no effort has been made to determine the distribution of G. juniperella it is
apparently confined to the southeastern U.S. It has been collected in Poplarville, Mis-
sissippi; Cairo, Georgia; N. Florida (Quincy, Monticello, and Jacksonville); and from
Tampa, Florida. All collections were made from juniper nursery stock and considering
the commerce in juniper between nurseries, G. juniperella is no doubt wide spread at
least south of the latitude of Cairo, Georgia. Cold temperatures appear to strongly
affect the distribution of this pest.


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Florida Entomologist 70(3)


TABLE 1. LIMITS OF LARVAL HEAD CAPSULE WIDTHS FOR 6 AND 7 INSTAR GROUP-
INGS OF GLYPHIDOCERA JUNIPERELLA ADAMSKI.

6 INSTAR GROUPING (SUMMER GENERATIONS)
Ratio of
Instar N Range (mm) Mean+SD (mm) Increase

1 19 .191-.248 .223.017
2 98 .250-.328 .283t.021 1.270
3 88 .330-.425 .376+.027 1.325
4 126 .430-.618 .517-.053 1.377
5 129 .629-.809 .700.047 1.353
6 182 .813-1.142 .908.053 1.296
Avg = 1.324
7 INSTAR GROUPING (OVERWINTERING GENERATION)
Ratio of
Instar N Range (mm) Mean+ SD (mm) Increase

1 80 .196-.247 .216.008
2 123 .250-.326 .279.015 1.2937
3 96 .330-.428 .376.025 1.3443
4 181 .430-.618 .521+.046 1.3872
5 143 .622-.759 .691_. 039 1.3255
6 156 .764-.9292 .833.051 1.2051
7 266 .933-1.235 1.02.058 1.2250
Avg = 1.2968


Field sampling was originally planned for 2 yr; however the extreme temperature
of January 1985, -20C in N. Florida, virtually wiped out populations of the moth. As
of December, 1985 populations were low even in juniper cultivars which normally had
high populations.
Two species of larval parasites and one pupal parasite were found during 1984 in
Florida. The pupal parasite in Florida was identified as a Brachymeria sp. (chalcidoid).
The 2 larval parasites are as yet unidentified. One of the larval parasites was frequently
found parasitizing 20-30% of larvae during the summer generation. A pupal parasite
Rubicundiella annulicornis (Ashmead) (Ichneumonidae) was collected from G.
juniperella in Poplarville, Mississippi.
G. juniperella exhibits a definite preference for low growing, prostrate cultivars of
juniper. Cultivars such as Juniperus horizontalis, 'Wiltonii' (Blue rug); J. chinensis
var. procumbens 'Nana'; J. horizontalis 'Prince of Wales'; and 'Andorra compacta,
were often heavily infested while other more upright cultivars were free of G.
juniperella.
Plant age and the relation of plant size and form to pot size are important factors in
preference. Low growing cultivars that have grown over the edge of the containers are
most frequently infested. This plant type usually has a large amount of dead material
on the media surface, making it ideal habitat for G. juniperella. Other upright growing
junipers may also become infested if needle clippings from pruning build up in the
container. G. juniperella larvae can be found feeding in the dead needles. However, in
these plants girdling of stems was not observed. We also have reared G. juniperella
larvae from early instars to adults using only dead, dried leaves of live oak trees,
(Quercus virginiana Mill). This and other observations suggest that G. juniperella is a


314


September, 1987











Schiffhauer & Mizell: Bionomics of a New Juniper Pest


'/t
it .


Fig. 4. Typical feeding damage of Glyphidocerajuniperella Adamski larvae on small
juniper twigs.


detritivore capable of developing on a variety of dead plant material. G. juniperella
does not appear capable of actually killing plants, but can cause considerable aesthetic
damage to junipers and requires control by nurserymen (Mizell and Schiffhauer 1987).
Further research into the factors which induce buildup of large populations is needed.
While sampling for G. juniperella, larvae of the moth, Oleuthreutes cespitana
Hubner (Tortricidae), was often found feeding on green needles either in the same
webbed masses or adjacent to those of G. juniperella. Larvae and adults of the moth
were found mainly in the spring and early summer in juniper but disappeared in fall.
These larvae in early instars can easily be mistaken for G. juniperella larvae. A larval
parasite, Orgilus sp. (Braconidae) was reared from 0. cespitana.

END NOTE

We thank Dr. Dale Habeck for help in identifying the lepidopteran larvae found in
container-grown junipers and Imperial Nursery for providing the plants for sampling.
Drs. Dale Habeck and Ronald Oettig provided helpful comments on a previous draft of
the manuscript. We thank Drs. L. Stange and V. Gupta, Florida Department of Agri-
culture and Consumer Services, Division of Plant Industry, for identifying the parasites
and David Adamski and Richard Brown, Mississippi State University, for identifying
the moth and providing the pupal parasite. Florida Agricultural Experiment Station
Journal Series No. 7215.

REFERENCES CITED

ADAMSKI, D. AND R. L. BROWN. 1987. A new nearctic Glyphidocera with description
of all stages (Lepidoptera: Blastobasidae: Symmocinae). Proc. Entomol. Soc.
Wash. 89: 329-43.


315











316 Florida Entomologist 70(3) September, 1987

DYAR, H. G. 1890. The number of molts of Lepidopterous larvae. Psyche 5: 420-2.
LYON, R. L. AND H. W. FLAKE, JR. 1966. Rearing Douglas-fir tussock moth larvae
on synthetic media. J. Econ. Entomol. 59: 696-8.
MIZELL, R. F. AND D. E. SCHIFFHAUER. 1987. Evaluation of insecticides for control
of Glyphidocera juniperella Adamski (Lepidoptera: Blastobasidae) in container-
grown juniper. Florida Entomol. 70: 000-000.
SCHMIDT, F. H. AND W. L. LAUER. 1977. Developmental polymorphism in Choris-
toneura spp. (Lepidoptera: Tortricidae). Ann. Entomol. Soc. America 70: 112-8.
SCHROEDER, W. J. 1970. Rearing the pecan bud moth on artificial diet. J. Econ.
Entomol. 63: 650-1.
SHOREY, H. H. AND R. L. HALE. 1965. Mass rearing of the larvae of nine noctuid
species on a simple artificial medium. J. Econ. Entomol. 58: 522-4.
SINGH, P. 1983. A general purpose laboratory diet mixture for rearing insects. Insect
Sci. Application 4(4): 357-62.
SOLOMON, J. D. 1973. Instars in the carpenterworm, Prionoxystus robiniae (Peck).
Ann. Entomol. Soc. America 66: 1258-60.
WATSON, T. F. AND P. H. JOHNSON. 1974. Larval instars of the pink bollworm, Pec-
tinophora gossypiella (Saunders). Ann Entomol. Soc. America 67: 812-4.





EVALUATION OF INSECTICIDES FOR CONTROL OF
GL YPHIDOCERA JUNIPERELLA
(LEPIDOPTERA: BLASTOBASIDAE)
IN CONTAINER-GROWN JUNIPER

R. F. MIZELL, III AND D. E. SCHIFFHAUER
Agricultural Research and Education Center, Monticello
Rt. 4 Box 63, Monticello, FL 32344

ABSTRACT

Glyphidocera juniperella Adamski (Lepidoptera: Blastobasidae) is a pest of con-
tainer grown juniper in the southeastern U.S. Larvae of this pest girdle stems and
damage the appearance of ornamental juniper. Pesticides covering a range of formula-
tions were tested for efficacy against G. juniperella in field and laboratory tests. An
entomophagous nematode, Neoaplectana carpocapsae Weiser, was also field tested for
control of G. juniperella. All formulations of Bacillus thuringiensis Berliner tested
gave good control, as did cypermethrin and acephate. Nematodes were ineffective.

RESUME
Glyphidocera juniperella Adamski (Lepidoptera: Blastobasidae) es una plaga de
juniperos en macetas en el sudeste de los Estados Unidos. Larvas de esta plaga circun-
dan los tallos y dafian la apariencia de juniperos ornamentales. Se probaron pesticides
que cubrian varias formulaciones para determinar su eficacia contra G. juniperella en
pruebas de campo y de laboratorio. El nemAtodo entomofago, Neoaplectana carpocap-
sae Weiser, se prob6 tambi6n en el campo para controlar a G. juniperella. Todas las
formulaciones de Bacillus thuringiensis Berliner que se probaron dieron buen control,
asi como cypermethrin y acephate. Los nematodos fueron inefectivos.











Mizell & Schiffhauer: Control of a New Juniper Pest


Containerized junipers are an important nursery crop in the southeastern U.S.
Glyphidocera juniperella Adamski is a recently discovered blastobasid moth whose
larvae damage juniper by girdling small stems. Larvae construct and feed in a large
mass of webbed frass and detritus near the media surface (Schiffhauer & Mizell 1987).
No information is available for chemical control of this pest and growers currently use
compounds based on efficacy against other pests. We evaluated the efficacy of several
formulations of insecticides registered for general use on ornamentals and several ex-
perimental chemicals, for control of G. juniperella larvae. This is the 1st report of
efficacy of insecticides for control of this insect.


MATERIALS AND METHODS

A field test was conducted in S. Georgia on infested juniper plants, Juniperus
horizontalis var. 'Prince of Wales', in 2 gal. pots. Eighteen liquid insecticide formula-
tions were applied to runoff on Nov. 1, 1984 with a Solo hand-pumped compressed-air
sprayer at 8.5 kg/cm2 pressure. Three granular and 1 bait formulations were applied to
pots by hand from premeasured containers. Four replicates of 5 plants each were tested
with each treatment and were arranged in a randomized block design. Treatments were
evaluated at 7, 18, or 25 days posttreatment depending upon formulation or mode of action:
(Group I) liquids and wettable powders-7 days, (Group II) baits and biologicals-18
days, (Group III) granulars-25 days. Separate control plants were evaluated on each
day treatments were evaluated. Evaluations were made by counting the number of live
G. juniperella larvae in each pot.
In a separate experiment, 5 replicates of 25 plants each, were treated with a suspen-
sion of entomophagous nematode, Neoaplectana carpocapsae Weiser. Juvenile nema-
todes were applied to 2 gal J. horizontalis 'Wiltonii' at a rate of 66,000/plant from 1 litre
spray bottles on June 22, 1984. Nematode treated plants were evaluated at 7, 14, and
21 days posttreatment.
Laboratory bioassays of efficacy were conducted with the 18 liquid insecticide formu-
lations used in the field study. Late instar larvae of similar size were collected from
the field and placed in diet cups partially filled with modified pinto bean diet (Shorey
& Hale 1965, Schroeder 1970). The formulations of biologicals (Group II) used were
dissolved in distilled water and applied to the diet surface at a rate of 2 ml per cup. The
remaining compounds were dissolved in acetone and topically applied to the thorax of
larvae at a rate of 1 Rl per larvae by means of a micropipette. Larvae were held 5 to a
cup and 5 replicates of 5 cups were tested with each treatment. Control replicates
consisting of 2 ml of distilled water on the diet surface, or 1 p.1 of topically applied
acetone were also used. Larvae in diet cups were checked daily for 5 days posttreatment
and number of dead and live larvae were counted. Laboratory data were analyzed using
the GLM procedure of the Statistical Analysis System (SAS 1982). Separate analyses
were run for each formulation group.


RESULTS AND DISCUSSIONS

Of the 13 compounds in Group I of the field tests, only the cypermethrin and
acephate treatments were significantly more effective than the control treatments
(Table 1). Group II contained the biologicals along with one bait, 5% acephate. All the
compounds in Group II were significantly more effective than the control. The 3 formu-
lations of Bacillus thuringiensis Berliner used were particularly effective in the field
tests. From Group III, comprised of granular insecticides, only disulfoton was signifi-
cantly more effective than the control.


317














Florida Entomologist 70(3)


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Mizell & Schiffhauer: Control of a New Juniper Pest


The laboratory tests were conducted to determine if failure in the field tests by the
insecticides was due to poor penetration of the plant foliage by the insecticides. Seven
of the 13 chemicals in Group I were significantly more effective than the control after
5 days. Of the 7 chemicals which differed from the control, cypermethrin was especially
effective. Four of the 5 compounds tested in Group II were significantly better than
the control. Only Bactospeine was not effective in the laboratory tests, although it
was effective in the field.
No differences were detected between the junipers treated with nematode suspen-
sions and controls. G. juniperella larvae infected with nematodes in the laboratory
produced very few Fi nematodes. The nematode treatments were applied in the field
on an overcast day with rainy periods, which should have been ideal conditions for
nematode infection of G. juniperella. Field and lab results indicate that N. carpocapsae
will not effectively control G. juniperella.
The compounds which were the most efficacious in the field proved the most effective
in the laboratory. All 4 formulations of B. thuringiensis were very effective in the field
as were cypermethrin and acephate. Some of the compounds such as carbaryl were very
effective in the laboratory tests but not in the field. G. juniperella larvae are difficult
to treat in the field because of the heavy webbing mass they produce under the foliage.
Presumably, the lack of control in the field by chemicals which were effective in the
lab, was due to poor penetration of the webbing mass. None of the granulars or the


TABLE 2. EFFICACY OF INSECTICIDES AGAINST GLYPHIDOCERA JUNIPERELLA
ADAMSKI ON PINTO BEAN DIET. GROUP I COMPOUNDS DISSOLVED IN
ACETONE AND TOPICALLY APPLIED TO LARVAE AT 1 p1 PER LARVAE.
GROUP II COMPOUNDS DISSOLVED IN DISTILLED WATER AND APPLIED
TO DIET SURFACE.

Mean no.
Group1 Formulation (lb ai/100 gal) live larvae/rep

I Control (acetone) 5.0 a2
Oxydemeton methyl 25C 1.0 4.8 a
Isozophos 4E 1.5 4.6 ab
Azinphos-methyl 50WP 0.5 4.0 abc
Carbophenothion 8E 1.0 4.0 abc
Malathion 50%EC 1.0 4.0 abc
Diazinon AG500 1.2 3.8 abc
Disulfoton 8E 0.3 3.4 bcd
Chlorpyrifos 2E 0.5 3.2 cde
Fluvalinate 2F 0.1 3.0 cde
Acephate 5% 1.7 2.4 de
Methomyl 1.8EC 0.4 2.2 de
Carbaryl 80S 1.3 2.0 e
Cypermethrin 2.5EC 0.075 0.0 f
II Control (Water) 4.4 a
Bactospeine 20WP3 0.75 3.4 ab
Diflubenzuron 25WP 1.2 2.6 b
Dipel 10WP3 1.5 1.2 c
Dipel 4L3 1.5 0.6 c
Thuricide 32B3 1.0 0.0 c

'Groups were evaluated separately.
2Means not followed by same letters are significantly different at Ps.05 as determined by DNMRT.
3Formulations of Bacillus thuringiensis Berliner.


319











320 Florida Entomologist 70(3) September, 1987

bait appeared to be effective in controlling G. juniperella. The biologicals, Bactos-
peine, Dipel, and Thuricide controlled G. juniperella larvae well and should be
recommended for grower use.

END NOTE

We thank Wight Nursery in Cairo, GA for providing the plants for the evaluation.
We thank Dr. Grover Smart and Nguyen Va Khuong for providing and applying the
nematode treatment and Imperial Nursery, Quincy, FL for providing the plants. Drs.
Dale Halbeck and Ronald Oettig made helpful comments on an earlier draft of the
manuscript. Florida Agricultural Experiment Station Journal Series No. 7212.

REFERENCES CITED

SAS. 1982. SAS Users Guide: Statistics SAS Institute, Inc. Carey, NC 22511.
SCHIFFHAUER, D. E., AND R. F. MIZELL, III. 1987. Biology of Glyphidocera
juniperella Adamski (Lepidoptera: Blastobasidae), a new pest of container-
grown junipers. Fla. Entomol. 70: 000-000.
SCHROEDER, W. J. 1970. Rearing the pecan bud moth on artificial diet. J. Econ.
Entomol. 63: 650-1.
SHOREY, H. H., AND R. L. HALE. 1965. Mass-rearing of the larvae of nine noctuid
species on a simple artificial medium. J. Econ. Entomol. 58: 524-4.





A SURVEY OF THE ECONOMICALLY IMPORTANT
FRUIT FLIES (DIPTERA: TEPHRITIDAE)
PRESENT IN CHIAPAS AND A FEW OTHER
FRUIT GROWING REGIONS IN MEXICO

MARTIN ALUJA
Department of Entomology
University of Massachusetts
Amherst, MA 01003 USA
MARIO CABRERA, EUGENIO RIOS, JORGE GUILLEN, HILARIO CELEDONIO,
JORGE HENDRICHS AND PABLO LIEDO
Program Mosca del Mediterraneo, DGSPAF-SARH
Apartado Postal 369
30700 Tapachula, Chiapas MEXICO

ABSTRACT

We report on the progress of a study that has as an objective the identification of
the economically important fruit flies (Diptera: Tephritidae) of the State of Chiapas,
Mexico. To date, the following genera and species have been identified: Anastrepha
acris, A. alveata, A. balloui, A. bicolor, A. chiclayae, A. distinct, A. fraterculus, A.
leptozona, A. ludens, A. obliqua, A. montei, A. pallens, A. robusta, A. striata, A.
serpentina and A. tripunctata acriss, alveata, balloui, leptozona and montei are new
records for Mexico); Toxotrypana curvicauda; Zonosemata cocoyoc; Hexachaeta sp.











Aluja et al.: Fruit Flies of Chiapas 321

(probably obscure); Molynocoelia sp. (near lutea) and Blepharoneura sp. We have also
identified the genus Richardia (Diptera: Richardidae) and the genus Xanthacrona
(Diptera: Otitidae). We provide information on how and where these specimens were
collected and discuss the significance of these findings. Finally, we report on the iden-
tification of a few specimens collected outside of the State of Chiapas, which include
among others A. spatulata, A. robusta and A. pallens.

RESUME

Se reportan los resultados de un studio cuyo objetivo es la identificaci6n de las
moscas de la fruta de importancia econ6mica (Diptera: Tephritidae) en el estado de
Chiapas, M6xico. Hasta la fecha se han identificado los siguientes g6neros y species:
Anastrepha acris, A. alveata, A. balloui, A. bicolor, A. chiclayae, A. fraterculus, A.
distinct, A. leptozona, A. ludens, A. obliqua, A. montei, A. pallens, A. robusta, A.
striata, A. serpentina y A. triangulata acriss, alveata, balloui, leptozona y montei son
nuevos reports para Mexico). Toxotrypana curvicauda; Zonosemata cocoyoc;
Hexachaeta sp. (probablemente obscura); Molynocoelia sp. (cerca de lutea);
Blepharoneura sp. Tambi6n se han identificado los generos Richardia (Diptera: Richar-
didae) y Xanthacrona (Diptera: Otitidae). Se provee informaci6n acerca de como y
donde se colectaron los especimenes y se discute sobre la importancia de estos descub-
rimientos. Finalmente, se report sobre la identificaci6n de algunos especimenes colec-
tados fuera del estado de Chiapas, que incluyen entire otros a A. pallens, A. robusta y
A. spatulata.



Even though fruit fly species (Diptera: Tephritidae) found in Mexico have been the
focus of continuous attention by researchers, regulatory entomologists and fruit growers
for most of the past 90 years, we still know relatively little about them. The early
literature is filled with anecdotal accounts on the behavior and control of these insects
(Herrera 1905, Crawford 1923) and more recent attempts to clear up some of the original
confusion about their biology, ecology and behavior are rare (Baker et al. 1944, Baker
1945, Christenson and Foote 1960, McFadden 1964, Shaw et al. 1970, Morgante et al.
1983, Malavasi et al. 1983, Aluja et al. 1983, Robacker and Hart 1985, Aluja 1985).
Taxonomy has been the field most intensely studied, resulting in extensive keys for
adult identification (Dampf 1933, Greene 1934, Stone 1942, Shaw 1962, Ramos 1975,
Steyskal 1977a, Foote 1980 and Norrbom 1985). Unfortunately, a challenge one still
faces when attempting to identify field collected material is the lack of appropriate
taxonomic keys for the immature stages since currently existing ones are out of date
or incomplete (Greene 1929, Philips 1938, Bush 1962 and Berg 1979).
Three economically important genera of fruit flies are found in Mexico: Anastrepha
Schiner, Rhagoletis Loew and Toxotrypana Gerstaecker. Stone (1942) reported 13
species of the genus Anastrepha: aphelocentema Stone, chiclayae Greene, distinct
Greene, fraterculus Wiedeman, lathana Stone, ludens Loew, mombinpraeoptans Sein,
robusta Greene, serpentina Wiedeman, spatulata Stone, striata Schiner, tripunctata
Wulp (also in Foote 1965) and zuelaniae Stone. Foote (1967) added A. triangulata
(described from Morelos by Shaw (1962)) to the list. Steyskal (1975) recognized mombin-
praeoptans as a synonym of obliqua Macquart and also placed the genera Lucumaphila
Stone, Phobema Aldrich and Pseudodacus Hendel in synonymy with Anastrepha
(Steyskal 1977b). This adds 4 more species to the original list by Stone (1942): dentata
Stone, sagittata Stone (formerly placed in Lucumaphila, Foote 1967), bicolor Stone,
pallens Coquillett (formerly placed in Pseudodacus, Foote 1967).
There are 8 species of the genus Rhagoletis Loew reported in Mexico: boycei Cres-
son, cingulata Loew, complete Cresson, juglandis Cresson, pomonella Walch, ramosae











322 Florida Entomologist 70(3) September, 1987

Hernandez-Ortiz, striatella Wulp, and zoqui Bush (Foote 1967, 1981, Hernandez-Ortiz
1985). Finally, Foote (1967) reports 1 species of the genus Toxotrypana : curvicauda
Gerstaecker.
We consider that the number of Anastrepha species present in Mexico is much
greater than the 18 previously reported. For example, a survey of tephritids conducted
many years ago in Panama, a country with vegetational and geographical conditions
that closely resemble certain regions of Mexico, yielded 54 species (Stone 1942). Only
a few efforts of this nature have been carried out so far in Mexico. Baker et al. (1944)
describe a series of intensive surveys throughout the country in which the partial distri-
butions of Anastrepha ludens, A. pallens, A. sagittata, A. serpentina, A. striata, A.
distinct, A. obliqua, A. fraterculus, A. chiclayae, A. aphelocentema and Toxotrypana
curvicauda are reported. Gonzales-Hernandez and Tejada (1980) report A. ludens, A.
serpentina and 2 unidentified species of Anastrepha in the State of Nuevo Leon. Duran-
Pompa et al. (1981) report A. ludens, A. serpentina and A. obliqua in the Municipio of
Texcoco, State of Mexico. Coronado (1964) and Vera-Graciano et al. (1979) report
Rhagoletis cingulata and R. pomonella for the same region. Finally, Rios-Martinez
(1961) identified 8 species of the genus Anastrepha in the State of Chiapas: chiclayae,
distinct, fraterculus, ludens, obliqua, serpentina, striata and tripunctata.
This paper partially reports the results of a study primarily conducted in the State
of Chiapas, in which the objectives were to study the taxonomy, biology, ecology and
behavior of the local fruit flies and to develop adequate integrated management pro-
grams for those species causing economic damage to local orchards. In this particular
case we report on the species identified so far in the Soconusco Region, the Mazapa de
Madero Valley and the Chahuites Region in the neighboring State of Oaxaca. We also
include the results of identifications made on a few specimens received from the States
of Nayarit, Sinaloa, Tabasco and Veracruz.

MATERIALS AND METHODS

Localities
The study area comprised the Soconusco region and Mazapa de Madero Valley in
the State of Chiapas, Mexico. The agriculturally important Soconusco Region is located
between 14'10' and 1520' northern latitude and 9210' and 9310' western longitude.
The region comprises 16 municipalities which cover an area of 5937 km2. Mean annual
temperature is 260 C (ranges between 19 and 320 C) and mean annual precipitation 3000
mm (ranges between 2000 and 5000 mm). Tapachula, the most important city in the
region, is easily accessed by railroad, road and airplane. The major part of our survey
was carried out in the municipalities of Tapachula, Suchiate and Mazatdn with mean
elevations of 137, 30 and 32 meters above the sea level.
The Mazapa de Madero Valley is located in the Sierra Madre de Chiapas approxi-
mately 105 km West from the city of Tapachula; it belongs to the municipality of Mazapa
de Madero. Agriculture is the main source of income for local people but it is carried
out on a much smaller scale than in the Soconusco Region. The valley has an area of
approximately 116.8 km2, a mean annual temperature of 200 C (ranges between 18 and
31 C) and a mean annual precipitation of 1500 mm (ranges between 1000 and 2000 mm).
It is at an elevation of 1040 meters above sea level.
We also include the identifications of a few specimens sent to us from the Chahuites
Region in the neighboring State of Oaxaca, the towns of Martinez de la Torre and
Papantla in the State of Veracruz, and the mango-growing regions in southern parts of
the States of Nayarit and Sinaloa and in central portions of the State of Tabasco.











Aluja et al.: Fruit Flies of Chiapas


Survey Techniques
We established a series of trapping routes in which a total of 750 McPhail traps
(McPhail 1937), designed for capturing adults, were distributed in the following fashion:
400 in the mango (Mangifera indica) orchards Buena Vista, San Francisco, El Pelon,
Quinta Irenne and El Vergel (all within the Soconusco Region). 80 traps were placed
in each orchard (50 in the main block of trees and 30 in the periphery, which in some
occasions included non-host trees). It is important to note that the Quinta Irenne or-
chard had citrus (Citrus spp.), guava (Psidium guajava), chico zapote (Manilkara ac-
hras), guanabana (Anona muricata) and chirimoya (Anona cherimola) trees in addition
to mango. Another 100 traps were placed in citrus, guava and chico zapote orchards.
These 500 traps were all distributed within the Soconusco Region.
Another 250 traps were distributed in the Mazapa de Madero Valley; most of them
were hung on mango trees, but some were also hung in citrus, papaya, and mamey
(Calocarpum zapota) trees.
Each trap was baited with Staley's hydrolized protein (PIB-7) and borax dissolved
in 250 ml of water (20 ml of protein and 10 g of borax). Traps were hung in the upper
3/4 of the tree canopy and checked every 7 days. When serviced, the contents of each
trap were sieved, the insects caught rinsed with clean water and then placed in a 25 ml
vial filled with 70% alcohol; the traps were then also rinsed and re-baited. A strict
record for each individual trap was kept to facilitate the processing of data.
The specimens received from outside of the State of Chiapas were also caught in
McPhail traps and handled in the same way as described above.

Identification
All the flies caught in the traps were individually identified in the laboratory. Iden-
tifications were made by the following people: Juan Garcia, Eugenio Rios, Jorge Guillbn
(MoscaMed Program) and Amparo Ramos, Guadalupe Siller (DGSV-SARH, Mexico
City) who identified A. acris, A. balloui, A. distinct, A. fraterculus, A. ludens, A.
obliqua, A. serpentina, A. spatulata, A. striata, A. robusta, T. curvicauda, Hexachaeta
sp. (probably obscura); Dr. Richard Foote (Systematic Entomology Laboratory, BBII,
Agricultural Research Service, Washington, D.C.; retired) who identified A. chiclayae,
A. leptozona, A. montei, Molynocoelia sp. (near lutea), Blepharoneura sp., Richardia
sp. and Xanthacrona sp. and Dr. Allen Norrbom (Systematic Entomology Laboratory)
who identified A. alveata, A. pallens, A. bicolor, A. tripunctata, Zonosemata cocoyoc
and confirmed the identification of A. balloui.


RESULTS

Results are summarized in Table 1. The tephritid genera Anastrepha, Toxotrypana,
Molynocoelia, Hexachaeta and Blepharoneura were identified in the Soconusco Region.
A few specimens of Richardia (Diptera: Richardidae) and Xanthacrona (Diptera:
Otitidae) were also collected in the same area. All specimens were collected in McPhail
traps hung in mango, citrus, chico zapote and one unidentified wild tree. It is interesting
to note that in 1 orchard (Quinta Irenne) eight species of Anastrepha were collected
(unpublished data). Figure 2 indicates the proportion (%) of each species collected in
the 5 commercial mango orchards studied.
In Mazapa de Madero only the genera Anastrepha, Toxotrypana and Zonosemata
were identified. Toxotrypana specimens emerged from host fruits (Carica papaya) and
Anastrepha specimens were collected in traps hung in mango trees. During 1 week in
May 1985, 6 species of Anastrepha were collected in a single trap distinctt, fraterculus,
ludens, obliqua, serpentina and an undetermined species).


323












Florida Entomologist 70(3)


TABLE 1. FRUIT FLY SPECIES IDENTIFIED IN THE STATE OF CHIAPAS AND A FEW
OTHER FRUIT GROWING REGIONS IN MEXICO, INDICATING LOCALITY
AND TREE TYPE WHERE TRAPS WERE HUNG.

Tree species on which
Locality Fruit Fly Species trap was hung


Soconusco, Chiapas


Mazapa de Madero,
Chiapas












Chahuites, Oaxaca










Nayarit


Anastrepha acris
A. balloui
A. distinct
A. chiclayae
A. fraterculus
A. leptozona
A. pallens
A. robusta
A. tripunctata
A. obliqua


A. striata
A. ludens

A. serpentina
A. alveata
Toxotrypana curvicauda
Molynocoelia sp.
near lutea
Hexachaeta sp.
probably obscure
Blepharoneura sp.

A. bicolor
A. chiclayae
A. distinct
A. fraterculus
A. ludens
A. montei
A. obliqua
A. serpentina
A. striata
Zonosemata cocoyoc
T. curvicauda

A. chiclayae
A. distinct
A. fraterculus
A. ludens
A. obliqua
A. pallens
A. robusta
A. serpentina
A. spatulata
A. striata
A. chiclayae
A. ludens
A. obliqua
A. serpentina


Mangifera indica
PI


M. indica, Manilkara
achras

M. indica, Citrus sp.
Manilkara achras

Unidentified wild tree
M. indica







M. indica


Emerged from Carica
papaya
M. indica
If






"r


September, 1987












Aluja et al.: Fruit Flies of Chiapas


TABLE 1. Con't

Tree Species on which
Locality Fruit Fly Species trap was hung

A. striata
Sinaloa A. chiclayae
A. serpentina
A. striata
Tabasco A. ludens
A. obliqua
A. serpentina
Martinez de la Torre
Veracruz A. distinct Citrus sp.
A. fraterculus
A. ludens "
A. obliqua
A. serpentina
A. striata
T. curvicauda
Papantla, Veracruz A. ludens
A. striata
A. serpentina


Finally, all the specimens received from localities outside of the State of Chiapas
belonged to the genera Anastrepha and Toxotrypana and were collected in traps hung
in mango and citrus trees.

DISCUSSION AND CONCLUSIONS

Our initial assumption that there must be more Anastrepha and in general more
tephritid species in Mexico than those previously reported in the literature, was con-
firmed. Five out of the 18 fruit fly species identified in this study are new records for
Mexico: Anastrepha acris, A. alveata, A. balloui, A. leptozona and A. montei. Note


TABLE 2. RELATIVE ABUNDANCE OF THE ANASTREPHA SPECIES COLLECTED IN
FIVE COMMERCIAL MANGO ORCHARDS IN THE SOCONUSCO REGION,
CHIAPAS, MEXICO FROM 1982 TO 1985.

Species Proportion (%)

A. obliqua 70.77
A. ludens 23.86
A. serpentina 1.65
A. distinct 1.51
A.fraterculus 0.47
A. acris 0.44
A. striata 0.25
A. balloui 0.03
A. leptozona 0.013
A. chiclayae 0.008
A. sp. 0.004











Florida Entomologist 70(3)


also that we confirmed the presence in the State of Chiapas of all the Anastrepha
species reported in a previous study by Rios-Martinez (1961).
Trapping a particular fruit fly species in a trap placed in a certain tree species does
not necessarily mean that it is a host. Being aware of this fact, we have conducted a
study in which we identified most of the true hosts of the fruit fly species reported here
(unpublished data). The fact that we identified up to 8 species of the genus Anastrepha
in 1 mango orchard is not surprising since all the commercial orchards in the study areas
are surrounded by native vegetation, including in many occasions, wild host plants.
Besides, it is common practice to plant a wide array of varieties and fruit species in the
same orchard (Aluja 1985, Aluja & Liedo 1986). The latter situation is bound to attract
a large number of fruit flies of different species.
Capturing 5 species in one trap during the same week is a very interesting result
which prompted us to conduct a series of behavioral studies to determine if there was
competition for mating sites or interspecific mating attempts. It was shown that in the
case of A. ludens and A. obliqua, the extremely different hours of the day when mating
activities take place is a strong enough barrier to reduce interspecific sexual activities
(Aluja et al. 1983).
We consider that the species which are new records for Mexico do not represent any
economic hazard to the commercial fruit orchards of the region. Only A. leptozona was
caught in large numbers (unpublished data) and when this was the case it was in areas
of native vegetation, not even near commercial orchards. A. acris was caught only
during a restricted part of the year (July-September) in 2 orchards (El Carmen and
San Francisco). This is puzzling, especially since all our efforts to locate the host plants
have been so far unsuccessful. Where these isolated populations spend the rest of the
year merits further investigation. A. montei has only been caught on 2 occasions (3
specimens). This species is reported to attack yuca, Manihot dulcis and esculenta (Stone
1942). The trap in which it was caught was near some yuca plants and it is therefore
possible that it was using orchard trees as resting or shelter sites. A. alveata and A.
balloui are also extremely rare.
This study has also expanded our knowledge on the distribution of some Anastrepha
species in Mexico. Information in this area is very limited. We are currently working
on an updated distribution of the most common tephritids in Mexico.
Due to lack of more recent information we have followed Baker's terminology (Baker
et al. 1944) and treated the specimens of A. fraterculus as the "Mexican form" of this
species. The debate as to whether the South American populations are distinct species
is an old one, but has not been resolved. Baker et al. (1944) and Baker (1945) pointed
out some differences and suggested further careful biological work to define the latter.
Mendes (1958) and Bush (1962) found some karyotypic differences between morpholog-
ically indistinguishable Mexican and Brazilian populations. Morgante et al. (1980) and
Malavasi and Morgante (1982) in studies carried out in Brazil, uncovered considerable
genetic heterogeneity among geographic populations and concluded that fraterculus
includes many unrecognized species. Malavasi and Morgante (1983) later found that the
level of genetic distance between populations stemming from different hosts within one
orchard was very small. Aluja et al. (1983) speculated that a similar phenomenon could
be acting in the case of A. ludens, A. serpentina and A. obliqua, species that also have
wide geographic distributions. In the cases of A. ludens and A. serpentina geographic
differences in the pattern of host exploitation were found and in the case of A. obliqua
there are some unconfirmed reports of behavioral differences between Brazilian and
Mexican populations (Aluja et al. 1983, Teles da Silva et al. 1985). It becomes clear
then, that morphological characters for both immature and mature forms are not suffi-
cient to unravel all the questions posed in relation to the taxonomic and phylogenetic
classification of neotropical tephritids. Genetic, ecological and behavioral studies are


326


September, 1987











Aluja et al.: Fruit Flies of Chiapas


327


also necessary. The efforts by Morgante et al. (1980), Malavasi and Morgante (1982,
1983) and the recent comprehensive work by Norrbom (1985) are certainly steps in the
right direction.
In conclusion we have provided evidence that the number of species of the genus
Anastrepha reported in Mexico should be increased by 5 and that the distribution of
some of these species is more widespread than the limited literature in this field indi-
cates. Our results, added to the literature reports, indicate that the species of the genus
Anastrepha present in Mexico are: acris, alveata, aphelocentema, balloui, bicolor, chic-
layae, dentata, distinct, fraterculus, lathana, leptozona, ludens, montei, obliqua, pal-
lens, robusta, serpentina, sagittata, spatulata, striata, triangulata, tripunctata and
zuelaniae.

ACKNOWLEDGMENTS

We thank Dr. Michael Peters, Dr. Ronald Prokopy, Danette Reynolds, Sylvia and
Dan Cooley, Marta Quezada (Univ. of Massachusetts, Amherst) and especially Dr.
Allen Norrbom (Systematic Entomology Laboratory, ARS, USDA) for useful comments
on earlier versions of this manuscript. We express our appreciation to Mr. Gomez, Ing.
Leal Cantu, Ing. Nifio, Mr. Renovales, Mr. Fernandez and Mr. Fierro for granting us
permission to work in their orchards. We would also like to express our deep apprecia-
tion to Ing. Jorge Gutierrez Samperio (DGSPAF) for his constant support and encour-
agement. We acknowledge partial financial support by Dr. J. R. Aluja during the pub-
lication process of this paper. This work was supported by the Mediterranean Fruit Fly
Program, DGSPAF, SARH, Mexico.

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MCFADDEN, M. W. 1964. Recent developments in our knowledge of the biology and
ecology of the Mexican fruit fly. Planning conference for research on tropical fruit
flies in Mexico and Central America. p. 1-10.
MCPHAIL, M. 1937. Relation of time of day, temperature and evaporation to attrac-
tiveness of fermenting sugar solutions to Mexican fruit flies. J. Econ. Entomol.
30: 739-749.
MENDES, L. O. T. 1958. Observacoes citologicas em "moscas das frutas". Bragantia
17: 29-38.
MORGANTE, J. S., A. MALAVASI AND G. L. BUSH. 1980. Biochemical systematics
and evolutionary relationships of Neotropical Anastrepha. Ann. Entomol. Soc.
Am. 73: 622-630.
- A. MALAVASI AND R. J. PROKOPY. 1983. Mating behavior of wild Anastrepha
fraterculus (Diptera: Tephritidae) on a caged host tree. Florida Entomol. 66:
234-241.
NORRBOM, A. L. 1985. Phylogenetic analysis and taxonomy of the cryptostrepha,
daciformis, robusta and schausi species groups of Anastrepha Schiner (Diptera:
Tephritidae). Unpubl. PhD Thesis. Pennsylvania State University. 355p.
PHILIPS, V. T. 1938. The biology and the identification of trypetid larvae (Diptera:
Tephritidae). Mem. Amer. Ent. Soc. 12: 1-154.
RAMOS, A. 1975. Guia ilustrada para la identificaci6n de adults de moscas (Diptera:
Tephritidae) que afectan a la fruta en Mexico y de species ex6ticas de importan-
cia cuarentenaria. DGSV-SARH. M6xico D.F. M6xico. 40p.
RIOS-MARTINEZ, E. 1961. Trypetidos capturados en el estado de Chiapas. Unpub-
lished thesis. Escuela Nacional de Agricultura. Chapingo, Mexico. 98p.


September, 1987











Aluja et al.: Fruit Fly Hosts of Chiapas


ROBACKER, D. C. AND W. C. HART. 1985. Courtship and territoriality of laboratory-
reared Mexican fruit flies, Anastrepha ludens (Diptera: Tephritidae), in cages
containing host and nonhost trees. Ann. Entomol. Soc. Am. 78: 488-494.
SHAW, J. G. 1962. Species of the spatulata group of Anastrepha (Diptera: Tep-
hritidae). J. Kansas Ent. Soc. 35: 408-414.
- F. LOPEZ AND D. L. CHAMBERS. 1970. A review of research done with the
Mexican fruit fly and the citrus blackfly in Mexico by the Entomology Research
Division. Bull. Entomol. Soc. Amer. 16: 186-193.
STEYSKAL, G. C. 1975. Anastrepha obliqua (Macquart) the prior name for Anastrepha
mombinpraeoptans Sein. U.S. Dept. Agr. Coop. Econ. Ins. Rpt. 25: 357-358.
- 1977a. Pictorial key to species of the genus Anastrepha (Diptera: Tephritidae).
Entomol. Soc. Washington Special Publ. 35p.
--. 1977b. Two new neotropical fruit flies of the genus Anastrepha, with notes on
generic synonymy (Diptera: Tephritidae). Proc. Entomol. Soc. Washington. 79:
75-81.
STONE, A. 1942. The fruit flies of the genus Anastrepha. USDA Misc. Publ. No. 439.
Washington. 112p.
TELES DA SILVA, M., J. POLLONI AND S. BRESAN. 1985. Mating behavior of some
fruit flies of the genus Anastrepha (Diptera: Tephritidae) in the laboratory. Rev.
Brasil. Ent. 29: 155-164.
VERA-GRACIANO, J., J. L. CARRILLO-SANCHEZ, M. C. SOSA, E. L. S. VEGA Y R.
A. P. HUERTA. 1979. La mosca y la palomilla del capulin en las regions de San
Miguel Tlaixpan, Chapingo y San Dieguito, M6xico. Folia Entomol. Mexicana.
42: 1-25.


-C- -- *-^-^ --- -- -- -C-- -- -C^



NATURAL HOST PLANT SURVEY OF THE
ECONOMICALLY IMPORTANT FRUIT FLIES
(DIPTERA: TEPHRITIDAE) OF CHIAPAS, MEXICO

MARTIN ALUJA
Department of Entomology
University of Massachusetts
Amherst, MA 01003 USA

JORGE GUILLEN, GERMAN DE LA ROSA, MARIO CABRERA, HILARIO CELEDONIO,
PABLO LIEDO AND JORGE HENDRICHS
Program Mosca del Mediterraneo
Apartado Postal 369
30700 Tapachula, Chiapas, Mexico

ABSTRACT

We provide a natural host-plant list of some of the fruit flies (Diptera: Tephritidae)
reported for the State of Chiapas. Out of 92 plant species sampled, 39 species represent-
ing 12 plant families were identified as natural host of Anastrepha distinct, A. frater-
culus, A. leptozona, A. ludens, A. obliqua, A. serpentina, A. striata and Toxotrypana
curvicauda.


329










Florida Entomologist 70(3)


RESUME

Se present una lista de las plants que han sido identificadas como hospederas
naturales de algunas de las species de moscas de la fruta (Diptera: Tephritidae) repor-
tadas en el estado de Chiapas, Mdxico. De 92 species vegetables muestreadas, 39 es-
pecies, pertenecientes a 12 families fueron identificadas como hospederas naturales de
Anastrepha distinct, A. fraterculus, A. leptozona, A. ludens, A. obliqua, A. serpen-
tina, A. striata y Toxotrypana curvicauda.


Fruit flies (Diptera: Tephritidae) are well known in Mexico for the devastating im-
pact they can have on commercial fruit production. Entire fruit growing regions have
been forced out of business due to heavy infestations of these widely distributed insects
(Aluja and Liedo 1986). Larvae-infested fruit usually drop from the tree and rot, render-
ing them useless for human consumption. The presence of these insects also elicits
neighboring regions and countries to apply severe quarantine restrictions to prevent
their introduction to fruit growing regions free of fruit flies. Even though economic
damage is basically restricted to semicommercial and commercial orchards and isolated
backyard trees, fruit flies infest a wide variety of wild native host plants and even small
populations serve as reservoirs from which commercial fruit orchards are invaded. Aluja
and Liedo (1986) proposed a series of ecologically based management practices such as
habitat manipulation, trap crops and orchard designs, where emphasis is placed on
trying to stop the continuous flow of populations from one host to another. They
nevertheless noted that one prerequisite that must be fulfilled before even attempting
to experimentally achieve this goal, is the complete identification of all cultivated and
wild host plants for each of the fruit fly species causing economic damage.
We define a natural host as a plant bearing fruit or any other tissue where female
fruit flies lay eggs in nature and the emerging larvae are able to survive and reach the
adult stage with varying degrees of fitness. Some host plant records in the literature
are misleading because they are based on oviposition induced under artificial conditions
on fruits presented to flies which had been deprived of an oviposition substrate for days.
Such practices have led researchers to report fruits such as prickly pear (Opuntia sp.),
banana (Musa sp.), squash (Cucurbita mexicana), walnut (Juglans regia), papaya
(Carica papaya) and garden bean (Phaseolus vulgaris) as hosts of Anastrepha ludens
(Baker et al. 1944, Ebeling 1950 and Weems 1963). Such reports are sometimes used
in quarantine protocols to limit the movement of these and other fruits from one country
to another, an unfortunate fact since it is almost impossible for these plants to serve as
hosts under natural conditions.
Some host plant records for fruit flies found in Mexico are commonly cited in the
literature but, unfortunately, many are very old and have not been confirmed since
their first publication (Arriaga-Martinez 1979, Baker et al. 1944, Bush 1957, 1962,
Coronado 1964, Greene 1934, McPhail and Berry 1936, Plummer et al., 1941, Ramos
1975, Shaw 1947, Stone 1942 and Weems 1963). We suspect that some host records may
be the result of misidentifications of both plant and fly species. Ballou (1945), Blanchard
(1937), Bricefo (1975), Caraballo (1981), Cuculiza and Torres (1975), Eskafi and Cunnin-
gham (1987), Fernandez-Yepez (1953), Guagliumi (1966), Herrera and Vifas (1977),
Jir6n and Zeled6n (1979), Korytkowski and Ojeda-Pefia (1968), Lima (1934), Lutz and
Lima (1918), McAlister (1936), Malavasi et al. (1980), Martorell (1939), Norrbom (1982),
Olarte (1980), Rosillo (1953), Wasbauer (1972) and Whervin (1974) also published records
for hosts that although not necessarily found in Mexico are nevertheless attacked by
fruit fly species occurring in that country. Norrbom (1985) provides an excellent sum-
mary of most of the published host records for all Anastrepha species.


September, 1987











Aluja et al.: Fruit Fly Hosts of Chiapas 331

In this paper, we provide an updated list of the natural host plants of some of the
fruit flies identified by Aluja et al. (1987) in the State of Chiapas.

MATERIALS AND METHODS

The study area comprised the Soconusco Region and the Mazapa Valley in the State
of Chiapas, Mexico. A detailed description of these areas is given in Aluja et al. (1987).
Fruits from known and potential host plants were collected in commercial and
semicommercial orchards, backyard gardens and areas covered with wild native vegeta-
tion from March 1982 to December 1985. Individual samples consisted of fallen fruit
(80%) and fruit still on the tree (20%) and ranged in number from 1 to 120 fruits
depending on the availability of a particular species or variety. The degree of ripeness
varied from very green to very ripe. Samples were individually labeled, placed in plastic
bags and transported to the laboratory in well cushioned and aereated styrofoam boxes
to avoid mortality of inmatures due to overheating and physical contact.
In the laboratory each bag was weighed and the number of fruits per sample re-
corded. Fruits were then immersed for 5 seconds in a 10% solution of sodium benzoate
and water (to inhibit growth of fungi and decomposing bacteria) and placed in "matura-
tion chambers". These are 18 by 33 by 25 cm styrofoam boxes, adapted to hold a wire
screen basket on which fruits were placed (Fig. 1). The bottom of each box was covered
with vermiculite or sawdust to provide an adequate pupation substrate for the third
instar larvae dropping from infested fruit. The cover of the "maturation chamber" had
a 10 by 28 cm screen that permitted air to flow freely but stopped any emerging fruit
flies or fruit fly parasitoids (see Aluja 1985 for details). It is important to note that when
collecting fruit from the tree, care should be taken to prevent the fruit from falling to
the ground. If the fruit ruptures, high larval mortality may be caused by the sodium
benzoate solution. Figure 2 shows a tool that is very useful for the collection of fruit
from tall branches.
Depending on the ripeness or decomposition stage of the fruit samples, these were
kept in the "maturation chambers" for a period of 4 to 14 days. Periodic inspections and
dissections of randomly selected samples determined when most of the larvae had left
the fruit. For each dissection we recorded the number and stage of remaining live or







c(-j--MESH t- 40 -I

FRUIT

25
WIRE MESH -S --


PUPATION MEDIA

Fig. 1. "Maturation Chamber" for the collection of fruit fly larvae and pupae from
infested fruit (dimensions are in cm).











Florida Entomologist 70(3)


Fig. 2. Fruit collection net with sharp hook for cutting of thick pedicels.

dead larvae. At the same time the sawdust was carefully inspected and sieved to collect
all pupae and larvae, which were placed in 500 ml plastic containers half filled with moist
soil. Containers were covered with a fine screen capable of allowing aereation but
stopping the emerging fruit flies and parasitoids from escaping. Water was regularly
added to the soil in order to maintain optimal humidity conditions. The laboratory in
which the plastic containers were kept had an average temperature of 28 3 C and
RH of 55 5%.
The containers were inspected daily and any fruit fly or parasitoid that had emerged
was removed. Part of this biological material was killed and identified and part used
for demographic studies. The identification process is described in Aluja et al. (1987).
Host plant identifications were based on Tejada (1980), Miranda (1976) and corroborated
by Mario Cabrera (Universidad Autonoma de Chiapas).

RESULTS

A total of 28,830 samples representing 92 species from 24 plant families were taken
(see Table 1). Of these, 39 species from 12 families turned out to be natural hosts of the
fruit fly genera Anastrepha and Toxotrypana (see Table 2). The number of hosts iden-
tified for each fruit fly species ranged from 1 to 14: A. ludens (14), A. serpentina (11),
A. obliqua (11), A. fraterculus (10), A. distinct (4), A. leptozona (2), A. striata (1) and
Toxotrypana curvicauda (1).

DISCUSSION AND CONCLUSIONS

Our results increase by 10 the number of natural hosts reported for the fruit flies of
Chiapas (Aluja 1985 reports 24 natural hosts).
It is important to note that the information provided here refers only to the positive
identification of a plant species as a natural host of fruit flies. It does not indicate the
degree of infestation nor the relative frequency of infestation. Aluja and Liedo (1986)
have shown that simply recording a plant as a host provides little information since
some hosts may be only occasionally exploited. Usually it is observed that a few fruit


332


September, 1987











Aluja et al.: Fruit Fly Hosts of Chiapas


TABLE 1. PLANT SPECIES SAMPLED IN THE SOCONUSCO REGION AND THE
MAZAPA DE MADERO VALLEY (CHIAPAS, MEXICO) TO DETERMINE
THEIR STATUS AS HOST PLANTS OF THE LOCAL FRUIT FLIES.

Common name
Spanish English Scientific name


Aceituno
Aguacate
Algodoncillo
Almendra
Anona
Anona colorada
Baricoco
Cachofla
Caco
Cafe
Caimito
Capulin
Caspirol
Chachalaco
Chalfim
Chayote
Chelel
Chico zapote
Chileamate
Chilindr6n
Chincuya
Cidra
Coroso
Cuajilote
Cuajinicuil
Cuernavaca
Cupap6
Encino (zapotillo)
Granada
Granadilla
Grosella
Guamuchil
Guandbana
Guaya
Guayaba
Higo
Huevo de mono
Hule silvestre
Jaboncillo
Jocote
Jobo de pava
Jushte
Lech6n
Lima chiche
Lima lim6n
Lombricero
Maluco
Mandarina
Mandarina china
Mandarina criolla


Wild olive
Avocado
?
Tropical almond
Sweetsop
Custard apple
?
?

Coco plum
Coffee
Star apple
Strawberry tree
Inga
?
Inga
Chayote
Inga
Sapodilla
?
?
?
Citron
Corozo palm fruit
Cuachilote
Inga
?
?

Pomegranate
Passion fruit
?
?
Soursop
Spanish lime
Guava
Fig
Inga
?
Soapberry
Purple mombin
Mombin
Breadnut
?

Sweet lime
?

Tangerine
Chinese tangerine
Creol tangerine


Simarouba glauca Dc.
Persea americana Mill.
?
Terminalia catappa L.
Annona squamosa L.
A. reticulata L.
Micropholis mexicana Gilly
?
Chrysobalanus icaco L.
Coffea arabica L.
Chrysophyllum cainito L.
Muntingia calabura L.
Inga lauriana Willd.
?
Inga micheliana Harms
Sechium edule (Jacq.) Swartz
Inga leptoloba Schl.
Manilkara achras Mill.
?
?
Annona purpurea Mets.
Citrus medical L.
Orbignya cohune (Mart.)
Paermenteria edulis D. C.
Inga spuria H. et B.
Solanum macranthum Dunal.
Cordia dodecandra Dc.
Comepia polyandra Rose
Punica granatum L.
Passiflora edulis Sims
?
Pithecolobium dulce Benth
Anona muricata L.
Melicocca bijuga L.
Psidium guajava L.
Ficus carica L.
Inga sp.
?
Sapindus saponaria L.
Spondias purpurea L.
Spondias mombin L.
Brosimum alicastrum Sw.
?
Citrus sp.
C. limetta Rissso (Christm.)
?
?
Citrus deliciosa Tencre
Citrus sp.
C. reticulata Blco.











Florida Entomologist 70(3)


TABLE 1. (Continued)

Common name
Spanish English Scientific name


Mango
Manzanilla
Mapahuite
Marachan
Marafi6n
Matapalo
Matasano
Matzfi
Melocot6n
Mel6n
Nance


Mango
Hawthorn
?
?
Cashew nut
?
White sapote
?
Peach
Musk melon
Nanche


Naranja agria
Naranja dulce
Nispero
Pacaya

Pan de la India
Pan de maria
Pan de palo

Papause
Papaya commercial
Papaya silvestre
Patashte

Paterna
Pepino silvestre
Pera de agua
Pifi6n
Pitanga
Pomarrosa
Pomelo

Poz61
Quishtan
Sangre de toro
Sikajal
Taculchacha
Tempisque

Tomate silvestre
Toronja
Toronjil
Tuna
Zapote amarillo
Zapote colorado
Zapote de agua
Zapote negro
Zunzapote


Sour orange
Orange
Loquat
Pacaya palm fruit

Jack fruit tree
?
Breadfruit

Ilama
Papaya
Wild papaya
?

Inga
Wild cucumber
?
?
Surinam cherry
Rose apple
Pummelo


Wild Tomato
Grapefurit
?
Prickly pear
Canistal
Mammee apple
Black sapote
Black sapote


Mangifera indica L.
Crataegus sp.
Trichilia sp.
?
Anacardium occidental L.
Ficus tecolutensis Standley
Casimiroa edulis Llave & Lex.
Cordia alba Roem & Schult.
Prunus persica (L.) Batsch
Cucumis melo L.
Byrsonima crassifolia (L.)
HBK
Citrus aurantium L.
C. sinensis (L.) Osbeck
Eriobotryajaponica Lindl.
Chamaedorea aguilariana
Standley & Steyerm.
Artocarpus heterophyllus Lam.
?
Artocarpus altilis (Perkins.)
Fosb.
Annona diversifolia Saff.
Carica papaya L.
Carica cauliflora JMacq.
Theobroma bicolor Humb. &
Bonpl.
Inga paterna Harms
Cucumis sp.
Syzygium malaceensis L.
Jatropha curcas L.
Syzygium uniflora L.
Syzygiumjambos L.
Citrus maxima (Burm.)
Merrill
Alchornea latifolia Sw.
Solanum wendlandii Hook.
?
?
?
Mastichodendron capiri var.
tempisque (A. DC.) Cronq.
Lycopersicon sp.
Citrus paradisi Macfady
Rheedia edulis Planch & Triana
Opuntia tuna (L.) Mill.
Pouteria campechiana Baehni
Calocarpum sapota Merr.
Pachira aquatica Aubl.
Diospyros ebenaster Retz.
Licania platypus Fritsch


334


September, 1987











Aluja et al.: Fruit Fly Hosts of Chiapas


335


TABLE 2. NATURAL HOST PLANTS OF SOME OF THE FRUIT FLIES OF CHIAPAS,
MEXICO. LISTED IN ORDER OF ECONOMIC IMPORTANCE.

Common
local name Scientific name
Fruit fly species of host plant of host plant


Anastrepha ludens














Anastrepha serpentina










Anastrepha obliqua


Anastrephafraterculus


Anastrepha distinct


Mango
Naranja dulce
Naranja agria
Pomelo
Mandarina criolla
Toronja
Tangerina
Cidra
Lima lim6n
Mandarina China
Naranja injerto
Matasano
Anona
Tempisque

Zapote colorado
Chico zapote
Caimito
Zapote amarillo
Baricoco
Pomelo
Naranja agria
Naranja dulce
Pozol
Nance
Mango
Mango
Guayaba
Pomarrosa
Pera China
Jobo de pava
Jocote
Jushte
Nispero
Poz61
Grosella
Manzanilla
Guayaba
Pomarrosa
Poz61
Pitanga
Tempisque


Almendra
Cafe
Mango
Grosella
Manzanilla
Chalim


Mangifera indica L.
Citrus sinensis L. (Osbeck)
C. aurantium L.
C. maxima (Burm.) Merrill
C. reticulata Blco.
C. paradisi Macfady
C. deliciosa Tenore
C. medical L.
C. limetta Risso (Christm.)
C. sp.
C. sp.
Casimiroa edulis Llave & Lex.
Annona squamosa L.
Mastichodendron capiri var.
tempisque (A. D.C.) Cronq.
Calocarpum sapota Merr.
Manilkara achras (Mill.)
Chrysophyllum cainito L.
Pouteria campechiana Baehni
Micropholis mexicana L.
Citrus maxima (Burm) Merrill
C. aurantium L.
C. sinensis (L.) Osbeck
Alchornea latifolia Sw.
Byssonima crassifolia (L.) HBK
Mangifera indica L.
Mangifera indica L.
Psidium guajava L.
Syzygiumjambos L.
S. malaceensis L.
Spondias mombin L.
S. purpurea L.
Brosium alicastrum (Ramon)
Eriobotryajaponica Lindl.
Alchornea latifolia Sw.
?
Crataegus sp.
Psidium guajava L.
Syzygiumjambos L.
Alchornea latifolia Sw.
Syzygium uniflora L.
Mastichodendron capiri var.
tempisque (A. D. C.) Cronq.
Terminalia catappa L.
Coffea arabica L.
Mangifera indica L.
?
Crataegus sp.
Inga micheliana Harms











Florida Entomologist 70(3)


TABLE 2. (Continued)

Common
local name Scientific name
Fruit fly species of host plant of host plant

Caspirol I. lauriana Willd.
Cuajinicuil I. spuria H. et B.
Paterna I. paterna Harms
Anastrepha leptozona Baricoco Micropholis mexicana Gilly
Manzanilla Crataegus sp.
Anastrepha striata Guayaba Psidium guajaba L.
Toxotrypana curvicauda Papaya Carica papaya L.



species are preferred in any given region. We therefore suggest that any future host
records also provide complete information as to how important the host is and what the
host utilization patterns are in nature (Aluja et al. unpublished information).
After carefully studying the host plant records reported in the literature, we have
concluded that the range of our sampling schemes need to be increased to consider more
species of families that at first would not appear to be an adequate substrate for larval
development. For example, the families Euphorbiaceae, Passifloraceae and Ebenaceae
are very common in the Soconusco Region, but were only sampled in low numbers. An
extended sampling effort may soon allow us to identify the natural hosts of A. alveata,
A. balloui, A. bicolor, A. chiclayae, A. montei, A. pallens and A. tripunctata, the
other fruit flies identified in the State of Chiapas by Aluja et al. (1987).
We consider that some of the hosts reported in the literature may be the result of
misidentifications or sampling errors. It seems unlikely that Manihot esculenta is a host
of A. obliqua as Guagliumi (1966) suggests. Caraballo (1981) reports Theobroma sp. as
a host of A. fraterculus which is a phenomenon not observed in the varieties of Theob-
roma sp. planted in Mexico. Most of the hosts reported by Korytkowski & Ojeda-Pefia
(1968) had never been reported before for the species they indicate.


ACKNOWLEDGMENTS

We gratefully acknowledge the authorization by Dr. Allen Norrbom (Systematic
Entomology Laboratory, ARS, USDA) to use some of the information contained in his
unpublished PhD thesis. In particular we refer to his extensive literature review on
host plants of some Anastrepha species. Dr. Norrbom also made many useful comments
to an earlier version of this paper. We want to thank Dr. Carrol Calkins (Insect Attrac-
tants, Behavior, and Basic Research Laboratory, ARS, USDA) for his important sup-
port during the publication process of this paper and Ing. Jorge Gutierrez Samperio
(DGSPAF) for his constant encouragement. Partial financial support during the publica-
tion process of this paper was provided by Dr. J. R. Aluja. This work was supported
by the Mediterranean Fruit Fly Program, DGSPAF, SARH, Mexico.


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Mexico. 241 p.


September, 1987












Aluja et al.: Fruit Fly Hosts of Chiapas 337

ALUJA, M., AND P. LIEDO. 1986. Future perspectives on integrated management of
fruit flies in Mexico. In: Pest control: operations and systems analysis in fruit fly
management. Proceedings of a NATO advanced research workshop. M. Mangel
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Tephritidae) present in Chiapas, Mexico. Florida Entomol. 70: 320-329.
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BALLOU, C. H. 1945. Notas sobre insects dafiinos observados en Venezuela (1938-
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BRICENO, A. 1975. Distribuci6n de las moscas de las frutas (Anastrepha spp., Diptera:
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Agron. (Univ. Zulia, Maracaibo) 3: 45-49.
BUSH, G. L. 1957. Some notes on the susceptibility of Avocados in Mexico to attack
by the Mexican fruit fly. Rio Grande Valley Hort. Soc. 7: 75-78.
- 1962. The cytotaxonomy of the larvae of some Mexican fruit flies in the genus
Anastrepha (Diptera: Tephritidae). Psyche 69: 87-101.
CARABALLO, J. 1981. Las moscas de frutas del g6nero Anastrepha Schiner, 1868
(Diptera: Tephritidae) de Venezuela. Unpubl. M.S. thesis, Universidad Central
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region de Texcoco, MWxico. Fit6filo 17: 9-18.
CUCULIZA, T. M., AND V. E. TORRES. 1975. Moscas de la fruta en las principles
plants hospederas del valle de Huasnuca. Revista Peruana Entomol. 20: 107-
114.
EBELING, W. 1950. Subtropical Entomology. Lithotype Process Co. San Francisco,
USA. 747. p.
ESKAFI, F. M., AND R. T. CUNNINGHAM. 1987. Host plants of fruit flies (Diptera:
Tephritidae) of economic importance in Guatemala. Florida Entomol. 70: 116-
123.
FERNANDEZ-YEPEZ, F. 1953. Contribuci6n al studio de las moscas de las frutas del
genero Anastrepha Schiner (Diptera: Trypetidae) de Venezuela. II Cong. Cien.
Nat. Afin. (Caracas) No. 7: 5-42.
GREENE, C. T. 1934. A revision of the genus Anastrepha based on a study of the
wings and on the length of the ovipositor sheath (Diptera: Trypetidae). Proc.
Entomol. Soc. Washington. 36: 127-179.
GUAGLIUMI, P. 1966. Insetti e Aracnidi delle plante comuni del Venezuela segnalanti
nel period 1938-1963. Rel. Monog. Agr. Subtrop. Trop. (Inst. Agron. L'Oltra-
mare, Firenze). No. 1-391.
HERRERA, A. J. M., AND L. E. VINAS. 1977. Moscas de la fruta (Diptera: Tep-
hritidae) en mangos de chulucanas, Piura. Revista Peruana de Entomol. 20: 107-
114.
JIRON, L. F., AND R. ZELEDON. 1979. El g6nero Anastrepha (Diptera: Tephritidae)
en las pricipales frutas de Costa Rica y su relaci6n con pseudomiasis hummana.
Rev. Trop. Biol. 27: 155-161.
KORYTKOWSKI, C., AND D. OJEDA-PE&A. 1968. Especies del genero Anastrepha
Schiner 1868 en el nor-oeste Peruano. Rev. Per. Entomol. 11: 32-70.
LIMA, A. DA COSTA. 1934. Moscas de frutas do genero Anastrepha Schiner, 1868
(Diptera: Trypetidae). Mem. Inst. Oswaldo Cruz 28: 487-575.
LUTZ, A., AND A. DA COSTA LIMA. 1918. Contribucao para o estudo das Tripanidas
(mosca de frutas) brasileiras. Mem. Inst. Oswaldo Cruz. 10: 4-16.











338 Florida Entomologist 70(3) September, 1987

MCALISTER, L. C. JR. 1936. Observations on the West Indian fruit fly at Key West
in 1932-33. J. Encon. Entomol. 29: 440-445.
MCPHAIL, M. AND N. O. BERRY. 1936. Observations on Anastrepha pallens (Coq.)
reared from wild fruits in the lower Rio Grande Valley of Texas during the spring
of 1932. J. Econ. Entomol. 29: 405-410.
MALAVASI, A., J. S. MORGANTE, AND R. A. ZUCCHI. 1980. Biologia de "Moscas
das-frutas" (Diptera: Tephritidae). I. Lista de hospederos e ocorencia. Revista
Brasil. Biol. 40: 17-24.
MARTORELL, L. F. 1939. Insects observed in the state of Aragua, Venezuela, South
America. J. Agr. Univ. Puerto Rico (Rio Piedras) 23: 219.
MIRANDA, F. 1976. La vegetaci6n de Chiapas. Ediciones del Gobierno del Estado.
Tuxtla Gutierrez, Chiapas, M6xico. 2 vols.
NORRBOM, A. L. 1982. A new host record for Rhagoletis striatella (Diptera: Tep-
hritidae). Melsheimer Entomol. Ser. 32: 11.
1985. Phylogenetic analysis and taxonomy of the cryptostrepha, daciformis,
robusta and schausi species groups of Anastrepha Schiner (Diptera: Tep-
hritidae). Unpubl. PhD Thesis. Penn, State University. 355 p.
OLARTE, W. E. 1980. DinAmica poblacional del complejo constituido por las moscas
de las frutas Anastrepha striata y Anastrepha fraterculus en el medio ecol6gico
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WASBAUER, M. 1972. An annotated host catalog of the fruit flies of America North
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Florida Dept. Agric.
WHERVIN, L. W. VAN. 1974. Some fruitflies (Tephritidae) in Jamaica. Pans. 20:
11-19.











Eger: Review of Tiridates 339

A REVIEW OF THE GENUS TIRIDATES STAL
(HETEROPTERA: PENTATOMOIDEA: SCUTELLERIDAE)

J. E. EGER, JR.
Project Leader, Agricultural Products Department
Dow Chemical U.S.A., 5100 West Kennedy Blvd., Suite 450
Tampa, FL 33609 USA
and
Research Associate, Florida State Collection of Arthropods
Florida Department of Agriculture & Consumer Service
Gainesville, FL 32602 USA

ABSTRACT

The genus Tiridates Stal is redescribed and compared to Agonosoma Laporte. The
two included species and a new subspecies, T. rubrocinctus schaffneri, are described,
figured, and keyed. Tiridates rubrocinctus sexmaculata Kirkaldy, T. rubrocinctus de-
color Kirkaldy, T. rubrocinctus decemmaculata Kirkaldy, and T. rubrocinctus mag-
nifica Kirkaldy, names proposed for StAl's (1862) varieties a-d, are placed in the
synonymy of T. rubrocinctus rubrocinctus (Herrich-Schaeffer). Lectotypes and paralec-
totypes are designated for T. rubrocinctus sexmaculata and T. rubrocinctus decolor
and a lectotype is designated for T. rubrocinctus decemmaculata.

RESUME

El g6nero Tiridates Stall se redescribe y compare con Agonosoma Laporte. Se
redescriben, figuran, y se le dan claves a las dos species incluidas y a la nueva subes-
pecie T. rubrocinctus. Tiridates rubrocinctus decemmaculata Kirkaldy, T. rubrocinctus
decolor Kirkaldy, T. rubrocinctus decemmaculata Kirkaldy, y T. rurocinctus magnifica
Kirkaldy, ombres propuestos para las variedades de a-d de Stal (1862), se ponen de
sin6nimos de T. rubrocinctus rubrocinctus (Herrich-Schaeffer). Se designan lectotipos
y paralectotipos para T. rurocinctus sexmaculata y T. rubrocinctus decolor, y un lec-
totipo es designado para T. rubrocinctus decemmaculata.


Among scutellerid genera in the subfamily Pachycorinae, Tiridates Stal, 1867, is
unusual in having both male and female external genitalia almost entirely concealed by
the sixth visible abdominal sternite. Certain species in several other genera of Neotrop-
ical Pachycorinae also have concealed male genitalia (Agonosoma Laporte, 1832; Crathis
StAl, 1861; Diolcus Mayr, 1864; Lobothyreus Mayr, 1864; and Symphylus Dallas, 1851).
Of these, only Agonosoma is similar to Tiridates in size (greater than 10 mm long) and
elongate oval shape. The sixth visible abdominal sternite also conceals the bulk of the
female genital plates in Agonosoma, but leaves more of the 8th and 9th paratergites
exposed than in Tiridates (Figs. 1, 4).
Several characters may be used to separate Tiridates from Agonosoma. The length
of the ostiolar rugae in Agonosoma is greater than 1/2 the distance from the ostiole to
the lateral margin of the metapleuron (Fig. 5). In Tiridates the rugae extend at most
1/3 of the distance from ostiole to lateral margin of the metapleuron (Fig. 2). The shape
of these rugae is also distinctive. In addition, the head of Tiridates is narrower and the
body more convex than in Agonosoma (Figs. 3, 6).
StAl (1867) proposed the genus Tiridates to include two species previously described
by Herrich-Schaeffer (1837), Pachycoris rubrocinctus and P. flavicinctus. A third











Florida Entomologist 70(3)


ti


Figs. 1-6. 1-3. Tiridates rubrocinctus rubrocinctus. 1.) Female abdomen, ventral
aspect. 2.) Meso- and metapleuron. 3.) Head. 4-6. Agonosoma trivittata (Panzer). 4.)
Female abdomen, ventral aspect. 5.) Meso- and metapleuron. 6.) Head. Dimensional
lines equal 1.0 mm.



species, T. mexicanus, also described by Herrich-Schaeffer (1837), has been considered
a senior synonym of T. flavicinctus by some authors. Because color patterns vary
considerably in this genus, several subspecific names persist for variously patterned
specimens of T. rubrocinctus.
This study was undertaken to characterize the genus and to clarify the status of
included taxa. All measurements are given in millimeters.


340


September, 1987











Eger: Review of Tiridates


Tiridates Stal 1867

Tiridates Stal, 1867, p. 494 (keyed, Pachycoris rubrocinctus and P. flavicinctus in-
cluded); Stal, 1870, p. 13-14; Distant, 1880, p. 18 (described, distribution); Schoute-
den, 1904, pp. 68-69 (described, keyed); Kirkaldy, 1909, p. 278, 365.

Medium-large (10.8-14.8 long), elongate oval, strongly convex above and below; dor-
sal surface glabrous, punctation fine, moderately dense, most dense on head and an-
terior border of pronotum. Ventral punctation dense, coarse on head and pronotum,
sparse medially on abdomen, becoming more dense laterally.
Head: convex, moderately declivent, gradually narrowing apically, apex narrowly
rounded; lateral margins of juga sinuous, borders rounded. Tylus slightly surpassing
juga; first antennal segment not reaching apex of head. First rostral segment longer
than bucculae. Bucculae abruptly curving dorsad at posterior limit, rounded anteriorly.
Antennae 5-segmented, arising midway down lateral surface of head and just anterior
of eyes, terminal two segments not distinctly flattened.
Thorax: anterior, anterolateral and posterior margins of pronotum slightly concave,
posterolateral margins convex; small tooth present at each anterolateral angle; an-
terolateral margins rounded dorsoventrally, not compressed or carinate. Pronotum
evenly rounded dorsally in lateral view. Prostethus entire. Tibiae sulcate. Ostiole aris-
ing nearer to metacoxae than to lateral margin of metapleuron. Ostiolar rugae auricu-
late, extending 1/3 or less of distance from ostiole to lateral margin of metapleuron,
roughly 3 to 5 times as long as width of ostiole (Fig. 2). All sterna sulcate, rounded
laterally, not distinctly carinate. Scutellum constricted basally; basal 1/2 to 1/3 of lateral
margins of hemelytra exposed.
Abdomen: lateral margins of connexiva evenly rounded. Abdominal sternites with
slight lateral submarginal impression; small rounded projection present laterally at pos-
terior extremity of each segment. Striated area present on third to fifth visible abdom-
inal sternites.
Pygophore concealed by sixth visible abdominal sternite, caudal face with distinct
projection in lateral aspect (Fig. 12); each lateral wall of pygophore with bifurcate
process, thecal plate attached to anterior arm of process (Fig. 11). Parameres hook-
shaped (Fig. 10). Theca with two elongate sclerotized processes; vesica elongate, finely
spiculate (Fig. 8).
Sixth visible abdominal sternite expanded to conceal female genital plates, posterior
margins of eighth and ninth paratergites narrowly exposed (Fig. 1). First gonocoxae
bilobed, sclerotized posterior lobe narrow, elongate (Fig. 21); anterior lobe membranous
with elongate sclerotized band along posterior margin. Ninth paratergites thin, moder-
ately elongate. Genital chamber with sclerotized groove and two large sclerotized plates
on floor of genital chamber. Spermathecal duct with large spherical dilation; pump
distinct, with well defined proximal and distal flanges; spermathecal bulb spherical,
connected to pump by long stout duct (Fig. 26).
Type Species: Pachycoris rubrocinctus Herrich-Schaeffer, 1837, by subsequent designa-
tion (Schouteden 1904). Kirkaldy (1909) also listed P. rubrocinctus as the type species
(p. 278), but subsequently (p. 365) designated Pachycoris flavicinctus Herrich-Schaef-
fer, 1837, as the type species. Kirkaldy (1909 p. XXXIV) believed that P. flavicinctus
should be the type species because Stal (1870) doubted the specific distinction of P.
rubrocinctus. However, at the time Stal described Tiridates (1867), both P. rub-
rocinctus and P. flavicinctus were included and both are available to be the type
species. Schouteden's designation of P. rubrocinctus was the first valid designation
making P. rubrocinctus the type species of the genus.
Distribution: Tiridates ranges from northern Mexico (Tamaulipas), south to Guatemala










Florida Entomologist 70(3)


9
J1


13
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September, 1987











Eger: Review of Tiridates


and Nicaragua. It is generally uncommon in collections, most of the material examined
being from Mexico.
Types: Dr. Martin Baehr of the Zoologische Staatssammlung, Munich has informed me
that the Herrich-Schaeffer types for Pachycoris flavicinctus, P. rubrocinctus, and P.
mexicanus could not be located and were probably destroyed by war action.
Comments: The female genitalia are unusual in Tiridates, having bilobed first gonocoxae
and sclerotized plates on the floor of the genital chamber. Bilobed first gonocoxae are
also found in Agonosoma (Scudder 1959).
Specimens of Tiridates exhibit considerable intra-specific variation in color patterns
and few useful taxonomic characters other than the genitalia. Some color patterns were
used in the following key and appeared to be consistent. However, dissection of genitalia
is recommended for definitive determination of species. The male aedeagus is diagnostic.
The sclerotized plates on the floor of the female genital chamber are somewhat variable,
but are usually useful for recognition of species.
Virtually nothing is known of the biology of species of Tiridates.

Key to species of Tiridates

1. Abdominal venter predominately dark brown to black, usually with basal
yellow to red macula, and scutellum bordered laterally and posteriorly by
continuous yellow to red vitta; thecal processes acute apically (Fig 8) ........... 2.
1'. Abdominal venter predominately yellow to red, with dark brown to black
transverse vitta present basally and longitudinal vitta present laterally; or
venter black and scutellum with lateral marginal vittae incomplete; apex
of each thecal process hatchet-shaped (Fig. 16) ...................... T. mexicanus.
2. Thecal lobes short, not encircling vesica (Fig. 8); Mexico, north and west of
the Isthmus of Tehuantepec ........................... T. rubrocinctus rubrocinctus.
2'. Thecal lobes long, encircling vesica (Fig. 13); Mexico, south and east of the
Isthmus of Tehuantepec into Nicaragua ................ T. rubrocinctus schaffneri.

Tiridates rubrocinctus rubrocinctus (Herrich-Schaeffer, 1837)

Pachycoris rubrocinctus Herrich-Schaeffer, 1837, p. 9, fig. 352; Germar, 1839, p. 103.
Agonosoma rubrocinctum: Dohrn, 1859, p. 4; Stal, 1862, p. 83 (vars. a-d described).
Agonosoma rubrocincta: Walker, 1867, p. 60.
Tiridates rubrocinctus: Stal, 1867, p. 494; Stal, 1870, p. 14; Distant, 1880, p. 19
(synonymy); Uhler, 1886, p. 2 (synonymy); Distant, 1889, p. 313; Lethierry and
Severin, 1893, p. 35 (synonymy); Schouteden, 1904, p. 69 (synonymy); Kirkaldy,
1909, p. 278.
Tiridates rubrocinctus decemmaculata Kirkaldy, 1909, p. 278 (=var. c Stil, 1862),
NEW SYNONYMY.




Figs. 7-14. 7-12. Tiridates rubrocinctus rubrocinctus. 7.) Aedeagus, dorsal aspect;
vesica (V); thecal process (TP); thecal lobe (TL). 8.) Aedeagus, lateral aspect. 9.) Proc-
tiger, caudal aspect. 10.) Right paramere, lateral aspect. 11.) Genital cup, caudal aspect,
omitting parameres and proctiger; anterior lobe of pygophoral process (A); posterior
lobe of pygophoral process (P). 12.) Genital cup, caudal margin, lateral aspect. 13-14.
Tiridates rubrocinctus schaffneri. 13.) Aedeagus, lateral aspect. 14.) Right paramere,
lateral aspect. Dimensional lines equal 1.0 mm.


343











Florida Entomologist 70(3)


Tiridates rubrocinctus decolor Kirkaldy, 1909, p. 278 (=var. b StAl, 1862), NEW
SYNONYMY.
Tiridates rubrocinctus magnifica Kirkaldy, 1909, p. 278 (=var. d Stal, 1862), NEW
SYNONYMY.
Tiridates rubrocinctus sexmaculata Kirkaldy, 1909, p. 278 (=var. a StAl, 1862), NEW
SYNONYMY.

Length of body 11.7-14.6. Head: broader than long (3.6-4.4 wide, 3.2 4.0 long),
entirely black or dark brown to broadly orange or red bordered. Venter of head entirely
black. Antennae dark brown to black; length of segments 1-5: 0.8-1.0; 0.6-0.8; 0.8-1.0;
1.3-1.7; 1.9-2.2. Rostrum dark brown to black, reaching 2nd to 3rd visible abdominal
sternite.
Thorax: pronotum 3.5-4.7 long, 6.5-8.4 wide; typically black or dark brown with
anterior, antero- and postero-lateral margins broadly yellow to red bordered and with
median longitudinal yellow to red vitta. Colored borders and median vitta occasionally
broken or absent, or expanded to cover entire pronotum. Dark brown maculae fre-
quently present in colored borders; these occasionally reduced or absent. Thoracic ven-
ter typically dark brown to black with orange to red macula occasionally present later-
ally on each metapleuron; coxae and small portion of adjacent pleura occasionally red or
yellow. Legs dark brown to black, occasionally with red to yellow band on femora.
Ostiolar rugae 0.6-0.9 long; distance from ostiole to metacoxae 1.2-1.7, distance from
ostiole to anterolateral corner of metapleuron 1.9-2.4. Scutellum 6.8-8.4 long, 6.5-8.5
wide; typically bordered on all sides by yellow to red vittae and with median transverse
yellow to red vitta; median longitudinal vitta sometimes also present. Vittae frequently
reduced, broken, or lacking, or vittae expanded to cover all of scutellum except 1 or 2
pairs of median black maculae.
Abdomen: venter dark brown to black, usually with red to yellow macula mesially
on 2nd to 5th visible sternites (rarely extending onto 6th sternite); this macula fre-
quently on fewer segments and occasionally lacking; broken lateral submarginal band
of poorly defined red to yellow maculae infrequently present. Mid-ventral length of
visible abdominal sternites 2-6: males 0.7-0.9, 0.5-0.6, 0.5-0.6, 0.6-0.7, 3.0-3.5; females
0.8-1.0, 0.7-0.8, 0.7-0.8, 0.7-0.8, 2.9-3.2.
Apex of posterior arm of pygophoral process truncate, apex of anterior arm rounded
(Fig. 11). Parameres and proctiger as in Figs. 9 and 10. Thecal processes elongate,
apices acute (Figs. 7, 8). Thecal lobes broadly rounded at apex, not encircling vesica.
Vesica thin, abruptly curving ventrad.
Posterior lobe of each first gonocoxa directed mesad; anterior lobe with thin, weakly
sclerotized band along posterior margin (Fig. 21). Plates on floor of genital chamber
moderately sclerotized, relatively flat; widely separated mesially (Fig. 23).
TYPES. The type of Pachycoris rubrocinctus is apparently lost. StAl (1862) described
four varieties of T. rubrocinctus as vars. a-d. Kirkaldy (1909) subsequently named each
variety. Thus, StAl's specimens are the type specimens for Kirkaldy's varieties (sub-
species) of T. rubrocinctus. Specimens of T. rubrocinctus from StAl's collection were
located in the Naturhistoriska Riksmuseet in Stockholm. StAl did not label his vars. a-d,




Figs. 15-20. Tiridates mexicanus. 15.) Aedeagus, dorsal aspect; vesica (V); thecal
process (TP). 16.) Aedeagus, lateral aspect. 17.) Proctiger, caudal aspect. 18.) Right
paramere, lateral aspect. 19.) Genital cup, caudal aspect, omitting parameres and proc-
tiger; anterior lobe of pygophoral process (A); posterior lobe of pygophoral process (P).
20.) Genital cup, caudal margin, lateral aspect. Dimensional lines equal 1.0 mm.


344


September, 1987







Eger: Review of Tiridates


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18


19


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Florida Entomologist 70(3)


/
Figs. 21-22. Female genital plates; first gonocoxae (Gxl); second gonocoxae (Gx2);
eighth paratergites (Pt8); ninth paratergites (pt9); sclerotized plates on floor of genital
chamber (SP); 21.) Tiridates rubrocinctus rubrocinctus. 22.) T. mexicanus.



but specimens agreeing with the descriptions of three of the four varieties were located.
Lectotypes and paralectotypes are designated as follows:

Tiridates rubrocinctus sexmaculata Kirkaldy
LECTOTYPE: Male, labelled: (a) "Mexico." (b) "Boucard." (c) "415, 86." (d)
"Riksmuseum, Stockholm".
PARALECTOTYPES: 2 females: (a) "Mexico." (b) "Salle." (c) "413, 86." (d)
"Riksmuseum, Stockholm"; and (a) "Mexico." (b) "Sall6." (c) "414, 86." (d)
"Riksmuseum, Stockholm".
Tiridates rubrocinctus decolor Kirkaldy
LECTOTYPE: Male, Labelled: (a) "Mexico." (b) "Tiridates rubrocinctus H.S." (c)
"417, 86." (d) "Riksmuseum, Stockholm".


September, 1987











Eger: Review of Tiridates 347




23

/ \ 7 1 ^-/--J (


SG



8 25







26 OF
Figs. 23-26. 23-25. Sclerotized plates in floor of genital chamber. 23.) T. rubrocinctus
rubrocinctus. 24.) T. rubrocinctus schaffneri. 25.) T. mexicanus. 26.) T. rubrocinctus
rubrocinctus, spermatheca and related structures; sclerotized groove in floor of genital
chamber (SG); dilation of spermathecal duct (D); proximal flange of spermathecal pump
(PF); distal flange of spermathecal pump (DF); spermathecal bulb (B). Dimensional
lines equal 1.0 mm.

PARALECTOTYPE: Female: (a) "Mexico." (b) "StAl." (c) "412, 86." (d)
"Riksmuseum, Stockholm".
Tiridates rubrocinctus decemmaculata Kirkaldy
LECTOTYPE: Female, labelled: (a) "Mexico." (b) "Stal." (c) "416, 86." (d)
"Riksmuseum, Stockholm". Apparently no paralectotypes.
Tiridates rubrocinctus magnifica Kirkaldy
No specimens examined agreed with Stal's description of this, var. d, although
it probably belongs in the synonymy of the nominate subspecies as did Stal's
other varieties. Specimens on which this variety was based are apparently not
extant.

Distribution: Mexico, states of Veracruz, Oaxaca, Guanajuato, and Tamaulipas.
Comments: Although there is considerable variation in the color patterns in this and
other species of Tiridates, all variations are apparently built on a typical pattern of
marginal, median longitudinal and median transverse vittae. These vittae may be re-
duced or absent, or expanded to cover virtually all of the dorsum.


Tiridates rubrocinctus schaffneri n. ssp.

Length of body 10.8-14.6. Length of head 2.9-3.8, width 3.5-4.4. Length of antennal
segments 1-5: 0.8-1.0, 0.6-0.9, 0.6-1.0, 1.2-1.6, 1.7-2.3. Pronotum 3.1-4.4 long, 6.0-8.2
wide. Scutellum 6.2-8.5 long, 5.8-8.3 wide. Mid-ventral length of visible abdominal ster-
nites 2-6: males 0.7-0.8, 0.5-0.6, 0.5-0.6, 0.6-0.7, 2.4-3.0; female 0.8, 0.7-0.8, 0.7-0.9,
0.7-0.8, 2.5-3.0.











Florida Entomologist 70(3)


Coloration as in T. rubrocinctus rubrocinctus. Head: more commonly entirely dark
brown to black, or red to yellow with dark markings restricted to small basal macula
and apex.
Abdomen: parameres more robust than in nominate subspecies, shank shorter (Fig.
14). Thecal lobes elongate apically, encircling vesica (Fig. 13). Plates on floor of genital
chamber moderately sclerotized; anterior margins curving ventrad, inner margins ar-
cuate; mesial separation narrow (Fig. 24).
TYPES.
HOLOTYPE: Male, labelled (a) "Valladolid, Yucatan. Gaumer." (b) "Distant Coll.; 1911-
383". Deposited in the British Museum of Natural History (BMNH).
PARATYPES: 3 males and 4 females: (a) "Temax, N. YucatAn, Gaumer." (b) "Distant
Coll.; 1911-383". (male, deposited in the BMNH); (a) "Chontales, Nicaragua, Janson"
(b) "Distant Coll. 1911-383". (male, deposited in the BMNH); (a) "HONDURAS, Cop.
3 mi. NE. Cucuyagua 3000', VII-25-1974, C. W. & L. O'Brien & Marshall" (b) "B.M.
1975-493". (male, deposited in the BMNH); "MEXICO: Yucatan; 13.3 mi. s. Valladolid;
July 30, 1980; Schaffner, Weaver, Friedlander". (female, deposited in the Texas A&M
University collection); "MEXICO: Quintana Roo; Nueva, X-Can; 17-VIII-1978; E. C.
Welling, coll." (female, deposited in the author's collection); "Coll. R. I. Sc. N. B.;
Mexique; Merida; 27-VII-1959; H. J. Bredo". (female, deposited in the Institut Royal
des Sciences Naturelles de Belgique); (a) "Chiquimula, Guatemala. Dec. 6, 1930, J. J.
White" (b) "J. C. Lutz Collection 1961" (c) "Tiridates decemmaculata Kirkaldy" (female,
deposited in the United States National Museum of Natural History).
Distribution: Mexican states of Yucatan and Quintana Roo south through Guatemala
and Honduras to Nicaragua.
Comment: The distribution of this and the nominate subspecies appears to be distinct,
being separated by the Isthmus of Tehuantepec. Genitalic differences are relatively
slight but consistent, suggesting that the two taxa may be distinct species. The lack of
other characters for separating the two and the allopatric distribution suggest sub-
specific status, however.
This subspecies is dedicated to Dr. J. C. Schaffner of Texas A&M University for his
contributions to the knowledge of Mexican Heteroptera and for the help and support
he has given me.

Tiridates mexicanus (Herrich-Schaeffer, 1837)

Pachycoris mexicanus Herrich-Schaeffer, 1837, p. 3, fig. 343; Germar, 1839, p. 89.
Pachycoris flavicinctus Herrich-Schaeffer, 1837, p. 8, fig. 351; Germar, 1839, p. 103.
Agonosoma mexicanum: Dohrn, 1859, p. 4.
Agonosoma flavocinctum: Dohrn, 1859, p. 4.
Agonosoma mexicana: Walker, 1867, p. 60.
Agonosomaflavicincta: Walker, 1867, p. 60.
Tiridatesflavocinctus: StAl, 1867, p. 494.
Tiridates mexicanus: Stal, 1870, pp. 13-14 (P. flavicinctus listed as junior synonym);
Distant, 1880, p. 19 (synonymy); Uhler, 1886, p. 2 (synonymy); Distant, 1889, p. 313;
Lethierry and Severin, 1893, p. 35 (synonymy); Schouteden, 1904, p. 69, pl. 4, fig.
13 (synonymy); Kirkaldy, 1909, p. 278.
T7",,,ha -flavicinctus: Kirkaldy, 1909, p. 278, 365.

Length of body 12.5-14.8. Head: 3.2-3.6 long, 3.8-4.4 wide, dark brown to black with
red to yellow markings usually present at least laterally and frequently most of dorsal
surface red to yellow. Antennae dark brown to black, first 1 or 2 segments occasionally
yellow to red; length of segments 1-5: 0.8-1.0; 0.6-0.8; 0.7-0.9; 1.4-1.6; 1.8-2.3. Rostrum


348


September, 1987











Eger: Review of Tiridates 349

brown to black, segments 1-2 frequently yellow to red; apex reaching 2nd to 3rd visible
abdominal sternite.
Thorax: pronotum broader than long (6.8-8.2 wide, 4.1-4.9 long), dark brown to
black with yellow to orange vittae along anterior, antero-, and posterolateral margins
and with median longitudinal vitta; vittae frequently reduced or expanded to cover
entire pronotum. Dark brown submarginal maculae frequently present in marginal vit-
tae. Thoracic venter typically dark brown to black, except following red to yellow
structures: coxae and large macula on adjacent pleura, lateral margins of pro-, meta-,
and occasionally mesopleura, margin of prostethus, posterior border of metapleura and
portions of evaporative area on metapleura. Legs dark brown to black except basal 2/3
to 5/6 of femora usually orange to red and tibiae infrequently with yellow or red median
band. Ostiolar rugae 0.4-0.7 long; distance from ostiole to metacoxae 1.3-1.6, distance
from ostiole to anterolateral corner of metapleuron 1.8-2.3. Scutellum 7.4-8.8 long, 6.7-
7.9 wide; typically bordered on all sides by broad yellow vittae and with median trans-
verse vitta, vittae frequently reduced or absent or expanded to cover nearly entire
scutellum. Fovea in each basal angle of scutellum and 2 small maculae on anterior
border dark brown to black, maculae absent in specimens lacking colored vitta on an-
terior margin.
Abdomen: venter orange to red with black markings as follows: basal transverse
vitta on first to third visible sternites, broad lateral longitudinal vitta (this vitta fre-
quently broken into series of maculae), broadly oval macula near posterior limit of last
sternite, 3 to 6 smaller maculae on sternites 4 to 6. Abdominal venter rarely entirely
dark brown to black or with basal red to yellow macula on 2nd to 5th visible segments.
Mid-ventral length of visible abdominal sternites 2-6: males 0.7-0.8, 0.7-0.9, 0.7-0.9,
0.6-0.7, 3.0-3.3; female 0.8-1.0, 0.9, 0.8-1.0, 0.8-0.9, 2.6-2.9.
Apices of both arms of each pygophoral process narrowly rounded (Fig. 19). Para-
meres and proctiger as in Figs. 17 and 18. Theca relatively short, without distinct lobes
(Figs. 15, 16). Thecal processes broad basally from dorsal aspect, narrowing apically to
hatchet-shaped apices. Vesica relatively broad, angled ventrad and curving caudad.
Posterior lobe of each first gonocoxa directed mesad and cephalad (Fig. 22); anterior
lobe with relatively broad, moderately sclerotized band along posterior margin. Plates
on floor of genital chamber strongly sclerotized, broadly concave, anterior and mesial
margins enlarged; narrowly separated mesially (Fig. 25).
Types: Apparently lost.
Distribution: South Mexico, states of Chiapas, Guerrero, Jalisco, Michoacan, Morelos,
Nayarit, Oaxaca, and Veracruz.
Comments: There appear to be 3 color morphs of T. mexicanus, all of which are appa-
rently sympatric and all of which have been collected together in series. Two of these
morphs were figured by Herrich-Schaeffer (1837) and described as Pachycoris
flavicinctus and P. mexicanus. I've seen only males of the former and both sexes of
the latter. A third morph is almost entirely black ventrally and is black dorsally with
red markings as follows: lateral band on head, anterior margin of pronotum, vittae along
lateral margins of pronotum and scutellum (these relatively thin and frequently broken),
submarginal vittae along lateral margins of pronotum, median longitudinal vitta on
pronotum, and median transverse vitta on scutellum. Specimens of this morph consisted
entirely of females.
Two small black maculae located just mesad of the fovea on the anterior margin of
the scutellum were always present in specimens of the first two morphs (the anterior
margin of the third morph was entirely black). No such maculae were present in speci-
mens of T. rubrocinctus.











Florida Entomologist 70(3)


ACKNOWLEDGEMENTS

I am indebted to the following individuals and institutions for the loan of material:
H. A. Brailovsky (Instituto de Biologia, Universidad Nacional Autonoma de Mexico);
P. Dessart (Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium); W.
R. Dolling (British Museum (Natural History), London, England); R. C. Froeschner
(United States National Museum of Natural History, Washington, D. C.); P. Lindskog
(Naturhistoriska Riksmuseet, Stockholm, Sweden); L. H. Rolston (Louisiana State Uni-
versity, Baton Rouge, Louisiana); J. C. Schaffner (Texas A&M University, College
Station, Texas); G. Schmitz (Mus6e Royal de 1'Afrique Central and Institut Royal des
Sciences Naturelles de Belgique, Tervuren, Belgium); and D. B. Thomas (United States
Department of Agriculture, Agricultural Research Service, Chiapas, Mexico). I also
thank F. J. D. McDonald (University of Sydney, Sydney, Australia) for comments on
the female genitalia, M. Baehr (Zoologische Staatssammlung, Munich) for the informa-
tion on Herrich-Schaeffer's types, and R. S. Peigler (Greenville, South Carolina), L. H.
Rolston, and J. E. McPherson (Southern Illinois University at Carbondale, Carbondale,
Illinois) for their comments and suggestions on the manuscript.

REFERENCES CITED

DISTANT, W. L. 1880-1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In: God-
man, F. D. and 0. Salvin, [Eds.]. Biologia Centrali-Americana. London, Vol. 1.
xx + 462 pp., 39 pls.
DOHRN, A. 1859. Catalogus Hemipterorum. Herrcke and Lebeling, Stettin, 112 pp.
GERMAR, E. H. 1839. Beitrage zur einer Monographie der Schildwanzen. Zeits. En-
tomol. 1: 1-146, 1 pl.
HERRICH-SCHAEFFER, G. A. W. 1837. Die Wanzenartigen Insekten. 4: 1-32.
KIRKALDY, G. W. 1909. Catalogue of the Hemiptera (Heteroptera). Vol. I. Cimicidae.
Berlin, xl + 392 pp.
LETHIERRY, L., AND G. SEVERIN. 1893. Catalogue general des Hemipteres-Pen-
tatomidae, 1. Bruxelles, x + 286 pp.
SCHOUTEDEN, H. 1904. Heteroptera. Fam. Pentatomidae. Subfam. Scutellerinae.
Wyts. Gen. Ins., fasc. 24, 98 pp., 5 pls.
SCUDDER, G. G. E. 1959. The female genitalia of the Heteroptera: morphology and
bearing on classification. Trans. Roy. Entomol. Soc. London 111: 405-467.
STAL, C. 1862. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stettin.
Entomol. Ztg. 23(1-2): 81-118, 273-281, 289-325, 437-462.
STAL, C. 1867. Bidrag till Hemipterernas systematik. Ofv. Kongl. Svenska Vetens.-
Akad. Forh. 24(7): 491-560.
STAL, C. 1870. Enumeratio Hemipterorum. Bidrag till en foreteckning 6fver alla hit-
tils kinda Hemiptera, jemte systematiska meddelanden. 1. Kongl. Svenska Vet-
ens.-Akad. Handl. 9(1): 1-232.
UHLER, P. R. 1886. Check-list of the Hemiptera-Heteroptera of North America.
Brooklyn Entomol. Soc., i + 32 pp.
WALKER, F. 1867. Catalogue of the specimens of Hemiptera: Heteroptera in the
collection of the British Museum. London. Part 1, pp. 1-240.


350


September, 1987











Schuster & Taylor: Abamectin on Tomato


RESIDUAL ACTIVITY OF ABAMECTIN AGAINST
LIRIOMYZA TRIFOLII (DIPTERA: AGROMYZIDAE)

D. J. SCHUSTER
Gulf Coast Research & Education Center
University of Florida
Bradenton, Florida 34203
AND
J. L. TAYLOR
Merck Sharp & Dohme Res. Labs.
P.O. Box 1893
Sanford, FL 32771

ABSTRACT

Abamectin was applied to tomato in the field and treated foliage was bioassayed in
the laboratory 0, 1, 3, 5 and 7 days after treatment for affects on Liriomyza trifolii
(Burgess). Surface residues on leaflets increased mortality of adults only when bioas-
sayed on the day of treatment. Oviposition and larval abundance were reduced even
when leaflets were bioassayed 7 days after treatment. Leaf puncturing (stippling) by
females was reduced for 3 days after treatment.

RESUME

Se aplico abamectin a tomatoes en el campo y se efectuaron bioensayos del follaje en
el laboratorio despues de 0, 1, 3, 5, y 7 dias de aplicaci6n para determinar sus efectos
en Liriomyza trifolii (Burgess). Los residues en la superficie de las hojas incrementaron
la mortalidad de los adults s6lo cuando se ensayaron en el dia en que el tratamiento
fue hecho. La oviposici6n y abundancia larvaria fueron reducidas afin cuando se en-
sayaron las hojas 7 dias despues de haber sido tratadas. El nimero de perforaciones en
las hojas hechas por las hembras fue reducido por 3 dias despu6s de la aplicaci6n del
insecticide.


Liriomyza trifolii (Burgess) is a pest of many vegetable and flower crops grown in
Florida. Damage is inflicted on foliage by leaf puncturing (stippling) by females and by
leafmining by larvae. Stipples may be utilized for oviposition or feeding. Females appa-
rently feed on exudates from stipples while males do not (Zoebisch & Schuster 1987).
Abamectin (MK 936) is an 80:20 mixture of avermectin Bla and Bib. Foliar applica-
tions of abamectin have resulted in less leafmining by L. trifolii on tomato in the field
(Schuster & Everett 1983). In the laboratory, abamectin induced adult and larval mor-
tality, inhibited oviposition and feeding and reduced egg hatch (Schuster & Everett
1983, Schuster & Taylor 1987).
The purpose of the present study was to evaluate the residual activity of abamectin
against L. trifolii on tomato.

MATERIALS AND METHODS

The experiment was conducted at the Gulf Coast Research & Education Center,
Bradenton, Fla. Plots consisted of single 4.6 m rows of 10 tomato plants cv. 'Sunny' set
5 Sept. 1984 on 20 cm high by 1.4 m wide beds of EauGallie fine sand covered with
black polyethylene plastic film. Prior to installing the plastic film, the beds were fumi-











Florida Entomologist 70(3)


gated with 335 kg/ha of a 98:2 mixture of methyl bromide and chloropicrin. Plots were
arranged in randomized complete blocks with four replications. Water was provided by
seepage irrigation. On 3 Oct., the fourth fully expanded leaf from the tops of at least
25 main or lateral stems in each plot were tagged and water or an abamectin preparation
(MK 936, 0.15 emulsifiable concentrate, 4.54 gm AI/378.5 liters) was applied at 748
liters/ha. When the foliage had dried and 1, 3, 5 and 7 days thereafter, 5 terminal
leaflets were excised from tagged leaves from each plot. The cut end of each petiole
was pushed through a small hole in parafilm stretched over the mouth of a water-filled
vial. Forty cylindrical cages were constructed from inverted 1-liter plastic containers
with screw caps. The side of each container had two 6-cm-diam holes covered with
organdy fabric. A single vial with a leaflet was attached to the bottom of each container
with modelling clay so that the leaflet would not touch the side of the container. The
tops were then attached and one pair of adults 5 24 h old was introduced into each cage
through small holes in the sides of the cages. After 24 h, surviving adults were transfer-
red to 30 ml plastic cups with a honey streak and mortality was observed 24 h later.
The leaflets were examined for the numbers of stipples and eggs and were held for an
additional 48 h to determine egg hatch. The leaflets were examined again 24 h later to
determine larval mortality. Mortality data were compared for significance by calculating
and squaring 'z' to obtain x2 (Fleiss 1981). All other data were analyzed by ANOVA
(SAS Institute 1982).

RESULTS AND DISCUSSION

The weather during the week-long sampling period was normal for the time of year.
The respective maximum and minimum temperatures (C) were as follows: 3 Oct., 26.7
& 14.5; 4 Oct., 27.8 & 15.0; 5 Oct., 28.4 & 16.1; 6 Oct., 28.9 & 16.1; 7 Oct., 29.5 & 17.2;
8 Oct., 30.0 & 17.2; 9 Oct., 30.0 & 17.2; and 10 Oct., 27.8 & 18.4. A trace of precipitation
was received on 9 & 10 Oct.


TABLE 1. PERCENT MORTALITY OF LIRIOMYZA TRIFOLII ADULTS CONFINED ON
LEAFLETS EXCISED FROM TOMATO PLANTS 0-7 DAYS AFTER THEY HAD
BEEN SPRAYED IN THE FIELD WITH EITHER AN ABAMECTIN PREPARA-
TION OR WATER. AFTER 24 H, THE ADULTS WERE TRANSFERRED TO
PLASTIC CUPS WITH HONEY FOR AN ADDITIONAL 24 H.

2411 48H
Evaluation 2
Treatment time Females Males Females Males

Abamectin Day 0 5.3aa 22.2a 63.2a 44.4ab
Water 0.Oa 5.3a 0.Oc 15.8bc
Abamectin Day 1 0.Oa 5.6a 5.3a 16.7a
Water 0.Oa 10.0a 0.Oa 15.0a
Abamectin Day 3 5.0a 0.Oa 10.0a 21.0a
Water 0.Oa 10.5a 0.Oa 10.5a
Ambamectin Day 5 0.Oa 0.Oa 0.Oa 15.0a
Water 5.0a 10.0a 10.0a 15.0a
Abamectin Day 7 0.Oa 5.3a 5.3a 5.3a
Water 0.Oa 0.Oa 0.Oa 0.Oa

aMeans within each time period followed by the same letter are not significantly different (X2; P = 0.05).


September, 1987











Schuster & Taylor: Abamectin on Tomato 353

TABLE 2. THE NUMBERS OF EGGS, FEEDING PUNCTURES STIPPLESS) AND SUB-
SEQUENT LARVAL ABUNDANCE OF LIRIOMYZA TRIFOLII EXPOSED 24 H
TO LEAFLETS EXCISED FROM TOMATO PLANTS 0-7 DAYS AFTER THEY
WERE SPRAYED IN THE FIELD WITH EITHER AN ABAMECTIN PREPARA-
TION OR WATER.

Eva n No./female No. larvae'
Evaluation
Treatment time Eggs Stipples Alive Dead

Abamectin Day 0 0.2ab 17.1a 0.Oa 0.Oa
Water 5.6b 160. lb 4.6b 2.2b
Abamectin Day 1 1.8a 73.6a 0. la 0.4a
Water 5.6a 174.0b 4.7b 1.3a
Abamectin Day 3 0.6a 80.8a 0.Oa 0.4a
Water 4.0b 162.7b 5. b 0.la
Ambamectin Day 5 0.9a 98.6a 0.la 0.Oa
Water 3. la 118. la 7. b 0.4a
Abamectin Day 7 3.9a 110.0a 0.Oa 0.6a
Water 8.0b 137.3a 8.0b 0.9a

aMortality data transformed V/X-+.5 prior to analysis. Means are presented in the original scale.
bMeans within columns for each time period followed by different letters are significantly different (F; P <0.05).


Mortality of adults induced by abamectin residues relative to the check was not
significantly different at any sampling when evaluated after the initial 24 h (Table 1).
When leaflets were bioassayed on the day of spraying, mortality of females 24 h after
the initial 24 h exposure was greater on leaflets sprayed with abamectin than on leaflets
sprayed with water. Mortality of females and males exposed to abamectin-treated leaf-
lets were similar. Abamectin-treated leaflets from the remaining samplings did not
induce significantly higher mortality of either sex relative to water-treated leaflets.
Oviposition and the number of surviving larvae were reduced by abamectin relative to
water even when leaflets were bioassayed 7 days after treatment (Table 2). Stippling
by females was reduced by abamectin until leaflets were bioassayed on the fifth day
following treatment.
These results indicate that surface residues of abamectin rapidly dissipate. Wright
et al. (1985) demonstrated translaminar movement of abamectin when applied to French
bean, cotton and chrysanthemum in the greenhouse. Our results support this finding
with abamectin applied to tomato in the field since the affects associated with surface
residues (adult mortality and stippling) persisted only 1 to 3 days while affects as-
sociated with internal residues (oviposition and larval mortality) persisted at least 7
days. The results further indicate that a single application of abamectin should provide
control of L. trifolii on tomato for at least a week.

ACKNOWLEDGMENT

Florida Agricultural Experiment Stations Journal Series No. 8030.

REFERENCES CITED

FLEISS, J. L. 1981. Statistical methods for rates and proportions, 2nd edition. Wiley
& Sons, Inc., New York, NY. 321 pp.











354 Florida Entomologist 70(3) September, 1987

SAS INSTITUTE. 1982. SAS user's guide: statistics. SAS Institute, Cary, N.C.
SCHUSTER, D. J., AND P. H. EVERETT. 1983. Response of Liriomyza trifolii (Dipt-
era:Agromyzidae) to insecticides on tomato. J. Econ. Entomol. 76: 1170-1174.
SCHUSTER, D. J., AND J. L. TAYLOR. 1987. Longevity and oviposition of adult
Liriomyza trifolii (Diptera:Agromyzidae) exposed to abamectin in the labora-
tory. J. Econ. Entomol. 80: (in press).
ZOEBISCH, T. G., AND D. J. SCHUSTER. 1987. Longevity and fecundity of Liriomyza
trifolii (Diptera:Agromyzidae) exposed to tomato foliage and honeydew in the
laboratory. Environ. Entomol. 16: (in press).
WRIGHT, D. J., A. LOY, A. ST. J. GREEN, AND R. A. DYAS. 1985. The translaminar
activity of abamectin (MK-936) against mites and aphids. Proc. 37th Intl. Symp.
Crop Protection, Ghent, pp. 595-601.


- p p-- -- p0 p pWp


ACOUSTIC AND MATING BEHAVIOR OF A
MEXICAN KATYDID, PTEROPHYLLA BELTRANI
(ORTHOPTERA:TETTIGONIIDAE)

KENNETH C. SHAW AND PATRICK GALLIART
Department of Zoology
Iowa State University
Ames, Iowa 50011

ABSTRACT

Males of the Mexican katydid, Pterophylla beltrani, sing day and night and form
aggregations at the ends of tree branches. Males of other pterophylline species sing
only at night and from singing sites which may be 15 m from nearest singing neighbors.
When singing alone (solo calling), P. beltrani males produce mostly 2-pulse (range 1-5)
phrases (a pulse is the sound produced during the closing phase of a wingstroke cycle).
Phrases are produced at a rate of 78-126/min and the pulse rate of 15-20/sec (25-30C)
is the fastest reported for the Pterophyllini. Acoustic interaction of male pairs (<1 m
apart) consists of soloing (singing two or more phrases prior to response of partner),
synchronizing (overlapping) phrases, and alternation of single or grouped phrases. Dur-
ing alternation, a katydid's phrase rate is slowed compared to the rate of solo and
synchrony, and phrases may be lengthened by one pulse. In contrast, when a katydid
solos during acoustic interaction the solo rate is faster than that of solo calling. When
antennating, or being antennated by, another male or female, P. beltrani males sing at
a rate which is equivalent to that of soloing during acoustic interaction. P. beltrani
males and females produce erratic "disturbance" sounds when handled. The nature of
acoustic interaction is compared with that of Pterophylla camellifolia males (Shaw
1968) and proximate mechanisms suggested, copulatory behavior is described, and pos-
sible mating strategies are discussed.

RESUME

Los machos de Pterophylla beltrani (Tettigonidae:Pseudophyllinae) cantan dia y
noche y forman agregaciones en las terminaciones de las ramas. Los machos de otras
species de Pterophylla cantan solamente de noche y desde sitios que podrian estar a
15 m distantes de los vecinos mas cercanos. Cuando un macho de P. beltrani canta
individualmente, este produce principalmente frases de dos pulsaciones (rango de 1-5)











Shaw & Galliart: Katydid Behavior 355

(la pulsaci6n es el sonido producido al cerrar las alas durante el ciclo complete de aper-
tura y cierre de las mismas). Las frases se produce con una frecuencia de 78-126/min,
siendo la frecuencia de pulsaci6n de 15-20/seg (25-30C) la mis rapida que ha sido repor-
tada para los Pterophyllini. La interacci6n acustica de pares de machos (distanciados a
< 1m) consist en el canto solitario (cantado 2 o mis frases antes que respond su
compafero), el canto de frases sincronizadas (traslapadas) y la alternaci6n de una o
varias frases. Durante la alternaci6n, la frecuencia de las frases es lenta comparada a
la frecuencia del canto solitario o del canto sincronizado, y las frases se pueden alargar
por una pulsaci6n. En contrast, cuando el P. beltrani canta solo durante la interacci6n
acustica la frecuencia de estos cantos solos es mas ripida que la del canto solitario
verdadero. Al tocar o ser tocados por las antenas de otro macho o una hembra, los
machos P. beltrani cantan a una frecuencia equivalent a la del canto solo durante la
interacci6n acistica. Machos y hembras de P. beltrani produce sonidos erraticos
cuando son molestados. La naturaleza de la interacci6n acustica se compare con la del
macho de Pterophylla camellifolia (Shaw 1968) y se sugieren mecanismos similares, se
describe el comportamiento copulatorio y se discuten las posibles estrategias de
apareamiento.


Pterophylla beltrani (Bolivar and C. Bolivar) was first described in 1942 (Bolivar &
Pieltain 1942). The authors suggested morphological similarities with Pterophylla
camellifolia Fabricius and Pterophylla robertsi Hebard but greater affinities with P.
robertsi. P. beltrani adults feed primarily on live oak (Quercus spp.) but females prefe-
rentially oviposit in the woody parts of "la nacahua" (Cordia boissieri A. DC.) and "la
hoja ancha" (Flourensia laurifolia DC.) (Barrientos & Ortega 1985, Barrientos et al.,
unpublished). Bolivar & Pieltain (1942) report P. beltrani in trees of Acer spp. in an
oak forest. The exact geographic distribution of P. beltrani is unknown; however, it has
been reported from the Mexican states of Nuevo Leon and Tamaulipas (Bolivar &
Pieltain 1942, Barrientos et al. 1984).
Little has been reported regarding the acoustic and reproductive behavior of P.
beltrani. According to Barrientos et al. (unpublished), males begin to sing two or three
days following eclosion and produce calling and "territorial or defensive" sounds. The
same authors also report that mating occurs between 0800 and 1000 h, copulation lasts
ca. 35 min, and involves the transfer of an external spermatophore. This paper describes
the courtship, copulation and the acoustical repertoire of P. beltrani including acoustical
interactions between pairs of males.


MATERIALS AND METHODS

Adult male and female specimens of P. beltrani were obtained with the aid of Ms.
Barrientos, Instituto de Investigaciones Alimentarias, Cd. Victoria, Tamaulipas,
Mexico. Males and females of P. beltrani were collected by climbing oak trees, shaking
katydid-bearing limbs, and plucking individuals from vegetation after they had jumped,
spread their tegmina and wings, and glided up to 30 m from their host trees. These
adults, collected on August 8, 1984, near the village of Alta Cumbre, Tamaulipas, and
some eggs and nymphs from Ms. Barrientos' laboratory were transported by car to
Iowa State University.
Eggs, nymphs and adults were maintained in Percival" environmental chambers on
a 12 L:12 D light regime, at temperatures of 290C L and 27'C D, and 40-60% R.H.
Adults and nymphs were provided with oak leaves (Quercus spp.) and water (cotton-
capped vials). Adults living into late fall were given lettuce and chicken mash. During
the summer and fall of 1984, sounds and behavior were recorded, described and analyzed
utilizing adults collected in the field and reared from nymphs. In 1985, adults were











Florida Entomologist 70(3)


reared in the laboratory from eggs brought from Mexico and from eggs laid by pregnant
females collected in Alta Cumbre or by females mated in the laboratory. Behavioral
observations were recorded from varying numbers of katydids placed in 26 x 30 x 52
cm glass terraria. Sounds were recorded from katydids in the terraria and from katydids
maintained in 9 x 10 x 17 cm wire screen cages. All sound recordings and behavioral
observations were made in an acoustic isolation chamber (4.6 x 5.3 x 2.4 m) (Industrial
Acoustics Company, Inc., Bronx, N.Y.). Sounds were recorded using a Panasonic WM-
1150 or a Bruel & Kjaer 1.3 cm microphone and a Sony TC-6300 or a Precision Data,
Inc. PI-6204 tape recorder. Sounds were analyzed using a Kay 7029A Sonograph
(sound spectrograph) and a Tectronix 5110 oscilloscope in conjunction with a Grass
Model C4R kymograph camera.
Thirty-nine different males were used to record the various song types. Males to be
recorded were moved from the environmental chamber to the acoustic chamber. After
a recording session, males were returned to the environmental chamber and placed on
a different shelf from those not yet recorded.
In order to compare singing rates (expressed as phrase periods, Fig. 1) of different
katydids in different contexts recorded at different temperatures, two types of statisti-
cal analyses were utilized. A mixed-effects ANOVA (Snedecor & Cochran 1980) allowed
the separation of variance resulting from differences among katydids (inherent and
temperature induced) from that resulting from differences among response types. LSD
tests (Steel & Torrie 1980) were then used to determine significant differences between
phrase periods of paired combinations of response types (Table 1). The second type of
analysis, analysis of covariance with temperature as the covariant, utilized regression
curves of response type and temperature to create adjusted means which were com-
pared by means of Student's t-test (Tables 2 and 3).

RESULTS

Field Observations

On Aug. 1984, K. Shaw, a graduate student (B. Smith), and L. Barrientos and her
co-worker (T. Reyes) visited a population of P. beltrani near the village of Alta Cumbre.
The katydids were in a montane forest of live oak (Quercus spp.). Males and females
formed large aggregates at the end of branches at or near the top of ca. 5-6 m high
trees. Many males were producing songs consisting of a continuous production of short
phrases (Fig. 1) during a hot (270C shade) afternoon (1300 to 1600 hr). Because of the
number of katydids singing, it was difficult to ascertain the nature of acoustic interac-
tion. P. beltrani males probably also sing at night. The colony was not visited at night
but males have sung at night in the darkened laboratory.

Solo calling

Detailed analyses of song parameters were made from solo calling songs (Fig. 1),
i.e., songs of isolated caged males or terrarium males in which only one male was
singing and other males were stationary 30 cm or more from the singing male. Sonag-
raphic analysis indicated that most of the sound energy of solo calling falls between less
than 1 to approximately 10 kHz (two to three phrases analyzed for five P. beltrani
males). Our analysis showed continuous but decreasing sound energy up to 40 kHz (the
upper frequency range of the microphone) but, as in many other tettigoniids, sound
energy probably occurs up to at least 100 kHz (Elsner & Popov 1978).
The calling sound consists of the repetitive production of 1- to 5-pulse phrases (see
End Note for definiftions) with 2-pulse phrases being, by far, the most common (Fig.


356


September, 1987












357


Shaw & Galliart: Katydid Behavior

PTEROPHYLLA BELTRANI


SOLO CALLING


0 C 0 C




I I I I I I I I I I I I I I I I I
0 0.16 0.32 0.48 0.64
TIME IN SECONDS


COURTSHIP





MALE DISTURBANCE


SI 1 II II I'llllll II II !'i Llr I Ill 111I h1a I


FEMALE DISTURBANCE





I I I I I I I I I I
0 1.6 3.2 4.8 6.4
TIME IN SECONDS

Fig. 1. Songs of P. beltrani. o-one or two sounds produced during wing opening;
c-sounds or pulses produced during wing closure; pl-phrase length; pi-phrase inter-
val. Phrase length + phrase interval = phrase period.


1). Of 53 males for which pulse number was determined, the distribution of modal
phrase pulse numbers was as follows (numbers in parentheses represents production of
phrase pulse numbers other than modal): 1-pulse, n=1; 2(1)-pulse, n=2; 2-pulse, n=39;
2(3)-pulse, n=6; 3(2)-pulse, n=2; 3(4)-pulse, n=2; 4(5)-pulse, n=l.
For nine katydids (9-11 phrases per katydid), mean phrase lengths, intervals and
periods (parameters indicated in Fig. 1) of 2-pulse phrases ranged from 84-117 (x =
105.8 + 25.1) msec, 380-682 (R = 492.5 104.6) msec, and 483-796 (R = 594.9 116.2)
msec respectively, resulting in phrase rates of 1.3-2.1 per sec (R = 1.7 per sec) (25-30C).
The mean length of the first pulse of katydids' 2-pulse phrases was significantly
longer than that of the second pulse (first pulse: 39-57 (R = 48.0 6.1) msec; second












358 Florida Entomologist 70(3) September, 1987

TABLE 1. COMPARISON OF MEAN PHRASE PERIODS (IN MSEC) OF RESPONSE
TYPES OCCURRING DURING ACOUSTIC INTERACTION OF PAIRS OF P.
BELTRANI MALES.

Response type Mean','

solo 346.3 a
overlap 421.5b
alternation 628.4 c
extended delay 1090.6 d

Two-way mixed effects ANOVA
katydid, F = 11.66, df = 21,621; P < 0.0001
response type, F = 298.05, df = 3,621; P < 0.0001

'Means identified by different letters are significantly different (LSD test, P < 0.05).
2Temperature range 25-30C.



TABLE 2. COMPARISON OF MEAN PHRASE PERIODS (IN MSEC) OF SOLOS OCCUR-
RING IN DIFFERENT SONG TYPES OF P. BELTRANI MALES.

Solo type Mean'.2

solo calling 573.3 a
solo after acoustic interaction 419.0 b
solo during acoustic interaction 345.1 b
courtship 325.5 b

Analysis of Covariance
Source df SS MS F

Temperature (covariate)" 1 22875 22875
Solo type 3 309658 103220 11.36 P < 0.01
Error 32 290810 9088

36 623343

'Means adjusted for temperature differences (25-30C).
2Means identified by different letters are significantly different (Student t-tests, P < 0.01).
'The effect of temperature after solo type was entered into the analysis was not quite significant at P < 0.05 (F
=3.08; P = 0.089).



TABLE 3. COMPARISON OF TEMPERATURE-ADJUSTED MEAN PHRASE PERIODS (IN
MSEC) OF SOLO CALLING AND RESPONSE TYPES OF ACOUSTIC INTERAC-
TION FOR P. BELTRANI MALES.

Differences between
temperature-adjusted means F' P

solo calling solo = 219.8 28.38 0.001
solo calling overlap = 143.1 10.92 0.0048
solo calling alternation = -45.4 0.49 0.4979
solo calling extended delay = -559.7 8.90 0.0088

'F-statistic for testing difference between adjusted means in analysis of covariance.


---











Shaw & Galliart: Katydid Behavior


pulse: 23-40 (R = 32.7 6.1) msec; t = 3.75-42.1, p < 0.01-0.001) and the mean interval
between pulses was two-to-four-fold shorter (9-22 (R = 17.2 4.2) msec) than the length
of the initial pulse. Not enough 3-pulse phrases were recorded to compare the magnitude
of the second intervals and their pulses, but oscillographs from song selections of three
3-pulse katydids suggested that the second interval is longer than the first and that the
third pulse is longer than the second.
Although the relation of the cycle of wing movement to sound production was not
analyzed, it is assumed that the most prominent sounds (pulses) are the result of wing
closure as occurs in P. camellifolia (Pierce 1948). In P. camellifolia, a short, relatively
soft sound may occur during the initial opening of the wings prior to the series of wing
movements associated with a multi-pulsed phrase. Although always shorter than pulses
assumed produced during wing closure, the sound level of presumed wing-opening
sounds was quite variable among males of P. beltrani. For some males, wing-opening
sounds were as intense as wing-closing pulses (Fig. 1). In addition, two wing-opening
sounds preceding the longer pulses of wing closure were not uncommon (Fig. 1). All
measurements of pulse and phrase length reported in this paper omitted the sounds
generated by wing opening because their existence and intensity varied among the
calling sounds of different katydids and even within the calling sound of an individual
(Fig. 1). If included, such sounds would increase phrase length by as much as 37 msec.

Acoustic Interaction

Acoustic interactions between pairs of males were recorded from males 0.15 to 1.0
m apart. From the conditions observed in the field, it is unlikely that acoustic interaction
at greater distances is common. To analyze acoustic interactions, varying lengths (12-72
phrases) of 11 paired interactions were taped and oscillographs made at a later date.
Acoustic interactions consisted of a combination of soloing, phrase alternation and
phrase overlap or synchrony. Pairs of katydids alternated single phrases (c, Fig. 1) or
sequences of two or more phrases (= solos) termed extended delay (d, Fig. 2). Alterna-
tion was intermittently interrupted by synchronous (S, Fig. 2) or almost synchronous
(0, Fig. 2) production of phrases.
During the 11 paired interactions, seven of the 22 males produced two different
phrase lengths. Six males produced 2- and 3-pulse phrases: one male produced 3- and
4-pulse phrases. All but one of the 70 (36%) longer phrases followed alternation or
extended delay; shortening of phrase length never occurred following alternation or
extended delay.
For purposes of analysis, the temporal relationships of the phrases of two acousti-
cally interacting males were classified into the following response types (Fig. 2): 1) solo:
one male sang two successive phrases and the partner was silent or the partner's phrase
synchronized with the first male's second phrase (a, Fig. 2): 2) overlap solo: the partner's
phrase was synchronized with or overlapped the other katydid's first and sometimes
second phrase (b, Fig. 2); 3) alternation: the partner's phrase fell between two succes-
sive phrases by the other male (c, Fig. 2): 4) extended delay: two or more partners'
phrases fell between two successive phrases by the other male (d, Fig. 2).
The sequence of response types was quite variable within and between katydid
pairs. Within paired interactions, one male usually sang more phrases than the other
(R = 61% [range: 51-67%] and 39% [range: 33-49%]) and, the mean percentage of solos
and overlap solos was 61% (range: 0-100%) while that of alternations and extended
delays was 39% (range: 0.-100%). Solos averaged 31% (range: 0-83%) while synchrony
or overlap affected 30% (range: 0-61%) of a katydid's phrases. Alternation averaged
17% (range: 0-72%) and extended delay averaged 22% (range: 0-58%) of response types.











360 Florida Entomologist 70(3) September, 1987

PTEROPHYLLA BELTRANI



ACOUSTIC INTERACTION






B a ? l S 1 2 S 2 S dS
B A AL A W i 1dll AiI


I i I I I I F I I I I I I I I
0 1.6 3.2 4.8 6.4

TIME IN SECONDS

Fig. 2. Types of acoustic interaction between pairs of P. beltrani males. A. two
males alternating phrases. B. two males alternating groups of solos. 1 and 2 indicate
phrases of two different katydids; S = synchronous overlap of phrases of two katydids;
zero = non-synchronous phrase overlaps. Small letters identify bracketed response
types: a-solo, b-overlap solo, c-alternation, d-extended delay.

The phrase periods of the four response types were significantly different from one
another such that solo < overlap solo < alternation < extended delay (Fig. 3, Table 1).
Thus, during acoustic interaction, anytime one katydid produced one or more phrases
during the period of two successive phrases of the other katydid, the other katydid's
phrase rate usually was slowed.
Comparison of the phrase periods of solos before, during and after acoustic interac-
tion indicated that males increase solo rate during acoustic interaction and maintain it
for some period following termination of the singing of an interacting partner (Table 2).
When compared to the slower rate of soloing during solo calling, the only response type
of acoustic interaction that is significantly slower is extended delay (Table 3).

Courtship and Copulation

Courtship was identified when a male lowered its abdomen and raised its tegmina.
A silent or singing male could be induced to court by antennating or being antennated
by a female or another male. Courtship soloing was significantly faster than solo calling,
but the mean phrase periods of courtship and soloing during and after acoustical interac-
tion were not significantly different (Fig. 3, Table 2). When courting, males showed
very rapid abdominal pumping which appeared to be an increase in respiratory rate;
however, the movements could have been associated with spermatophore formation (T.
Walker, personal communication).
Receptive females climbed onto the back of courting males. Although females be-
came active and began to move toward a calling male, they did not hesitate to attempt
to mate with a silent male that they contacted during movement. Some silent and all
singing males assumed the courtship posture immediately after being antennated by a
female. Non-receptive males rejected females by kicking them with their hind legs and
moving away. Some females induced unreceptive males to kick and retreat by biting
them on their hind legs.








Shaw & Galliart: Katydid Behavior







I


1800



1600



1000



800-



600 -


400 -



200-


I I I I I I 5
0 1 2 3 4 5 6
RESPONSE TYPE


Fig. 3. Range of mean phrase periods (n = 7-19 katydids) for solo calling, the re-
sponse types of acoustic interaction, soloing after acoustic interaction and courting.
0-solo calling, 1-solo, 2-overlap solo, 3-alternation, 4-extended delay, 5-solo
after acoustic interaction, 6-courting.

After a female climbed onto the back of a courting male, successful copulation was
initiated by the male slipping the female's ovipositer into the terminal slit of his subgen-
ital plate (Fig. 4) while simultaneously grasping the base of the female's ovipositor with
the proximal hooks on the dorsal arms of his cerci (Fig. 4). Apparently, the pushing of
the female's abdomen dorsally with his subgenital plate aids in exposing the female's











Florida Entomologist 70(3)


d
/


I-


Fig. 4. Genitalic region of P. beltrani male. d-proximal hook on dorsal arm of
cercus, s-subgenital plate slit, v-ventral arm of cercus. Wings have been removed.


genital opening. The very prominent ventral curved arms of the male's cerci (Fig. 4)
hooked around the female's abdomen. After engagement of the genitalia, the male
loosened his tarsal hold on the substrate and hung head down from the female. The
male appeared to be supported by his hind legs grasping the female's, the cereal hold
on the base of the female's ovipositer, and genitalic attachment. In the copulation de-
scribed here in detail, the male intermittently grasped the branch below with his
forelegs, however, the front legs usually were held at an angle of approximately 900 to
the body.
Copulation consisted of accordion-like movements of the abdomen which resulted in
the male rocking to and fro. During copulation, the male continued the rapid abdominal
pumping initiated during courtship. The tip of the spermatophore was visible after ca.
1 min of copulation. For ca. 10 min, the strong, pumping movements of the male's abdo-
men occurred at a rate of ca. 10/min; then, they increased dramatically to ca. 120/min.
During these very rapid abdominal contractions, the male's body and legs flailed back-
and-forth. The male ceased the contractions every 10-15 sec and tightly tensed its whole
body before resuming the contractions within a few sec. After 2-3 min, the male alter-
nated 2-3 sec of rapid abdominal pumping with equivalent lengths of cessation of pump-
ing while tensing his body.
After 3 to 4 minutes, abdominal pumping assumed a low intensity, very rapid vibra-
tion and the female began to crawl across the branch dragging the male with her. After
ca. 7 min of this, during which the female appeared to be attempting to disengage from
the male, the couple separated terminating a 25 min copulation period. Of five copula-
tions observed, copulation time ranged from 25 to 38 min.
The external spermatophore was relatively small (estimated to be considerably less
than 10% of wet body weight; see Gwynne 1983), and consisted of a narrow throat and


September, 1987


P t.
:


%i" "











Shaw & Galliart: Katydid Behavior


two sperm sacs. After separation, the male continued abdominal contractions at a rate
considerably faster than the female (male: ca. 375/min; female: 60-80/min). Within to 2
min following separation, the male groomed his tarsi by drawing them through his
mouthparts, and in another 3 min bent his head ventrally and posteriorly to groom his
genital area.
After ca. 6 min from separation, the female bent her head ventrally and posteriorly
and began to eat the spermatophore; she continued to eat the spermatophore until it
was completely consumed after 6.5 min. After this, the female groomed the ovipositer
with her mouthparts.

Disturbance Sound.

Males and females of P. beltrani may produce very erratic disturbance sounds (Fig.
1) when grasped. It is assumed that such sounds would be produced when the insects
are grasped by predators. Sound components and intervals were very variable (Fig. 1).
There was considerable variation in ease of elicitation of the disturbance sound.
Males did not always produce these sounds and females were even less likely to do so.
During one afternoon's attempt to record the disturbance sound, two of five males and
none of eight females produced sounds when grasped and manipulated for a few seconds.
On another afternoon, 15 females were handled before the last one emitted the sound.


DISCUSSION

Bolivar & Pieltain (1942) suggested morphological similarities between P. beltrani
and P. camellifolia. Since we have described the songs of P. beltrani and since songs
of P. camellifolia have been described and analyzed in some detail (Shaw 1968), how
do they compare?
Three populations of P. camellifolia have been differentiated based upon differences
in morphology and calling song (Alexander 1968, Shaw 1968, Shaw & Carlson 1969,
North & Shaw 1979). These populations have been termed northern (occurs from New
England to the northern edge of the Appalachians and west to at least midway between
the Mississippi and Missouri rivers), southeastern (occurs south and east of the Appalac-
hians into northern Florida) and western (currently only reported from central and
southeastern Iowa but males have been heard in northwestern Missouri (unpublished)).
Alexander (1968) and Shaw & Carlson (1969) suggest the possibility of a fourth popula-
tion in Georgia, Alabama and Louisiana.
The range and mode of phrase pulse numbers of P. beltrani and northern P. camel-
lifolia are the same whereas those of the southeastern and western P. camellifolia
populations are greater (Table 4). P. beltrani songs exhibit the fastest pulse rate, the
shortest phrases and the fastest phrase rate (Table 4). P. beltrani songs are unusual in
the loud but shorter sounds produced during wing opening (Fig. 1). A short, usually
softer sound, may be produced by P. camellifolia males proceeding only the initial pulse
of a phrase.
Bolivar & Pieltain (1942) indicated that morphologically P. beltrani was more similar
to P. robertsi than P. camellifolia. The only description of the song of P. robertsi was
Hebard's (1941) statement that "the song was noted to be four or five rather rapid
notes, 'cha-cha-cha-cha-cha'." If the chas are phrases, P. robertsi produces a very unique
song consisting of groupings (sentences) of five phrases. If chas are pulses, then Hebard
is describing a 5-pulse phrase. Hebard reports P. robertsi males calling day and night
and he describes a nocturnal song produced when temperatures might be cool enough
for humans to differentiate individual pulses within a 4- or 5-pulse phrase. Barrientos











Florida Entomologist 70(3)


TABLE 4. COMPARISON OF PTEROPHYLLA CALLING SONGS.

Phrase
Species or Pulses/ Length Phrases/
Population Pulse No. sec (msec) min Temp C

Range Mode
P. beltrani 1-5 2 15-20 106 78-126 25-30
P. camellifolia1 1-5 2 6-8 250 60 27
(northern)
P. ,i Hi..f,,l. 5-22 8 6-8 1160 30 25
(western)
P. camellifolia" 3-8 4 10-11 300 58 27
(southeastern)

'Shaw 1968
'North and Shaw 1979
'Alexander 1968, Shaw 1968


(personal communication) has collected specimens in the same area (near Ciudad Vic-
toria, Tamaulipas, Mexico) at different times and these have been identified by D.
Nickle of the U.S.D.A. Systematic Entomology Laboratory as P. beltrani on one occa-
sion and P. robertsi on another. Barrientos is currently attempting to clarify the distri-
bution and taxonomy of P. beltrani and P. robertsi.
The field acoustical behavior of P. beltrani is very different from that of any of the
P. camellifolia populations. P. camellifolia sings at night and adjacent males alternate
phrases at nearest neighbor distances (NNDs) of 8-15 m (Shaw 1968). In contrast, P.
beltrani males sing day and night and, when observed, formed dense aggregations (tens
to hundreds of males and females in contact or within antennal contact of one or more
neighbors) at the end of tree top branches. When P. camellifolia males are less than 1
m apart, they produce aggressive sounds which typically involve alternation of longer
phrases (typically 1-4 pulses longer than the phrase of solo calling although up to 30
additional pulses have been reported (Alexander 1960)). Some males may not lengthen
their phrases but produce two or more solos at increased phrase rates (phrase intervals
decrease by as much as 50% (Shaw 1968)); these bursts of solos are usually alternated
with a long phrase of another male.
All of our recordings of P. beltrani males were at NNDs of 1 m or less. Some males
increased phrase length but never by more than one pulse. This is similar to phrase
length changes by some P. camellifolia males at 8-15 m. Although phrase length differ-
ences were great between and within species (e.g., 2-pulse P. beltrani phrase-106
msec; 2-pulse P. camellifolia phrase-250 msec; 6-pulse P. camellifolia aggressive
phrase-820 msec), phrase intervals of solo calling of both species were similar (Table
5). The degree of shortening of solo phase intervals elicited by close range acoustic
interaction also was similar (Table 5). These similarities in phrase intervals probably
represent similarities in recovery times of the two species' "acoustic pacemakers" (Shaw
1968).
The proximate mechanisms of acoustic interaction in P. beltrani appear to similar
to those of P. camellifolia (Shaw 1968) as well as several other species of katydids
(Busnel et al. 1956, Jones 1966a, 1966b). Alternation and extended delay are probably
the result of one katydid being inhibited from singing while its partner is singing. The
shortening of solo phrase interval could be the result of a post-inhibitory excitatory
phenomenon and/or excitatory effects of non-acoustic stimuli. With P. camellifolia


364


September, 1987











Shaw & Galliart: Katydid Behavior


TABLE 5. COMPARISON OF MEAN SOLO PHRASE INTERVALS (IN MSEC) OF P. BEL-
TRANI AND P. CAMELLIFOLIA MALE.

P. camellifolia (27-28.3oC)'
Solo type N2 Range Mean

solo calling song 6 474-651 541
solo during alternation 9 372-603 481
solo during aggressive calling 4 200-240 213

P. beltrani (25-300C)
Solo type N Range Mean

solo calling song 9 379-682 493
solo during acoustic interaction 16 119-486 235
solo during courting 7 154-497 253

'Shaw 1968
2N = no. of katydids; 10-20 phrases/katydid


males, Shaw (1968) could elicit an increase in solo rate and, for a few males, a 1-pulse in-
crease in phrase length in response to electronically-produced sound phrases; however,
increasing the length and sound level of the electronically-produced phrases failed to
elicit the magnitude of increase in solo rate and phrase length characteristic of acoustic
interaction when males are <1 m from one another. These results suggested that non-
acoustic stimuli emanating from a neaby male were responsible for the enhanced excitat-
ory effect. The rapid solo rate of P. beltrani's courtship song, which can be produced
when no other males are singing, reinforces the probable role of non-acoustic stimuli.
That the phrase periods following phrase overlap were longer than solo phrase
periods suggests some inhibition or delay. See Shaw (1968) for a discussion of an increas-
ing inhibitory effect as the interval between the initiation of a katydid's phrase and a
"stimulus" phrase are increased.
Because of the probable high energy cost of singing (over 50% of a tree cricket's
daily respiratory budget is devoted to calling (Prestwich & Walker 1981)), Alexander
(1975) has suggested that an increase in population density of a singing orthopteran
species should result in a decrease in time of male calling and an increase in time of
male searching. A corollary to this would be an expected decline in the energy utilized
in high intensity aggressive acoustic interactions. It is possible that the failure to pro-
duce long phrases when in close proximity to other males represents some energy
savings for P. beltrani males when compared to P. camellifolia males; but, a similarity
in production of solo bursts at a similarly reduced chirp interval during acoustic interac-
tion and during courting, the latter not occurring in P. camellifolia, suggests little if
any energy savings. Since P. beltrani males are heard singing day and night and because
of their tendency to aggregate, it would appear that males of this species would be
continually bombarded with potential stimuli for eliciting calling and courting. However,
it is unlikely that males sing continuously. In our laboratory, calling was sporadic and
not all males called when one male did initiate calling. With 5 or 6 males in the terrarium
and an equivalent number in screen cages within 1 m of the terrarium, there would be
hours during which no male called and this included times when there were one to two
virgin females in the terrarium. When a male did begin to call, he frequently was joined
by one to three other males but rarely did all the males sing.
These results and studies indicating the high energy cost of singing (MacNally &
Young 1981, Prestwich & Walker 1981) suggest that males must limit their calling time.











Florida Entomologist 70(3)


Therefore, each male should distribute his calling time to maximize his opportunities
for achieving a mating. If receptive females are available equally throughout a 24 hour
period, then male calling should be equally distributed over this period (at least over
the periods in which calling would not be affected by such factors as temperature
change, predator activity, etc.) (Walker 1983). Barrientos et al. (unpublished) report
that most matings occur between 0800 and 1000 hrs. This suggests that calling may
peak at some time prior to 0800 with the probability of a male calling at other times
dependent upon the factors mentioned above, plus any other factors that might affect
female receptivity at other periods. A need for field work is obvious.
No courtship song has been reported for P. camellifolia. However, the senior author
has observed courtship and copulation in P. camellifolia on two occasions. There is no
recollection of males calling during courtship; males stopped calling and lowered their
abdomen as soon as they were antennated by a male or female. In the case of P.
camellifolia, it is unlikely that another male would be nearby. For P. beltrani, is it
likely that many nearby males would be calling and in order to compete, it may be
important that a male continues to call while courting. The data presented here show
that not only does the male call, but he calls at the fastest solo rate possible.
The disturbance sounds of both species are similar in consisting of sound pulses
produced at irregular rates. So-called disturbance, alarm or defense signals which are
produced "by arthropods held in the hand, or disturbed in various manners such as
pinching, probing or restraining are known in almost every order of insect" (Alexander
1967). Such signals are commonly noiselike (i.e., possess a very wide frequency spec-
trum) and, if they consist of a sequence of pulses, are usually presented at an irregular
or erratic rate.
Disturbance sounds may aid insects by startling the would-be-predator, by acting
as an aposematic signal or by mimicking an aposematic signal (Haskell 1961, Masters
1979). There is some experimental evidence that disturbance sounds do deter predators
(reviewed by Masters 1979).
If disturbance sounds are an effective anti-predator device, why don't all P. beltrani
males and females produce them every time they are handled, or at least the first time
they are handled? Why are females less likely to produce disturbance sounds than
males?

END NOTE

Pulse-" . any sound that seems unitary by the methods being used to analyze it"
(Alexander 1967). In this paper, pulse refers to the sound presumedly made during the
closing stroke of a wing movement cycle. However, the songs of some P. beltrani males
include one or two prominent shorter sounds apparently made during the opening por-
tion of a wing movement cycle. This combination of sounds produced during both open-
ing and closing phrases of a wing movement cycle has been termed a phonatome (Walker
and Dew 1972).
Phrase-a sound consisting of one or more pulses or phonatomes.

ACKNOWLEDGEMENTS

We would like to thank L. Barrientos and her colleagues from the Instituto de
Investigaciones Alimentarios, Cd. Victoria, Tamaulipas, Mexico for all their help in
providing us with insects, with information about rearing P. beltrani, and for sending
us copies of published and unpublished manuscripts. Thanks also to Dr. David Cox and
Mr. Steve Rathbun of the Department of Statistics, Iowa State University, for the
statistical analyses.


September, 1987











Shaw & Galliart: Katydid Behavior


367


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1968. Arthropods. Pages 167-216 in T. A. Sebeok (ed.). Animal Communica-
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BUSNEL, R.-G., B. DUMORTIER, AND M.-C. BUSNEL. 1956. Recherches sur le com-
portement aqcoustique des Ephippigeres (Orthopteres, Tettigoniidae). Bull. Biol.
1-68.
ELSNER, N., AND A. POPOV. 1978. Neuroethology of acoustic communication. Adv.
Insect Physiol. 13: 229-355.
GWYNNE, D. T. 1983. Male nutritional investment and the evolution of sexual differ-
ences in Tettigoniidae and other Orthoptera. Pages 337-366 in D. T. Gwynne and
G. K. Morris, (eds.). Orthopteran mating systems: sexual competition in a diverse
group of insects. Westview Press: Boulder Colo., 376 pp.
HASKELL, P. T. 1961. Insect sounds. Quadrangle Books, Inc.: Chicago, Ill., 189 pp.
HEBARD, M. 1941. The group Pterophyllae as found in the United States (Tet-
tigoniidae:Pseudophyllinae). Trans. Amer. Entomol. Soc. 67: 197-220.
JONES, M. D. R. 1966a. The acoustic behavior of the bush cricket, Pholidoptera
griseoaptera, I. Alternation, synchronism and rivalry between males. J. Exp.
Biol. 45: 15-30.
1966b. The acoustic behavior of the bush cricket, Pholidoptera griseoaptera,
II. Interaction with artificial sound signals. J. Exp. Biol. 45: 31-44.
MACNALLY, R., AND D. YOUNG. 1981. Song energetic of the bladder cicada, Cys-
tosoma saundersii. J. Exp. Biol. 90: 185-196.
MASTERS, W. M. 1979. Insect disturbance stridulation: its defensive role. Behav.
Ecol. Sociobiol. 5: 187-200.
NORTH, R. C., AND K. C. SHAW. 1979. Variation in distribution, morphology and
calling song of two populations of Pterophylla camellifolia (Orthoptera:Tet-
tigoniidae). Psyche 86: 363-374.
PIERCE, G. W. 1948. The songs of insects. Harvard Univ. Press: Cambridge, Mass.,
329 pp.
PRESTWICK, K. N., AND T. J. WALKER. 1981. Energetics of singing in crickets:
effect of temperature in three trilling species (Orthoptera:Gryllidae). J. Comp.
Physiol. B, 143: 203-260.
SHAW, K. C. 1968. An analysis of the phonoresponse of males of the true katydid,
Pterophylla camellifolia (Fabricius) (Orthoptera:Tettigoniidae). Behaviour 31:
203-260.
SHAW, K. C., AND 0. V. CARLSON. 1969. The true katydid, Pterophylla camellifolia
(Fabricius) (Orthoptera:Tettigoniidae) in Iowa: two populations which differ in
behavior and morphology. Iowa State J. Sci. 44: 193-200.











Florida Entomologist 70(3)


SNEDECOR, G. W., AND W. G. COCHRAN. 1980. Statistical methods, 7th ed. Iowa
State U. Press: Ames, Ia, 507 pp.
STEEL, R. G. D., AND J. H. TORRIE. 1980. Principles and procedures of statistics: a
biometrical approach, 2nd ed. McGraw-Hill:New York, 633 pp.
WALKER, T. J. 1983. Diel patterns of calling in nocturnal Orthoptera. Pages 45-72 in
D. T. Gwynne and G. K. Morris (eds.). Orthopteran mating systems: sexual
competition in a diverse group of insects. Westview Press: Boulder, Colo., 376
pp.
WALKER, T. J., AND D. DEW. 1972. Wing movement of calling katydids: fiddling
finesse. Science 178: 174-176.





A DESCRIPTION OF THE LARVA AND PUPA OF RUPELA
ALBINELLA, A PEST OF RICE IN LATIN AMERICA
(LEPIDOPTERA: PYRALIDAE: SCHOENOBIINAE)

STEVEN PASSOA' AND D. H. HABECK
Department of Entomology and Nematology
University of Florida
Gainesville, Florida 32611

ABSTRACT

The larva and pupa of Rupela albinella (Cramer) (Lepidoptera; Pyralidae: Scho-
enobiinae) are described. Differences between larvae and pupae of this rice pest and
other known Schoenobiinae species are discussed.

RESUME

Se describe la larva y la pupa de Rupela albinella (Cramer) (Lepidoptera: Pyralidae:
Schoenobiinae). Se discuten las differencias entire las larvas y las pupas de esta plaga
del arroz y de otras conocidas species de Schoenobiinae.


The subfamily Schoenobiinae (Pyralidae) contains many pests of rice throughout the
world (Kranz et al. 1977, Yano 1968). The most common member of this complex in
Latin America is Rupela albinella (Cramer), which occurs from Mexico to Brazil (Hein-
rich 1937). Its usual Spanish common name, at least in Honduras, is novia del arroz (=
rice sweetheart) (Passoa 1983, Koone & Banegas 1958) although Hummelen (1974) called
it the white rice borer. Outbreaks have been reported from the Caribbean (Gooding
1980); Central America (King & Saunders 1984) and the northern portions of South
America, including Peru and Colombia (Torre 1961, Cheaney & Jennings 1975).
A morphological description of the larva and pupa of R. albinella is presented here
to facilitate identification of the immature stages. King and Saunders (1984) illustrated
the superficial appearance of the egg, larva and adult in color. Cheaney & Jennings
(1975) presented photographs of the larval damage.


'Present address: Department of Entomology, 320 Morrill Hall, University of Illinois, Urbana, Illinois 61801.


September, 1987


368











Passoa & Habeck: Description of Rupela albinella


MATERIAL AND METHODS

The larva was described from 18 specimens preserved in 75% alcohol. In addition,
2 individuals were slide mounted and stained with Chlorazal E Black. A compound
microscope was used to examine fine details of the mouthparts and chaetotaxy. Three
pupae were preserved in alcohol. Setal nomenclature follows Hinton (1946) whereas
pupal structures were named using Mosher (1916).
Material examined. COLOMBIA: Cali, IV-1983, no collector recorded, stem borer
in rice, larval slides #314-316 and genitalia slide #299 S. Passoa collection (2 pupae, one
with an adult that died before emergence in pupal shell; 9 larvae); ECUADOR: El Oro
Prov., Arenilles, 13-V-1986, V. Gonzaga and D. Habeck, boring in rice stem (9 larvae,
1 pupa).


Fig. 1-3. Rupela albinella immature stages. 1, anterior half of larva, lateral view,
scale line = 1.4 mm; 2, ventrolateral view of pupal head, scale line = 1.0 mm; 3, larval
anal shield, lateral view, scale line = .5 mm.


369









Florida Entomologist 70(3)


\L2
Os
L1

SV SV1



Dl
D2

oSD1
SD2 0 S


SV1 \'
SV2
A8 V11

4


AD1
3D2
\SD2
SD1
SL2
L

SV1v

cx V1
MS



D1SD1


L1
L2

SV1

A9 V1"


D1 D2
\SD1

\L1
L2"
L3
A SV1

A1 \V1


AF2
AF1


Fig. 4-7. Rupela albinella larval features. 4, chaetotaxy of the prothorax,
mesothorax, Al, A6, A8, and A9 respectively, lateral view; 5, skin texture (400x); 6,
epicrania, front, and labrum; 7, mandible, ventral view. See Fig. 8-14 legend for ab-
breviations.
RESULTS
Larval Description. Preserved larvae with reddish-brown head; cream-colored body;
and pale tan prothoracic and anal shields. Skin texture spinose, each spine stellate in


M3 L3
M3 L3


September, 1987











Passoa & Habeck: Description of Rupela albinella


cross section (Fig. 5); crochets in a uniordinal circle (sometimes weakly biordinal); ma-
ture larva approximately 30 mm long with a 1.5 mm head capsule width; head appears
reduced relative to the body size.
Head (Fig. 6): Front extends only 1/2 distance to epicranial notch; AF setae closely
spaced, AF2 just above front, AF1 slightly below; frontal pores lie midway between a
slanted line connecting Fl and C2; C2 slightly longer than C1; P1 three times length of
P2; A2 and L1 shorter than Al and A3; front with sclerotized flange extending from
clypeus to frontal area.
Six stemmata present; stemmata 1-3 evenly spaced; stemmata 3 and 4 contiguous;
stemma 6 posterodorsad of stemma 5.
Labrum (Fig. 6): M1 longer than M2 or M3; L1 shorter than L2 or L3, the latter
two setae almost subequal in length.
Mandible (Fig. 7): With three teeth, each tooth with a molar ridge; anterior mandibu-
lar seta minute and peg-like.
Hypopharyngeal complex (Fig. 9): With elongate spinneret equal to three times
length of basal segment of labial palpus; distal area with long thin spines dorsally;
proximolateral and proximodorsal areas also spined; stipular setae very long, subequal
in length to the spinneret.
Thorax (Fig. 1, 4, 8): Prothoracic shield with slight raised area on the posterior
margin; XD2 closer to SD1 than to XD1; D1 slightly posterodorsad of XD1; D2 and XD1
in a horizontal line; SD2 anterodorsad of spiracle; L1 twice length of L2; distance be-
tween SV1 and SV2 subequal to the distance between L1 and L2; V1 posterior to coxae,
close to the midline; membranous sac present anterior to coxae.
Mesothorax: D2 twice length of D1; SD1 twice length of SD2; all three L setae
arranged in a slanted line; SV group unisetose; V1 posterior to coxa and twice the
distance from the midline as VI on prothorax.
Abdomen (Fig. 3, 4): First abdominal segment with all setae posterior to spiracle
except SD2; D2 posteroventrad of Dl; SD2 anterodorsad of spiracle; SD1 ventrad of
Dl; L2 shorter than L1 or L3; SV group unisetose and dorsad of V1.
Sixth abdominal segment: D2 posteroventrad of Dl; SD1 ventrad of Dl; SD2 an-
terodorsad of spiracle; LI longer than L2; L3 much closer to proleg than to L1; SV
group trisetose, subequal in length, and arranged in an equilateral triangle; V1 below
posterior edge of proleg.
Eighth abdominal segment: D and SD setae as in A6; L setae much closer together
on A8 than A6 and arranged almost in a horizontal line; SV group bisetose; V1 near the
midline.
Ninth abdominal segment: D2 about three times length of Dl, the latter seta closer
to SD1 than to D2; L group bisetose, L2 half length of L1; position of L2 variable,
usually anteroventrad of L1, but sometimes ventrad; SV1 dorsad of V1, both setae
posteroventrad of L1.
Anal shield (Fig. 13): SD2 half the length of D1 and anteroventrad of the latter seta;
SD1 much closer to SD2 than D2.

Pupal Description. Pupa with a slight shouldered appearance, white to golden tan;
length approximately 20 mm.
Head (Figs. 2, 12): Vertex without setae; labrum U-shaped; frons rounded, almost
resembling a tubercle; labial palps thin and elongate, their length about equal to vertical
diameter of labrum; maxillary palps absent; maxillae very short, length equal to only
one third that of the prothoracic coxa; pilifers present; postgenae obvious; antennae
extending two thirds the distance to caudal wing margin.
Thorax (Figs. 10, 12): Prothoracic coxa very wide and long, its width subequal to
width of the head, its length twice the vertical diameter of head; prothoracic legs extend











Florida Entomologist 70(3)


8


Fig. 8-14. Rupela albinella immature stages. 8, larval prothoracic coxae and mem-
branous sac, ventral view; 9, larval hypopharyngeal complex, lateral view; 10, pupal
mesothoracic spiracle, dorsal view; 11, A1-A4 of pupa, dorsal view; 12, pupa, ventral
view; 13, larval anal shield, lateral view; 14, terminal abdominal segment of pupa,
lateral view. Abbreviations: a-antenna; Al, A2, A3-anterior setae; AF1, AF2-adfrontal
setae; C1, C2-clypeal setae; cx-coxa; Dl, D2-dorsal setae; Fl-frontal seta; L1, L2, L3-
lateral setae; M1, M2, M3-medial setae; mc-mesothoracic coxa; m-metathoracic coxa;
ml-mesothoracic leg; mtl-metathoracic leg; MS-mesothorax; P-prothorax; p-prothoracic
leg; pc-prothoracic coxa; pr-proleg; P1,P2-posterior setae; S-spiracle; sc-prothoracic
membranous sac; SD1, SD2-subdorsal setae; SV1, SV2, SV3-subventral setae; XD1,
XD2-extra prothoracic dorsal setae; Vl-ventral seta.


372


September, 1987











Passoa & Habeck: Description of Rupela albinella


cephalad of the maxillae; antennae and prothoracic legs extend an equal distance to
caudal wing margin; mesothoracic coxae exposed; mesothoracic legs about one third
longer than prothoracic legs, the anterior end projecting cephalad of maxillae;
metathoracic legs extend to A5 in female or A8 in the male; mesothoracic spiracle
absent, or possibly hidden, by a deep pit; skin texture of thorax smooth.
Abdomen (Fig. 11, 14): Spiracles oval, not projecting, those of A2 and A4-8 with
furrows, spiracle of A3 hidden by wing covers so that only a few furrows are visible;
proleg scars present on A5 and A6 ventrally; cremaster absent, terminal segment
rounded; skin texture of abdomen smooth except for wrinkled areas around spiracles
and a pit on the anterior dorsum of Al.

Host. Rice is the only known host (Arregoces, et al. 1980). The record of R. albinella
on sugarcane needs confirmation (see discussion).


DISCUSSION

Schoenobiinae larvae (based on 4 genera) may be separated from other pyralid sub-
families by their membranous sac located anterior to the prothoracic coxae on the mid-
line (Fig. 1). In addition, Allyson (1976) mentioned the reduced L2 seta on Al-8 as
another unusual feature.
R. albinella may be separated from other known genera of the Schoenobiinae by
the sinuous lateral margins of the anal shield; the elongate stipular setae subequal in
length to the spinneret; bisetose SV group on A8; uniordinal crochets arranged in a
circle on the prolegs; and by the presence of V1 on the thoracic segments. Uniordinal
crochets may be found on some Asian Tryporyza (Rothschild 1967) but R. albinella
differs from this genus, as well as some Old World species, by the presence of V1 on
the thoracic segments. Asian species of Schoenobiinae lack Vl on the thorax (Yoshiyasu
1985). Donacaula maximella (Fernald), a Nearctic species, differs from R. albinella in
its multiple membranous sacs (one on the prothorax and mesothorax between the
coxae). An undetermined species of Donacaula in Florida, collected on Paspalum, is
readily separated from R. albinella by its pigmented cranial tonofibrillary platelets.
The head of the rice sweetheart is unpigmented. Among Rupela spp., R. albinella
closely resembles R. leucatea (Zeller) because both species have a sinuate anal shield.
However, the foodplants are different. R. leucatea feeds on Echinochloa polystacha
(Heinrich 1937) whereas the host of R. albinella is rice. The anal shield of R. horridula
Heinrich resembles the vast majority of pyralids which have the lateral margins evenly
curved. Thus, it may be separated from the above two Rupela spp. by the lack of a
sinuate anal shield.
In spite of the above differences, R. albinella shares a series of unusual features
with other members of the Schoenobiinae. These include the posterior placement of the
D and SD setae behind the spiracle on Al-8 (illustrated by Hasenfuss (1960) for a
European Schoenobius); the reduction of the L2 seta on the abdominal segments (men-
tioned by Allyson (1976) as a characteristic of Schoenobiinae larvae); and the peg-like
anterior mandibular seta (found on D. maximella). The bisetose L group on A9 has
usually been considered a characteristic of the Nymphulinae (Bollmann 1955, Hasenfuss
1960, Yoshiyasu 1985) but it is also found in some Schoenobiinae larvae such as
Tryporyza (Rothschild 1967), D. maximella, and R. albinella.
Pupae of the Schoenobiinae (based on 4 genera) may be distinguished from other
pyralid subfamilies by the exposed mesothoracic coxae, pit-like mesothoracic spiracle,
lack of both a cremaster and maxillary palpi, and the projecting piothoracic legs which
extend cephalad of the maxillae (Fig. 2).











374 Florida Entomologist 70(3) September, 1987

The pupa of R. albinella differs from other known schoenobiines by the spiracle of
A3 which is hidden by the wings; its spiracular furrows; shape of the frons; and by the
abdominal spiracles which do not project outward. A rounded frons is not found on
Tryporyza (Rothschild 1967) or Donacaula maximella. Breniere's (1979) figure of Sci-
rphophaga does show a tubercle-like rounded frons but, unlike the rice sweetheart, the
abdominal spiracles are strongly projecting outward. Projecting prothoracic pupal legs
were considered a distinguishing feature of Pterophoridae pupae in North America
(Mosher 1916) and Asia (Nakamura 1981) but they were also found in the Schoenobiinae
pyralids examined during this study. The lack of a cremaster and the long exarate
metathoracic legs of R. albinella are also found in the many Nymphulinae pupae illus-
trated by Yoshiyasu (1985).
The record on R. albinella on sugarcane by Wille (1952) in Peru needs confirmation.
At first glance this record would appear to result from confusion between Diatraea
(common on sugarcane) and R. albinella. However, Wille (1952) noted the eggs of the
species he observed were covered with scales. This is a distinctive characteristic of
Rupela; the eggs of Diatraea are naked. The species involved was probably Rupela but
not albinella. Except for Wille's (1952) record, R. albinella has never been reared from
sugarcane.
All Schoenobiinae larvae examined during this study (Rupela, Schoenobius, Sci-
rphophaga (= Tryporyza?) and Donacaula) have a membranous sac between the
prothoracic coxae but this structure may not be homologous to other cervical glands
illustrated by Peterson (1962) or discussed by Razowski (1973). The membranous sac
of the Schoenobiinae may occur on each segment of the thorax, for example D.
maximella (although the metathoracic sac appears to be vestigial) and Scirphophaga
intacta Snellen, whereas the cervical (jugular) gland mentioned by Razowski (1973) is
restricted to the prothorax. Furthermore, the cervical gland has a smooth texture and
may be retracted into a slit in the integument when not in use (Peterson 1962). The
Schoenobiinae membranous sac has a spinose texture (see Hasenfuss (1960) for typical
example) and there is no integumental slit. Therefore the sac is visible all the time,
even in specimens which were not inflated by a syringe. The function of the Scho-
enobiinae membranous sac remains unknown, future research in this area is needed to
contribute information about possible homologies with other Lepidopteran glands.

ACKNOWLEDGEMENTS

The authors wish to thank CIAT (Centro Internacional de Agricultura Tropical,
Colombia) and S. Allyson (Canadian National Collection) for the gift of Schoenobiinae
larvae and pupae. Dr. R. Hodges and A. Solis provided space and other assistance
during a visit to the Smithsonian Institute. The comments of S. Allyson, Dr. G. Godfrey
and Dr. M. Berenbaum were greatly appreciated. Florida Agricultural Experiment
Station Journal Series no. 7816.

REFERENCES

ALLYSON, S. 1976. North American larvae of the genus Loxostege Hubner (Lepidopt-
era: Pyralidae: Pyraustinae). Canadian Entomol. 108(1): 89-104.
ARREGOCES, O., E. CASTRO, C. CARDONA, J. GONZALEZ, J. A. REYES, J. GARCIA,
AND J. RAIGOSA. 1980. Barrenadores del tallo del arroz en America Latina y
su control. Cali, Colombia: Centro Int. Agric. Trop. Guia de Estudio. 31 p.
BOLLMAN, H. G. 1955. Die raupen mitteleuropaischer Pyraustinae (Lepidoptera-
Pyralidae). Beitr. Entomol. 5(5-6): 521-639.
BRENIERE, J. 1979. Reconnaissance des principaux lepidopteres du riz de L'Afrique
de L'Ouest. Agron. Tropicale 31: 213-231.











Passoa & Habeck: Description of Rupela albinella


CHEANEY, R. L., AND P. R. JENNINGS. 1975. Problems en cultivos de arroz en
America Latina. Cali, Colombia: Centro Int. Agric. Trop. 91 p.
GOODING, E. G. B., editor. 1980. Pest and pesticide management in the Caribbean.
Volumes I-III. Barbados, West Indies: Consortium Int. Crop Prot./U.S. Agency
for Int. Devel. 558 p.
HASENFUSS, I. 1960. Die larvalsystematik der Zunsler (Pyralidae). Abh. Larvalsys-
temik Insekt. 5: 1-263.
HEINRICH, C. 1937. Moths of the genus Rupela (Pyralididae: Schoenobiinae). Proc.
U.S. Nat. Mus. 84(3019): 355-386.
HINTON, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. En-
tomol. Soc. Lond. 97: 1-37.
HUMMELEN, P. J. 1974. Relations between two rice borers in Surinam, Rupela al-
binella (Cr.) and Diatraea saccharalis (F.), and their hymenopterous larval para-
sites. Meded. Landbhoogesch. Wageningen 74(1): 1-88.
KING, A. B. S., AND J. L. SAUNDERS. 1984. The invertebrate pests of annual food
crops in Central America. Overseas Devel. Admin. London, England, 166 p.
KOONE, H. D., AND A. D. BANEGAS. 1958. Entomologia economic Hondurena.
Minist. Recur. Nat. Bol. Tech. (Tegucigalpa) 6: 1- 138.
KRANZ, J., H. SCHMUTTERER, AND W. KOCH (eds.). 1977. Diseases, pests, and
weeds in tropical crops. Berlin: P. Parey, 666 p.
MOSHER, E. 1916. A classification of the Lepidoptera based on characters of the pupa.
Bull. Ill. St. Lab. Nat. Hist. 12(2): 1-159.
NAKAMURA, M. 1981. Key to the classification of the Japanese lepidopterous pupae.
Tyo to Ga 32(1-2): 1-12.
PASSOA, S. 1983. Lista de los insects asociados con los granos basicos y otros cultivos
selectos en Honduras. Ceiba 25(1): 1-97.
PETERSON, A. 1962. Larvae of insects. An introduction to Nearctic species. Part I.
Lepidoptera and plant infesting Hymenoptera. Ann Arbor, Michigan, 315 p.
RAZOWSKI, J. 1976. Lepidoptera of Poland. 1-General part. Springfield, Virginia:
U.S. Nat. Tech. Inform Serv., 157 p.
ROTHSCHILD, G. H. L. 1967. Descriptions of larval and pupal stages of four lepidop-
terous rice borers in Malaysian Borneo (Sarawak). Bull. Entomol. Res. 57(3):
343-352.
TORRE, G. DE LA. 1961. La mariposa blanca Rupela albinella Cram., un nuevo insecto
en el arroz. Rev. Peru. Entomol. Agric. 4(1): 74-76.
WILLE, J. E. 1952. Entomologia agricola del Peru. Lima, Peru: Minist. Agric. 543 p.
YANO, K. 1968. Key to the rice stem borers from Asia (Lepidoptera). Mushi 42(7):
81-87.
YOSHIYASU, Y. 1985. A systematic study of the Nymphulinae and the Musotiminae
of Japan. Sci. Rep. Kyoto Pref. Univ. Agric. 37: 1-162.











376 Florida Entomologist 70(3) September, 1987

A NEW SPECIES OF NEMAPALPUS
(DIPTERA; PSYCHODIDAE; BRUCHOMYIINAE)
FROM NORTHEASTERN COLOMBIA

J. BRUCE ALEXANDER
Department of Entomology and Nematology, 3103 McCarty Hall,
University of Florida, Gainesville, FL 32611


ABSTRACT

Nemapalpus patriciae n.sp. is described and illustrated from male and female speci-
mens collected at Arboledas, Northeastern Colombia. It is the the first species of this
widespread genus to be recorded from Colombia, and is the 26th Nemapalpus species
to be described. On the basis of the structure of the male genitalia and presence of tufts
of long setae on tergites 3-7, it is suggested that N. patriciae should be placed in the
sziladyi species group.


RESUME

Se described e ilustran Nemapalpus patriciae sp. n. de muestras de machos y hem-
bras colectadas en Arboledas, al noreste de Colombia. Es la primera especie de este
extendido genero de ser registrada de Colombia, y es la 26 especie de Nemapalpus de
ser descrita. Basado en la estructura de la genitalia del macho y la presencia de
mechones de setas largas en las tergitas 3-7, se sugiere que N. patriciae debe de in-
cluirse en el grupo de species "sziladyi".


The genus Nemapalpus Macquart 1838 (the variant spelling Nemopalpus Macquart
1839 has been commonly accepted, but Duckhouse & Lewis (1980) pointed out that by
priority the original spelling should be adopted) shows a disjunct distribution, with most
of the 26 known species occurring in geologically ancient parts of the world, such as
South Africa, the Canary Islands, South America, Hispaniola, Australia and New Zea-
land (Stuckenberg 1962). One exception is N. nearcticus Young, found in Northern
Florida (Young 1974). This distribution pattern may reflect the great age of the genus,
with present-day species representing the remnants of a group that was formerly much
more widespread and diverse (Quate 1961). The family Psychodidae, which includes
Nemapalpus and the haematophagous phlebotomine sand flies, is known to have arisen
by the Permian (250 myr ago) and fossil psychodids dating from this time closely resem-
ble modern species. Two fossil species of Nemapalpus have been recovered from
Oligocene Baltic amber (Edwards 1921) and members of this genus, together with others
in the Bruchomyiinae, are thought to be the most primitive of living psychodids (Young
1974).
Very little is known about the biology of Nemapalpus. Adults of most species are
generally found resting in the same habitats as phlebotomine sand flies during the day
and, like them, occur both in semi-arid areas of the Old World, such as the Canary
Islands, and forested areas of the Neotropics. Adult Nemapalpus may be collected from
cracks in rocks, buttress roots and lower tree trunks during daylight. It is not known
whether these insects are diurnal or nocturnal, nor is it known whether they feed as
adults. Unlike sand flies, the females are not haematophagous.
Larvae of Nemapalpus have been found in the same habitat as larval sand flies in
Panama (Hanson 1968) and are morphologically very similar. They can be reared in the











Alexander: New Colombian Psychodidae 377

laboratory on the same types of medium used for sand fly colonies (Alexander unpub-
lished, Endris et al. 1983) and probably feed on leaf litter and other detritus in the wild.
No species of Nemapalpus has been recorded previously from Colombia, although
examples are known from most parts of the Neotropics, including Venezuela (Ortiz &
Scorza 1963). Collections from the Municipality of Arboledas, Norte de Santander
(7039'N, 7248'W) during 1985-6 revealed the presence of three species, including one
previously recorded only from Guatemala and Mexico, and two new species, one of
which is described here for the first time.


Nemapalpus patriciae Alexander, new species
(Fig. 1-8)

MALE: General coloration dark brown. Head infuscated, eyes black and large, separated
at narrowest distance by 3 facet diameters. Proboscis length 0.10-0.12 mm, (n=8).
Length of palpal segments (in mm): 1(0.06-0.07, n=5); 2(0.10-0.12, n=7); 3(0.13-0.16,
n=7); 4(0.17-0.18, n=6); 5(0.52-0.57, n=5). Approximately 15 peg-like sensory rods
(Newstead's scales) at proximal end of palpal segment 3.
Antennae with 3 segments, dimensions (in mm) as follows: scape (0.08-0.10, n=7);
pedicel (0.06-0.07, n=8); flagellomere 1(0.44-0.51, n=8); 2(0.39-0.44, n=8); 3(0.40-0.45,
n=8); 4(0.40-0.43, n=8); 5(0.39-0.41, n=8); 6(0.36-0.41, n=8); 7(0.34-0.38, n=8); 8(0.31-
0.36 n=8); 9(0.30-0.33, n=8); 10(0.29-0.31, n=7); 11(0.25-0.31, n=6); 12(0.25-0.26, n=5);
13(0.22-0.25, n=4); 14(0.20-0.21, n=3). Ascoids mushroom-shaped, 1 on inner, distal
part of each flagellomere.
Wing length 3.9-4.1 mm, (n=7); width 0.9-1.0 mm (n=8); wing venation as shown,
cross vein r-m proximal to fork of MI-M2. Mesonotum and pleurae infuscated, with
sparse, fine setae on pleura. Halteres pale brown.
Legs dark, except for tarsi which are covered in closely-appressed silvery-white
scales. Short, erect, black setae sparsely distributed on all segments of legs. Dimensions
of segments as follows (in mm): foreleg coxa (0.53-0.61, n=8); trochanter (0.12-0.15,
n=8); femur (1.38-1.53, n=6); tibia (2.68-3.23, n=6); tarsus (3.03-3.21, n=6); midleg
coxa (0.46-0.56, n=7); trochanter (0.13-0.15, n=7); femur (1.53-1.66, n=6); tibia (3.29-
3.40, n=6); tarsus (2.96-3.26, n=6); hindleg coxa (0.49-0.53, n=8); trochanter (0.13-0.18,
n=8); femur (1.61-1.84, n=6); tibia (3.49-3.82, n=6); tarsus (2.93-3.59, n=6).
Abdomen infuscated. Tergites 3-7 each bearing paired verrucae from which emerge
tufts of long, whitish setae, anterior to smaller tufts of shorter setae. Long, whitish
setae also diffusely distributed on tergite 2, and scattered, short setae present on all
abdominal segments. Dorsum of tergites 1-3 divided by groove along midline.
Genital pump length approximately 0.38 mm, one-third length of partially-coiled,
annulated sperm ducts. Ducts end in funnel-like expansions, in which a few fine spines
are visible. Genitalia inverted and complex. Aedeagus tubular, bifurcating at tip into
spinose, membranous projections as shown. Gonostylus ending in single sclerotized
curved spine. Paramere triramous and well sclerotized. Each of two lateral processes
further divided into 2 flattened arms, with shorter of these partially overlapping
outer,longer arm, which bears outward-pointing spine near tip.
FEMALE: Size and coloration as for male. Lengths of proboscis, palpal segments,
and antennal segments 1 & 2 same as in male. Other antennal segments missing on
allotype specimen.
Leg dimensions (in mm) as follows (allotype only): Foreleg coxa 0.57; trochanter
0.14; femur 1.45; tibia 2.70; tarsus 2.93; Midleg coxa 0.49; trochanter 0.12; femur, tibia
and tarsus missing; Hindleg coxa 0.51; trochanter 0.18; femur 1.78; tibia and tarsus
missing.












Florida Entomologist 70(3)


-- 15





4


Fig. 1-8. Nemapalpus patriciae n.sp. 1) Male head; 2) Abdominal segment of male,
showing setal tufts; 3) Male wing; 4) Male genitalia, lateral view; 5) Aedeagus; 6) Male
genitalia, ventral view; 7) Female genitalia, lateral view; 8) Spermatheca. Figs. 2, 4, 6
& 7 to same scale. All measurements in mm.


Abdomen lacking paired tufts of long setae. Single spermatheca 0.92 mm long, 0.35
mm wide, with dense covering of small spines over entire surface and groove running
four-fifths of length, curving down to join lower surface. No other distinctive features
visible.


September, 1987











Alexander: New Colombian Psychodidae 379

TYPE DATA: This species is named in memory of my mother, Patricia Mary Alexan-
der, who died in Edinburgh, Scotland on September 18, 1985, after a long illness.
The type locality is a coffee plantation at altitude 900 m, in the Eastern Cordillera
of lie Colombian Andes. Specimens were obtained using a flight trap or by aspirating
resting adults from trunks of large trees which provide shade for coffee bushes.
HOLOTYPE: Male, flight trap, finca La Esperanza, Vereda (district) Siravita, 2.5 km
E of Arboledas, Norte de Santander, Colombia, 24-VII-1986.
ALLOTYPE: Female, tree trunk, finca La Esperanza, 22-VII-1986.
PARATYPES: 1 male. flight trap at finca El Tejar, Siravita, 24-VII-1985: 1 male, tree
trunk. La Esperanza, 10-VI-1986: 1 male, tree trunk, La Esperanza, 30-VI-1986: 3
males, flight trap, La Esperanza, 23-VII-1986: 1 male, flight trap, La Esperanza. 24-
VII-1986.
Holotype, allotype and paratypes all mounted in Canada balsam on slides. Holotype
and allotype to be deposited in Florida State Collection of Arthropods. Paratypes in
collections of author, Dr. D. G. Young (University of Florida, USA), U.S. National
Museum, Washington, USA, Dr. D. Duckhouse (University of Adelaide, Australia), Dr.
A. Morales (Instituto Nacional de Salud, Bogota, Colombia) and British Museum (Nat-
ural History), London, UK.

DISCUSSION

This species brings the number of described Neotropical Nemapalpus species to 14.
It differs from N. dampfianus Alex., 1940 in having pleural setae, small spines in the
expanded portion of the sperm ducts and complex parameres. The latter two features
also distinguish N. patriciae from N. pallipes (Shannon and Del Ponte, 1927); N. pilipes
Tonnoir, 1922; N. immaculatus Freeman, 1949; N. brevinervis Barretto and d'An-
dretta, 1946; and N. dissimilis (Barretto and d'Andretta, 1946).
The new species differs from N. antillarum Fairchild, 1952 in general coloration,
and the male style of the latter species terminates in paired, sclerotized blades, rather
than a single spine; N. arroyi de Leon, 1950 has very similar female terminalia to the
new species but again differs in structure of the male terminalia. Males of N. arroyi
lack tufts of long setae on tergites 3-7; male terminalia of N. mopani de Leon, 1950
show similarities to the new species but the style bears three spines, rather than a
single one; males of N. moralesi de Leon, 1950 also lack long setae on tergites 3-7 and
the style bears four finger-like lobes; N. sziladyi Tonnoir,1940 is the species which
resembles N. patriciae most closely, particularly in the structure of the parameres and
style. The legs of both species are clothed in shining appressed scales and dark erect
setae. The species differ in that the spinose, bifurcate aedeagus of N. sziladyi is mar-
kedly shorter and broader and the style bears a blade-like process as well as a strong
spine; N. torrealbai Ortiz and Scorza, 1962 is also similar to the new species in the male
genital armature but is distinguishable by the presence of a small blade-like process in
addition to the strong spine on the style. Finally, N. yucatanensis Vargas and Diaz
Najera, 1958 resembles N. patriciae in the structure of the parameres but the four-lobed
style is quite unlike that of the new species.
On the basis of the structure of the male genitalia and the presence of paired tufts
of long setae on tergites 3-7, N. patriciae should probably be grouped with N. anti-
llarum, N. sziladyi, N. torrealbai, N. mopani, N. yucatanensis and the sole North
American species, N. nearcticus. Ortiz & Scorza (1963) have proposed the name
* ..i:;/ ,i group" for all Nemapalpus species in which the males bear setal tufts; on this
basis alone, the new species would also be grouped with N. dissimilis, in which the
style is simple, and N. immaculatus, in which the parameres are virtually absent and
tufts of erectile hairs are restricted to tergites 6 & 7. Tonnoir (1940) suggested that the











Florida Entomologist 70(3)


erectile hairs of Nemapalpus might function as disseminators of a sex pheromone, and
the fact that they are restricted to males may be evidence that they play some part in
the courtship of certain species, although no detailed physiological or behavioral studies
have been made of this to date. It is not known whether the long setae of N. patriciae
can be erected or flattened at will, although they are morphologically similar to those
of other sziladyi group species. The presence of a groove on tergites 1-3 may be further
evidence that the setae are erectile.
Specimens of Nemapalpus patriciae were caught or seen on the following dates:-
24-VII-1985 (4 males, flight trap); 10-VI-1986 (1 male, tree trunk); 16-VI-1986 (1 seen,
tree trunk); 24-VI-1986 (1 seen, tree trunk); 30-VI-1986 (1 male, tree trunk); 5-VII-1986
(1 seen, tree trunk): 12-VII-1986 (1 female, tree trunk); 22-VII-1986 (1 female, tree
trunk, 1 seen, tree trunk. & 1 male, flight trap); 23-VII-1986 (3 males, in flight trap);
24-VII-1986 (2 males, flight trap); 12-VIII-1986 (1 male, flight trap); 2-IX-1986 (1 male,
flight trap). Males outnumbered females 14 to 2 in collections. Sampling from tree
trunks was done between 08.30 and 11.30 each morning during June, July and August
of 1985, November 1985, June and July 1986, and October and November 1986. Flight
traps were in place on finca El Tejar from 17-VI-1985 to 25-VII-1985 and on finca La
Esperanza from 17-VII-1986 to 20-IX-1986.
This species is structurally similar to phlebotomine sand flies, although considerably
larger and more agile, and like the 17 sand fly species which occur in the same habitat
at Arboledas, shows a sex ratio skewed markedly in favor of males in both resting site
and flight trap collections (Alexander, unpublished). It is not known why this is so, since
presumably N. patriciae shows a 50:50 sex ratio, as in phlebotomine sand flies and most
other Diptera.
A single specimen of another undescribed Nemapalpus species was caught in the
flight trap at El Tejar on 24-VII-1985, and two male Nemapalpus mopani (de Leon)
were found on tree trunks at finca El Encanto, Vereda El Uvito, approximately 1km
from Arboledas. This species has not been recorded from Colombia before, but is known
from Mexico and Guatemala (Fairchild 1952).


ACKNOWLEDGEMENTS

I thank Oscar J. Jaimes and Jesus A. Parada for assistance with field work, and Dr.
D. G. Young and Margo A. Duncan for help in preparation of the manuscript. This work
was supported by grant 5-P01-A120108 from the National Institutes of Health. This is
Florida Agricultural Experimental Station Journal No. 8356.


REFERENCES CITED

DUCKHOUSE, D. A., AND D. J. LEWIS. 1980. Family Psychodidae, p. 93-105. In R.
J. Crosskey, Catalogue of the Diptera of the Afrotropical region. British
Museum, (Natural History), London, 1437 p.
EDWARDS, F. W. 1921. A note on the subfamily Bruchomyiinae (Diptera Nematoc-
era). Ann. Mag. Nat. Hist. 7: 437-9.
ENDRIS, R. G., P. V. PERKINS, D. G. YOUNG, AND R. N. JOHNSON. 1982. Tech-
niques for laboratory rearing of sand flies (Diptera: Psychodidae). Mosquito News
42: 400-7.
FAIRCHILD, G. B. 1952. Notes on Bruchomyia and Nemopalpus (Diptera, Psy-
chodidae). Ann. Entomol. Soc. America 45: 259-80.
HANSON, W. J. 1968. The immature stages of the subfamily Phlebotominae in Panama
(Diptera: Psychodidae). Ph.D thesis, Univ. Kansas, Lawrence, Kansas. 104 p.


September, 1987











Baranowski & Slater: New Lygaeidae


ORTIZ, I., AND J. V. SCORZA. 1963. Notas biological y taxonomicas sobre algunos
Phlebotominae (Diptera, Psychodidae) de Rancho Grande, Venezuela. Acta Biol.
Venezuela 3: 341-61.
QUATE, L. W. 1961. Zoogeography of the Psychodidae (Diptera). Eleventh Int.
Congr. Entomol. Wien 1960. 1: 168-73.
STUCKENBERG, B. R. 1962. The South African species of Nemopalpus (Diptera:
Psychodidae). Ann. Natal Mus. 15: 201-18.
TONNOIR, A. L. 1940. Sur un remarquable organe sexuel secondaire chez males du
genre Nemopalpus Macq. avec description d'une espece nouvelle et d'une autre
peu connue (Dipt., Psychodidae). Sixth Congr. Intern. Entomol. Madrid, 1935,
pp. 203-13.
YOUNG, D. G. 1974. Bruchomyiinae in North America with a description of Nemopal-
pus nearcticus n.sp. (Diptera: Psychodidae). Florida Entomol. 57: 109-13.





NEW GENERA AND SPECIES OF ANTILLOCORINI FROM
TRINIDAD AND BRAZIL (HEMIPTERA:LYGAEIDAE)

RICHARD M. BARANOWSKI
University of Florida, IFAS
Tropical Research and Education Center
Homestead, Florida 33031
AND
JAMES A. SLATER
Section of Systematic and Evolutionary Biology
University of Connecticut
Storrs, Connecticut 06268

ABSTRACT

Arimacoris pileacola, NEW GENUS and NEW SPECIES from Trinidad, and Gem-
macoris NEW GENUS (TYPE SPECIES G. nitens, new species), from Trinidad and G.
albidus, NEW SPECIES from Brazil are described. The generic relationships are dis-
cussed. Feeding habits of A. pileacola are discussed and the nymphs described. Illustra-
tions are given for A. pileacola and G. nitens.

RESUME

Se described Arimacoris pileacola, genero nuevo y especie nueva de Trinidad, y Gem-
macoris, genero nuevo (especie tipo, G. nitens, nueva especie), de Trinidad, y la especie
nueva G. albidus del Brazil. Se discuten las relaciones genericas. Se discuten los habitos
de alimentaci6n de A. pileacola y se described sus ninfas. Se presentan ilustraciones de
A. pileacola y de G. nitens.


The tribe Antillocorini is represented by many taxa and is much more diverse and
the relationships more complex than previously suspected. Slater, Sweet and
Baranowski (1977) and Slater (1980) have described a number of new taxa; the latter
paper discusses the phylogeny and provides a cladogram and a key to the Western
Hemisphere genera. Slater (1982) suggests that the Antillocorini may well be a
paraphyletic tribe.











Florida Entomologist 70(3)


Recent collecting in Trinidad has provided us with additional material which is, in
part, the subject of the present paper. These taxa do not possess a combination of
derived features that allow placement in any existing genus and are accordingly de-
scribed as new. It should be pointed out however that generic relationships of some of
the Neotropical antillocorines will remain somewhat ambiguous until a careful study of
the genus Botocudo is accomplished. This genus, as presently constituted, seems to us
to be doubtfully monophyletic since a number of species do not possess the character
states used by Slater (1980) in his key and cladogram. We hope to comment further on
"Botocudo" in a later contribution.

Biology

One of the interesting features of the Trinidad species described below is that they
were taken above ground on Pilea microphylla (L.) Liebm. (Urticaceae) and that
Arimacoris pileacola Baranowski & Slater NEW SPECIES, at least, definitely feeds on
the seeds of this plant and breeds on it.
Rhyparochromines are mostly oligophagous (Sweet 1960, 1964) primarily feeding on
seeds in fallen litter on the ground. Sweet (1964) also indicated that there was not a
well-marked host specificity in the New England species he studied. Slater (1972) clas-
sified the lygaeids feeding on seeds as, 1) arboreal seed predators, 2) obligatory terres-
trial seed predators, 3) frequent facultative terrestrial seed predators and 4) accidental
terrestrial seed predators. At that time members of the Antillocorini were known only
to feed in seed litter on the ground.
Slater, Sweet and Baranowski (1977) provided what is probably the first record of
members of the Antillocorini as arboreal seed feeders. Three species of Bathydema (B.
socia Uhler, B. jamaicensis Slater and Baranowski, B. maculosa Slater and
Baranowski) were reported to feed on the seeds of Pilea species while the seeds were
still on the plants. It was suggested that this adaptation-feeding on seeds still on the
plant-is effective in moist tropical habitats where seeds falling to the ground either
germinate or mold quickly. The ground litter habitat is thus not suitable for geophilous
seed feeding lygaeids (see Sweet (1964 p 26-27)).
Recently a series of Paurocoris wygodzinskyi Slater, collected by Dr. J. Eger, was
brought to our attention. Adults and nymphs were collected on Pilea microphylla and
were observed feeding on the seeds. Eggs were deposited singly in the floral parts in
much the same manner as are the eggs of A. pileacola. The fact that A. pileacola is
known to breed only on P. microphylla, and P. wygodzinskyi was collected on and
breeds on the same host plant, while species of Bathydema are known to feed on more
than one species of Pilea, but have not been collected on P. microphylla, suggests that
Paurocoris and Arimacoris are closely related genera.
In addition to the two species described below a species of the Ozophora pallescens
complex also definitely breeds on Pilea microphylla and a specimen of a species of
Botocudo has been taken on this plant. These additional rhyparochromines that live on
mature seeds above ground strengthens our suggestions of the selective advantage of
utilizing seeds (in moist tropical habitats) before they fall into the litter.
While most of the known Pilea feeders are antillocorines, and these are all small
bugs the limiting factor may be the size of the insects rather than the taxonomic group.


Arimacoris Baranowski and Slater NEW GENUS

Type Species: Arimacoris pileacola Baranowski and Slater, NEW SPECIES.
Body short, stout, broadly elliptical. Head, pronotum, scutellum, corium, lateral and


382


September, 1987











Baranowski & Slater: New Lygaeidae


ventral surfaces, and corium laterad of radial furrow pruinose; inner portion of corium
and entire clavus subshining. Entire dorsal surface punctate; clavus with 3 rows of
punctures. Head short and broad, only slightly declivent anteriorly. Eyes sessile, in
contact with antero-lateral pronotal angles. Ocelli set slightly behind posterior margins
of eyes. Pronotum with lateral margins of anterior lobe narrowly but sharply carinate.
No transverse pronotal impression. Posterior pronotal margin concave before base of
scutellum. Scutellum lacking a median carina. Lateral corial margins broadly explanate.
Inner half of apical corial margin deeply and abruptly concave. Membrane translucent.
Gular trough nearly reaching base of head. Fore femora mutic. Metathoracic scent
gland auricle strongly curving posteriorly, apex blunt. Evaporative area occupying only
inner 1/3 of metapleuron, outer margin convex. Second antennal segment clavate seg-
ments III and IV fusiform. Well developed, equally sized scent gland scar openings
present between abdominal terga 3-4, 4-5 and 5-6 (Fig. 1B). The scent gland configura-
tion is similar to that of Cligenes distinctus (Dist.) (Fig. 1C). Inner laterotergites on














A



B












D


C

Fig. 1. A. Arimacoris pileacola, Baranowski and Slater n. sp. abdominal trichobot-
hria; B. (same) abdominal scent gland openings; C. Cligenes distinctus (Dist.) abdominal
scent gland openings; D. (same) abdominal trichobothria.


383











Florida Entomologist 70(3)


segments 4, 5 and 6. Trichobothria of sternum 5 linear, the 2 posterior trichobothria
close together, the posterior one placed well behind spiracle (Fig. 1A). Abdominal spira-
cles 2 and 4 placed on lateral sternal shelf, spiracle 3 placed below shelf.
Remarks: Slater (1980), following his description of the genus Paurocoris, mentioned
an undescribed species from Trinidad that appeared to be somewhat transitional be-
tween Paurocoris, Botocudo and CIy', i. .. Arimacoris is based upon this species, de-
scribed below.
The trichobothrial arrangement of Arimacoris is transitional in that the two poste-
rior trichobothria of sternum 5 are essentially linear (the posterior one is placed very
slightly more ventral than the preceding one) (Fig. 1A). However, the posterior
trichobothrium is placed well behind the spiracle of sternum 5. A polarity sequence can
be conceived from a plesiomorphic condition in which the two posterior trichobothria of
sternum 5 are located one above the other and both behind spiracle 5 (as in Paurocoris),
to the condition described above for Arimacoris, to a condition where the two posterior
trichobothria are completely linear and have "migrated" forward of the spiracles as in
Cligenes (Fig. 1D).
Arimacoris may also be distinguished from Paurocoris by: 1) the spiracle of sternum
4 (Fig. 1A) placed on the sternal shelf rather than below it, 2) the dorsal surface of the
head pruinose rather than shining, 3) the lateral margins of the posterior portion of the
pronotum not sharply carinate, 4) the strongly posteriorly curved metathoracic scent
gland auricle, and 5) the strongly produced explanate margins of the coria.
Arimacoris will key to couplet 4 in Slater's (1980) key to the genera of Western
Hemisphere Antillocorini but will not pass through this couplet. The characters are
contradictory in that the trichobothria of abdominal sternum 5 are linear in position but
are not all located anterior to the spiracle. It should also be noted here that couplet 5,
which separates Cligenes from Botocudo by the former possessing a deep median groove
on the prosternum, is true of the type species of Cligenes (distinctus Distant) but not
of C. subcavicola Scudder, Darlington & Hill.
All measurements that follow are given in mm.

Arimacoris pileacola Baranowski and Slater, NEW SPECIES
(Fig. 2)

Head light brown, tylus yellowish. Pronotum and scutellum uniformly dark brown
except anterior pronotal "collar," antero-lateral margins of pronotum and apex of scutel-
lum yellowish. Clavus straw colored with slightly darkened posterior 1/2. Proximal 1/2,
lateral edge and apex of corium straw colored, remaining portion dark brown. Lateral
edges of abdominal terga also straw colored. Legs nearly uniformly yellowish. Proximal
2/3 of fore femura testaceous. Antennal segments I, II, III, dull yellow, segment IV
testaceous.
Head short, broad, tylus extending to distal 1/3 of antennal segment I. Vertex
convex. Head length 0.20, width 0.40, interocular space 0.32. Pronotum broadly
trapezoidal, lateral margins along anterior 1/2 narrowly explanate. Pronotum length
0.32, width 0.66. Scutellum length 0.50 width 0.36. Hemelytra with corial margins
slightly sinuate, arcuately rounded distad of level of claval commissure. Claval commis-
sure length 0.14. Mid-line distance apex clavus-apex corium 0.30. Mid-line distance apex
corium-apex membrane 0.16. Labium extending to mesocoxae. Labial segments length
I 0.14, II 0.16, III 0.10, IV 0.10. Antennal segments length I 0.14, II 0.18, III 0.18, IV
0.24. Total body length 1.34.
HOLOTYPE: S TRINIDAD, W.I. St George Co. Arima-Blanchisseuse Rd. 7.75 mi.
post 10-X-1978 (R. M. Baranowski). In United States National Museum of Natural
History Type No. 100251.


384


September, 1987










Baranowski & Slater: New Lygaeidae


I


Fig. 2. Arimacoris pileacola Baranowski and Slater, n.sp., dorsal view.

PARATYPES: TRINIDAD 1 6, 3 9 Arima-Blanchisseuse Rd 15-1-1976 (F. D. Ben-
nett); 2 d, 2 9 Arima-Blanchisseuse Rd, 7.5 mi mark 7-VII-1976 (R. M. Baranowski);
7 6, 12 9 same 10.25 mi mark 9-VIII-1975; 15 6, 42 9 same 7.5 mi post 9-VIII-1975;
8 6, 8 9 same 9-IX-1977; 14 6, 9 9 same 7.75 mi post 16-VIII-1978; 8 S, 10 9 same
29-VII-1978; 1 9 same 10.25 mi mark 6-VII-1975; 3 6, 5 9 same 23-VIII-1982; 4 d, 9
9 St. George. Co. Arima-Blanchisseuse Rd. C2/11 31-VIII-1980 (R. M. Baranowski) 21
6, 13 9 St. George Co, Arima-Blanchisseuse Rd. 7.75 mi mark, 10-X-1978 (R. M.
Baranowski) 9 6, 26 9 same 26-IX-1978; 2 6, 3 9 same 24-VII-1978; 6 6, 10 9 same
15-VII-1978; 4 6, 2 9 same 19-VIII-1983; 6 6, 6 9 same 12.5 mi post 23-IX-1979; 9 6,
8 9 same 10.25 mi post 17-VIII-1980; 1 6, 2 9 same C3/11, 12-VIII-1981; 5 3, 8 9
same, 7.75 mi post 23-VIII-1982 (R. M. Baranowski, J. A. & E. Slater, R. Clayton, M.
H. Hassey) 12 6, 20 9 St. George Co. Arima-Blanchisseuse Rd. Simla 1-IX-1983; 3 S,


*< *r., ,*/*: '"'










Florida Entomologist 70(3)


8 9 same 2-IX-1983; 11 6, 16 9 same 14-IX-1983. In United States National Museum
of Natural History, British Museum (Natural History), Florida State Collection of Ar-
thropods, J. A Slater and R. M. Baranowski collections.
Remarks: Arimacoris pileacola occurs in a very unusual habitat for a rhyparoc-
hromine lygaeid, occurring exclusively on Pilea microphylla, even though a second
species, P. inaequalis (Poiret) Weddell, is often closely associated with P. microphylla.
A. pileacola has been observed feeding on the seeds on the plant in the laboratory, and
because most rhyparochromine lygaeids are exclusively seed feeders, this species is
probably also. P. microphylla commonly occurs on man-made structures, most fre-
quently on the vertical rock or concrete surfaces of culverts, bridges, etc. A. pileacola is
most readily collected by placing a net below the small Pilea plants growing from a
vertical rock face and "brushing" or shaking the minute insects off the plants into the
net from which they can be collected with an aspirator.
It is striking, as the paratype series indicates, that A. pileacola is known from a
very restricted area in Trinidad covering only a few miles in the northern range along
the Arima-Blanchisseuse Rd, Arima Valley, St George Co. Despite extensive collecting
on P. 11,,, ,,l.::,ll/l in the Aripo valley, a few miles east of the Arima valley and in the
Caura Valley, west of the Arima Valley no specimens have been taken. In addition A.
pileacola was not collected on P. microphylla along the north coast road from Blanchis-
seuse to Port of Spain, in the Wallerfield area where extensive patches of P. micro-
phylla occur on shaded abandoned paved runways and roads, nor culverts along roads
in the central range.
In the laboratory A. pileacola deposited its eggs singly in the floral parts of its host
plant. This was also true of Bathydemajamaicensis (Slater, Sweet, Baranowski, 1977).


Immature Stages:

Fifth instar nymph (in alcohol)
Body short, stout, ovoid. Head cream colored, eyes and ocelli red. Pronotum and
wing pads tan. Abdomen cream colored with posterior edge of terga 2-5 narrowly red-
dish. Antennal segments I, II whitish, segments III, IV reddish brown. Femora
whitish, tibiae and tarsi dusky. Head length 0.28, width 0.36, interocular space 0.24.
Pronotum length 0.28, width 0.60, wing pad length 0.46. Labial segments length I 0.18,
II 0.18, III 0.10, IV 0.14. Antennal segments length I 0.12, II 0.16, III 0.14, IV 0.24.
Abdomen length 0.70. Total body length 1.42.

Fourth instar nymph (in alcohol)
Similar in shape and color to 5th instar. Head length 0.22, width 0.32, interocular
space 0.20. Pronotum length 0.22, width 0.50, wing pad length 0.20. Labial segments
length I 0.14, II 0.14, III 0.10, IV 0.12. Antennal segments length I 0.12, II 0.12, III
0.12, IV 0.20. Abdomen length 0.60. Total body length 1.10.

Third instar nymph (in alcohol)
Similar in shape and color to 4th instar. Head length 0.18, width 0.30, interocular
space 0.20. Pronotum length 0.20, width 0.44, wing pad length 0.16. Labial segments
length I 0.10, II 0.12, III 0.10, IV 0.10. Antennal segments length I 0.10, II 0.10, III
0.10, IV 0.20. Abdomen length 0.56. Total body length 1.10.

Second instar nymph (in alcohol)
Similar in shape to 3rd instar. Head and thorax pale tan, abdomen cream colored
suffused with reddish, especially along posterior margin of terga. Legs uniformly cream
colored. Head length 0.18, width 0.26, interocular space 0.20. Pronotum length 0.14,


386


September, 1987











Baranowski & Slater: New Lygaeidae


width 0.32. Labial segments I 0.10, II 0.10, III 0.10, IV 0.10. Antennal segments length
I 0.08, II 0.10, III 0.10, IV 0.14. Abdomen length 0.42. Total body length 0.80.

First instar nymph (in alcohol)
Similar in shape to 2nd instar. Head, thorax, legs, antennal segments I, II, III
dusky tan, abdomen and antennal segment IV whitish. Head length 0.14, width 0.22,
interocular space 0.16. Pronotum length 0.10, width 0.28. Labial segments length I
0.08, II 0.08, III 0.08. IV 0.08. Antennal segments I 0.06, II 0.08, III 0.06, IV 0.14.
Abdomen length 0.30. Total body length 0.50.

Egg (in alcohol)
Cream colored, elongate oval. Length 0.52, width 0.30 at its widest point anteriorly,
somewhat tapering posteriorly. Anterior end almost flat; only 2 micropylar projections
evident.

Gemmacoris Baranowski and Slater NEW GENUS

Type Species: Gemmacoris nitens Baranowski & Slater NEW SPECIES
Body stout, tapering anteriorly not broadly elliptical. Dorsal and ventral surfaces
almost entirely shining and polished. Pruinose only narrowly across base of head be-
tween ocelli. Pronotum and hemelytra pruinose or subshining. Dorsal surface nearly
completely punctate except for large, swollen, impunctate calli of anterior pronotal lobe.
Clavus with 3 rows of punctures. Head elongate, somewhat porrect. Eyes sessile in
contact with antero-lateral pronotal angles. Ocelli anterior to behind posterior margin
of compound eye. Lateral margins of pronotum obtusely carinatee" (weakly so), strongly
impressed and sinuate in area of transverse impression which is obsolete laterally and
absent mesally. Posterior margin of pronotum evenly, shallowly concave. Scutellum
distally with a prominent median ridge. Lateral corial margins narrowly explanate,
sinuate at level of middle of scutellum. Inner portion of apical corial margin deeply
concave. Membrane translucent. Gular trough remote from base of head, terminating
near level of middle of eye. Meso and metepisternum moderately to strongly swollen.
Metathoracic scent gland auricle curved prominently caudad. Evaporative area occupy-
ing only inner half or less of metapleuron, outer margin straight for most of length.
Second antennal segment clavate, segments 3 and 4 fusiform. Well-developed, equal
sized scent gland opening scars present between abdominal terga 3-4, 4-5 and 5-6. Inner
laterotergites on segments 4, 5 and 6 (Fig. 3 A, C) Two posterior trichobothria of
sternum 5 not linear, located nearly dorso-ventrally and anterior to spiracle 5 (Fig. 3B,
D). Spiracles 3 and 4 located below sternal shelf.
Remarks: This genus is readily recognizable by the combination of partially shining
polished body surface; sinuate, slightly obtusely carinate lateral pronotal margins, well-
developed scent gland opening scars between abdominal terga 3-4, 4-5, 5-6; backwardly
curved metathoracic scent gland auricle, the position of the abdominal spiracles and the
trichobothria on sternum 5. As noted above, Arimacoris has almost achieved a linear
arrangement but with the posterior trichobothrium still placed well behind the spiracle.
In Gemmacoris the two posterior trichobothria are not linear. The "middle" trichobot-
hrium is slightly anterior to the posterior one and both are placed well ahead of the
spiracle. The position of the trichobothria in both Gemmacoris and Arimacoris suggests
that the "advanced" linear condition with all three trichobothria anterior to the spiracle
and arranged linearly (Fig. 1D) could have been achieved independently in two ways:
1) by initial development of a linear sequence as in Arimacoris (Fig. 1A) and a sub-
sequent migration of the posterior pair of trichobothria to a position anterior to the
spiracle, and 2) by migration of the two posterior trichobothria to a position anterior of
the spiracle while still in a dorso-ventrad position relative to one another, as in Gem-











Florida Entomologist 70(3)


Fig. 3. A. Gemmacoris nitens, Baranowski and Slater, n.sp. abdominal trichobot-
hria; B. (same) abdominal scent gland openings: C. Gemmacoris albius, Baranowski
and Slater, n.sp. abdominal scent gland openings; D. (same) abdominal trichobothria.

macoris (Fig. 3A, D), with subsequent development of the linear position. This possibil-
ity of independent attainment of a linear sequence anterior to the spiracle must be
considered and it raises the question of whether the "advanced" condition should be
considered a synapomorphy or a homoplasy.


September, 1987











Baranowski & Slater: New Lygaeidae


Key to Species of Gemmacoris

1. Fourth antennal segment yellowish brown, concolorous with preceding
segments; pronotum subshining; scutellum pruinose only basally; tylus
extending onto distal 1/3 of 1st antennal segment (Trinidad) .......... nitens n.sp.
1'. Fourth antennal segment white, strongly contrasting with brown coloration
of preceding segments; pronotum dull, strongly contrasting with shining
texture of head; scutellum with entire distal 1/2 pruinose; tylus extending
anteriorly only to middle of 1st antennal segment (Brazil) ........... albidus n.sp.

Gemmacoris nitens Baranowski and Slater, NEW SPECIES
(Fig. 4)

Head reddish brown, tylus slightly lighter, sparsely covered with short fine golden
hair. Ocelli conspicuous, located slightly behind hind margin of compound eyes. A single
long seta located just anterior to each ocellus. Pronotum and scutellum reddish brown,
coarsely punctate except for pronotal calli. Clavus pale anteriorly gradually changing
to dark brown posteriorly. Corium straw colored except for dark brown apical and a
dark brown irregular median band. Ventral surface reddish brown. Legs nearly un-
iformly yellowish except for proximal 2/3 of femora darker. Antennae light brown,
proximal end of III, IV, distal end of III slightly darker.
Head elongate, porrect. Tylus extending to distal 1/3 of antennal segment I. Vertex
convex. Head length 0.42, width 0.48, interocular space 0.28. Lateral margins of pro-
notum weakly carinate, strongly impressed and sinuate in area of transverse impression
which is obsolete laterally and absent mesally. Posterior margin of pronotum evenly,
shallowly concave. Pronotum length 0.48, width 0.82. Scutellum length 0.46, width 0.46.
Lateral corial margins narrowly explanate, sinuate at level of middle of scutellum. Inner
portion of apical corial margin deeply concave. Claval commissure length 0.22. Mid-line
distance apex clavus-apex corium 0.48. Mid-line distance apex corium-apex membrane
0.30. Labium extending to mesocoxae. Labial segments length I 0.28, II 0.26, III 0.20,
IV 0.22. Antennal segments length I 0.28, II 0.30, III 0.24, IV 0.34. Total body length
2.20.
HOLOTYPE: 9 TRINIDAD, W.I. Arima-Blanchisseuse Rd. mi mark 7.75, 23-VIII-
1982 (R. M. Baranowski). In United States National Museum of Natural History Type
No.100253.
PARATYPES:TRINIDAD 1 9 3 mi N Cumuto, 14-VI-1973 (R. Baranowski, F.
O'Rourke, V. Picchi, J. Slater); 1 9 St. Augustine (N. A. Weber). In J. A. Slater and
American Museum of Natural History collections.

Gemmacoris albidus Baranowski and Slater, NEW SPECIES

Head light brown, tylus slightly lighter, sparsely covered with short fine hairs.
Ocelli located slightly behind hind margin of eyes. Pronotum and scutellum brown;
punctate except for pronotal calli. Clavus straw colored except for slightly darker apex.
Corium straw colored except for brown apical 1/4, a brown spot on lateral margin and
on inner margin at level of claval apex. Membrane reduced, overlapping but not extend-
ing beyond apex of corium. Exposed dorsal surface of abdomen and entire ventral
surface brownish. Legs yellowish except for anterior femora and proximal 2/3 of anterior
tibiae darker. Antennal segments I, II, III brown, IV white.
Head elongate, porrect. Tylus extending to middle of antennal segment I. Vertex
convex. Head length 0.42, width 0.50, interocular space 0.30. Lateral pronotal margin
weakly carinate anteriorly, strongly impressed in area of transverse impression which


389









39


September, 1987


'0 Florida Entomologist 70(3)



---'

t' .




.it -. f F. i
r
t .



i'


I ,

/



I .
y7""








II
~Wyi *.'^^^ '''w~S~

*-**^, rB


N
rv!.

.t.


i'

,
N


I,


Fig. 4. Gemmacoris nitens Baranowski and Slater, n.sp., dorsal view.

is obsolete laterally and absent mesally. Posterior margin of pronotum evenly, shallowly
concave. Pronotum length 0.48, width 0.74. Scutellum length 0.38, width 0.40. Lateral
corial margins slightly sinuate at level of posterior 1/2 of scutellum. Inner portion of
apical corial margin deeply concave. Claval commissure length 0.20. Mid line distance
apex clavus-apex corium 0.28. Labium extending to mesocoxae. Labial segments length


-- --;


i
81"
r'
.


N+











Baranowski & Slater: New Lygaeidae


I 0.30, II 0.34, III 0.18, IV 0.20. Antennal segments length I 0.30, II 0.40, III 0.36, IV
0.40. Total body length 2.40.
HOLOTYPE. S BRAZIL, Parana. Bocaiura, 1000 m., 25'08 29'04, May 1964 (F.
Plaumann). In United States National Museum of Natural History No. 100254.
PARATYPES: 2 9, (same data as holotype). In J. A. Slater and P. D. Ashlock
collections.

ACKNOWLEDGEMENTS

We thank Dr. Richard Howard (Harvard Univ.) for determining the species of Pilea,
Dr. P. D. Ashlock (Univ. of Kansas) and Dr. R. T. Schuh (American Museum of Natural
History) for the loan of material, Dr. F. D. Bennett (Commonwealth Institute of Biolog-
ical Control, Trinidad) for assistance in many ways over the years, Ms Mary Jane
Spring (Univ. of Connecticut) and Mrs Kathy Schmidt (American Museum of Natural
History) for preparation of the illustrations and Drs. R. C. Froeschner (USNMNH), B.
J. Harrington (Univ. of Wisconsin) and A. G. Wheeler (Commonwealth of Penn. Dept.
of Agric.) for their helpful reviews.
This is No. 5289 of the Florida Agricultural Experiment Station Journal Series. The
senior author is a Research Associate of the Florida State Collection of Arthropods,
Florida Department of Agriculture and Consumer Services, Gainesville.

REFERENCES CITED

SLATER, J. A. 1972. Lygaeid bugs (Hemiptera:Lygaeidae) as seed predators of figs.
Biotropica (Washington) 4:3: 145-151.
1980. Systematic relationships of the Antillocorini of the Western Hemisphere
(Hemiptera:Lygaeidae). Syst. Ent. (London) 5: 199-226.
1982. Lilliputocorini, a new tribe with six new species of Lilliputocoris and a
cladistic analysis of the Rhyparochrominae (Hemiptera, Lygaeidae). Amer. Mus.
Novitates, 2754: 1-23.
-- M. H. SWEET, AND R. M. BARANOWSKI. 1977. The systematics and biology
of the genus Bathydema Uhler (Hemiptera:Lygaeidae). Ann. Ent. Soc. Amer.
70: 343-358.
SWEET, M. H. 1960. The seed bugs: a contribution to the feeding habits of the
Lygaeidae. (Hemiptera:Heteroptera). Ann. Ent. Soc. Amer. 53: 317-321.
1964. The biology and ecology of the Rhyparochrominae of New England
(Hetroptera:Lygaeidae) Part I. Ent. Amer. 43 (n.s.): 1-124.




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