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Title: damselflies (Zygoptera) of Texas
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of the
FLORIDA STATE MUSEUM
Biological Sciences
Volume 16 1972 Number 2




THE DAMSELFLIES (Zygoptera)
OF TEXAS



. {, Clifford Johnson


UNIVERSITY OF FLORIDA


GAINESVILLE










Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL
SCIENCES, are published at irregular intervals. Volumes contain about 300
pages and are not necessarily completed in any one calendar year.


















OLIVER L. AUSTIN, JR., Editor
FRED G. THOMPSON, Managing Editor


Consultantsfor this issue:
HARRY K. CLENCH
DENNIS R. PAULSON


















Communications concerning purchase or exchange of the publication and all
manuscripts should be addressed to the Managing Editor of the Bulletin,
Florida State Museum, Museum Road, University of Florida, Gainesville,
Florida 32601.


Publication date: 18 January, 1972


Price: $1.25
















THE DAMSELFLIES (ZYGOPTERA) OF TEXAS




CLIFFORD JOHNSON,



0 SYNOPSIS: This report presents an identification guide to adult damselflies oc-
curring in Texas. Illustrated characters, a guide to morphological terminology,
and short text support the diagnostic keys. The text gives geographical range
Sand habitat preferences for each group. Distribution data appear by county
for each species and reveal patterns of convergence between east and west
S faunas.



TABLE OF CONTENTS

INTRODUCTION .......... ..................... ............. ....... 56
ACKNOW LEDGEMENTS ........................... .................... 57
M ETH ODS . ............................... ....... ...... ......... 57
KEY TO THE FAM ILIES ............................................ 62
L ESTIDA E .......... ........................ ................... .... 63
-Archilestes ....................................... ......... 63
Lestes ............................................ .......... 65
CALOPTERYGIDAE .................................................. 69
C alop teryx ................................................... 69
H etaerina .................................................... 71
3 PROTONEURIDAE ................................................... 73
C OENAGRIONIDAE .................................................. 74
A rg ia ................................................. ... 78
Enallagma .................................................. 93
Ischnu ra .................................................... 100
Smaller Genera .................................. ............ 107
DIscussioN ................................... ....... ........... 111
LITERATURE CITED ................................................ 115
APPENDIX ......... ............................................... 117


The author is Associate Professor in the Department of Zoology, University of
Florida, Gainesville, Florida, 32601. Manuscript accepted 22 March 1971 Ed.

Johnson, Clifford. 1971. The Damselflies (Zygoptera) of Texas. Bull. Florida
State Mus., Biol. Sci., Vol. 16, No. 2 pp. 55-128.
55





75&3(S


57 B E L D SL UE M
56 BULLETIN FLORIDA STATE MUSEUM


Vol. XVI No. 2


INTRODUCTION
This study provides a species list for the damselfly fauna of Texas,
diagnostic keys for adult identification, and current knowledge of their
distribution. The taxonomy of U. S. damselflies is well known, and the
latest description of a species occurring in the state appeared in 1924.
Table 1 lists 53, possibly 54, species inhabiting Texas with description
dates and authorities.
Hagen's (1861) A Synopsis of North American Neuroptera was the
first major publication to include Texas material. A significant number of
Texas records appears in the Odonata section of Biologia Centrali-Amer-


TABLE 1. THE DAMSELFLY FAUNA OF TEXAS

LESTIDAE Needham 1903


Archilestes grandis (Rambur) 1842
Lestes alacer Hagen 1861
Lestes disjunctus australis Walker 1952
Lestesforficula Rambur 1842


Lestes inaequalis Walsh 1862
Lestes sigma Calvert 1901
Lestes vigilax Hagen 1862
Lestes simplex IHagen 1861'


CALOPTERYGIDAE Selys 1853


Calopteryx dimidiata Burmeister 1839
Calopteryx maculata (Beauvois) 1805


Hetaerina americana (Fabricius) 1798
Hetaerina titia (Drury) 1773


PROTONEURIDAE Tillyard 1926


Neoneura aaroni Calvert 1903


Protoneura cara Calvert 1903


COENAGRIONIDAE Kirby 1890


Anomalagrion hastatum (Say) 1839
Argia apicalis (Say) 1839
Argia barretti Calvert 1902
Argia bipunctulata (Hagen) 1861
Argia fumipennis violacea (Hagen) 1861
Argia hinei Kennedy 1918
Argia immunda (Hagen) 1861
Argia lugens (Hlagen) 1861
Argia moesta (Hagen) 1861
Argia munda Calvert 1902
Argia nahuana Calvert 1902
Argia plana Calvert 1902
Argia rhoadsi Calvert 1902
Argia sedula (Hagen) 1861
Argia tibialis (Rambur) 1842
Argia translate Hagen 1865
Enallagma basidens Calvert 1902
Enallagma civil (Hagen) 1861
Enallagma divagans Selys 1876
Enallagma dubium Root 1924
lQuestionable status: see text.


Enallagma durum (Hagen) 1861
Enallagma exsulans (Hagen) 1861
Enallagma geminatum Kellicott 1895
Enallagma novaehispaniae Calvert 1907
Enallagma praevarum (Hagen) 1861
Enallagma signatum (Hagen) 1861
Enallagma traviatum Selys 1876
Enallagma vesperum Calvert 1919
Ischnura barberi Currie 1903
Ischnura demorsa (Hagen) 1861
Ischnnra denticollis (Burmeister) 1839
Ischnura kellicotti Williamson 1898
Ischnura posita (Hagen) 1861
Ischnura prognatha (Hagen) 1861
Ischnura ramburii (Selys) 1850
Ischnura verticalis (Say) 1839
Hesperagrion heterodoxum (Selys) 1868
Nehalennia integricollis Calvert 1913
Telebasis salva (Hagen) 1861
Teleallagma daeckii (Calvert) 1903









JOHNSON: TEXAS DAMSELFLIES


icana by Calvert (1901-1908), the Catalogue of the Odonata of North
America by Muttkowski (1910), and the Handbook of Dragonflies of
North America by Needham and Heywood (1929). The following regional
studies for the state supplement these contributions. Williamson (1914)
reported on collections in central and south Texas, while Tucker (1908),
Ferguson (1940, 1942), and Harwell (1951) provided distributional notes
from north-central and east Texas. Tinkham (1934) and Gloyd (1958) re-
ported on the Texas fauna occurring west of the Pecos River. Gloyd's pa-
per also includes important taxonomic revisions and is essential to any
student of the area. Shorter references to Texas species appeared in Cal-
vert (1893), Gloyd (1932), Johnson (1961, 1962, 1963), and Donnelly
(1964). Isolated data on Texas material also exist in taxonomic works such
as Williamson (1912, 1917). These studies present an incomplete list of the
state's fauna and are long out-of-print.


ACKNOWLEDGEMENTS

Data in this report represent collective efforts of many students and rec-
ords from numerous collections. Thomas Donnelly contributed unpublished
state records of Enallagma dubium, Ischnura kellicotti, I. prognatha, and Ne-
halennia integricollis, in addition to other distribution data. Leonora K. Gloyd
provided the unpublished state record for Argia rhoadsi, distribution data
from the Williamson Collection at the University of Michigan, and several
smaller collections. Her invaluable comments on the genus Argia deserve spe-
cial mention. Dennis R. Paulson, George H. Bick, and Claron Bjork contri-
buted distribution data from private collections. Lois O'Brian, Horace C.
Burke, Kenneth W. Stewart, and James Sublette supplied collections for study
from Texas Technological, Texas A. and M., North Texas State, and Eastern
New Mexico Universities respectively. B. Elwood Montgomery assisted with
early literature citations and usage of taxonomic categories. Oliver S. Flint, Jr.
answered questions on material at the U. S. National Museum, and Minter J.
Westfall, Jr. provided data from the Florida State Collection of Arthropods. A
state-wide treatment of Texas damselflies would have been impossible without
the collective assistance of all the above.


METHODS

The paper's principal objective provides an identification guide for Texas
damselflies. Text discussions compare species characters with a view to reduc-
ing confusion in determinations rather than giving descriptions in formal mon-
ograph style. The order of families follows Fraser (1954) where lestids repre-
sent a primitive stock and two lines of descent include (1) calopterygids, and
(2) protoneurids and coenagrionids.
A statement on geographical range and group characteristics proceeds
each genus and species key. The keys require a basic familiarity with numerous
structures and veins. Correct orientation of dorsal and lateral views for struc-
tural characters is essential, and the required orientation appears throughout


1972









58 BULLETIN FLORIDA STATE MUSEUM


the keys. Structural characters naturally vary somewhat within a species;
where such variability affects diagnoses, a species identification occurs at more
than one key couplet. Line drawings of structural traits omit hairs and color
patterns. An expanded explanation in the proceeding text supplements difficult
key characters. Keys are reversible by the number in parenthesis at each
couplet.
The Needham system of vein terminology was chosen for use in the keys
over the Fraser-Tillyard system simply to conform with the majority of North
American Odonata literature. Figure 1 shows body structures with the follow-
ing usage of symbols: prothorax, Ti; mesothorax, T2; metathorax, Tg; pter-
othorax, T2 + Ts; middle and hind prothoracic lobes, MPL and HPL; meso-
stigmal plates, MP; median carina, MC; humeral suture, H; metapleural su-
ture, LS; abdominal segments 1 and 2, abd. seg. 1 and 2; compound eyes, CE;


D



S\ RAMBURII\ r/BALIS

FIGURE 1. A, thorax in left lateral view (prothorax disarticulated); B, C,
head in left lateral and dorsal (anterior end uppermost) views; D, E, tibial
spines in selected coenagrionids. Structures identified in text.


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


ocelli, 0; vertex, V; postclypeus, PC; labrum, Lr; labium, La; and antennae,
Ant. Body length refers to the total length from anterior tip of the head to the
apex of abdominal segment 10. Abdominal and body length exclude appen-
dages. Greatest width across the compound eyes refers to a line drawn over
the width of the head connecting points x' and x" shown in Figure 1.
The following characters identify males of all species. The genital fossa
accommodating the penis lies conspicuously in the sternum of abdominal seg-
ment 2 and the anterior part of segment 3. The abdominal appendages con-
sist of a superior and inferior pair just posterior to segment 10. Argia-type
appendages shown in Figure 2 A and B exemplify these structures. Additional
structures torii, torifer, and torealea) occur on the 10th abdominal segment in


A B


ACF


MDS
FIGURE 2. Generalized Argia-type, male abdominal appendages, tori shown
in black; A left lateral and, B, dorsal (anterior end lowermost) views; C, D,
left lateral views of terminal abdominal structures in female coenagrionids;
E, dorsal (anterior end uppermost) view of mesostigmal plates and associated
structures in a typical coenagrionid female. Structures identified in text.








60 BULLETIN FLORIDA STATE MUSEUM


Argia males, and an explanation of their structure appears under that genus.
Abdominal male appendages occasionally dry or become accidentally displaced
to an atypical position. Usually such conditions are obvious, but may lead to
error in determinations if not recognized. Females lack a genital fossa and pos-
sess an ovipositor at the terminal end of the abdomen (Fig. 2 C, D). Determin-
ations of females frequently use the mesostigmal plates or laminae, MIP, (Fig.
1 A; Fig. 2 E). The text for the Coenagrionidae contains a description of these
structures.
Figures 3 and 4 show wing venation with the following usage of symbols.

A An N P
A .-,-


M;_2
M1-2
FIGURE 3. Basal half of fore wings in: A, Calopterygidae, Iletaerina ameri-
cana; B. Lestidae, Lestes vigilax; C, Coenagrionidae, Argia tibialis. Venation identi-
fied in text.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


Longitudinal veins are costal, C; cubital veins 1 and 2, Cul and Cu2; com-
bined 1st and 2nd branches of the median vein, MI-2; four medial branches,
MI to M4; radial vein, R; the radial sector, Rs; and the subcostal, Sc. Spe-
cialized cross veins are the arculus, A; nodus, N; and subnodus, Sn. The anal
crossing, Ac, appears also as a cross vein. Other identified cross veins are the
ante- and postnodal cross veins, An and Pn. Postnodal cross veins consist of
the first cross vein distal to the nodus, N. and all cross veins in the series
outward to and including the cross vein just proximal to the brace vein, b,







PM2
OF Rs






SpM



4M2











S4. Basal half of fore wings in protonerids: A, Protone c ; B,
X2 CuI M 4

C P M






MOES TA

D




HASTATUM
SEDULA
FIGURE 4. Basal half of fore wings in protoneurids: A, Protoneura cara; B,
Neoneura aaroni; C, D, and E, stigma variations in selected coenagrionids,
venation as in Fig. 3.









62 BULLETIN FLORIDA STATE MUSEUM


under the stigma, st. Specific cells are the discoidal cell or quadrangle, Q; ante-
nodal postquadrangular cells, ANC; and median space. MS. Subquadrangle
cells, Xi and X2, just posterior to the quadrangle, Q, form their longitudinal
margins by veins Cul, Cu2, and the wing's posterior margin, PM. Where
Cu2 is absent in some protoneurids, X2 is also absent. The contour of the
wing base is narrow in all Texas damselflies other than calopterygids, and
the slender stalk is the petiole, P. The following reference points identify the
above veins. The arculus, A, forming the distal margin of the medial space, MS,
and the notch-like nodus, N, on the anterior wing margin are distinct struc-
tures. Counting four longitudinal veins (including C) from the anterior wing
margin rearward at a level just distal to A identifies M1-3 in all Texas Zygopt-
era with petiolate wings. Ms is the first branch of the medial vein separating
from the M1-3 stem distal to the arculus. The quadrangle, Q, has as its basal
(nearest the body) side the posterior portion of A, and M4 is, completely or in
part, the forward margin.
Color patterns of the pterothorax and abdomen are alternating dark and
pale stripes or bands. Pale areas occur in a wide range of colors; the dark
stripes, rings, or bands are usually brown, black or metallic bronze. Pale
areas are largely absent in some species, and the dark pattern then consists of
metallic greens, blues or bronze. A pale antehumeral stripe borders the mid-
dorsal thoracic stripe on each side. A dark humeral stripe borders each ante-
humeral stripe laterally. The basic pattern in dorsal view appears in Figure 2
E. An additional dark stripe often occurs on each metapleural suture. The mid-
dle abdominal segments are predominantly pale or dark, with narrow dark
apical or pale basal transverse rings or bands on each segment. Pale seg-
ments may possess only a dark stripe on each dorsolateral side. These stripes
may be constricted about midsegment and, if the constriction is complete, two
elongate spots result; the postbasal and apical spots. The terminal abdominal
segments of many males are distinct with extensive, pale dorsolateral surfaces.
The head pattern typically consists of a pair of pale postocular spots, PS, often
a pale postoccipital bar, PB, and a variable facial pattern (Fig. 1 C). Much va-
riation exists on these basic patterns, and texts for specific keys give, where
needed, additional explanation. Color characters require recognition of general
and mature specimens. A recently emerged, winged adult is a general speci-
men, nd its exocuticle is still soft and the wings are fragile. Sexual maturity
develops after a variable period depending on the species, and frequently in-
volves a change in color. See Walker (1953) for an introduction to odonate mor-
phology.
Distributional data represent collective contributions of all sources listed
in the Acknowledgements, acceptable published records, and material col-
lected by the author. Each record of doubtful validity was omitted. Specimens
available for confirmation or determinations by an authority constitute the
locality records. Specific cases of questionable data appear in the Discussion.
Distribution data by county for each species follow their respective keys and a
cross-listing for records appears in the Appendix. This shows the parts of
Texas where additional data are necessary.

KEY TO TH E FAMILIES

1 a) Numerous antenodal and several quadrangle cross veins; wings not
petiolate (Fig. 3 A); wings pigmented with some black, brown or


Vol. XVI No. 2









)72 JOHNSON: TEXAS DAMSELFLIES 63

red in males, black, brown or nonpigmented in females-...........
................. Calopterygidae
b) Two antenodal and no quadrangle cross veins; wings petiolate (Fig.
3 B, C; Fig. 4 A, B); wings nonpigmented or with translucent brown
or amber............................................ 2
2 (1) a) Vein M3 separating from MI-2 nearer the arculus than nodus (Fig. 3
B); stigma nearly twice as long as wide ............. Lestidae
b) Vein M3 separating from MI-2 nearer the nodus than the arculus
(Fig. 3 C; Fig. 4 A, B); stigma not twice as long as wide ........ 3
3 (2) a) Vein Cu2 absent or rudimentary; vein Cul short forming anterior
border to only 3 (rarely 4) cells distal to arculus (Fig. 4 A, B) .....
.......................................... Protoneuridae
b) Vein Cu2 and Cul well developed, both enclosing several cells distal
to arculus (Fig. 3 C)........... . .... ..... Coenagrionidae


LESTIDAE

Two genera represent the family in Texas, Archilestes and Lestes. One
species of Archilestes exists in the state, with other species occurring west
and south into Mexico. Lestes occurs in both eastern and western hemi-
spheres, and six (possibly seven) species inhabit Texas.
Lestids are characteristic about lake margins, ponds, or slow streams
in frequently isolated colonies. They fly rather slowly and perch on emer-
gent vegetation or trees with half-spread wings. The latter habit is typical
of lestids, while other perched damselflies usually fold the wings together
over the body if not engaged in a behavioral display. Clear petiolated
wings separate lestids from calopterygids, and large body size together
with the spread-wing perching trait distinguish the group from most coen-
agrionids. The long slender coenagrionid, Teleallagma daeckii, has non-
lestid perching habits and pale bluish or tan body color.

KEY TO THE GENERA
1 a) Vein M2 branches from M1 one cell (occasionally 1.5 to 2 cells) distal
to the nodus; hind wing length greater than 33 mm .............
........................................... A rchilestes grandis
b) Vein M2 branches from M1 several cells distal to the nodus (Fig. 3
B); hind wing length less than 33 mm ................. Lestes


Archilestes Selys, 1862

Archilestes grandis is the largest damselfly in the Texas fauna (59-64
mm in body length), and the above key identifies both sexes. Figure 5 A
illustrates the male abdominal appendages. The species occurs in associa-
tion with both streams and ponds, otherwise its biology is unknown.
Kennedy (1915) described behavior and habitat for the western species,
A. californica.









64 BULLETIN FLORIDA STATE MUSEUM


DISTRIBUTION RECORDS FOR TEXAS
Bell, Brewster, Crosby, Dallas, Denton, Hays, Jeff Davis, Lubbock, Mon-
tague, Presidio, Travis, and Uvalde counties.

TABLE 2. BODY-LENGTH RANGES OF SPECIES IN MM GROUPED FROM SMALL TO LARGE.


Lestidae


L. disjunctus 34-44 mm
L. alacer 37-43 mm
L. forficula 38-40 mm
L. sigma 40-42 mm



Calopteryx
C. dimidiata 37-46 mm
C. maculata 37-57 mm


Neoneura aaroni 32-34 mm



A. bipunctulata 27-30 mm
A. alberta 28-31 mm
A. sedula 29-34 mm
A.fumipennis 29-34 mm
A. nahuana 32-34 mm
A. hinei 33-36 mm
A. immunda 33-36 mm
A. rhoadsi 33-37 mm


E. basidens 22-27 mm
E. geminatum 22-28 mm
E. dubium 24-27 mm
E. traviatum 27-32 mm
E. divagans 29-33 mm
E. civil 29-39 mm
E. praevarum 30-33 mm


Ischnura


I. posita 18-29 mm
I. verticalis 20-33 mm
I. denticollis 23-28 mm
I. demorsa 24-28 mm


Calopterygidae


Protoneuridae


Coenagrionidae
Argia


L. simplex 42 mm (one male)
L. vigilax 42-47 mm
L. inaequalis 46-51 mm
Archilestes grandis 59-64 mm



Hetaerina
H. americana 39-50 mm
H. titia 40-51 mm


Protoneura cara 35-37mm


Smaller Genera
Anomalagrion hastatum 20-25 mm Hesperagrion heterodoxum 26-29 mm
Nehalennia ,,It gr,...,ll 20-25 mm Teleallagma daeckii 39-44 mm
Telebasis salva 25-28 mm


A. apicalis 35-37 mm
A. tibialis 35-38 mm
A. translate 36-40 mm
A. plan 36-40 mm
A. munda 38-40 mm
A. barretti 41-43 mm
A. moesta 41-46 mm
A. lugens 44-49 mm


Enallagma


E. signatum 30-35 mm
E. vesperum 30-35 mm
E. carunculatum 30-36 mm
E. exsulans 30-38 mm
E. durum 31-38 mm
E. novaehispaniae 33-37 mm



I. kellicotti 26-32 mm
I. ramburii 29-35 mm
I. barberi 31-36 mm
I. prognatha 34-38 mm


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


Lestes Leach, 1815

MALES.-A dorsal view study of abdominal appendages reveals their
distinctive structural traits. The medial margins of the superior abdominal
appendages typically possess a basal tooth followed distally by a differ-
entiated apical tooth (pointed projection) or by one or two lobes
(broad, rather convex, non-pointed projections). These lobes have either
serrated or smooth medial margins. Shape (straight or sigmoid) and length
of the inferior abdominal appendages are useful characters. The superior
appendages are shorter than the inferior appendages (L. inaequalis),
or the apical lobe or tooth on the medial margins of each superior appen-
dage is anterior or posterior to distal ends of the inferior appendages. Ser-
ration on the medial margins of the superior appendages varies. A serrated
margin occurs on the basal lobe of L. vigilax while its apical lobe is
smooth, and the space between the basal and apical teeth of L. disjunctus
is smooth or with variable serration. The basal tooth of the superior appen-
dage in L. disjunctus varies from slender (as in Fig. 5D) to more blunt and
the inferior appendages of that species project straight to the rear or
toward the midline crossing each other to form a figure X. Lestes alacer
is variable in the shape of the serrated lobe on the medial margin of the
superior appendages, and the inferior appendages project straight toward
the rear or slant slightly toward the midline. This variation in L. alacer
raises a taxonomic problem with L. simplex. The description of both L.
alacer and L. simplex appeared in the same paper by Hagen (1861) and
Texas records of L. simplex exist (Calvert, 1901-1908; Williamson, 1914).
L. K. Gloyd examined the Texas specimens (from Clifton, Bosque County)
determined as L. simplex by Williamson (1914) and states (Pers. comm.,
1969) "Specimens from Clifton, Texas, appear to me to be same as L. alacer
from the type locality." Figure 5 H illustrates a specimen from Mexico de-
termined as L. simplex by P. P. Calvert. As shown in that figure, the major
differences from L. alacer are shape of the serrated lobe and stronger slant
of the inferior appendages toward the midline. This specimen possesses a
broad dark band oriented obliquely across each side of the thorax just
above the metapleural suture. Such bands are absent or much less de-
veloped in L. alacer. Variation in L. alacer approaches L. simplex char-
acters, and the female of L. simplex is unknown. All material available to
the author or checked by authorities failed to provide Texas specimens of
L. simplex, and its status (at least in Texas) remains questionable. Body
length ranges, grouped from smallest to largest species including both
sexes, appear in Table 2.
The largest Texas specimen of L. disjunctus seen by the author was 39
mm; the larger specimens are more northern records (Walker, 1953).
FEMALES.-Lestes females have indistinct structural differences, thus
diagnosis largely uses color patterns. Females of L. alacer, L. forficula,
and L. sigma are more modified in appearance by age than other Texas
species. Females of L. sigma are rather nondescript individuals and may per-


1972








66 BULLETIN FLORIDA STATE MUSEUM


haps be confused with females of L.forficula. Females of L. sigma and L.forfi-
cula have generally more pruinescence than other species.

KEY TO THE MALES
1 a) In dorsal view, inferior abdominal appendages greater in length than
superior abdominal appendages (Fig. 5 B) ............. inaequalis
b) In dorsal view, superior abdominal appendages greater in length
than inferior abdominal appendages (Fig. 5 C-H) ............ 2
2 (1) a) In dorsal view, inferior abdominal appendages sigmoid in shape,
slender in apical half, and with apices divergent (Fig. 5 C) ... sigma
b) In dorsal view, inferior abdominal appendages not sigmoid, may be
slender or stout in shape, apices not divergent (Fig. 5 D-H) ...... 3











B C D


A ^
INAEQUALIS SIGMA DISJUNCTUS
GRANDIS












E F G H

VIGIL AX FORFICULA ALACER SIMPLEX
FIGURE 5. Dorsal view of left superior and inferior male abdominal appen-
dages in Archilestes grandis and Lestes species following sequence of deter-
mination in key.


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


3 (2) a) In dorsal view, medial margins of superior abdominal appendages
with a distinct basal tooth and a more blunt apical tooth, the two
teeth separated by variable degree of serration (Fig. 5 D); labrum
pale blue ........................................ disjunctus
b) In dorsal view, medial margins of superior abdominal appendages
with a distinct basal tooth, no differentiated apical tooth, but mar-
gins with one serrated lobe or two lobes-one smooth, one serrated-
distal to basal tooth (Fig. 5 E-H); labrum bluish in L. vigilax .... 4
4 (3) a) In dorsal view, medial margins of superior abdominal appendages
with two lobes distal to basal tooth-the more basal lobe somewhat
serrated, the distal lobe smooth; inferior abdominal appendages
long and slender, extending posteriorly beyond distal lobe of superior
appendages (Fig. 5 E); dorsum of pterothorax with metallic green
stripe ................................................ vigilax
b) In dorsal view, medial border of superior abdominal appendages
with one differentiated, distinctly serrated lobe distal to basal
tooth (Fig. 5 F-H).......................................... 5
5 (4) a) In dorsal view, inferior abdominal appendages long, extending be-
yond posterior level of serrated lobes on medial margins of superior
appendages (Fig. 5 F); narrow metallic green stripes on dorsum of
pterothorax but obscured by age with pruinescence..... forficula
b) In dorsal view, inferior abdominal appendages shorter, extending
approximately to posterior level or less of serrated lobes on medial
margins of superior appendages (see text) (Fig. 5 G,H); broad black
or dark bronze stripe on dorsum of pterothorax, often obscured by
pruinescence ..................... ....... ................. 6
6 (5) a) In dorsal view, medial margins of superior abdominal appendages
with well-developed serrated lobe distinctly differentiated on its
posterior end; inferior abdominal appendages rounded at apices, not
curved mesially, (Fig. 5 G), see text. ...................... alacer
b) In dorsal view, medial margins of superior abdominal appendages
with serrated lobe not distinctly expanded and not terminating pos-
teriorly in a distinct notch; inferior abdominal appendages slightly
slanted towards each other, their apices bluntly pointed (Fig. 5 H),
see text.......................................... ... simplex

KEY TO THE FEMALES
1 a) Middorsal and humeral dark stripes only slightly distinguishable
from grayish or yellowish-brown of pterothorax (irregular, elongate
greenish spots may overlay middorsal and humeral stripes but lost
with age; distinct blackish spot in posterior half of each antehumeral
area, and an elongate blackish spot parallel and posterior to each
humeral suture (latter spot may be obscured early by pruinescence
and whole dorsal thoracic surface may become blackish with old
age); abdomen predominantly dark brown without strongly con-
trasting pale areas...................................... sigma
b) Distinct and wide, dark middorsal and humeral thoracic stripes
bordering narrow, pale antehumeral stripes, or only distinct, wide,
dark middorsal thoracic stripe, humeral stripes absent, or thoracic
dorsum pale brown to gray and black spots not present in ante-
humeral areas but two narrow metallic green stripes on each side,









68 BULLETIN FLORIDA STATE MUSEUM


one stripe near the middorsal carina, another stripe (smaller and
more irregular) just posterior to each humeral suture; abdomen may
be predominantly brown or have contrasting dark and pale pattern
..................... ... ................................ 2
2 (1) a) Dorsum of pterothorax brown to gray without typical middorsal and
humeral dark stripes, two narrow metallic greenish stripes on each
side of thorax, one stripe just lateral and parallel to the middorsal
carina, one stripe (shorter and more irregular) posterior and paral-
lel to the humeral suture.............................. forficula
b) Dorsum of pterothorax with wide distinct dark middorsal stripe;
no combination of above characters ........................... 3
3 (2) a) Middorsal stripe wide and dark bronze to blackish in color, not reach-
ing laterally to humeral suture, no dark humeral stripe, remainder of
thoracic sides pale tan to brown, devoid of dark pattern lateral or
posterior to the humeral sutures..........................alacer
b) Middorsal and humeral stripes wide and dark bronze to greenish in
color, antehumeral areas typically more narrow than either of above
stripes, the humeral dark stripes often somewhat paler than mid-
dorsal stripe, occurring largely posterior to the humeral sutures .. 4
4 (3) a) Dark stripes on thoracic dorsum typically metallic green; hind wing
length 27 mm or greater; distance across compound eyes usually 6
mm or greater...................................... inaequalis
b) Dark stripes on thoracic dorsum typically dull bronze or greenish-
black; hind wing length usually less than 27 mm; distance across
compound eyes usually less than 6 mm. ......................... 5
5 (4) a) A dense cross vein reticulation developed between the principal long-
itudinal veins near apical margin of each wing, numerous small cells
produced typically occurring as a double row about the apex .. cigilax
b) No or very few extra cross veins developed between principal veins
that converge at the wing's apex, the associated cells becoming
gradually smaller, extra interpolated cells few and typically limited
to wing margin ..................................... disjunctus
Female L. simplex unknown, see text.




DISTRIBUTION RECORDS FOR TEXAS
Lestes alacer: Blanco, Bosque, Brazos, Caldwell, Crosby, Gonzales, Hill,
Jeff Davis, Kimble, Lubbock, Matagorda, Reeves, and San Patricio counties.
Lestes disjunctus: Aransas, Blanco, Brazos, Brewster, Colorado, Dallas,
Hardin, Harris, Hunt, Jeff Davis, Jim Wells, Lamar, Lubbock, Montgomery,
Nacogdoches, San Jacinto, San Patricio, Tarrant, Travis, Williamson, and Wil-
son counties.
Lestes forficula: Bexar, Brazos, Cameron, Hidalgo, Kleberg, San Patricio,
and Starr counties.
Lestes inaequalis: Angelina and Harrison counties.
Lestes sigma: Cameron, Gonzales, Kleberg, San Patricio, Starr, and Vic-
toria counties.
Comments on a questionable species for Texas, L. congener, appear in the
Discussion.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


CALOPTERYGIDAE

Two genera, Calopteryx and Hetaerina, represent the family in Texas.
Calopteryx species occur widely in the northern hemisphere, and Hetaer-
ina species inhabit only North and South America, reaching their highest
diversity in southern latitudes. Broad, non-petiolated wings and body
colors of brown, metallic greens, blue and bronze distinguish these dam-
selflies. Sexual dimorphism exists in wing pigments; males possess the
brighter colors and females usually have much less wing pigment.

KEY TO THE GENERA
1 a) Median space without cross veins; coloration metallic green or blu-
ish, little or no pale area; wing pigments blackish, red colors absent,
stigma absent in males, present and distinctly white in females ....
........... ....................................... Calopteryx
b) Median space with several cross veins; body pattern with distinct
pale areas adjacent to dark metallic stripe or diffuse brownish body;
male fore wings possess basal red area with or without adjacent
brownish area, small stigma usually in both sexes ...... Hetaerina

Calopteryx Leach, 1815

Color pattern differences given in the keys also allow field identifica-
tion. Female C. maculata have fully pigmented wings as the male, but
they are usually darker in the apical fourth and possess a distinct white
stigma or pseudo-stigma (Tillyard, 1917). Teneral C. maculata have only
smoky-gray wing colors. Female C. dimidiata occur in two forms; the
wings have little to no pigment or the apical fourth is pigmented similar
to the male pattern (Johnson and Westfall, 1970). Both female morphs
have distinctive white stigmas.

KEY TO THE MALES
1 a) Wings translucent brown (in general specimens) to opaque black for
full length; ventral surface of abdominal segments 9 and 10 whitish;
inferior abdominal appendages two-thirds or greater length of su-
perior abdominal appendages (Fig. 6 C) ............. maculata
b) Wings with basal five-sixths to three-fourths area clear, apical
portion distinctly marked with brown or black pigment (Fig. 6 G);
ventral surface of abdominal segments 9 and 10 black; inferior ab-
dominal appendages less than two-thirds length of superior ab-
dominal appendages (Fig. 6 D) ........................ dimidiata

KEY TO THE FENIALES
1 a) Wings possessing brownish-black pigment over full length, often
with greater intensity in apical fourth; ventrolateral surface of pter-
othorax blackish ..................................... maculata
b) Wings devoid of brownish-black pigment, or pigment restricted to
apical fourth or less (Fig. 6 H); ventrolateral surface of pterothorax
pale................................................ dimidiata


1972









70 BULLETIN FLORIDA STATE MUSEUM


AMERICANA TITIA


TITIA


MACULATA DIMIDIATA


FEMALES


D/MIDIATA
MALE






FEMALE





FIGURE 6. A-D, dorsal (anterior ends uppermost) views of left superior and
inferior abdominal appendages in calopterygids; left lateral views of thoracic
patterns of females of the Hetaerina titia complex; E, H. tricolor; F. H. titia;
wing patterns (fore wing only) in Calopteryx dimidiata: A, male; B, female.


Both species are stream forms with C. dimidiata having less ecological
tolerance. See Johnson and Westfall (1970) for references to ecology.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


DISTRIBUTION RECORDS FOR TEXAS
Calopteryx dimidiata: Hardin and San Jacinto counties.
Calopteryx maculata: Anderson, Angelina, Bastrop, Bowie, Cherokee,
Collin, Dallas, Denton, Grayson, Gregg, Grimes, Hemphill, Houston, Marion,
Montgomery, Nacogdoches, Robertson, Rusk, Shelby, Walker, and Wood
counties.


Hetaerina Hagen, 1853

Sequential development of wing patterns during general stages may
lead to incorrect determinations. In H. titia the color patterns are pale
brown in both wing pairs soon after emergence, and only hind wing spots
in males of H. americana are initially brown. Both fore and hind wing
spots become deep red with age in H. americana, and the fore wings of
mature H. titia males have both red and brown areas. Hind wing spots in
male H. titia remain brown but veins within the spots may be red. The
wings of H. titia females are rarely clear, but are more typically diffuse
brown. The wings of H. americana females have no wing pigment or pos-
sess diffuse brown to orange basal spots. The female condition is not an
age effect, and the variation in H. americana is similar to that mentioned
for Calopteryx dimidiata, possibly representing a sex-limited dimorphism.
KEY TO THE MALES
1 a) Red spot at base of fore wing bordered distally by no pigment or
brownish-black area extending variable distance toward wing tip,
hind wing brown and varying from basal spot to entire wing (Fig. 7);
pigmentation paler in general specimens .......... titia complex
b) Red spot at base of fore and hind wings, may reach distally to nodus;
apical wing areas nonpigmented; red color limited to fore wing in
general specimens with hind wing spot brown ........ americana

KEY TO THE FEMALES
1 a) Abdomen brown on ventrolateral surface, slightly lighter, if any, in
color than dorsum; dorsum of head and abdomen dark brown; thorax
brown with broad metallic green stripe on either side of median
carina, or stripe isolated into two elongated spots (Fig. 6 E, F)..........
................. .. ................... ....... titia com plex
b) Abdomen pale-colored on ventrolateral surface, and distinctly con-
trasting with metallic green dorsum; dorsum of head and abdomen
metallic green, latter usually with a pale basal ring per segment;
dorsum of thorax with broad metallic green stripe on either side of
median carina ................................... . .americana
Hetaerina titia as recognized in the above key is a highly polymor-
phic species or includes two species, H. titia and H. tricolor. These two
taxa now appear as synonyms; however, several observations are incon-
sistent with this interpretation (Johnson, 1963). The male abdominal ap-
pendages are highly variable in H. americana (Calvert, 1901-1908). Fig-
ure 6 A and B illustrates appendages of both H. americana and H. titia.








72 BULLETIN FLORIDA STATE MUSEUM


se D
FIGURE 7. Selected male patterns in left fore and hind wings of the Hetaerina
titia complex; H. tricolor characterized by type A.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


DISTRIBUTION RECORDS FOR TEXAS
Hetaerina americana: Baylor, Bexar, Blanco, Bosque, Brazos, Brewster,
Caldwell, Cherokee, Childress, Colorado, Comal, Cooke, Crosby, Dallas, Den-
ton, Fayette, Gillespie, Goliad, Gonzales, Gregg, Grimes, Hays, Hill, Jeff Davis,
Jim Wells, Kendall, Kerr, Kimble, Limestone, Llano, Lubbock, Medina, Men-
ard, Palo Pinto, Pecos, Presidio, Randall, Real, Reeves, Robertson, San Ja-
cinto, San Patricio, Sutton, Travis, Uvalde, Val Verde, Victoria, Williamson,
Wilson, and Zavala counties.
Hetaerina titia (and tricolor): Angelina, Bexar, Bosque, Brazos, Caldwell,
Colorado, Comal, Dallas, Denton, Fayette, Goliad, Gonzales, Grimes, Guada-
lupe, Hays, Jackson, Jim Wells, Kendall, Kimball, McLennan, Polk, Presidio,
San Jacinto, San Patricio, Robertson, Travis, Uvalde, Victoria, and Webb counties.
Comments on two questionable species for Texas, H. sempronia and H.
vulnerata, appear in the Discussion.

PROTONEURIDAE
Two genera, Protoneura Selys 1857 and Neoneura Selys 1860, repre-
sent the family in Texas, each genus by one species. Higher diversity char-
acterizes the family in the neotropical region. The key gives characteristic
venation and diagnostic traits in male appendages and female mesostig-
mal plates. The sexes of P. cara have similar stripe and color patterns
(pale orange colors bordered by bronze stripes). Initially, P. cara appears
similar to Enallagma signatum while in flight. Neoneura aaroni males
develop a copperish-red thoracic dorsum at maturity, while their females
are light brown without distinctive pattern. Immature general males and
all females of N. aaroni are similar to several species while in flight.

KEY TO THE SPECIES
1 a) One subquadrangular cell in each wing (Fig. 4 A); well developed
dark bronze stripe pattern and light yellow to orange antehumeral
areas in both sexes; superior abdominal male appendage in lateral
view as long or slightly longer than inferior appendage (Fig. 16 I);
females' mesostigmal plates' medial corners raised and curved an-
teriorly (Fig. 11 N, 0); body length 35-37 mm ....... Protoneura cara
b) Two subquadrangular cells in each wing (Fig. 4 B, xi and x2); ma-
ture males with copperish-red on dorsum of mesothorax and an-
terior of head and face; females light tan with reduced stripe pattern;
superior abdominal male appendage in lateral view slightly shorter
than inferior appendage (Fig. 16 J); females' mesostigmal plates
possessing posteriorly projecting lobes (Fig. 11 M); body length
32-34 mm................................... Neoneura aaroni
Stream habitats are typical for both species. Williamson (1914) de-
scribed the habitat of N. aaroni as ". . deep pools in small streams with
drift or overhanging bushes near at hand."

DISTRIBUTION RECORDS FOR TEXAS
Neoneura aaroni: Caldwell, Goliad, Gonzales, Medina, Nueces, San Pa-
tricio, and Victoria counties.
Protoneura cara: Hidalgo, Kendall, Medina, Uvalde, and Val Verde coun-
ties.








74 BULLETIN FLORIDA STATE MUSEUM


COENAGRIONIDAE

Eight genera including forty species represent the family in Texas.
Anomalagrion, Hesperagrion, and Teleallagma are monotypic genera;
Nehalennia and Telebasis occur in the state with a single species in each
genus; Argia, Enallagma, and Ischnura have 15, 12, and 8 species respec-
tively. Enallagma and Ischnura have their highest species diversity in
North America although both genera are almost cosmopolitan in distri-
bution. Nehalennia occurs in North America and the palearctic, and
the remaining five genera have neotropical and nearctic distributions.
A separate generic key for males and females avoids unduly long key
couplets. An external exudate, pruinescence, having a white or bluish-
gray color, may obscure body patterns in old individuals. A drop of ace-
tone or alcohol temporarily reduces this effect.
Female determination in the Coenagrionidae relies largely on the
mesostigmal plates. A generalized dorsal view of mesostigmal plate mor-
phology appears in Figure 2 E following Walker (1953). The middorsal
thoracic carina, MC, bifurcates at the anterior end of the mesothorax and
typically terminates into ridges or flattened plates (rami), RM. This mid-
dorsal area consists of a median pit, MPT, between the two rami of the
dorsal carina, an anterior carina, ACF, forming a forward, transverse mar-
gin and two lateral carinae, LCF. These carinae or ridges collectively
constitute the frame. A mesostigmal plate, MP, occurs laterally on each
side of the frame. A mesothoracic spiracle (mesostigma) occurs inconspi-
cuously below the anterior margin of each plate. The plates may have
lobes projecting rearward from the posterior margins, and ridges oriented
transversely or obliquely across the plate, however, such distinctive struc-
tures are often absent. Color pattern and size identify many females; how-
ever, the structural characteristics of the plates give more reliable determ-
inations. Study of the plates requires a strong light source and at least a 20
X magnification. In addition, a forward flexure of the head and prothorax
is helpful, as the hind lobe of the prothorax typically overlies the plates.
Species identified in these keys are members of monotypic genera
or the only representatives of their genus in Texas. Additional data ap-
pear for such species under Smaller Genera.

KEY TO THE GENERA: MALES
1 a) Majority of spines on 2nd and 3rd tibiae long, distance between
spines approximately one-half of spine length (Fig. 1 E); dorsoapical
margin of abdominal segment 10 with torifer and usually distinct tori
(Fig. 2 A, B) ........................................... Argia
b) Majority of spines on 2nd and 3rd tibiae short, distance between
spines greater than one-half of spine length (Fig. 1 D); dorsoapical
margin of abdominal segment 10 without torifer and tori but may
possess bifid prominence or spine, if above tibial spines long and
thoracic dorsum metallic green then dorsoapical margin of abdom-
inal segment 10 serrated and body length less than 30 mm ..........2


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


2 (1) a) Thoracic dorsum solid metallic green to bronze and abdominal dor-
sum predominantly greenish-black with some blue on segments 8, 9,
and 10; dorsoapical margin of abdominal segment 10 serrated, not
elevated into bifid prominence and abdominal appendages in lateral
view as in Fig. 16 K; ave. body length 24-27 mm .....................
...................................... Nehalennia integricollis
b) No such combination of characters .......................... .3
3 (2) a) Abdominal length 34 mm or greater; Cu2 terminating near midway
point between nodus and origin of M2; petiole of wing usually extends
distally to anal crossing; dorsal thoracic stripe narrow and humeral
stripes reduced to elongate spots; superior abdominal appendages in
lateral view with distinct ventrally-directed lobe (Fig. 17 H); ave.
body length 40-44 mm. ....................... Teleallagma daeckii
b) No such combination of characters ........................... 4
4 (3) a) Thoracic dorsum with black dorsal stripe having distinct lateral
tooth in posterior half and finely divided by pale-colored carina;
antehumeral areas and thoracic sides reddish brown, abdominal
dorsum red; abdominal appendages in lateral view as in Fig. 17 I;
ave. body length 26-30 mm. ..................... Telebasis salva
b) No such combination of characters ........................... 5
5 (4) a) Abdominal segment 10 without spine on dorsoapical margin and su-
perior abdominal appendage distinctly bifid, upper lobe curved dor-
soposteriorly reaching above margin of 10th segment and lower arm
transversely expanded and directed ventrally (Fig. 17 F, G); anter-
ior margin of stigma usually shorter than posterior margin; see text
for color pattern variation; ave. body length 29-33 mm ............
......... ........................ Hesperagrion heterodoxum
b) No such combination of characters ........................... 6
6 (5) a) Stigma of fore wing removed from wing margin (Fig. 4 E); abdom-
inal segment 10 elevated into spine, superior abdominal appendage
with distinct dorsoposteriorly directed arm in lateral view (Fig. 17
J); ave. body length 23-27 mm ............... Anomalagrion hastatum
b) Stigma of fore wing not removed from wing margin; abdominal seg-
ment 10 and appendages not as in Fig. 17 J .....................7
7 (6) a) M2 separating from M1-2 near 5th and 4th postnodals or beyond in the
fore and hind wings respectively; abdominal segment 10 not elevated
on dorsoapical margin; stigma similar in fore and hind wings........
................................................. Enallagma
b) M2 separating from MI-2 near the 4th (or 3rd) and 3rd (or 2nd) post-
nodals in the fore and hind wings respectively; abdominal segment
10 may or may not have elevation on dorsoapical margin developed
into spine-like process; stigma color may differ between fore and
hind wings in well-matured individuals. ................... .... 8
8 (7) a) Abdominal segment 10 elevated on dorsoapical margin into spine-
like process bifid at tip (Fig. 16 A-D, F); if process absent or poorly
developed, inferior abdominal appendage extends posteriorly beyond
apical level of superior appendage and latter appendage with dis-
tinct ventrally-directed lobe (Fig. 16 E); stigma may differ in fore
and hind wings ....................................Ischnura
b) Abdominal segment 10 not elevated and inferior abdominal appen-
dage shorter than superior appendage, or if not, superior abdominal








76 BULLETIN FLORIDA STATE MUSEUM


appendage with ventrally-directed lobe; stigma similar in fore and
hind wings ........................................ Enallagma

KEY TO THE GENERA: FEMALES
1 a) Majority of spines on 2nd and 3rd tibiae long, distance between spines
approximately one-half of spine length (Fig. 1 E); if body length
less than 30 mm, then pale antehumeral stripe bordered by distinct
dark middorsal and humeral stripes. ................... .. Argia
b) Majority of spines on 2nd and 3rd tibiae short, distance between
spines greater than one-half of spine length (Fig. 1 D); if spines
longer, then body length less than 30 mm and thoracic dorsum metal-
lic green without antehumeral pale stripes. ..................... 2
2 ( 1) a) Thoracic dorsum solid metallic green to bronze, abdominal dorsum
predominantly greenish-black; abdominal segment 8 without vulvar
spine, such as Fig. 2 C; mesostigmal plates in dorsal view as in Fig.
15 V; ave. body length 24-27 mm .......... Nehalennia integricollis
b) No such combination of characters ........................... 3
3 ( 2) a) Abdominal length 34 mm or greater; Cu2 terminating near midway
point between nodus and origin of M2; petiole of wing usually ex-
tends distally to anal crossing; stripe pattern reduced or absent, pale
colors brownish; abdominal segment 8 with vulvar spine, such as
Fig. 2 D ................................... Teleallagma daeckii
b) No such combination of characters ........................... 4
4 ( 3) a) Thoracic dorsum with dark dorsal stripe having distinct lateral tooth
in posterior half and finely divided by pale-colored carina; antehu-
meral areas, thoracic sides, and abdominal dorsum brown; hind lobe
of prothorax with two tooth-like projections directed anteriorly (Fig.
11 J, K); ave. body length 26-30 mm............... Telebasis salva
b) No such combination of characters .......................... 5
5 ( 4) a) Anterior lateral margin of mesostigmal plates raised into a distinct
ridge, truncated sharply on medial end (Fig. 15 W); thoracic dorsum
completely pale, or with well-developed middorsal stripe, humeral
stripes absent or faint, or dorsum predominantly dark with antehu-
meral areas separated into two pale spots on each side; black not
predominating on all abdominal segments 3-9, see additional color
notes in text; stigma usually shorter on anterior margin than on pos-
terior margin; M2 separation typically near 5th and 4th postnodals in
fore and hind wings respectively; fore wing length 19-21 mm or
greater; ave. body length 29-33 mm. ...... Hesperagrion heterodoxum
b) No such combination of characters ........................... 6
6 ( 5) a) Sternum of abdominal segment 8 with a vulvar spine (Fig. 2 D).....7
b) Sternum of abdominal segment 8 without a vulvar spine (Fig. 2 C)..
. .. . . . . . . . . . . . . . . . . . . . . . . . . . . 13
7 ( 6) a) Distinct dorsal and humeral blackish thoracic stripes bordering pale
antehumeral stripes; antehumeral colors include light tan, blues,
and green ......................... .... .. ............. 8
b) Distinct dorsal thoracic stripe present as solid line or finely divided
on the carina, or absent; pale thoracic area lateral to dorsal stripe
(if present) orange, brown, or bluish; never with distinct humeral
stripe (latter may be represented by small isolated spots) .......11
8 ( 7) a) M2 separating from M1-2 near the 5th and 4th postnodals or beyond


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


on the fore and hind wings respectively............... Enallagma
b) Mz separating from M1-2 near the 4th and 3rd (or 2nd) postnodals
on the fore and hind wings respectively. ...................... 9
9 (8) a) Body length less than 30 mm; mesostigmal plates with low even
ridges on anterior and posterior margins, slightly expanded at pos-
terior lateral corner (Fig. 15 U); postocular spots obscured and no
dark stripe on metapleural suture........... Anomalagrion hastatum
b) Body length 30 mm or greater, or one or more of the following traits
present: mesostigmal plates raised vertically on edge or with distinc-
tive ridges (Fig. 15 J, N, Q, S); postocular spots distinct; dark stripe
on metapleural suture; prominence on mesothorax just posterior to
each mesostigmal plate (Fig. 15 B); humeral stripe divided longi-
tudinally by inserted pale line ............................... 10
10 ( 9) a) Postocular spots triangular in shape occupying most of postocular
area and their apices directed anteriorly, or spots circular in shape
and abdominal segment 8 partly blue; mesothoracic dorsum poster-
ior to mesostigmal plates without raised knobs; dark humeral stripe
(if present) never divided longitudinally by pale line........ Ischnura
b) Postocular spots elongated transversely with or without connection
to pale postocular bar (bar may be absent), if spots circular in shape
abdominal segment 8 largely black; either raised knobs on mesothor-
acic dorsum posterior to mesostigmal plates, or humeral stripes long-
itudinally divided by inserted pale line ................... Enallagma
11 ( 7) a) M2 separating from M1-2 near the 5th and 4th postnodals or beyond
on the fore and hind wings respectively. ..................... 12
b) M2 separating from M1-2 near the 4th and 3rd (or 2nd) postnodals on
the fore and hind wings respectively. ................... ... 13
12 (11) a) Dorsum of thorax with dorsal black stripe bordered by orange later-
ally which becomes dark brown or greenish with age (dorsal stripe
obscured in older individuals); transverse postocular spots and bar
broadly confluent with pale of rear head area but spots, bar and
rear head areas obscured by black at early age.............. Ischnura
b) Dorsum of thorax with or without dorsal black stripe, pale areas lat-
eral to stripe pale blue or bluish-brown; distinct bluish postocular
spots with well defined borders, often with pale postocular bar;
these patterns not obscured with age ................. Enallagma
13 ( 6) a) Mesostigmal plates with low even ridges on anterior and posterior
(11) margins, slightly expanded at posterior lateral corner (Fig. 15 U);
body length less than 30 mm; dark middorsal thoracic stripe; hum-
eral stripe faint, or absent; pale antehumeral stripe not divided into
spots; abdominal segments 8, 9 and 10 black; no dark stripe on meta-
pleural suture. ......................... Anomalagrion hastatum
b) Mesostigmal plates with one of following attributes present; raised
vertically on edge, with transverse diagonal ridges, or with distinct
ridge development on anterior or posterior margins (Fig. 15 N-S); if
these traits absent (such as Fig. 15 M, T) one or more of the following
traits present: body length 30 mm or greater, antehumeral pale areas
divided into 2 spots (requires close examination in mature specimens
due to presence of pruinescence), or abdominal segments 8, 9 and 10
largely pale and dark stripe on metapleural suture.......... Ischnura








78 BULLETIN FLORIDA STATE MUSEUM


Argia Rambur, 1842

MALES.-Reliable species diagnosis in males requires examination of
abdominal appendages and the tenth abdominal segment. Diagrams of
the terminal abdominal morphology for a typical male appear in Figure
2 A, B. A depressed V or U-shaped area with apex pointing anteriorly and
bordered by raised ridges occurs in the dorsal surface of the tenth abdom-
inal segment and forms part of its apical margin. The area is the torifer,
tor. The posterior border of the torifer separates into two ridges by an in-
dentation at the midline. These ridges are perpendicular or oblique to
the midline, and each ridge bears a pad-like structure, the torus, t. The tori
in dorsal view are either circular or elongate structures. The elongate
tori are swollen pads or form thin, rim-like borders of the torifer ridges.
If the tori are narrow, posterior borders to convex torifer ridges, they are
obliquely rather than transversely elongate. Torus shape and transverse
distance between medial corners of the two tori are useful in species
identification. Torus width refers to its transverse axis, and length is the
distance through the pad from front to rear. Tori exist with width and
length approximately equal or with width exceeding length. In dorsal
view, shape of the tori varies slightly in some species. For example, each
transversely elongate torus shown for A. plana in Fig. 9 P has the two
transverse borders essentially parallel. In other specimens, each torus is
slightly ovoid-shaped. Construction of the key takes these variations into
consideration. The tori are whitish or similar to the color of the torifer
ridge, a change apparently associated with aging. Figures show the tori
in solid black.
Between the torus-bearing ridges on the torifer's posterior surface at
the midline are two swellings or tubercles, the toreale, to. Description of
the toreale generally refers to their shape seen in dorsal view. The toreale
project posteriorly, are typically small, and their apical tips may termin-
ate anterior, on line with, or posterior to the rear margin of the tori.
The relative length of the toreale measured in this fashion has value in spe-
cies identification. The toreale of A. bipunctulata are disproportionately
long, reaching rearward for at least two-thirds length of the superior ab-
dominal appendage, SA. The toreale are whitish or similar to the torifer's
color.
The inferior abdominal appendage, I A, in lateral view bears a dor-
sally-directed superior lobe, SL, and in all but four Texas species (A.
moesta, A. lugens, A. rhoadsi, and A. bipunctulata) an inferior lobe, IL.
The latter lobe arises from the appendage's lower posterior margin, and
its shape in lateral view has diagnostic value. It is blunt (deeper than long
at its base) or tapers to the apex (as long or longer than depth at base). On
the dorsal or dorsolateral surface of the inferior abdominal appendage
and just anterior to base of superior lobe, a tooth or blunt prominence
occurs in A. rhoadsi, A. hinei, and A. tibialis. In dorsal view, the inferior
lobes of A. sedula typically curve inward; however, if the appendages are


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


widely spread, they appear to point directly rearward. Other Texas argi-
ans have lobes pointing rearward and the medial margins taper outward.
The superior lobe in lateral view is blunt or ends in a pointed tooth-like
tip; however, this portion of the appendage varies (in lateral view) in
some species. The lateral profile view of the inferior abdominal appen-
dage in A. nahuana, A. plan and A. immunda in particular may vary
from figures given for these species. Their inferior lobe appears more blunt
in some individuals than others; however, the key should effectively sep-
arate these species bearing in mind this possible variation.
The apical and medial margins of the superior abdominal appendage
in dorsal view are convex, bifid, etc. These appendages in some species
are difficult to see clearly when intact on the specimen. An oblique dor-
solateral view facing the specimen from a posterior angle often reveals
the most definitive structure. Figures illustrate appendages in this position
if the key uses the trait. The superior abdominal appendages have less
diagnostic value in lateral view; however, appendages in A. munda and,
to a lesser extent, A. hinei when viewed laterally reveal portions of their
ventral surfaces. Appendages of other species when viewed laterally reveal
ventral surfaces only when in atypical positions. A tooth, usually black,
typically occurs at the apical or subapical surface of the appendage and a
blunt process may arise from the medial surface. The apical tooth is later-
ally visible or hidden behind the superior lobe of the inferior abdominal
appendage, depending on articulation of the appendages.
Argian species occur over a range of 20 mm in body length, and this
trait is helpful in diagnosing species near the limits of length distribution.
Body length ranges, grouped from smallest to largest species including
both sexes, appear in Table 2. Adult size probably varies with environ-
mental conditions and duration of the larval stage; consequently, small
variations from the measurements in Table 2 may occur.
Color and stripe patterns are not well-suited for species determina-
tion in an argian fauna of 15 or 16 species. An experienced observer can
identify individuals in the field with such characters; however, color
tones are particularly unreliable in a key. Colors fade in museum speci-
mens, have reversible changes in life for some species and pruinescence
effect adds further differences. Stripe patterns show little change if any
with aging (except through pruinescence); nevertheless, limited taxo-
nomic value lies in thoracic stripes due to intraspecific variability. The ab-
dominal stripe and spot patterns are less variable.
Distinct dark middorsal and humeral stripes occur in A. bipunctu-
lata, A. rhoadsi, A. moesta, A. tibialis, A. translate and A. sedula. Their
humeral stripes are broader than the suture and the stripe usually forks
somewhat at its posterior third in A. tibialis, A. sedula, and A. rhoadsi,
and either forks or encloses a pale spot in A. bipunctulata. A forked stripe
bifurcates as its name implies. The humeral stripe of A. moesta is largely
posterior to the suture and often obscured by pruinescence. A pale stripe
occurs within the posterior half of the humeral stripe in A. translate. The








80 BULLETIN FLORIDA STATE MUSEUM


middorsal stripe is as wide or wider than the antehumeral stripe in all
above species excepting A. tibialis.
The middorsal stripe is usually narrower or no wider than the pale
antehumeral area in A. plana, A. hinei, A. munda, A. immunda, A. fiimi-
pennis and A. nahuana. Humeral stripes of these species are narrow with
a posterior bifurcation or an interrupted stripe results in the posterior
third of its length. An interrupted stripe is broad at its anterior half, con-
fined to the humeral suture or lost in the posterior two-thirds and reap-
pears as a spot at the upper end of the suture. Forked stripes typically oc-
cur in A. immunda, A. fimipennis and A. nahuana, while interrupted or
terminated stripes occur in A. munda, A. hinei and A. plan.
The following three species are intermediate or distinct from above
groups. The dark middorsal and humeral stripes of A. barretti are distinct
and narrower than the pale antehumeral areas. The humeral stripe is not
forked or interrupted yet margins of the stripes are not parallel. The dark
middorsal stripe of A. apicalis is absent or confined to the carina, and the
humeral stripe exists as an anterior elongate spot; however, exceptional
individuals of A. apicalis may occur. Such males of A. apicalis have a full-
length humeral stripe of varying width; the females may have both broad
middorsal and humeral stripes. In Florida, this variation in A. apicalis
correlates with geographic distribution (Johnson and Westfall, 1970).
Such variants were absent in Texas material examined by the author;
however, patterns approaching the Florida types rarely occur in Texas
(Gloyd, per comm.). The dark middorsal stripe of A. lugens is no wider
than the carina, and a dark line occurs laterally in each antehumeral
area becoming confluent with the middorsal stripe at the latter's posterior
end. The humeral stripe forks.
Antehumeral areas are typically bluish in A. rhoadsi, A. sedula, A.
nahuana, and A. barretti; violet or purple in A. lugens, A. tibialis, A. tran-
slata, A. immunda, A. fumipennis; and bluish-violet in A. hinei and A.
munda. This area is blue or violet in A. plan and A. apicalis, cream or
tan colored in A. bipunctulata and A. moesta, and usually obscured in
the latter. Lower sides of the pterothorax have paler colors in all species.
Pale abdominal colors are blue in A. barretti and A. nahuana; blue or
bluish-violet in A. rhoadsi, A. sedula, A. immunda, A. munda, A. hinei
and A. plan; violet in A. fnumipennis; and blue anteriorly, violet
posteriorly in A. bipunctulata. Other species have inconspicuous pale
areas on the abdomen. Wings are occasionally diffuse pale brown in A.
sedula, apparently characteristically amber in A. rhoadsi, but remain
transparent in both species. Other species and most A. sedula possess
clear wings.
FEMALES The females of many species have small structural differ-
ences; consequently, A. alberta, A. apicalis, A. moesta, and A. nahuana
have two and A. sedula has three routes for determination in the key.
Structural characters used in female diagnosis are as follows. The anter-
ior carina (as in Fig. 2 E) is visible or hidden in the lateral view of a meso-


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


stigmal plate. The carina in lateral view appears as a dorsally-directed
projection in the anterior fourth or third of the plate (see figures of A.
apicalis, A. tibialis, and A. munda). The dorsal thoracic carina bifurcates
or divides at the anterior end of the mesothorax into pear-shaped, ovally
flattened pads or into two narrow and erect ridges terminating between
the mesial borders of the mesostigmal plates. Both pads and ridges are
the rami of the dorsal thoracic carina. Location of the bifurcation occurs
at a point (1) approximately even with or just rearward of the posterior
borders of the mesostigmal plates, or (2) distinctly posterior to the plates.
The mesostigmal plates have their posterior margins smoothly
rounded or expanded rearward into lobes. If lobes exist, they are (1) dis-
tinct thumb-like projections, with length about equal to or greater than
width of the lobe's base, or (2) broad-based, convex projections. The lat-
ter category of projections appear as lobes to some workers and only as
raised prominences to others. Couplet 8 in the key requires a decision
on the presence of a lobe; however, both alternates in the couplet identify
species with weakly developed lobes. A distinct depression may occur in
the mesothoracic surface underlying each posterior lobe of the mesostig-
mal plates. The depression is a vertical-sided pit in A. munda, and a cone-
shaped opening in A. lugens and A. moesta. Shallow depressions occur
in other species but have no semi-circular margins. Females of A. hinei
and A. fumipennis are similar in structure, and the following character
supplements the key. A small knob or tubercle exists at the base of the
posterior lobe of the mesostigmal plates in A. fumipennis, best seen in
oblique posterior view. A strong, concentrated light source and good
magnification usually verify its presence. The tubercle is absent in A. hinei
and this species is absent from most of Texas. Distributions for body size
fall within the measurements in Table 2. Pale colors in females are
typically brown or shades of tan, and reduced stripe patterns frequently
exist compared with conspecific males. See note on A. apicalis under males.
Dark stripe and spot patterns on abdominal segments vary between many
species. The fifth and sixth segments are easiest to score for these patterns,
and Figure 12 illustrates the major variations. Female A. immunda fre-
quently possess pale, small grayish flecks or spots in the antehumeral
areas; female A. sedula may have grayish dots at the base of hairs on the
thoracic dorsum, but the mesostigmal plates are very different from A.
immunda.
The keys include A. alberta, a likely addition to the Texas fauna as
indicated in the Discussion.

KEY TO THE MALES
1 a) Tdreale long in dorsal view, reaching ;3 length of superior abdominal
appendage (Fig. 8 B); inferior abdominal appendage in lateral view
not bifid (Fig. 8 A); body length less than 30 mm; fore wing length
usually less than 17 mm; usually 3 antenodal postquadrangular cells
in fore wing; dorsum of abdominal segments 3-6 blue with black api-
cal rings ................ ................... ..bipunctulata









82 BULLETIN FLORIDA STATE MUSEUM


b) Toreale short in dorsal view, reaching at most slightly posterior to
apical margin of tori; inferior abdominal appendage in lateral view
bifid, or if not, 5 or more antenodal postquadrangular cells in fore
wing; body length greater than 30 mm; fore wing length greater than
17 mm; abdominal color pattern variable................... .. 2
2 ( 1) a) In lateral view, apical margin of inferior abdominal appendage
rounded, blunt, without inferior lobe (Fig. 8 C, E, G); wings amber
or body length greater than 40 mm. ........................... 3
b) In lateral view, apical margin of inferior abdominal appendage with
superior and inferior lobes, latter lobe may occur only as an angulate




A C E G






A ~ p72 F H



BIPUNCTULATA RHOADSI MOESTA LUGENS




I M









IMMUNDA TIBIALIS TRANSLATE


a c e


b d f
FIGURE 8. A-N, left lateral and dorsal right side (anterior end to the left)
views of male abdominal appendages in Argia species, tori shown in black;
a-f, dorsal patterns (anterior end to the left) of abdominal segments 5 and
6 in male argians.


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


blunt process arising from lower apical margin of the inferior abdom-
inal appendage and directed posteriorly (Fig. 8 I, K, M; Fig. 9); if
wings brownish, inferior abdominal appendage with distinct tapered
inferior lobe................................. ............. 5
3 (2) a) In dorsal view, tori transversely elongated pads, at least twice as
wide as long (Fig. 8 D); inferior abdominal appendage in lateral
view with a dorsally-directed tooth just anterior to superior lobe
(Fig. 8 C); typically 4 antenodal postquadrangular cells in fore wing;
dorsum of abdominal segments 8, 9, and 10 pale blue, black if present
reduced and limited to lateral sides of segments; wings light amber;
body length less than 40 mm. .......................... rhoadsi
b) In dorsal view, tori are circular knobs (Fig. 8 F) or reduced to thin
pads confined to apical ridge of torifer (Fig. 8 H); inferior abdom-
inal appendage in lateral view with a single small dorsally-directed
tooth on apex of superior lobe (Fig. 8 E, G); 5 or more antenodal post-
quadrangular cells in fore wing; dorsum of abdominal segments 8, 9
and 10 largely black; wings clear; body length greater than 40 mm..
........................................................ 4
4 ( 3) a) In dorsal view, tori are circular knobs (Fig. 8 F); distinct bluish-
white pruinescence on dorsum of head and thorax in nonteneral indi-
viduals; dark middorsal thoracic stripe wider than dorsal carina ...
.................................................... moesta
b) In dorsal view, tori are thin pads confined to apical ridge of torifer
(Fig. 8 H); bluish-white pruinescence greatest on lower sides of
pterothorax; dark middorsal thoracic stripe largely confined to ca-
rina with a parallel dark stripe on either side in antehumeral area.
............. . ..... ..................... ..... lugens
5 ( 2) a) Dorsum of abdominal segments 5 and 6 with an anterior to pos-
terior pattern sequence of pale, dark, pale, dark (Fig. 8 a, b); infer-
ior lobe of inferior abdominal appendage in lateral view rounded,
deeper at its base than long and apical margin of tori extends pos-
teriorly beyond tips of toreale (Fig. 8 I, J).............. immunda
b) Dorsum of abdominal segments 5 and 6 pale with black apical rings
or black with pale basal rings (Fig. 8 e-f); inferior lobe of inferior
abdominal appendage in lateral view distinctly pointed, as long or
longer than deep at its base or, if not, apical margin of tori not ex-
tending posteriorly beyond tips of toreale .................... 6
6 ( 5) a) Dorsum of abdominal segments 5 and 6 predominantly black with
pale basal rings (Fig. 8 c-e) ............................... 7
b) Dorsum of abdominal segments 5 and 6 predominantly pale with
black apical rings (Fig. 8 f) ............................... 11
7 ( 6) a) Dorsum of abdominal segment 8 black or mostly black; torus not
wider than long (Fig. 8 L, N) ............................ 8
b) Dorsum of abdominal segment 8 pale, (very reduced black area if
any); torus wider than long (Fig. 9 B, D; Fig. 18 B)............ 9
8 ( 7) a) In lateral view, inferior lobe of inferior abdominal appendage not
projecting posteriorly well beyond superior lobe (Fig. 8 K); abdom-
inal segments 9 and 10 pale.......................... tibialiss
b) In lateral view, inferior lobe of inferior abdominal appendage pro-
jecting posteriorly well beyond superior lobe (Fig. 8 M); abdom-
inal segments 9 and 10 partly black................... translate


1972








84 BULLETIN FLORIDA STATE MUSEUM


9 ( 7) a) In dorsal view, inferior lobes of the inferior abdominal appendage
have inward curved apical tips and tori obliquely elongate on
apical margin of torifer (Fig. 9 B) .......................sedula
b) In dorsal view, inferior lobes of the inferior abdominal appendage
have laterally divergent curvature from medial base to apical tips
or lobes directed straight to the rear; tori distinct, raised, bean-
shaped pads (Fig. 9 D; Fig. 18 B)...........................10
10 ( 9) a) In dorsal view, superior abdominal appendage's apical margin
oblique to longitudinal axis of abdomen (Fig. 18 B); distinct dark
middorsal and humeral stripes; body length 31 mm or less. .alberta
b) In dorsal view, superior abdominal appendage's apical margin ap-
pears slightly trifid or convex without oblique alignment to abdom-
inal axis (Fig. 9 D); thoracic stripe pattern typically reduced but
see text; body length 35 mm or greater. .................. apicalis


A CE H

A


FZ\;






SEDULA APICALIS BARRETTE NAHUANA


J N 1. N L' Q












MUNDA HINEI PLAN A FUMIPENNIS

FIGURE 9. Male abdominal appendages in Argia species, tori shown in black,
F, 0, and R are disarticulated superior abdominal appendages in posterior
oblique view, otherwise orientation same as Fig. 8.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


11 ( 6) a) In dorsal view, apical end of superior abdominal appendage bifid,
oblique dorsolateral view of medial surface often required to con-
firm structure (Fig. 9 F, G); distance between medial corners of
tori greater than width of one torus; black stripe on lateral sides of
abdominal segments 8, 9 and 10 .......................barretti
b) In dorsal view, superior abdominal appendage with apical margin
convex, transversely or obliquely straight, or slightly sigmoid, and
ventrally or medially-directed prominence (tooth) may occur on
medial margin; distance between medial corners of tori less than
width of one torus, or if not, sides of abdominal segments 8, 9 and
10 without black stripe................................. 12
12 (11) a) In dorsal view, looking directly downward at midline, medial mar-
gin of superior abdominal appendage with blunt tooth, same appen-
dage with convex apical margin (Fig. 9 I); pale thoracic and abdom-
inal colors blue .................................... nahuana
b) In dorsal view, looking directly downward at midline, no distinct
tooth or process arising from medial margin of superior abdominal
appendage, any such process typically seen only from oblique dor-
solateral view and associated with pale thoracic and abdominal col-
ors of violet or bluish-violet................................. 13
13 (12) a) Tori in dorsal view thin, nonraised, obliquely elongated pads on con-
vex apical margin of torifer (Fig. 9 K); superior abdominal appen-
dage in lateral view, appears hollowed-out on ventral surface giving
dome-shaped dorsal margin (Fig. 9 J), same appendage possesses
pointed apical tooth; toreale tips not posterior to apical margin of
tori ........................... ........ .. .. .......... m u nda
b) Tori in dorsal view, transversely elongated pads (Fig. 9 M, P), or
obliquely elongated (Fig. 9 S) and body length 34 mm or less; if ven-
tral surface of appendage visible in lateral view (slightly in hinei),
toreale tips project posterior to apical margin of tori ............... 14
14 (13) a) Inferior abdominal appendage in lateral view with a small promin-
ence or swelling projecting from its dorsal or dorsolateral surface
just anterior to base of appendage's superior lobe (Fig. 9 L); su-
perior abdominal appendage in lateral view typically exposing ven-
tral surface of its small medial projection (Fig. 9 L), same appendage
in dorsal view with convex apical margin (white hairs may obscure
margin) (Fig. 9 M)...................................... hinei
b) Inferior abdominal appendage in lateral view without prominence on
dorsal surface anterior to base of superior lobe (Fig. 9 N, Q), but an-
terior margin of superior lobe may be convex; superior abdominal
appendage in lateral view exposing little if any of appendage's me-
dial structure (Fig. 9 N, Q), same appendage in dorsal view with
apical margin transversely or obliquely straight or slightly sigmoid
(Fig. 9 P, S).................. ..... ..................... 15
15 (14) a) No black stripe on lateral sides of abdominal segments 8, 9 and 10; in
oblique dorsolateral view, a single blackish tooth projects ventrad
from superior abdominal appendage's subapical surface (Fig. 9 0)
. .. . ... .. .. .. . .. .. .. . ... .. .. .. .. .. p lan a
b) Black stripe on lateral sides of abdominal segments 8, 9 and 10; in
oblique dorsolateral view, a tooth projects ventrad from superior
abdominal appendage's subapical margin and a blunt tooth projects









86 BULLETIN FLORIDA STATE MUSEUM


ventrad from medial margin (Fig. 9 R) ................ fitmipennis


KEY TO THE FEMALES
1 a) Wings distinctly amber or pale brown, yet translucent .............. 2
b) Wings clear, devoid of color.................................. 3
2 ( 1) a) In dorsal view, distinct, posteriorly-directed lobes on mesostigmal
plates; wings amber. .............................. .. rhoadsi





A S L A

SEDULA AP/CA L /S


MOESTA TIBIALIS


IMMUNDA NAHUANA


LUGENS MUNDA


TRANSLATE BARRETTE
FIGURE 10. Dorsal, left side (anterior end uppermost) and left lateral views of
mesostigmal plates in female Argia species.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


b) In dorsal view, distinct, mesially-directed lobes on mesostigmal
plates, raised into upturned ridges (Fig. 10 A, B); wings pale brown
................................................... sedula
3 (1) a) Three antenodal postquadrangular cells in fore wing; no thumb-
like lobes projecting posteriorly from rear margin of mesostigmal
plates, (lobe-like mesial corner of plates' rear margins directed me-
sially and pale antehumeral stripe narrower than dark middorsal
stripe in A. bipunctulata) ............... ... ................ 4
b) Four or more antenodal postquadrangular cells in fore wing, or if
less, thumb-like lobes arising from posterior margins of each meso-
stigmal plate are directed rearward and pale antehumeral stripe
wider than dark middorsal stripe. ............................. .5
4 ( 3) a) Abdominal segment 8 pale dorsally; segments 5 and 6 black dorsally
with pale basal rings (Fig. 12 a, m).................. bipunctulata
b) Abdominal segment 8 predominantly black dorsally; segments 5 and
6 with dorsal black constricted at midsegment, occasionally separ-
ated into 2 spots ....................................... alberta
5 (3) a) Pterothorax without distinct dark stripes, middorsal and humeral
stripes absent or reduced to sutures, often irregular spot for hu-
meral stripe................................................ 6
b) Pterothorax with distinct dark middorsal or humeral stripes or both
stripes; middorsal stripe may be wide, narrow, subdivided by pale
carina, or restricted to carina; a dark stripe may occur in each ante-
humeral pale area; humeral stripe forked, interrupted, or termin-
ated in posterior half, if stripe entire its longitudinal borders may or
may not be parallel .......................................... 8
6 ( 5) a) In dorsal view, thoracic carina bifurcates into raised ridges diverg-
ing between posterior medial corners of mesostigmal plates, plates
without posterior lobes (Fig. 10 C); anterior carina usually visible in
lateral view of plate (Fig. 10 D); humeral stripe variable, usually
represented by elongate anterior spot, or absent........... apicalis
b) In dorsal view, thoracic carina bifurcates between posterior medial
corners of mesostigmal plates as flat, pear-shaped pads and plates
with distinct posterior lobes (Fig. 10 A, E); anterior carina not vis-
ible in lateral view of plate (Fig. 10 B, F). ..................... .7
7 ( 6) a) Stigma surmounts 1 cell (Fig. 4 D), rarely 1.5 cells; lobes of meso-
stigmal plates directed mesially and raised into elevated ridges (Fig.
10 A, B); abdominal color pattern usually a pale uniform tan (Fig.
12 1, u) ......... ......... .... ..................... sedula
b) Stigma surmounts 1.5 to 2 cells (Fig. 4 C); lobes of mesostigmal
plates directed posteriorly; in dorsal view, a low ridge curves over
mesial half of each above lobe (oblique posterior view often re-
quired) (Fig. 10 E, F); abdominal color pattern with distinct lateral
dark brown spots or stripes (Fig. 12 j, q). ................. moesta
8 ( 5) a) In dorsal view, no lobes projecting rearward beyond posterior bor-
der of mesostigmal plates (Fig. 10 C, G, I, K; Fig. 18 C)........... 9
b) In dorsal view, distinct lobes arising from posterior border of me-
sostigmal plates (Fig. 10 A, M, O, Q, S; Fig. 11 A, C, F)........13
9 ( 8) a) In lateral view of mesostigmal plate, anterior carina visible (Fig. 10
D, H); black pattern on dorsolateral surface of abdominal seg-
ments 5 and 6 continuous stripe; not constricted along midsegment









88 BULLETIN FLORIDA STATE MUSEUM


(Fig. 12 a, b, c, m, p)...................................... 10
b) In lateral view of mesostigmal plate, anterior carina not visible (or
barely so) (Fig. 10 J, L; Fig. 18 D); black pattern on dorsolateral
surface of abdominal segments 5 and 6 broken into apical and post-
basal spots, or if spots connected, resulting stripe constricted about
midsegment (Fig. 12 h-k, o, r, more black on alberta); humeral
stripe variable ............................................ 11
10 ( 9) a) In dorsal view, thoracic carina bifurcates into raised ridges diverg-
ing between posterior medial comers of mesostigmal plates (Fig. 10
C) ............................................... apicalis
(very few A. apicalis will be carried this far in the key, see text)
b) In dorsal view, thoracic carina bifurcates between posterior medial
corners of mesostigmal plates as flat pear-shaped pads (Fig. 10 G);
humeral stripe typically forked.......................... tibialis
11 ( 9) a) Thoracic carina in dorsal view bifurcates into rami just posterior to
mesostigmal plates (Fig. 10 K) ........................ nahuana
b) Thoracic carina in dorsal view bifurcates into widely diverging rami
well behind mesostigmal plates (Fig. 10 I; Fig. 18 C)............ 12
12 (11) a) Mesial posterior borders of mesostigmal plates raised into broad-
based rims or 'lobes' with their ventral surfaces slightly visible in
lateral view (Fig. 18 D); antehumeral areas brownish with no small
gray spotting; body length usually less than 32 mm.......alberta
b) Mesial posterior borders of mesostigmal plates occur as blunt, low
prominences with their ventral surfaces not visible in lateral view
(Fig. 10 J); antehumeral areas brownish, purple, or bluish and
often with pale grayish spots; body length usually greater than 32
mm ............................................... immunda
13 ( 8) a) In dorsal view, lobe on posterior border of each mesostigmal plate
with a low, curved ridge over mesial half of its base (Fig. 10 E); a
shallow pit beneath each lobe but lobes not converging toward mid-
line; elongated dark stripe rarely in each antehumeral area.moesta
b) In dorsal view, no ridge over base of lobe on posterior border of
mesostigmal plates, if distinct pit beneath lobes, lobes are either
distinctly diverging from or converging toward midline (Fig. 10 A,
K, M, O, Q, S; Fig. 11 A, C, F; Fig. 18 C) ......................14
14 (13) a) In dorsal view, thoracic carina bifurcates well behind mesostigmal
plates into widely diverging rami (Fig. 18 C); no deep pits posterior
to plates; 'lobes' arising from rear mesial border of plates more like
up-turned edges ...................................... alberta
b) In dorsal view, thoracic carina bifurcates just posterior to, or even
with, rear border of mesostigmal plates (Fig. 10 A, K, M, O, Q, S;
Fig. 11 A, C, F) .................... ...................... 15
15 (14) a) In dorsal view, lobes of mesostigmal plates directed mesially, and
raised into elevated ridges (Fig. 10 A); ventral surface of upturned
lobe exposed in lateral view (Fig. 10 B); dark stripe pattern poorly
defined, borders of stripes obscured...... . .......... sedula
(very few A. sedula will be carried this far in the key)
b) In dorsal view, lobes of mesostigmal plates not directed mesially, or
if so, little or no ventral surface of lobe visible in lateral view; dark
stripes wide or narrow but well defined from bordering pale areas
........................................................ 16


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


16 (15) a) In dorsal view, posterior lobes of mesostigmal plates distinctly di-
verging (Fig. 10 M); 5 (or 6) antenodal postquadrangular cells in
fore wing; middorsal stripe largely restricted to carina, elongated
dark stripe in each antehumeral area often confluent with middorsal
stripe posteriorly ........................................ lugens
b) In dorsal view, posterior lobes of mesostigmal plates not diverging;
4 (or less) antenodal postquadrangular cells in fore wing; middorsal
stripe variable, no dark elongated bands in antehumeral areas..17
17 (16) a) Distinct deep pit occurs in dorsum of pterothorax just posterior to
each mesostigmal plate, lobe of each plate overlying pit from dorsal
view (Fig. 10 0); anterior carina visible in lateral view (Fig. 10 P);
middorsal stripe narrow, often divided by pale carina; humeral
stripe narrow, usually interrupted; dark pattern on dorsolateral
surface of abdominal segments 5 and 6 in 2 spots or a stripe con-
stricted about midsegment (Fig. 12 f, g, n, o).............. munda
b) No deep pit posterior to mesostigmal plates, shallow indentation
with sloping margin may occur; anterior carina not visible in lateral
view; middorsal and humeral stripes variable ................. 18
18 (17) a) In dorsal view, posterior lobes of mesostigmal plates slightly con-
stricted at base (Fig. 10 Q); in lateral view, lobe raised into thin
linear projection (Fig. 10 R); middorsal stripe wider than one ante-
humeral area; humeral stripe wide with inserted pale stripe in pos-
terior half; abdominal segments 5 and 6 black dorsally (thin pale
middorsal line may occur) with pale basal rings (Fig. 12 a-c, m, p)
.................................. ............ translate
b) In dorsal view, posterior lobes of mesostigmal plates not basally
constricted (Fig. 10 K, S; Fig. 11 A, C, F); middorsal stripe nar-
rower than antehumeral area; humeral stripe narrow, sides non-
parallel, forked, or interrupted; abdominal segments 5 and 6 vari-
able in pattern........................................... 19
19 (18) a) In dorsal view, apex of posterior lobe on mesostigmal plates directed
mesially, lines drawn through longitudinal axes of two lobes inter-
cept in obtuse angle at midline (Fig. 10 S); dark pattern on abdom-
inal segments 5 and 6 in nonconstricted stripe (Fig. 12 d, e, n, s)...
.................................................... barretti
b) In dorsal view, posterior lobe or border of mesostigmal plates
broadly rounded, directed upward or rearward into low ridge (Fig.
10 K; Fig. 11 A); dark pattern on abdominal segments 5 and 6 in 2
spots or stripe constricted at midsegment (Fig. 12 i-k, o, r, t), or
lobes on mesostigmal plates thumb-like and directed slightly mesad
with lines drawn through longitudinal axes of 2 lobes intercepting
in acute angle at midline (Fig. 11 C, F). ..................... .20
20 (19) a) In dorsal view, posterior lobe of mesostigmal plates broad-based,
rounded (Fig. 11 A); in lateral view, lobe lies close over dorsum of
pterothorax (Fig. 11 B); abdominal segment 8 pale dorsally...plana
b) In dorsal view, posterior margin of mesostigmal plates smoothly
curved, no lobe-like projection extending rearward (Fig. 10 K), or
with distinct, thumb-like lobes arising from posterior margin of
plates (Fig. 11 C, F); abdominal segment 8 with black markings dor-
sally.................................................... 21
21 (20) a) In dorsal view, posterior margin of mesostigmal plate scarcely lobe-








90 BULLETIN FLORIDA STATE MUSEUM


like, and barely extending rearward over pterothorax (Fig. 10 K);
dorsooblique view reveals 'lobe' as low, thin, broad ridge.nachuana
b) In dorsal view, posterior lobes of mesostigmal plates exist as dis-
tinct thumb-like projections (Fig. 11 C, F) ................... 22
22 (21) a) In posterior dorsooblique view, posterior lobe of mesostigmal
plate column-like (Fig. 11 E) ............................. hinei




AB
PLAN



C D F GV






E
"E FUMIPENNIS
H/NEI



SKL


DENTICOLLIS SALVA VERTICALIS




M :
AARONI


CARA
FIGURE 11. A-H, mesostigmal plates in female argians, E and H posterior
oblique views, otherwise orientation same as Fig. 10, I-L, female structures in
identified species: I, J, and L, left lateral views of prothorax; K, dorsal view of
prothorax; M, N, dorsal views and 0, left lateral view of mesostigmal plates.


Vol. XVI No. 2







1972 JOHNSON: TEXAS DAMSELFLIES 91

b) In posterior dorsooblique view, posterior lobe of mesostigmal
plate transversely flattened, slightly concave on ventral surface
(Fig. 11 H ) .................................... fum ipennis

Argia species typically inhabit stream-riverine conditions; however,
lake shores, springs, and small seepage areas often support populations.
Argia bipunctulata usually occurs about small seepage sites, and ecologi-
cal studies will likely reveal specific habitats for other species. Unlike most
damselflies, many argians prefer bare soil and rocks for perch sites.










a b c d e f








g h i j k I




m n o



p q r

-no

St u

FIGURE 12. a-1, dorsal patterns (anterior end uppermost) and m-u, left lateral
patterns of abdominal segments 5 and 6 in female argians.









92 BULLETIN FLORIDA STATE MUSEUM


DISTRIBUTION RECORDS FOR TEXAS
Argia apicalis: Angelina, Bexar, Bosque, Brazos, Caldwell, Cameron,
Chambers, Cherokee, Collin, Colorado, Cooke, Dallas, Denton, Falls, Fannin,
Fayette, Goliad, Gonzales, Grayson, Gregg, Grimes, Guadalupe, Harris,
Harrison, Hidalgo, Hunt, Jim Wells, Karnes, Kendall, La Salle, Leon, Liberty,
Limestone, Lubbock, Marion, Matagorda, McLennan, Medina, Montgomery,
Morris, Orange, Panola, Robertson, Rusk, San Jacinto, San Patricio, Shackel-
ford, Tarrant, Travis, Victoria,'Walker, Williamson, and Wilson counties.
Argia barretti: Comal, Hays, Kimble, Travis, and Uvalde counties.
Argia bipunctulata: Cherokee, Gregg, Panola, San Jacinto, and Wood coun-
ties.
Argia fumipennis: Austin, Bexar, Blanco, Bosque, Brewster, Cooke, Gil-
lespie, Gregg, Grimes, Harrison, Jeff Davis, Kendall, Kinney, Marion, Mont-
gomery, Presidio, Reeves, Robertson, Rusk, San Jacinto, Travis, Uvalde, Val
Verde, and Williamson counties.
Argia hinei: Brewster and Jeff Davis counties.
Argia immunda: Bexar, Bosque, Blanco, Brazos, Brewster, Caldwell,
Comal, Crosby, Dallas, Denton, Guadalupe, Gillespie, Grimes, Hays, Hidalgo,
Jeff Davis, Jim Wells, Kendall, Kerr, Kimble, Medina, Polk, Reeves, Robert-
son, Rusk, San Saba, Tom Green, Travis, Uvalde, Val Verde, Williamson,
Wilson, and Wood counties.
Argia lugens: Brewster, Crosby, Jeff Davis, Lubbock, Presidio, and Reeves
counties.
Argia moesta: Anderson, Angelina, Baylor, Bexar, Blanco, Bosque, Brazos,
Brewster, Burnet, Caldwell, Cherokee, Collin, Comal, Cooke, Crosby, Dallas,
Denton, Fannin, Fayette, Gillespie, Goliad, Gonzales, Grayson, Gregg, Grimes,
Guadalupe, Hays, Hidalgo, Hill, Howard, Jeff Davis, Jim Wells, Kendall,
Kerr, Kimble, Liberty, Limestone, Llano, Marion, Matagorda, Maverick,
McLennan, Medina, Montgomery, Palo Pinto, Pecos, Polk, Presidio, Randall,
Reeves, San Jacinto, San Patricio, Starr, Tom Green, Travis, Uvalde, Val
Verde, Victoria, Walker, Webb, Williamson, Wilson, and Zavala counties.
Argia munda: Jeff Davis county.
Argia nahuana: Blanco, Bosque, Brewster, Collin, Cooke, Crosby, Dallas,
Hays, Hill, Jeff Davis, Kendall, Kimble, Menard, Presidio, Reeves, Robertson,
San Saba, Tom Green, Travis, Uvalde, and Williamson counties.
Argia plan: Blanco, Bosque, Brewster, Collin, Crosby, Culberson, Dallas,
Hill, Jeff Davis, McLennan, Presidio, San Saba, and Wood counties.
Argia rhoadsi: Cameron county.
Argia sedula: Austin, Bee, Bexar, Blanco, Bosque, Brazos, Burnet, Cald-
well, Cameron, Cherokee, Collin, Colorado, Comal, Cooke, Dallas, Denton,
Dimmit, Fayette, Gillespie, Goliad, Gonzales, Grayson, Grimes, Guadalupe,
Harris, Hays, Hidalgo, Hill, Jeff Davis, Jim Wells, Kendall, Kerr, Kimble,
Kinney, Liberty, Limestone, Lubbock, Matagorda, McLennan, Medina, Men-
ard, Montgomery, Pecos, Polk, Presidio, Reeves, Rusk, San Jacinto, San
Patricio, Starr, Sutton, Robertson, Tom Green, Travis, Uvalde, Val Verde,
Victoria, Webb, Williamson, Wilson, and Zavala counties.
Argia tibialis: Angelina, Brazos, Franklin, Gregg, Grimes, Hardin, Hen-
derson, Jackson, Leon, Liberty, Matagorda, Montgomery, Morris, Nacog-
doches, Orange, Polk, Rusk, San Jacinto, Tyler, Victoria, Walker, and Wood
counties.
Argia translate: Bexar, Blanco, Bosque, Brazos, Caldwell, Collin, Comal,


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


Cooke, Dallas, Denton, Fayette, Gillespie, Gonzales, Grimes, Guadalupe,
Hays, Hidalgo, Hill, Howard, Kendall, Kimble, Limestone, McLennan, Me-
dina, Palo Pinto, San Patricio, Travis, Uvalde, Val Verde, Williamson, and
Wilson counties.


Enallagma Charpentier, 1840

MALES Identification of males requires examination of the superior
abdominal appendages for an attached tubercle-like structure. The key
recognizes the presence of the tubercle only if it forms part of the appen-
dage's lateral profile (Fig. 13 A; Fig. 18 E). Medial projections not detect-
able in lateral view are not tubercles in this sense, (E. durum for example).
Tubercles are usually paler in color than the appendage but darken with
age.
Stripe patterns in Enallagma species are similar between the sexes;
however, pale areas are more colorful in adult males than females. Species
with blue colors in males are E. basidens, E. civil, E. divagans, E.
durum, E. exsulans, E. geminatum, E. praevarum, and E. traviatum.
Purple and blue colors exist in males of E. novaehispaniae. Species with
yellowish, orange, or reddish colors in the mature males are E. signa-
tum, E. vesperum, and E. dubium, the latter species possessing reddish
color. Teneral individuals may differ in color of the pale areas; for
example, E. signature is light blue in the immature stage. Such general
effects are of short duration, and Enallagma species typically change less
with age than argians and female ischnurans. Polymorphic variation oc-
curs in the postocular spot patterns of some species (Johnson, 1964).
Body size aids in recognizing species. Body length ranges, grouped
from smallest to largest species including both sexes, appear in Table 2.
Teleallagma daeckii has the general appearance of a pale bluish Enal-
lagma; however, it is typically greater than 40 mm in body length.

FEMALES Identification of females requires study of the prothor-
acic dorsum, mesostigmal plates, anterior dorsum of the mesothorax,
antenodal postquadrangular cell number, and stripe patterns. The middle
lobe of the prothorax possesses a pair of shallow pits on its dorsum in sev-
eral species. This trait requires close attention; for instance, the pits of
E. dubium occur anteriorly on a black middle lobe and casual observa-
tion may miss them. The pits are shallow, rather wide depressions in E.
signatum and they do not appear pit-like. The mesostigmal plates possess
distinctive structural differences for several species and their dorsal view
is most useful. The dorsal view of prothoracic pits and mesostigmal plates
usually requires a forward flexure of the specimen's head. The dorsum of
the mesosthorax posterior to the rear margin of the mesostigmal plates
possesses, in several species, a pair of elevated knob or ridge-like projec-
tions (Fig. 15 B, C). These structures, visible in dorsal view, are most dis-
tinct when viewed at an oblique lateral angle. Cross veins between the


1972








94 BULLETIN FLORIDA STATE MUSEUM


longitudinal veins M14 and Cul and from the quadrangle outward to the
level of the subnodus form the antenodal postquadrangular cells.
Female stripe patterns are similar to conspecific males; however, pale-
colored areas occur in shades of tan or brown, or these areas are occasion-
ally bluish typical of the conspecific male. Walker (1953) suggested that
this variation represents sex-limited dimorphism similar to Ischnura. Pale
and dark color patterns on the terminal abdominal segments are often
obscured with a coating of silt if the females have recently oviposited.
Acetone or alcohol and a fine brush will remove such material.
Females of E. carunculatum, E. civil, and E, praevarum are similar
and their separation requires care. The mesostigmal plates of E. civil
have no ridge or prominence crossing the plate's midsection in an anter-
ior to posterior direction. The lateral anterior margin of each mesostig-
mal plate curves distinctly upward. Pale colors are frequently greenish
or bluish-brown rather than tan. Females of E. carunculatum and E. prae-
varum have a ridge or prominence diagonally crossing the mesostigmal
plate in an anterior to posterior direction developing a depression re-
stricted to the anterior medial half of each plate. The depressions differ
as described in the key. Pale colors in E. praevarum are often light tan
to brown while E. carunculatum is greenish or bluish-brown. These color
traits are not characteristic of all individuals, possibly change with age,
and fade in museum specimens. The rear margin of the hind lobe of the
prothorax is typically concave in E. praevarum and convex in E. carun-
cultatum and E. civil; however, occasional E. praevarum possess such
lobes with a squarish rear margin. Also, occasional E. carunculatum and
E. civil have lobes with a squarish margin.
The keys include E. carunculatum, a likely species for Texas as indi-
cated in the discussion.

KEY TO THE MALES
1 a) Superior abdominal appendage in lateral view with distinct tuber-
cle lying between dorsal and ventral lobes of the appendage or pro-
truding from apical end of appendage (Fig. 13 A; Fig. 18 E) ......2
b) Superior abdominal appendage in lateral view without such a tu-
bercle ................................................... 3
2 ( 1) a) Tubercle of superior abdominal appendage overlaid in lateral view
by dorsal lobe of appendage (Fig. 13 A) ................... civil
b) Tubercle of superior abdominal appendage not overlaid in lateral
view by dorsal lobe of appendage (Fig. 18 E) ........ carunculatum
3 (1) a) Superior abdominal appendage in lateral view notched on dorso-
apical margin with only distinct ventral lobe (Fig. 13 B), or appen-
dage bifid with dorsal and ventral lobes, the latter equal to or
greater in length than the dorsal lobe (Fig. 13 C-E) ............. 4
b) Superior abdominal appendage in lateral view elongate with small
tooth or distinct ventral lobe directed downward (Fig. 13 F-H, J-L),
or appendage bifid with divergent dorsal and ventral lobes, the lat-
ter shorter in length than the dorsal lobe (Fig. 13 I)............. 7
4 (3) a) Inferior abdominal appendage in lateral view greater in length


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


than superior abdominal appendage ........................... .5
b) Inferior abdominal appendage in lateral view equal to or less than
length of superior abdominal appendage ......................6
5 ( 4) a) Dorsum of abdominal segments 3-6 black except narrow blue basal
rings (Fig. 14 B); body length 28 mm or less; ventral lobe of su-
perior abdominal appendage curved dorsally (Fig. 13 B) ..........


C/VIL E


EXSULANS


GEM/NATUM DURUM


O/VAGANS


DUBIUM


BAS/DENS PRAEVARUM


TRAVIA TUM VESPERUM


NOVAEHISPANIAE


SIGNA TUM


FIGURE 13. Male abdominal appendages in Enallagma species following se-
quence of determination in key. Orientation same as lateral views in Fig. 8.


1972







96 BULLETIN FLORIDA STATE MUSEUM


................................................. gem inatu m
b) Dorsum of abdominal segments 3-6 blue except black spots in apical
half (Fig. 14 C); body length 31 mm or greater; ventral lobe of su-
perior abdominal appendage curved mesially (Fig. 13 C) with small
tubercle seen in dorsal view........................... durum
6 ( 4) a) Superior abdominal appendage with two lobes directed poster-


A


zjII


C/VILE


B GEMINATUM


C DURUM


D EXSULANS


E DIVAGANS


F DUBIUM


BASIDENS


H PRAEVARUM


I NOVAEHISPANIA E


J TR VIA LTU


K VESPERUM


L SIGNATUM
FIGURE 14. Dorsal (anterior end to the left) views of male abdominal pat-
terns in Enallagma species following sequence of determination in key.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


iorly in lateral view (Fig. 13 D); dorsum of abdominal segment 9
blue or purple, segment 8 partly black (Fig. 14 D)........ exsulans
b) Superior abdominal appendage in lateral view with rounded knob
for dorsal lobe and ventral lobe directed ventroposteriorly (Fig. 13
E); dorsum of abdominal segments 8 and 9 blue (Fig. 14 E)........
......................... ......................... divagans
7 ( 3) a) Dorsum of abdominal segments 8, 9, and 10 black (Fig. 14 F); body
length 28 mm or less; first 2 antennal segments black; antehumeral
stripe narrower than humeral stripe ................... dubium
b) Dorsum of abdominal segments 8 or 9 or both predominantly blue
or yellowish-orange (Fig. 14 G-L); no such combination of other
above characters ........................................... 8
8 ( 7) a) Dorsum of abdominal segments 8 and 9 predominantly blue or pur-
ple (Fig. 14 G-J) ........................................9
b) Dorsum of abdominal segment 9 predominantly blue or orange,
segment 8 black (Fig. 14 K, L) ............................... 13
9 ( 8) a) Dorsum of abdominal segments 4 and 5 with basal half (or third in
some E. praevarum) pale blue (bluish-brown in some museum
specimens) (Fig. 14 G, H) .................................. 10
b) Dorsum of abdominal segments 4 and 5 with basal half predom-
inantly black (Fig. 14 I, J)................... ............. 11
10 ( 9) a) Humeral stripe longitudinally divided by pale stripe; body length
27 mm or less ....................................... basidens
b) Humeral stripe not longitudinally divided by pale stripe; body
length 30 mm or greater. ................. ......... praevarum
11 ( 9) a) Superior abdominal appendage in lateral view distinctly bifid with
divergent lobes, the inferior lobe extending posteriorly beyond
level of inferior abdominal appendage (Fig. 13 I); pale body color
purple (or blue and purple); prothorax predominantly black......
............... ................. .. novaehispaniae
b) Superior abdominal appendage in lateral view not distinctly bifid
(Fig. 13 H, J), the inferior lobe not extending posteriorly beyond
level of inferior abdominal appendage; pale body color blue; pro-
thorax predominantly blue or black..........................12
12 (11) a) Inferior lobe of superior abdominal appendage scarcely visible in
lateral view (Fig. 13 J); prothorax with distinct dorsal and two lat-
eral pale blue spots on middle lobe; postclypeus without black
markings; distinct pale spots bordering ocelli region..... traviatum
b) Inferior lobe of superior abdominal appendage distinctly visible in
lateral view (Fig. 13 H); prothorax predominantly black; post-
clypeus with distinct black transverse band; very small or no pale
spots bordering ocelli region ....................... praevarum
13 ( 8) a) Dorsum of abdominal segment 9 blue; humeral stripe reduced to
hair line and usually restricted to suture for most of its length. ...
...................... ............... . .. esperum
b) Dorsum of abdominal segment 9 orange or yellow in mature indi-
viduals, (in general specimens compare abdominal appendages
and dark stripe pattern); humeral stripe distinct and not restricted
to suture............................................ signatum


1972








98 BULLETIN FLORIDA STATE MUSEUM


KEY TO THE FEMALES
1 a) Middle lobe of prothorax with pair of depressions or pits on the
dorsal or dorsolateral surface ............................... 2
b) Middle lobe of prothorax without pair of pits. .................. 4
2 ( 1) a) Dorsum of mesothorax without pair of elevated, knob-like projec-
tions just rearward of mesostigmal plates' posterior margins (Fig.
15 A); body length 30 mm or greater. .................. vesperum
b) Dorsum of mesothorax with pair of elevated, knob-like projections
posterior to mesostigmal plates (Fig. 15 B, C)................... 3
3 ( 2) a) Body length 27 mm or less; fore wing length 16 mm or less; post-
nodal veins 7 or less; middle prothoracic lobe black....... dubium
b) Body length 30 mm or greater; fore wing length 19 mm or greater;


[ A
VESPERUM



EXULNS
EXSULANS


/IVAGANS


M *!
POSITA



IQ 9
DEMORSA




HASTATUTA


B
DUBIUM



GF I
GEMINATUMA


BASIDENS




DENTICOLLIS



F R 1


SIGNATUM




NOVAEHISPANIAE



K V
C/VIL E




BARBER


FDII
DURUM



H
TRA VIA TUM




PRAEVARUM



P
RAMBURII



FT:


PROGNATHA VERTICALS KELLICOTTI




INTEGRICOLLIS HETERODOXUM


FIGURE 15. Female mesostigmal plates in Enallagma species, A-L; in Isch-
nura species, M-T; in Anomalagrion, Nehalennia, and Hesperagrion, U, V,
and W. Enallagma and Ischnura figures follow sequence of determination in
respective keys. Orientation same as dorsal views of Fig. 10.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


postnodal veins 8 or greater; middle pfothoracic lobe with pale
spot pattern ............ .................... ..... signatum
4 ( 1) a) Antenodal postquadrangular cells in wings 4 to 5 ...............5
b) Antenodal postquadrangular cells in wings 2 to 3 ...............6
5 ( 4) a) Dark humeral stripe distinct and not divided longitudinally .....
.................................... .. duru m
b) Dark humeral stripe longitudinally divided by pale brownish line
overlying humeral suture. ............................. exsulans
6 ( 4) a) Abdominal segment 8 pale excepting short middorsal black stripe,
or black with distinct basal, apical or lateral rings or spots...... 7
b) Abdominal segment 8 black over entire dorsum excepting very nar-
row apical ring ............................................. 10
7 ( 6) a) Abdominal segment 8 with two pale lateral spots separated by mid-
dorsal black stripe along entire length of segment; body length 28
mm or less........................................ geminatum
b) Abdominal segment 8 without such a pale-dark color pattern; body
length 29 mm or greater (E. traviatum rarely less).............. 8
8 ( 7) a) Abdominal segment 9 black dorsally; dorsum of abdominal segment
8 pale with black apical band (often wedge-shaped) over apical
third ..................... .................. novaehispaniae
b) Abdominal segment 9 blue (pale); black if present confined to
basal border ............................................. 9
9 ( 8) a) Lateral apices of mesostigmal plates prominently elevated (Fig. 15
H); middorsal thoracic stripe usually divided longitudinally by
pale dorsal carina .................................. traviatum
b) Lateral apices of mesostigmal plates not distinctly elevated from
mesothoracic border (Fig. 15 I); middorsal thoracic stripe not long-
itudinally divided (for full length) by pale carina......... divagans
10 ( 6) a) Humeral stripe divided longitudinally for part of its length by in-
serted or overlying pale stripe ........................... .11
b) Humeral stripe well developed and not longitudinally divided by
pale color pattern......................................... 12
11 (10) a) Body length less than 30 mm; distinct elevated projections on an-
terior and posterior borders of mesostigmal plates (Fig. 15 J)......
................... ................... .......... basidens
b) Body length 35 mm or greater; mesostigmal plates without distinct-
elevations (Fig. 15 E). ............................ ... exsulans
12 (10 a) In dorsal view, lateral half of anterior margin of each mesostigmal
plate curved upward, posterior margins of plates with little if any
elevation; a shallow, trough-like transverse depression exists unin-
terrupted from mesial to lateral ends of each plate (Fig. 15 K), (see
text) ................................................ civil
b) In dorsal view, lateral half of each mesostigmal plate elevated into
prominence bordered mesially by diagonal ridge (Fig. 15 L), or
each plate with elevated ridge oriented diagonally between poster-
ior mesial and anterior lateral borders (Fig. 18 F); a circular or
ovoid depression developed in anterior, medial half of each plate
(Fig. 15 L; Fig. 18 F) ....................................... 13
13 (12) a) In dorsal view, depression in anterior medial half of each meso-
stigmal plate circular (Fig. 15 L); margin of prothoracic hind lobe









100 BULLETIN FLORIDA STATE MUSEUM


slightly concave (see text) ........................... praecaru
b) In doisal view, depression in anterior medial half of each meso-
stigmal plate ovoid, axis of depression oriented diagonally between
posterior mesial and anterior lateral borders of plate (Fig. 18 F);
margin of prothoracic hind lobe typically convex (see test).........
................................. .. carunculatum
General collecting experience suggests considerable habitat specifi-
city between Enallagma species but ecological studies on the subject are un-
available.

DISTRIBUTION RECORDS FOR TEXAS
Enallagma basidens: Bexar, Blanco, Bosque, Bowie, Burnet, Caldwell, Cam-
eron, Cherokee, Collin, Colorado, Cooke, Dallas, Denton, Fannin, Franklin, Gil-
lespie, Grayson, Gregg, Grimes, Hidalgo, Hill, Hunt, Jim Wells, Kendall, Kinney,
Liberty, Lubbock, Marion, Medina, Montgomery, Nueces, Reeves, Robertson,
San Jacinto, Travis, Uvalde, Val Verde, Victoria, Wilson, Wise, Wood, and Zavala
counties.
Enallagma civil: Aransas, Bexar, Blanco, Bosque, Brazos, Brooks, Brew-
ster, Burnet, Cameron, Childress, Collin, Crosby, Dallas, Denton, Dimmit,
Franklin, Gillespie, Grayson, Guadalupe, Hidalgo, Hill, Houston, Howard, Hunt,
Jackson, Jeff Davis, Jim Wells, Kennedy, Kimble, Kleberg, Lubbock, McLen-
nan, Matagorda, Maverick, Pecos, Presidio, Raines, Reeves, Robertson, Runnels,
San Jacinto, San Patricio, Starr, Travis, Uvalde, Van Zandt, Victoria, Webb,
Wilbarger, Willacy, and Wood counties.
Enallagma divagans: Austin, Brazos, Franklin, Lamar, Gregg, Grayson,
San Jacinto, and Walker counties.
Enallagma dubium: Harris and San Jacinto counties.
Enallagma durum: Matagorda and San Patricio counties.
Enallagma exsulans: Angelina, Bosque, Brazos, Caldwell, Gregg, Grimes,
Hays, Hill, Kendall, Liberty, Marion, Nacogdoches, Robertson, Rusk, San Jac-
into, Tom Green, Val Verde, and Wilson counties.
Enallagma geminatum: Harris, Matagorda, Panola, and San Jacinto counties.
Enallagma novaehispaniae: Comal, Hays, Hidalgo, Uvalde, Val Verde, and
Wilson counties.
Enallagma praevarum: Blanco, Gillespie, Jeff Davis, Presidio, Reeves, Sut-
ton, and Uvalde counties.
Enallagma signatum: Anderson, Austin, Burnet, Colorado, Denton, Grimes,
Harris, Hays,. Hunt, Liberty, Marion, Matagorda, San Jacinto, San Patricio,
Uvalde, Victoria, Walker, Wilson, and Wise counties.
Enallagma traviatum: Grayson, Harrison, Liberty, Marion, and Rusk
counties.
Enallagma vesperum: San Jacinto and Wood counties.
Comments on a questionable species for Texas, E. boreale, appear in the
Discussion.

Ischnura Charpentier, 1840

MALES.- Male ischnurans have species-specific color patterns and
distinctive abdominal appendages. The following observations generally
eliminate errors in their identification. The apical margin of abdominal


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


segment 10 raises into a prominence usually pointed toward the apex in
I. barber, I. ramburii, I. denticollis, I. verticalis, and I. posita; however,
this prominence is distinctly smaller than the spine developed on the same
margin in 1. prognatha and I. demorsa. Only I. kellicotti has a 10th ab-
dominal segment without a raised apical margin. The superior abdominal
appendage of I. ramburii in lateral view may appear slightly bifid with a
blunt lateral lobe, depending on articulation of the appendage. The bifid
nature of I. prognatha's superior appendage in lateral view is distinct with
slender lobes or arms. The bifid condition of the inferior abdominal append-
ages of I. demorsa is diagnostic for that species in Texas, and is visible in
both lateral and dorsal view; however, the latter view must have the longi-
tudinal axis of the 10th segment perpendicular to the line of vision to in-
sure seeing the bifid condition.
The antehumeral pale stripes (totally absent in I. denticollis) are more
narrow in width than the middorsal and humeral dark stripes. In I. posita,
each stripe typically separates into an elongated anterior and a circular
posterior spot having an exclamation mark-pattern. Nonetheless, infre-
quent individuals of I. posita occur with the spots connected resulting in
a medially constricted stripe. Large samples of I. ramburii and I. verticalis
occasionally have individuals with their antehumeral stripes separated
into two spots, and similar variation may occur in other ischnurans.
The eighth and ninth abdominal segments (and also the 10th segment
in I. kellicotti) possess a pattern of blue and black: a reduced or absence of
abdominal blue color is characteristic of I. posita. The variation between
species in this trait largely involves the extent of black on lateral sides of
the eighth and ninth abdominal segments. Black pattern on the sides of
segment eight in I. demorsa varies from total absence to a wide line, the
latter being the typical condition. The key uses only such abdominal pat-
terns as a primary diagnostic difference in couplet six involving species
without conflicting variation to our knowledge. Use of all color patterns
requires caution. Variation of pattern on the ninth abdominal segment of
I. ramburii was the basis for the taxonomic recognition of Ischnura credula
or I. ramburii credula by different authorities. The variation has geo-
graphic correlation in some areas (Paulson, 1966); however, the two types
are widely sympatric.
Ischnurans frequently have shape and color differences in the stigma
of fore and hind wings of males. This difference is absent in I. posita, oc-
curs in color although weakly developed in I. demorsa, I. denticollis, I.
barberi, I. verticalis, I. ramburii, and is distinct for color in I. kellicotti.
Both shape and color differences characterize fore and hind stigmas of
male I. prognatha.
Range in body length within Ischnura species has a seasonal correla-
tion. Larger specimens characterize early season collections, and smaller
individuals appear in late summer or fall. The wide range in body length of
adults probably indicates different generations experiencing different lar-
val durations. Table 2 gives the species from smallest to largest minimal


1972








102 BULLETIN FLORIDA STATE MUSEUM


body length including both sexes.
FEMALES. -The female key relies largely on structure of mesotigmal
plates, as color patterns have a complex variation involving a sex-limited
female dimorphism. One form, the andromorph, possesses the species-spe-
cific color pattern shared with the male, and the second form, the hetero-
morph, differs in color and usually stripe pattern. These morphs develop
soon after emergence. Some species retain distinctiveness of the morphs
throughout life, while morphs of other species superficially change with
age by pruinescence, giving all females an outwardly similar appearance.
A few species have only one of the above morphs. All species in Texas pos-
sess both morphs except I. prognatha and I. posita. These species possess
only the andromorph in I. posita and the heteromorph in I. prognatha.
Age changes, other than pruinescence, further complicate use of color.
Whereas male and female I. posita are similar in pattern following emer-
gence, females soon develop a bluish-gray dorsum of both thorax and ab-
domen through pruinescence. If this effect causes difficulty at couplet 2,
a drop of acetone applied to the thorax usually reveals the basic pattern
temporarily. Female I. prognatha pass through a color change typical of
most heteromorphic females. Soon after emergence, a dark dorsal thoracic
stripe exists bordered by orange thoracic sides, and humeral stripes are ab-
sent. The orange areas become tan to dark brown, often greenish, with age,
but females are always distinct from males. The heteromorphic female I.
denticollis have dark dorsal and humeral stripes with tan pale areas. The
andromorphic I. denticollis females (apparently very rare) are similar to
conspecific males in having no pale antehumeral areas. Heteromorphic
female I. ramburii are basically similar to I. prognatha females but the mid-
dorsal thoracic stripe is typically wider and pale areas more brownish. Het-
eromorphic female I. barber are variable having no thoracic stripes or a
well developed middorsal and faint humeral stripes. Both female morphs
in I. ramburii and I. barberi develop little pruinescence and retain
their distinctness through life. Heteromorphic female I. verticalis, I. prog-
natha and I. ramburii are generally similar in thoracic pattern immediately
following emergence. Females of I. verticalis develop pruinescence over
most of the thorax and abdomen resulting in a bluish-gray color. Within
two days, 1. verticalis females of both morphs are similar in general appear-
ance and distinguishable only by close examination and application of
acetone to the thorax. The morph pattern in I. kellicotti and I. demorsa
differs as follows. Both female morphs possess the thoracic stripe pattern
shared with males; however, andromorphs have the typical bluish (I. kelli-
cotti), or greenish (I. demorsa) pale-colored areas characteristic of males
while heteromorphs have a brown or orange color. This difference exists
unchanged by age in I. kellicotti but is largely lost in I. demorsa similar to
changes in I. verticalis. Pruinescence-effect and color changes with age
occurring in females of I. demorsa, I. posita, I. verticalis, and Anomalagr-
ion hastatum produce individuals similar in outward appearance. Struc-
tural characters are necessary for their reliable diagnosis.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


A statement of additional variations may eliminate confusion with
other groups. The M2 vein of the fore wing originates closer to the 4th and
5th postnodals in males and females respectively for I. prognatha while
both sexes in Ischnura usually have an M2 origin nearer the 4th postnodal.
Some species are also sexually dimorphic for postnodal vein number
(Johnson, 1969). The vulvar spine is variable in some Ischnura species, and
its diagnostic value reduced. All females of I. posita examined or reported
were without spines, and it is probably absent in this species.

KEY TO THE MALES
1 a) Inferior abdominal appendage in lateral view serrated and extend-
ing posteriorly beyond apical level of superior abdominal appen-
dage (Fig. 16 A); dorsum of abdominal segments 8 and 9 black
rarely with small bluish area); antehumeral pale areas usually di-
vided into 2 spots having an exclamation mark pattern......posita
b) Inferior abdominal appendage in lateral view not serrated (Fig. 16
B-H); dorsum of abdominal segments 8 or 9 or both predominantly
blue; antehumeral pale areas absent or as continuous stripes (very
rarely divided into 2 spots) .................................. 2
2 (1) a) Dorsum of mesothorax solid black with metallic green lustre, pale
antehumeral areas absent; abdominal appendages in lateral view
no greater in length than ,' of segment 10's length (Fig. 16 B); ab-
dominal segments 8 and 9 blue dorsally with distinct black lateral
stripes ............................................. denticollis
b) Dorsum of mesothorax with distinct pale antehumeral areas; no
such combination of above characters ...................... .. 3
3 (2) a) Superior abdominal appendage in lateral view bifid with distinct
mesial lobe directed ventrad, lateral lobe of same appendage di-
rected posteriorly; apical dorsal margin of abdominal segment 10
prolonged into distinct spine (Fig. 16 C); abdominal segment 8
black dorsally; body length greater than 30 mm ........ prognatha
b) Superior abdominal appendage in lateral view not bifid (slightly so
in I. ramburii depending on articulation of appendages) (Fig. 16
D-H); apical dorsal margin of abdominal segment 10 not raised
into distinct spine (except in I. demorsa which is always less than
30 mm in body length); abdominal segment 8 predominantly blue
dorsally .................................................. 4
4 (3) a) Inferior abdominal appendage in lateral view bifid with lobe on
ventral margin curved mesially, in dorsal view, both lobes of in-
ferior abdominal appendage project posteriorly beyond superior
appendage (Fig. 16 D; Fig. 17 A); abdominal segment 8 blue dor-
sally, usually black stripe laterally; segment 9 completely blue ....
.................................................... demorsa
b) Inferior abdominal appendage not bifid, confirm with lateral and
dorsal views (Fig. 16 E-H; Fig. 17 B-E); abdominal segments 8 and
9 not as above....................... ......... ......... .. 5
5 (4) a) In dorsal view, each superior abdominal appendage long as or
longer than wide (Fig. 17 B); in lateral view, dorsally-directed lobe
of inferior abdominal appendage may be hidden by overlying su-
perior appendage (Fig. 16 E); abdominal segments 8 and 9 blue


1972








104 BULLETIN FLORIDA STATE MUSEUM


dorsally with black lateral stripe, segment 10 black dorsally with 2
blue spots; pale areas of pterothorax bluish; postocular pale spots
large and triangular................................. kellicotti
b) In dorsal view, each superior abdominal appendage wider than
long (Fig. 17 C-E); abdominal segments 8 and 9 variable, segment
10 black; pale areas of pterothorax greenish; postocular spots
small and circular ......................................... 6
6 (5) a) Abdominal segment 8 blue dorsally with distinct black lateral stripe
or rectangular mark; in dorsal view, apical margins of superior ab-
dominal appendages taper laterally from base to apex (Fig. 17 C)..
.................................. .. vcerticalis



A B C



POS/TA DENTICOLLIS PROGNATHA



D E F



DEMORSA KELLICOTTI VERTICAL/S



G H



BARBER RAMBURII









CARA AARONI INTEGRICOL L IS
FIGURE 16. Male abdominal appendages in Ischnura species, A-H; in pro-
toneurids, I and J; in Nehalennia, K. Orientation same as lateral views of
Fig. 8.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


b) Abdominal segment 8 completely blue (except occasional very nar-
row basal rings); in dorsal view, superior abdominal appendages
with concave or sigmoid apical margins but not distinct lateral di-
vergence (Fig. 17 D, E) .................... .... ........... 7
7 (6) a) In lateral view, inferior abdominal appendage with dorsal margin
concave and apical tip curved dorsally (Fig. 16 G); abdominal seg-
ment 9 completely blue ................................barberi
b) In lateral view, inferior abdominal appendage with dorsal margin
not concave, apical tip of appendage directed posteriorly or me-
sially (Fig. 16 H); abdominal segment 9 with variable amount of dor-
sal black markings ............................. ramburii






A B C


DEMORSA KELLICOTTI VERTICALIS





D E \


BARBER


RA MBURII


HETERODOXUM


DAECKII SALVA HASTATUM
FIGURE 17. Dorsal views of male abdominal appendages in Ischnura species,
A-E; lateral and dorsal views in Hesperagrion, F and G; lateral views in
Teleallagma, Telebasis and Anomalagrion, H, I and J. Lateral view orien-
tation same as Fig. 8.


1972









106 BULLETIN FLORIDA STATE MUSEUM


KEY TO THE FEMALES
1 a) Prothoracic middle lobe in lateral view with distinct tooth-like pro-
jection on dorsolateral surface (actually pair of projections present,
one on either side) (Fig. 11 I); mesostigmal plates in dorsal view
appear to stand on edge and tilt forward (Fig. 15 N).....denticollis
b) Prothoracic middle lobe smoothly convex on dorsolateral surface,
no tooth-like projection present (Fig. 11 L); mesostigmal plates in
dorsal view not standing on edge but variable in degree of ridge for-
mation (Fig. 15 M, O-T)..................................... 2
2 (1) a) Each antehumeral pale area divided into elongate anterior and cir-
cular posterior spots; mesostigmal plates in dorsal view with trans-
verse depressions produced by low ridges on anterior and posterior
margins (Fig. 15 M); dorsum of abdominal segments 8 and 9 black
..... ......... .... ........... ........ p osita
b) Each antehumeral pale area occurs as a narrow continuous stripe
bordered by dark middorsal and humeral stripes humerall stripe
may be absent); mesostigmal plates with unequally developed
ridges on anterior and posterior margins (if no pronounced ridge
differences, dorsum of abdominal segments 8 and 9 predominantly
pale or body length 29 mm or greater) ....................... 3
3 (2) a) In dorsal view, posterior margins of mesostigmal plates meet at mid-
line forming a continuous ridge transversely crossing mesothorax
along a straight or slightly curved line, no indentation of this mar-
gin at midline about apical ends of rami arising from median thor-
acic carina (Fig. 15 0, P) ..................................4
b) In dorsal view, posterior margins of mesostigmal plates not contin-
uous at midline and not transversely crossing mesothorax along
straight or slightly curved line; distinct concave indentation at mid-
line between plates' posterior margins, indentation occupied by
distinct rami of median thoracic carina ....................... .5
4 (3) a) In dorsal view, distinct oblique ridge crossing each mesostigmal
plate from anterior lateral corner to posterior margin near midline,
attaining greatest height near lateral edge (Fig. 15 O).....barberi
b) In dorsal view, no distinct ridge on mesostigmal plates, posterior
lateral margins slightly raised and expanded, and plates' anterior
margins with variable degree of sigmoid shape (Fig. 15 P).ramburii
5 (3) a) In dorsal view, mesostigmal plates' anterior margins developed
into distinct ridges having truncated medial borders, a distinct
tooth or tubercle arising at the plates' posterior medial corners
(Fig. 15 Q); body length less than 30 mm; (lobe on anterior margin
and the blunt, raised prominence of posterior medial corner of each
plate in I. ir..-,,.itl.. much less distinctive than similarly-placed
structures on I. demorsa's plates, and associated with individuals
greater than 30 mm in body length) ..................... demorsa
b) In dorsal view, mesostigmal plates' anterior margins not developed
into ridge as above (lateral half lobe-like in I. prognatha), and
tooth-like projection absent from posterior medial corner of each
plate or as blunt prominence (Fig. 15 R-T) .....................6
6 (5) a) In dorsal view, mesostigmal plates with convex lobe over lateral
half of anterior margin (Fig. 15 R); body length greater than 34
mm ...................................... ....... prognatha


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


b) In dorsal view, mesostigmal plates with no lobe on anterior mar-
gins; body lengths 33 mm or less ............................ 7
7 (6) a) In dorsal view, posterior margin of mesostigmal plate raised into a
distinct ridge for greater part of its length (Fig. 15 S); dorsum of ab-
dominal segment 8, 9 and 10 black or gray (except in general
stages of andromorphic individuals); pruinescence largely obscur-
ing pattern in mature individuals. .................... .verticalis
b) In dorsal view, posterior margins of mesostigmal plates not de-
veloped into a ridge (Fig. 15 T); dorsum of abdominal segments 8,
9 and 10 predominantly pale; pruinescence absent or weakly de-
veloped in all ages.................................. kellicotti
Habitat differences occur between ischnuran species (Johnson, 1966),
but this behavior rarely aids species identification. The close association of
I. kellicotti with lily pads, Nuphar, has been described (Johnson and West-
fall, 1970).

DISTRIBUTION RECORDS FOR TEXAS
Ischnura barberi; Crockett, Pecos and Ward counties.
Ischnura demorsa: Brewster, Culberson, Jeff Davis, Presidio, and Reeves
counties.
Ischnura denticollis: Brewster, Garza, Jeff Davis, and Presidio counties.
Ischnura kellicotti: San Jacinto county.
Ischnura posita: Anderson, Bosque, Brazos, Burnet, Cameron, Cherokee,
Collin, Colorado, Cooke, Dallas, Denton, Grayson, Gregg, Harrison, Hays,
Hunt, Jim Wells, Liberty, Marion, Menard, Polk, Rusk, San Jacinto, Sutton,
Robertson, Upshur, Val Verde, Victoria, Walker, Wilson, and Wood counties.
Ischnura prognatha: San Jacinto county.
Ischnura ramburii: Angelina, Aransas, Brazos, Brazoria, Burnet, Cald-
well, Cameron, Chambers, Colorado, Dallas, Denton, Galveston, Goliad, Gon-
zales, Gregg, Harris, Harrison, Hidalgo, Jackson, Jefferson, Jim Wells,
Kennedy, Kimble, Kleberg, Liberty, Live Oak, Lubbock, Marion, Matagorda,
Polk, Panola, Refugio, Rusk, San Jacinto, San Patricio, Robertson, Tom Green,
Travis, Val Verde, Victoria, Walker, and Williamson counties.
Ischnura verticalis: Cooke, Grayson, and Lubbock counties.
Comments on two questionable species for Texas, I. damula and I. per-
parva, appear in the Discussion.


SMALLER COENAGRIONID GENERA

Anomalagrion Selys, 1857

Anomalagrion hastatum: Females of this small damselfly are similar
to female Ischnura posita, verticalis or possibly demorsa. Female A. hastatum
pass through a color change similar to heteromorphic ischnurans. Early
stages following emergence have a dark dorsal thoracic stripe bordered by
orange. Dark humeral stripes are absent. The orange areas become brown,
and whitish pruinescence develops along the lower lateral sides of the
pterothorax. The changes produce a pale, narrow lateral border to the dor-









108 BULLETIN FLORIDA STATE MUSEUM


sal stripe; at this stage faint humeral stripes appear. Abdominal segments
1-5 and 6-10 are predominantly orange and black respectively; however,
aging changes the whole abdominal dorsum to a bluish-gray. A vulvar
spine occurs in some females; however, it is always quite small if present.
Males are distinct in the field with their slender, yellow-orange abdomen.
Lentic habits with emergent vegetation are characteristic.

DISTRIBUTION RECORDS FOR TEXAS:
Aransas, Austin, Blanco, Bosque, Brazos, Brazoria, Brooks, Burnet, Cam-
eron, Chambers, Collin, Colorado, Cooke, Dallas, Denton, Goliad, Gregg,
Guadalupe, Hardin, Harris, Harrison, Hidalgo, Hunt, Jim Wells, Kendall, Kle-
berg, Lamar, Madison, Marion, Matagorda, Montgomery, Orange, Panola,
Polk, Reeves, Rusk, San Jacinto, San Patricio, Robertson, Travis, Upshur,
Uvalde, Victoria, Walker, Williamson, and Wise counties.


Hesperagrion Calvert, 1902

Hesperagrion heterodoxum.-Abdominal appendages and mesostigmal
plates in the male and female respectively identify this species; however,
color and stripe patterns are exceptionally variable. Calvert (1901-1908) as-
sumed the differences represented aging effects and described several
stages from tenerals to mature adults. I cultured numerous larvae through
metamorphosis and observed color patterns in breeding adults in south-
western New Mexico providing additional evidence. All males shortly
after emergence are brownish-yellow other than a dark middorsal thoracic
stripe and a dark longitudinal line on the dorsum of abdominal segments
4-7. At maturity the yellowish areas of the abdomen turn to orange-red on
segments 8, 9 and 10: segments 1, 2 and 3 are orange or greenish-yellow.
The dorsum of the head is black excepting large red postocular spots, the
face is pale tan excepting a black transverse band on the postclypeus, the
prothorax is dorsally black and the dorsum of the pterothorax is black lat-
erally to at least the humeral suture, and the pale antehumeral areas exist as
two isolated spots (rarely connected). Pale areas of the pterothorax are blue
or cream in color. The femur and tibia develop black stripes laterally. Fe-
males at emergence occur in one of two patterns. Most females are brown-
ish-yellow without any dark stripe pattern. Suture lines are somewhat
darker, especially on the head; the postocular region is frequently dark
brown but without definite postocular spots. Other females at emergence
possess the color and stripe pattern described above for general males. Cal-
vert's description of the variation is not clear relative to the condition he
associated with maturity for females; however, two female types exist at
maturity in H. heterodoxum as with many Ischnura species.
At maturity, heteromorphic females remain brownish-yellow without
any dark pattern (excepting actual suture lines). The color becomes fully
brown or tan and loses all yellow attributes. Such females mate, oviposit,
and judging from the condition of wings and exocuticle, they reach ad-


Vol. XVI No. 2








JOHNSON: TEXAS DAMSELFLIES


vanced age for a damselfly without further change in color or pattern. An-
dromorphic females are similar to males at maturity except that abdominal
segments 9 and 10 are black, and segment 9 is blue dorsally with a black
apical band while segments 1-7 are dorsally black. Reddish postocular
spots develop but the surrounding black color spreads with age and the
spots are lost. Females emerging with a male-like pattern become andro-
morphic types at maturity. These andromorphic females also mate and ovi-
posit. Males and andromorphic females do, in fact, change considerably in
color pattern from general stage to maturity. If only museum specimens
were available for comparison, color patterns at emergence unknown, and
maturity, judged by mating behavior, unassociated with color patterns,
recognition of dimorphic females would be unlikely.
Males are not similar to other species in Texas, whereas heteromorphic
females are similar in flight to several coenagrionids. Characteristic habi-
tats are small streams in the xeric southwest.

DISTRIBUTION RECORDS FOR TEXAS
Brewster, Culberson, Jeff Davis, and Presidio counties.



Nehalennia Selys, 1850

Nehalennia integricollis:-Both sexes of N. integricollis have a solid
greenish-bronze thoracic dorsum without pale antehumeral regions. The
abdominal segments are dorsally dark-green or bronze excepting some blue
on segments 8, 9 and 10 in the males. Lentic habitats are typical for the
species.

DISTRIBUTION RECORDS FOR TEXAS
Montgomery and San Jacinto counties.



Teleallagma Kennedy, 1920

Teleallagma daeckii:-A long, slender abdomen, greater than 34 mm
in length, and pale body colors distinguish this damselfly. Males are pale
blue, females are brown to green-yellow, and both sexes have reduced dark
stripe patterns. These characters effectively separate T. daeckii from other
Texas coenagrionids. Perching habit (wings folded together over the back)
and color pattern eliminate confusion with lestids. Margins of swamps and
lentic water describe our current knowledge of preferred habitats.


DISTRIBUTION RECORD FOR TEXAS
Montgomery County.


1972








110 BULLETIN FLORIDA STATE MUSEUM


Telebasis Selys, 1865

Telebasis salva:-The males are the only damselflies in Texas with a
bright red abdomen, brownish-red thorax, and a middorsal dark thoracic
stripe having a tooth-like notch laterally on each side in the posterior half.
Females possess the characteristic middorsal dark stripe, but have brown
bodies and are less distinct when observed in flight. Habitats include both
lentic and lotic situations.

DISTRIBUTION RECORDS FOR TEXAS
Bexar, Blanco, Bosque, Brazos, Brewster, Burnet, Caldwell, Crosby, Cul-
berson, Dallas, Denton, Gillespie, Grayson, Grimes, Hays, Hidalgo, Hill, Jeff
Davis, Jim Wells, McCulloch, Menard, Reeves, Robertson, Travis, Uvalde, Val
Verde, Victoria, Williamson, and Wilson counties.


D


ARGIA ALBERTA


ISCHNURA
DA MUL A


F
ENALLAGMA
CARUNCULATUM


HETAER/NA
VUL NERA TA


LESTES
CONGENER


FIGURE 18. Possible additions to Texas fauna. Male structures, A, B, E, G-I;
female structures, C, D, F. Orientation same as proceeding figures.


Vol. XVI No. 2


~









JOHNSON: TEXAS DAMSELFLIES


DISCUSSION

QUESTIONABLE RECORDS AND SYNONYMS. The literature carries
several references to species occurring in Texas without adequate docu-
mentation, and Table 3 lists these species. The case of Lestes simplex ap-
pears under that genus. Erroneous determinations account for the follow-
ing records. Calvert's (1893) references to Lestesforcipatus was actually L.
disjunctus (see Walker, 1953), and Tinkham's (1934) record for L. un-
guiculatus was actually a specimen of Argia plana (see Gloyd, 1958).

TABLE 3. UNSUBSTANTIATED RECORDS FOR THE TEXAS DAMSELFLY FAUNA

Erroneous or questionable literature records for Damselflies in Texas. See
Discussion Text.

Lestes congener Hagen 1861 Hetaerina sempronia Hagen 1853
Lestes dryas Kirby 1890 Hetaerina vulnerata Hagen 1853
Lestesforcipatus Rambur 1842 Amphiagrion saucium (Burmeister) 1839
Lestes simplex Hagen 1861 Enallagma cyathigerum (Charpentier) 1840
Lestes unguiculatus Hagen 1861 Ischnura perparva Selys 1876

Probable additions to the Texas Damselfly Fauna. See Text.

Lestes congener Hagen 1861 Enallagma carunculatum Morse 1891
Argia alberta Kennedy 1918 Ischnura damula Calvert 1901
Enallagma boreale Selys 1876


Donnelly, who described Enallagma westfalli in 1964, now considers it
as E. traviatum westfalli (Donnelly, pers comm., 1970). References to other
species for Texas exist, but the localities given are only 'Texas', determina-
tions are questionable, and actual specimens are unavailable for examina-
tion. These records include Lestes congener listed for Texas in several
publications, but all such records trace to Hagen (1861). Calvert (1901-
1908) listed Ischnura perparva from Texas (near San Antonio) on the basis
of one male. The species has otherwise a northwestern distribution. All
references to I. perparva from Texas trace to this one record, and I am con-
sidering it unsubstantiated until confirmed by additional collecting.
Calvert (1901-1908) listed Hetaerina sempronia for Texas on the basis of
three males reported in 1875 but not actually seen by Calvert. The vari-
ability of H. titia may account for these records. Some H. titia appear
similar to H. sempronia in wing pattern. Currently no firm record of H.
sempronia is available for Texas. Tucker (1908) reported on Odonata
from Piano, Texas, (north of Dallas) including H. vulnerata. Wing patterns
are quite similar in both H. vulnerata and H. americana; however, the
abdominal appendages in dorsal view of americana have a distinct lobe
on the medial margins and the same margins in vulnerata are without
lobes (Fig. 6 A and Fig. 18 G). Hetaerina vulnerata occurs in southwest
New Mexico and south in Mexico and this geographical pattern casts


1972








112 BULLETIN FLORIDA STATE MUSEUM


strong doubt on the central Texas record. I assume Tucker had H. ameri-
cana, a highly variable species with at least five synonyms (see Needham
and Heywood, 1929).
An unpublished thesis on the Odonata of Texas submitted to Trinity
University, San Antonio, in 1952 by P. N. Albright lists, among other
species, Lestes congener, L. dryas, L. unguiculatus, Amphiagrion sauc-
ium, Enallagma cyathigerum, and Hetaerina sempronia. Albright's speci-
mens are not available, and the source of his determinations is not clear.
Acceptance of these species for Texas requires documentation by specimens
with verified determinations.
From the above questionable species, Lestes congener may exist in
Texas as it reaches southern New Mexico and Ischnura damula exists as a
probable species on the same basis. Lestes congener males are distinct from
other Texas lestids in structure of abdominal appendages. Each inferior ab-
dominal appendage in dorsal view extends posteriorly no further than the
midpoint of the serrated lobe on the superior appendage (Fig. 18 I). Each
antehumeral pale area in Ischnura damula exists as two spots, and the su-
perior abdominal appendage in lateral view is deeper at its base than the
appendage is long. (Fig. 18 H). Enallagma boreale occurs at low eleva-
tions in central New Mexico, and its appearance in west Texas is pos-
sible. Descriptions of these three species are in Walker (1953).
Bick and Bick (1957) summarized the Odonata distributions in Okla-
homa. Argia alberta and Enallagma carunculatum occurred respectively in
Harmon and Cimarron Counties, Oklahoma, adjoining Texas in the pan-
handle area. Enallagma carunculatum occurs also in Roosevelt County,
New Mexico, adjoining Texas. These two northern species may well have
populations in Texas and the keys include them. Ischnura barberi occurred

TABLE 4. SYNONYMS FOR TEXAS DAMSELFLIES APPEARING IN POST-1900 LITERATURE


Species:


Synonym:


Calopteryx dimidiata Agrion dimidiatum (Burmeister) 1839
Calopteryx maculata Agrion maculatum Beauvois 1805
Hetaerina titia Hetaerina tricolor (Burmeister) 18391
Argia fumipennis violacea Argia violacea (Hagen) 1861
Argia lugens Hyponeura lugens Hagen 1861
Argia moesta Argia putrida Hagen 1861 and
Argia intruda Williamson 1912
Argia munda Argia vivida munda Calvert 1902 and
Argia rita Kennedy 1919
Argia nahuana Argia agrioides nahuana Calvert 1902
Argia plana Argia vivida plan Calvert 1902
Enallagma novaehispaniae Enallagma coecum novae-hispaniae Calvert 1907
Ischnura barberi Ischnura utahensis Muttkowski 1910
Ischnura ramburii Ischnura ramburii credula Hagen 1861 and
Ischnura credula Hagen 1861
Teleallagma daeckii Telagrion daeckii Calvert 1903 and
Enallagma daeckii (Calvert) 1903
1 See discussion under the genus Hetaerina.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


in Jackson County, Oklahoma, adjoining Texas far east of its easternmost
Texas county record. Data for I. barberi are probably very incomplete.
Louisiana records (Bick, 1957) reveal no species west of the Mississippi
River not also occurring in Texas. Damselfly distribution in the Mexican
states adjacent to the Texas border is not well known.
Most synonyms appeared prior to 1900 and rarely occur in the liter-
ature. Synonyms published after 1900 appear in Table 4. Needham and
Heywood (1929) placed the genus Neoneura in the family Coenagrionidae.
The family name Agriidae often appears for Calopterygidae, while Coen-
agriidae frequently appears for Coenagrionidae. The name Agrionidae has
a confusing history representing at different times both Calopterygidae
and Coenagrionidae. Minor spelling variations of names occurred with
usage (examples, Lestes inequalis; Ischnura ramburi; Teleallagma daecki;
and Heterina).
GEOGRAPHICAL DISTRIBUTION. The wide range of habitats within
Texas and its geographic location produce convergence of typically
eastern, western, and neotropical faunas. Many species consequently
have range limits in the state and the following distributional patterns
result.
Statewide distribution, where suitable habitat occurs, exists
for the following damselflies. Species occurring across most of the
United States, parts of Canada and south into Mexico are Hetaerina
americana, Argia moesta, A. sedula, and Enallagma civil. Argia
fumipennis violacea and Enallagma basidens occur widely in the east,
north, south, and west to at least Arizona. Telebasis salva occurs widely
in the west, Kansas and Louisiana to the north and east respectively.
The following species have their western limits of distribution
within Texas, also occur north or both north and south into Mexico,
and have their westernmost Texas county records in parentheses.
Lestes disjunctus (Jeff Davis), L. inaequalis (Angelina), L. vigilax
(Walker), Calopteryx dimidiata (San Jacinto), C. maculata (Hemp-
hill), Hetaerina titia (Presidio), Argia apicalis (Lubbock), A. bipunc-
tulata (Wood), A. tibialis (Victoria), Enallagma divagans (Grayson),
E. dubium (Harris), E. durum (San Patricio), E. exsulans (Val
Verde); E. geminatum (Matagorda), E. signatum (Uvalde), E. travia-
tum (Grayson), E. vesperum (Wood). Ischnura kellicotti (San Jacinto), I.
posita (Val Verde), I. prognatha (San Jacinto), I. ramburii (Lubbock). I.
verticalis (Lubbock), Anomalagrion hastatum (Reeves), Nehalennia
integricollis (Montgomery), and Teleallagama daeckii (Montgomery).
Three of these eastern species have more western populations than indi-
cated above. Smith and Pritchard (1956) report I. ramburii in California.
I can trace no confirmation for the California record, and the species is ap-
parently absent in xeric parts of west Texas and the southwest although
abundant farther south in Mexico. Ischnura posita has apparently reached
the Hawaiian Islands yet it has failed to colonize western North America
(Zimmerman, 1948). Calvert (1903) gives one record of I. verticalis from








114 BULLETIN FLORIDA STATE MUSEUM


New Mexico. That species may occur over much of north Texas judging
from Bick and Bick's (1957) report of the Odonata of Oklahoma.
The following species have their eastern limits of distribution within
Texas, also occur north or both north and south into Mexico, and have
their easternmost county records in parentheses. Archilestes grandis (Dal-
las), Lestes alacer (Matagorda), Argia lugens (Crosby), A. nahuana (Rob-
ertson), A. plan (Wood), Enallagma praevarum (Blanco), Ischnura
barberi (Crockett), I. demorsa (Brewster), and I. denticollis (Garza).
The following species reach Texas from the south or southwest and
have their northernmost county records in parentheses. Lestes forficula
(Brazos), L. sigma (Gonzales), Neoneura aaroni (Caldwell), Protoneura
cara (Kendall), Argia barretti (Kimble), A. hinei (Brewster), A. munda
(Jeff Davis). A. rhoadsi (Cameron), Enallagma novaehispaniae (Hays),
and Hesperagrion heterodoxum (Brewster).
Two species occurring in Texas are apparently absent from New Mex-
ico and Louisiana but range north and south of the state. These species
with eastern and westernmost county records respectively are Argia im-
munda (Rusk, Jeff Davis) and A. translate (Brazos, Howard).
SEASONAL DISTRIBUTION.--The seasonal distribution or adult flight
season of most species extends over spring and summer months andmay
persist into cool weeks of early fall. Many dragonflies (Anisoptera) ap-
parently have an obligate diapause associated with a massed, synchronized
spring or early summer emergence. Such species consist of a homogenous
adult age group, growth patterns require at least a full year's cycle, often
longer, and adults vanish from the scene when their average life ex-
pectancy expires. Most damselflies appear to differ from this pattern by
having growth controlled in a faculative manner. As soon as temperature
conditions in the spring permit, emergence begins and continues through-
out much of the summer.
Emergence in north Texas may therefore be several weeks later than
for the same species in southern parts of the state. The populations
have a heterogeneous age structure, and growth rates may permit more
than one generation in a year. The lestids, calopterygids, and larger coena-
grionids appear to require, for most species and habitats, a year's life cycle.
The smaller coenagrionids may have two to three generations in a year.
Life cycles are typically longer in species restricted to streams than in
forms characteristic of lentic habitats. The date of emergence will vary
from year to year for a given habitat as local climate varies. In areas where
warm springs occur (ex. Palmetto State Park), adults fly earlier than in sur-
rounding habitats having lower temperatures. Adults in southern coun-
ties will likewise exist later into the fall season than counterparts to the
north and may have twice the generation number a year.
For these reasons, a comprehensive knowledge of flight seasons will
require more ecological data than currently available. A few exceptions
to these generalizations exist. The form of Hetaerina titia known as tri-
color is characteristically a spring form in central Texas (Johnson, 1963).


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


The records for Enallagma divagans suggest it to be a spring to early
summer species and notes for Archilestes grandis and Lestes congener
point to late summer and fall flight seasons.



LITERATURE CITED

Albright, P. N. 1952. Contributions to the knowledge of the Odonata of Texas
with particular attention to the area surrounding San Antonio. M. S.
Thesis. Trinity Univ., San Antonio, Texas.
Bick, G. H. 1957. The Odonata of Louisiana. Tulane Stud. Zool. 5: 71-135.
Bick, G. H. and J. C. Bick. 1957. The Odonata of Oklahoma. Southwestern Nat.
2: 1-18.
Calvert, P. P. 1893. Catalogue of the Odonata of the vicinity of Philadelphia,
with an introduction to the study of this group of insects. Trans. Amer.
Ent. Soc. 22: 152a-272.
1901-1908. Odonata. In: Biologia Centrali-Americana, Vol. 50: Neur-
optera. xxx + 420 pp.
1903. A list of insects of Beulah, New Mexico, edited by Henry
Skinner. Trans. Amer. Ent. Soc. 29: 42-43.
Donnelly, T. W. 1964. Enallagma westfalli, a new damselfly from eastern Texas,
with remarks on the genus Teleallagma Kennedy. Ent. Soc. Wash. 66: 103-109.
Ferguson, A. 1940. A preliminary list of the Odonata of Dallas County, Texas.
Field and Laboratory. 88: 1-10.
1942. Scattered records of Texas and Louisiana Odonata with addi-
tional notes on the Odonata of Dallas County, Field and Laboratory. 10:
145-149.
Fraser, F. C. 1954. The origin and descent of the order Odonata based on the
evidence of persistent archaic characters. Proc. R. Ent. Soc. London (B).
23: 89-94.
Gloyd, L. K. 1932. Four new dragonfly records for the United States. Ent.
News. 43: 189-190.
1958. The dragonfly fauna of the Big Bend Region of Trans-Pecos
Texas. Occ. Pap. Mus. Zool. Univ. Mich. 593: 1-29.
Hagen, H. 1861. Synopsis of the Neuroptera of North America, with a list of
the South American species. Smithsonian Misc. Coll. 4: 1-347.
Harwell, J. E. 1951. Notes on the Odonata of Northeastern Texas. Tex. J. Sci.
3: 204-207.
Johnson, C. 1961. Breeding behavior and oviposition in Hetaerina americana
(Fabricius) and H. titia (Drury) (Odonata: Agriidae). Canad. Ent. 93:
260-266.
-- 1962. A description of territorial behavior and a quantitative study
of its function in males of Hetaerina americana (Fabricius) (Odonata:
Agriidae). Can. Ent. 94: 178-190.
.1963. Interspecific territoriality in Hetaerina americana (Fabri-
cius) and H. titia (Drury) (Odonata: Calopterygidae) with a preliminary
analysis of the wing color pattern variation. Canad. Ent. 95: 575-582.
1964. Polymorphism in the damselflies. Enallagma civil (Hagen)
and E. praevarum (Hagen) Amer. Midl. Naturalist. 72: 408-416.


1972









116 BULLETIN FLORIDA STATE MUSEUM


1966. Environmental modification of the behavior for habitat
selection in adult damselflies. Ecology. 47: 674-676.
1969. Genetic variability in ischnuran damselflies. Amer. Midl.
Naturalist. 81: 39-46.
Johnson, C. and M. J. Westfall, Jr. 1970. Diagnostic keys and notes on the
damselflies (Zygoptera) of Florida. Bull. Florida State Mus. 15: 45-89.
Kennedy, C. II. 1915. Notes on the life history and ecology of the dragonflies
(Odonata) of Washington and Oregon. Proc. U. S. Nat. Mus. 49: 259-345.
Muttkowski, R. A. 1910. Catalogue of the Odonata of North America. Bull. Pub.
Mus. City of Milwaukee. 1: 1-207.
Needham, J. G. and H. B. Heywood. 1929. A Handbook of the Dragonflies of
North America, C. C. Thomas. Springfield, Ill. and Baltimore, Md. VIII +
378 pp.
Paulson, D. R. 1966. New Records of Bahamian Odonata. Quart. J. Florida
Acad. Sci. 29: 97-110.
Smith, R. F. and A. E. Pritchard. 1956. Odonata. Chapter 4 in Aquatic Insects
of California edited by R. L. Usinger. Univ. Calif. Press, Berkeley. IX +
508 pp.
Tillyard, R. J. 1917. The Biology of Dragonflies (Odonata or Paraneuroptera).
Cambridge Univ. Press. 380 pp.
Tinkham, E. R. 1934. The dragonfly fauna of Presidio and Jeff Davis Counties
of the Big Bend Region of Trans-Pecos, Texas. Can. Ent. 66: 213-218.
Tucker, E. S. 1908. Incidental captures of neuropterous insects at Plano,
Texas. Psyche. 15: 97-100.
Walker, E. M. 1953. The Odonata of Canada and Alaska. Vol. 1. Univ. Tor-
onto Press, Toronto; xi + 292 pp.
Williamson, E. B. 1912. The dragonfly Argia moesta and a new species (Odonata).
Ent. News. 23: 196-203.
S1914. Dragonflies collected in Texas and Oklahoma. Ent. News.
25: 411-415, 444-455.
1917. The Genus Neoneura. Trans. Amer. Ent. Soc. 43: 211-246.
Zimmerman, E. C. 1948. Insects of Hawaii. Vol. 2 Univ. of Hawaii Press,
Honolulu. X + 475 pp.


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES

APPENDIX
Zygoptera known from Texas Counties


Anderson Co.
Calopteryx maculata
Argia moesta
Enallagma signatum
Ischnura posita
Andrews Co. (no records)
Angelina Co.
Lestes inaequalis
Hetaerina titia
Argia apicalis
Argia moesta
Argia tibialis
Enallagma exsulans
Ischnura ramburii
Aransas Co.
Lestes disjunctus
Calopteryx maculata
Enallagma civil
Ischnura ramburii
Anomalagrion hastatum
Archer Co. (no records)
Armstrong Co. (no records)
Atascosa Co. (no records)
Austin Co.
Argia fumipennis
Argia sedula
Enallagma divagans
Enallagma signatum
Anomalagrion hastatum
Bailey Co. (no records)
Bandera Co. (no records)
Bastrop Co.
Calopteryx maculata
Baylor Co.
Hetaerina americana
Argia moesta
Bee Co.
Argia sedula
Bell Co.
Archilestes grandis
Bexar Co.
Lestesforficula
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia fumipennis
Argia immunda


Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Telebasis salva
Blanco Co.
Lestes alacer
Lestes disjunctus
Hetaerina americana
Argia fumipennis
Argia immunda
Argia moesta
Argia nahuna
Argia plan
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Enallagma praevarum
Anomalagrion hastatum
Telebasis salva
Borden Co. (no records)
Bosque Co.
Lestes alacer
Hetaerina americana
Hetaerina titia
Argia apicalis
Argiafumipennis
Argia immunda
Argia moesta
Argia nahuana
Argia plan
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Enallagma exsulans
Ischnura posita
Anomalagrion hastatum
Telebasis salva
Bowie Co.
Calopteryx maculata
Enallagma basidens
Brazoria Co.
Ischnura ramburii
Anomalagrion hastatum


1972.









118 BULLETIN FLORIDA STATE MUSEUM


Brazos Co.
Lestes alacer
Lestes disjunctus
Lestesforficula
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Argia tibialis
Argia translate
Enallagma civil
Enallagma divagans
Enallagma exsulans
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Brewster Co.
Archilestes grandis
Lestes disjunctus
Hetaerina americana
Argiafumipennis
Argia hinei
Argia immunda
Argia lugens
Argia moesta
Argia nahuana
Argia plan
Enallagma civil
Ischnura demorsa
Ischnura denticollis
Hesperagrion heterodoxum
Telebasis salva
Briscoe Co. (no records)
brooks Co.
Enallagma civil
Anomalagrion hastatum
Brown Co. (no records)
Burleson Co. (no records)
Burnet Co.
Argia moesta
Argia sedula
Enallagma basidens
Enallagma civil
Enallagma signatum
Ischnura posita
Ischnura ramburii


Anomalagrion hastatum
Telebasis salva
Caldwell Co.
Lestes alacer
Hetaerina americana
Hetaerina titia
Neoneura aaroni
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma exsulans
Ischnura ramburii
Telebasis salva
Calhoun Co. (no records)
Callahan Co. (no records)
Cameron Co.
Lestesforficula
Lestes sigma
Argia apicalis
Argia rhoadsi
Argia sedula
Enallagma basidens
Enallagma civil
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Camp Co. (no records)
Carson Co. (no records)
Cass Co. (no records)
Castro Co. (no records)
Chambers Co.
Argia apicalis
Ischnura ramburii
Anomalagrion hastatum
Cherokee Co.
Calopteryx maculata
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia bipunctulata
Argia moesta
Argia sedula
Enallagma basidens
Ischnura posita


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


Childress Co.
Hetaerina americana
Enallagma civil
Clay Co. (no records)
Cochran Co. (no records)
Coke Co. (no records)
Coleman Co. (no records)
Collin Co.
Calopteryx maculata
Argia apicalis
Argia moesta
Argia nahuana
Argia plan
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Ischnura posita
Anomalagrion hastatum
Collingsworth Co. (no records)
Colorado Co.
Lestes disjunctus
Hetaerina americana
Argia apicalis
Argia sedula
Enallagma basidens
Enallagma signatum
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Comal Co.
Hetaerina americana
Hetaerina titia
Argia barretti
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma novaehispaniae
Comanche Co. (no records)
Concho Co. (no records)
Cooke Co.
Hetaerina americana
Argia apicalis
Argiafumipennis
Argia moesta
Argia nahuana
Argia sedula
Argia translate


Enallagma basidens
Ischnura posita
Ischnura verticalis
Anomalagrion hastatum
Coryell Co. (no records)
Cottle Co. (no records)
Crane Co. (no records)
Crockett Co.
Ischnura barberi
Crosby Co.
Archilestes grandis
Lestes alacer
Hetaerina americana
Argia immunda
Argia lugens
Argia moesta
Argia nahuana
Argia plana
Enallagma civil
Telebasis salva
Culberson Co.
Argia plan
Ischnura demorsa
Hesperagrion heterodoxum
Telebasis salva
Dallam Co. (no records)
Dallas Co.
Archilestes grandis
Lestes disjunctus
Calopteryx maculata
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia immunda'
Argia moesta
Argia nahuana
Argia plan
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Dawson Co. (no records)
Deaf Smith Co. (no records)
Delta Co. (no records)


1972









120 BULLETIN FLORIDA STATE MUSEUM


Denton Co.
Archilestes grandis
Calopteryx maculata
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Enallagma signatum
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
De Witt Co. (no records)
Dickens Co. (no records)
Dimmit Co.
Argia sedula
Enallagma civil
Donley Co. (no records)
Duval Co. (no records)
Eastland Co. (no records)
Ector Co. (no records)
Edwards Co. (no records)
Ellis Co. (no records)
El Paso Co. (no records)
Erath Co. (no records)
Falls Co.
Argia apicalis
Fannin Co.
Argia apicalis
Argia moesta
Enallagma basidens
Fayette Co.
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia moesta
Argia sedula
Argia translate
Fisher Co. (no records)
Floyd Co. (no records)
Foard Co. (no records)
Fort Bend Co. (no records)
Franklin Co.
Argia tibialis


Enallagma basidens
Enallagma civil
Enallagma divagans
Freestone Co. (no records)
Frio Co. (no records)
Gaines Co. (no records)
Galveston Co.
Ischnura ramburii
Garza Co.
Ischnura den ticollis
Gillespie Co.
Hetaerina americana
Argia fimipennis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Enallagma praevarum
Telebasis salva
Glasscock Co. (no records)
Goliad Co.
Hetaerina americana
Hetaerina titia
Neoneura aaroni
Argia apicalis
Argia moesta
Argia sedula
Ischnura ramburii
Anomalagrion hastatum
Gonzales Co.
Lestes alacer
Lestes sigma
Hetaerina americana
Hetaerina titia
Neoneura aaroni
Argia apicalis
Argia moesta
Argia sedula
Argia translate
Ischnura ramburii
Gray Co. (no records)
Grayson Co.
Calopteryx maculata
Argia apicalis
Argia moesta
Argia sedula
Enallagma basidens


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JOHNSON: TEXAS DAMSELFLIES


Grayson Co. Continued
Enallagma civil
Enallagma divagans
Enallagma traviatum
Ischnura posita
Ischnura verticalis
Telebasis salva
Gregg Co.
Calopteryx maculata
Hetaerina americana
Argia apicalis
Argia bipunctulata
Argiafumipennis
Argia moesta
Argia tibialis
Enallagma basidens
Enallagma divagans
Enallagma exsulans
Ischnura posita
Ischnura ramburii
Anomalgrion hastatum
Grimes Co.
Calopteryx maculata
Hetaerina americana
Hetaerina titia
Argia apicalis
Argiafumipennis
Argia immunda
Argia moesta
Argia sedula
Argia tibialis
Argia translate
Enallagma basidens
Enallagma exsulans
Enallagma signatum
Telebasis salva
Guadalupe Co.
Hetaerina titia
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma civil
Anomalagrion hastatum
Hale Co. (no records)
Hall Co. (no records)
Hamilton Co. (no records)
Hansford Co. (no records)


Hardeman Co. (no records)
Hardin Co.
Lestes disjunctus
Calopteryx dimidiata
Argia tibialis
Anomalagrion hastatum
Harris Co.
Lestes disjunctus
Argia apicalis
Argia sedula
Enallagma dubium
Enallagma geminatum
Enallagma signatum
Ischnura ramburii
Anomalagrion hastatum
Harrison Co.
Lestes inaequalis
Argia apicalis
Argiafumipennis
Enallagma traviatum
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Hartley Co. (no records)
Haskell Co. (no records)
Hays Co.
Archilestes grandis
Hetaerina americana
Hetaerina titia
Argia baretti
Argia immunda
Argia moesta
Argia nahuana
Argia sedula
Argis translate
Enallagma exsulans
Enallagma novaehispaniae
Enallagma signatum
Ischnura posita
Telebasis salva
Hemphill Co.
Calopteryx maculata
Henderson Co.
Argia tibialis
Hidalgo Co.
Lestesforficula
Protoneura cara
Argia apicalis
Argia immunda


1972









122 BULLETIN FLORIDA STATE MUSEUM


Hidalgo Co. Continued
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma civile
Enallagma novaehispaniae
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Hill Co.
Lestes alacer
Hetaerina americana
Argia moesta
Argia nahuana
Argia plan
Argia sgdula
Enallagma basidens
Enallagma civil
Enallagma exsulans
Telebasis salva
Hockley Co. (no records)
Hood Co. (no records)
Hopkins Co. (no records)
Houston Co.
Calopteyx maculata
Enallagma civil
Howard Co.
Argia moesta
Argia translate
Enallagma civil
Hudspeth Co. (no records)
Hunt Co.
Lestes disjunctus
Argia apicalis
Enallagma basidens
Enallagma civil
Enallagma signatum
Ischnura posita
Anomalagrion hastatum
Hutchinson Co. (no records)
Irion Co. (no records)
Jack Co. (no records)
Jackson Co.
Hetaerina titia
Argia tibialis
Enallagma civil
Ischnura ramburii
Jasper Co. (no records)


Jeff Davis Co.
Archilestes grandis
Lestes alacer
Lestes disjunctus
Hetaerina americana
Argiafumipennis
Argia hinei
Argia immunda
Argia lugens
Argia moesta
Argia munda
Argia nahuana
Argia plana
Argia sedula
Enallagma civil
Enallagma praevarum
Ischnura demorsa
Ischnura denticollis
Hesperagrion heterodoxum
Telebasis salva
Jefferson Co.
Ischnura ramburii
Jim Hogg Co. (no records)
Jim Wells Co.
Lestes disjunctus
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Enallagma basidens
Enallagma civil
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Johnson Co. (no records)
Jones Co. (no records)
Karnes Co.
Argia apicalis
Kaufman Co. (no records)
Kendall Co.
Hetaerina americana
Hetaerina titia
Protoneura cara
Argia apicalis
Argiafumipennis
Argia immunda


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JOHNSON: TEXAS DAMSELFLIES


Kendall Co. Continued
Argia moesta
Argia nahuana
Argia sedula
Argia translate
Enallagma basidens
Enallagma exsulans
Anomalagrion hastatum
Kenedy Co.
Enallagma civil
Ischnura ramburii
Kent Co. (no records)
Kerr Co.
Hetaerina americana
Argia immunda
Argia moesta
Argia sedula
Kimble Co.
Lestes alacer
Hetaerina americana
Argia barretti
Argia immunda
Argia moesta
Argia nahuana
Argia sedula
Argia translate
Enallagma civil
Ischnura ramburii
King Co. (no records)
Kinney Co.
Argiafumipennis
Argia sedula
Enallagma basidens
Kleberg Co.
Lestesforficula
Lestes sigma
Enallagma civil
Ischnura ramburii
Anomalagrion hastatum
Knox Co. (no records)
Lamar Co.
Lestes disjunctus
Enallagma divagans
Anomalagrion hastatum
Lamb Co. (no records)
Lampasas Co. (no records)
La Salle Co.
Argia apicalis
Lavaca Co. (no records)


Lee Co. (no records)
Leon Co.
Argia apicalis
Argia tibialis
Liberty Co.
Argia apicalis
Argia moesta
Argia sedula
Argia tibialis
Enallagma basidens
Enallagma exsulans
Enallagma signatum
Enallagma traviatum
Ischnura posita
Ischnura ramburii
Limestone Co.
Hetaerina americana
Argia apicalis
Argia moesta
Argia sedula
Argia translate
Lipscomb Co. (no records)
Live Oak Co.
Ischnura ramburii
Llano Co.
Hetaerina americana
Argia moesta
Loving Co. (no records)
Lubbock Co.
Archilestes grandis
Lestes alacer
Lestes disjunctus
Hetaerina americana
Argia apicalis
Argia lugens
Argia sedula
Enallagma basidens
Enallagma civil
Ischnura ramburii
Ischnura verticalis
Lynn Co. (no records)
Madison Co.
Anomalagrion hastatum
Marion Co.
Calopteryx maculata
Argia apicalis
Argiafumipennis
Argia moesta
Enallagma basidens










124 BULLETIN FLORIDA STATE MUSEUM


Marion Co. Continued
Enallagma exsulans
Enallagma signatum
Enallagma traviatum
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Martin Co. (no records)
Mason Co. (no records)
Matagorda Co.
Lestes alacer
Argia apicalis
Argia moesta
Argia sedula
Argia tibialis
Enallagma civil
Enallagma durum
Enallagma geminatum
Enallagma signatum
Ischnura ramburii
Anomalagrion hastatoum
Maverick Co.
Argia moesta
Enallagma civil
McCulloch Co.
Telebasis salva
McLennan Co.
Hetaerina titia
Argia apicalis
Argia moesta
Argia plan
Argia sedula
Argia translate
Enallagma civil
McMullen Co. (no records)
Medina Co.
Hetaerina americana
Neoneura aaroni
Protoneura cara
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Menard Co.
Hetaerina americana
Argia nahuana
Argia sedula


Ischnura posita
Telebasis salva
Midland Co. (no records)
Milam Co. (no records)
Mills Co. (no records)
Mitchell Co. (no records)
Montague Co.
Archilestes grandis
Montgomery Co.
Lestes disjunctus
Calopteryx maculata
Argia apicalis
Argia fimipennis
Argia moesta
Argia sedula
Argia tibialis
Enallagma basidens
Anomalagrion hastatum
Nehalennia intergricollis
Teleallagma daeckii
Moore Co. (no records)
Morris Co.
Argia apicalis
Argia tibialis
Motley Co. (no records)
Nacogdoches Co.
Lestes disjunctus
Calopteryx maculata
Argia tibialis
Enallagma exsulans
Navarro Co. (no records)
Newton Co. (no records)
Nolan Co. (no records)
Nueces Co.
Neoneura aaroni
Enallagma basidens
Ochiltree Co. (no records)
Oldham Co. (no records)
Orange Co.
Argia apicalis
Argia tibialis
Anomalagrion hastatum
Palo Pinto Co.
Hetaerina americana
Argia moesta
Argia translate
Panola Co.
Argia apicalis
Argia bipunctulata


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


Panola Co. Continued
Enallagma geminatum
Ischnura ramburii
Anomalagrion hastatum
Parker Co. (no records)
Parmer Co. (no records)
Pecos Co.
Hetaerina americana
Argia moesta
Argia sedula
Enallagma civil
Ischnura barberi
Polk Co.
Iletaerina titia
Argia immunda
Argia moesta
Argia sedula
Argia tibialis
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Potter Co. (no records)
Presidio Co.
Archilestes grandis
Hetaerina americana
Hetaerina titia
Argia fu m ipen n is
Argia lugens
Argia moesta
Argia nahuana
Argia plan
Argia sedula
Enallagma civil
Enallagma praevarum
Ischnura demorsa
Ischnura denticollis
Ilesperagrion heterodoxum
Raines Co.
Enallagma cicile
Randall Co.
Hetaerina americana
Argia moesta
Reagan Co. (no records)
Real Co.
Hetaerina americana,
Red River Co. (no records)
Reeves Co.
Lestes alacer
Hetaerina americana


Argiafumipennis
Argia immunda
Argia lugens
Argia moesta
Argia nahuana
Argia sedula
Enallagma basidens
Enallagma civil
Enallagma praevarum
Ischnura demorsa
Anomalagrion hastatum
Telehasis salva
Refugio Co.
Ischnura ramburii
Roberts Co. (no records)
Robertson Co.
Calopteryx maculata
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia fumipennis
Argia immunda
Argia nahuana
Argia sedula
Enallagma basidens
Enallagma civil
Enallagma exsulans
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Rockwall Co. (no records)
Runnels Co.
Enallagma civil
Rusk Co.
Calopteryx maculata
Argia apicalis
Argia fumipennis
Argia immunda
Argia sedula
Argia tibialis
Enallagma exsulans
Enallagma traviatum
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Sabine Co. (no records)
San Augustine Co. (no records)


125









BULLETIN FLORIDA STATE MUSEUM


San Jacinto Co.
Lestes disjunctus
Calopteryx dimidiata
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia bipunctulata
Argiafumipennis
Argia moesta
Argia sedula
Argia tibialis
Enallagma basidens
Enallagma civil
Enallagma divagans
Enallagma dubium
Enallagma exsulans
Enallagme geminatum
Enallagma signatum
Enallagma vesperum
Ischnura kellicotti
Ischnura posita
Ischnura prognatha
Ishnura ramburii
Anomalagrion hastatum
Nehalennia integricollis

San Patricio Co.
Lestes alacer
Lestes disjunctus
Lestesforficula
Lestes sigma
Hetaerina americana
Hetaerina titia
Neoneura aaroni
Argia apicalis
Argia moesta
Argia sedula
Argia translate
Enallagma civil
Enallagma durum
Enallagma signatum
Ischnura ramburii
Anomalagrion hastatum

San Saba Co.
Argia immunda
Argia nahuana
Argia plan
Schleicher Co. (no records)
Scurry Co. (no records)


Shackelford Co.
Argia apicalis
Shelby Co.
Calopteryx maculata
Sherman Co. (no records)
Smith Co. (no records)
Somervell Co. (no records)
Starr Co.
Lestesforficula
Lestes sigma
Argia moesta
Argia sedula
Enallagma civil
Stephens Co. (no records)
Sterling Co. (no records)
Stonewall Co. (no records)
Sutton Co.
Hetaerina americana
Argia sedula
Enallagma praevarum
Ischnura posita
Swisher Co. (no records)
Tarrant Co.
Lestes disjunctus
Argia apicalis
Taylor Co. (no records)
Terrell Co. (no records)
Terry Co. (no records)
Throckmorton Co. (no records)
Titus Co. (no records)
Tom Green Co.
Argia immunda
Argia moesta
Argia nahuana
Argia sedula
Enallagma exsulans
Ischnura ramburii
Travis Co.
Archilestes grandis
Lestes disjunctus
Hetaerina americana
Hetaerina titia
Argia apicalis
Argia barretti
Argia fumipennis
Argia immunda
Argia moesta
Argia nahuana
Argia sedula


Vol. XVI No. 2









JOHNSON: TEXAS DAMSELFLIES


Travis Co. Continued
Argia translate
Enallagma basidens
Enallagma civil
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Trinity Co. (no records)
Tyler Co.
Argia tibialis
Upshur Co.
Ischnura posita
Anomalagrion hastatum
Upton Co. (no records)
Uvalde Co.
Archilestes grandis
Hetaerina americana
Hetaerina titia
Protoneura cara
Argia barretti
Argiafumipennis
Argia immunda
Argia moesta
Argia nahuana
Argia sedula
Argia translate
Enallagma basidens
Enallagma civil
Enallagma novaehispaniae
Enallagma praevarum
Enallagma signatum
Anomalagrion hastatum
Telebasis salva
Val Verde Co.
Hetaerina americana
Protoneura cara
Argiafumipennis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma exsulans
Enallagma novaehispaniae
Ischnura posita
Ischnura ramburii
Telebasis salva
Van Zandt Co.
Enallagma civil


Victoria Co.
Lestes sigma
Hetaerina americana
Hetaerina titia
Neoneura aaroni
Argia apicalis
Argia moesta
Argia sedula
Argia tibialis
Enallagma basidens
Enallagma civil
Enallagma signatum
Ischnura posita
Ischnura ramburii
Anomalagrion hastatum
Walker Co.
Calopteryx maculata
Argia apicalis
Argia moesta
Argia tibialis
Enallagma divagans
Enallagma signatum
Ischnura posita
Ischnurar ramburii
Anomalagrion hastatum
Waller Co. (no records)
Ward Co.
Ischnura barberi
Washington Co. (no records)
Webb Co.
Hetaerina titia
Argia moesta
Enallagma civil
Wharton Co. (no records)
Wheeler Co. (no records)
Wichita Co. (no records)
Wilbarger Co.
Enallagma civil
Willacy Co.
Enallagma civil
Williamson Co.
Lestes disjunctus
Hetaerina americana
Argia apicalis
Argiafumipennis
Argia immunda
Argia moesta
Argia nahuana
Argia sedula


1972









128 BULLETIN FLORIDA STATE MUSEUM


Williamson Co. Continued
Argia translate
Ischnura ramburii
Anomalagrion hastatum
Telebasis salva
Wilson Co.
Lestes disjunctus
Hetaerina americana
Argia apicalis
Argia immunda
Argia moesta
Argia sedula
Argia translate
Enallagma basidens
Enallagma exsulans
Enallagma novaehispaniae
Enallagma signatum
Ischnura posita
Telebasis salva
Winkler Co. (no records)
Wise Co.
Enallagma basidens
Enallagma signatum
Anomalagrion hastatum
Wood Co.
Calopteryx maculata
Argia bipunctulata
Argia immunda
Argia plan
Argia tibialis
Enallagma basidens
Enallagma civil
Enallagma vesperum
Ischnura posita
Yoakum Co. (no records)
Young Co. (no records)
Zapata Co. (no records)
Zavala Co.
Hetaerina americana
Argia moesta
Argia sedula
Enallagma basidens


Vol. XVI No. 2




S-70. 80


v, /



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