BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 15 Number 2 DIAGNOSTIC KEYS AND NOTES ON THE DAMSELFLIES (ZYGOPTERA) OF FLORIDA Clifford Johnson and Mintef J. Westfall, Jr. \/821/ UNIVERSITY OF FLORIDA Gainesville 1970 Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM are pub- lished at irregular intervals. Volumes contain about 300 pages and are not neces- sarily completed in any one calendar year. WALTER AUFFENBERG, Managing Editor OLIvER L. AusTIN, JR., Editor Consultants for this issue: LEWIS BERNER HARRY K. CLENCH Communications concerning purchase or exchange of the publication and all manuscripts should be addressed to the Managing Editor of the Bulletin, Florida State Museum, Seagle Building, Gainesville, Florida 32601. Published 10 April 1970 Price for this issue $.80 DIAGNOSTIC KEYS AND NOTES ON THE DAMSELFLIES ( ZYGOPTERA) OF FLORIDA CLIFFORD JOHNSON AND MINTER J. WESTFALL, JR.1 SYNOPSIS: This study presents a current species list and identification guide to the -45 species of damselflies (.Zygoptera) occurring in Florida, a guide to mor- phological terms, and a short text improving accuracy of determinations. Illus- trated characters of each species and. sex accompany the keys. Color patterns, behavior traits, and habitat preferences serving as identifying characters in the field support the keys. The text provides general distribution within the state for eacb species, and references to larval descriptions. Attention is directed to problems in damselfly ecology. TABLE. OF CONTENTS INTRODUCTION' 45 . COENAGRIONIDAE 59 ACKNOWLEDGMENTS - - 46 ATgia - 63 METHODS AND MATERIALS 46 Enallagma 70 KEY To THE FAMILIES . _ 51 Ischnura 79 LESTIDAE, Lestes 51 SMALLER GENERA 83 CALOPTERYGIDAE 54 Calopteryx 56 DIscussION 85 Hetaerina 57 LITERATURE CITED 87 INTRODUCTION This report presents a current list and identification guide to adult damselHies in Florida. The 45 species identified in the state are listed in Table 1 with date of original description and authority. Literature citations for species de5criptions and synonyms have not been in- cluded, as taxonomic history is outside the paper's objective. Table 1 shows that more than 73 per cent of the state's 45 species were known prior to 1900 ( Lestes disiunctus Selys originally recognized in. 1862) and only two new species and two subspecies have been described since 1930. This decline in appearance of new taxa reflects a stabiliz- ing taxonomy rather than inactivity of systematists. 1The authors are associate professor and professor respectively in the Department of ·Zoology, University of Florida, Gainesville. Manuscript received 12 November 1969. J6hnson, Clifford and Minter J. Westfall, Jr. 1970. Diagnostic keys and notes on the damselflies ( Zygoptera) of Florida. Bull. F16rida State Mus., Vol. 15, No. 2, pp 45-89. 46 BULLETIN FLORIDA STATE MUSEUM Vol. 15 Needham and Heywood ( 1929), long out of print, contains taxo- nomic keys for North American species, and Byers ( 1930) specifically keys the Florida Odonata. The subsequent 40 years have significantly increased our knowledge of the fauna, and the above keys often prove unreliable for diagnosis. ACKNOWLEDGMENTS We wish to recognize the assistance of all individuals who have contributed to collections studied. Dennis R. Paulson's extensive field work in south Florida has added immeasurably to our southern distribution records. The assistance of the University of Florida students who have tested the keys deserves special recognition for their efforts have contributed measurably to the ' keys' success. The assistance of Paul Laessle in preparing the figures for publication is kindly acknowledged. METHODS -AND MATERIALS Materials used for this guide include Odonata in the Florida State Collection of Arthropods and other collections at the University of Florida in addition to many published and unpublished field observa- tions. Preceeding each genus and species key is a brief statement on geographical range, characteristics, and habitat preferences. An ex- panded explanation is given for difBcult key characters where exper- ience has proved necessary. The keys have been developed using students unfamilar with Odonata and progressively modified until such students, without the aid of a determined collection for comparison, repeatedly reached correct identifications. Success with the keys re- quires proper orientation of dorsal and latefal views of· the structures in question. Inattention to this requirement was found to be the largest single cause for incorrect diagnosis; consequently the required orientation is given through6ut the keys. Abdominal appendages in males occasionally dry or accidentally become lodged in an atypical position. This condition is usually obvious, but may lead to difficulty when not recognized. Some variati6n from the figures must be ex- pected. Where such variability has affected diagnosis, the keys have been expanded and several lengthy key couplets are required. Males of all species are identified by the following characters: The genital fossa accommodating the penis lies conspicuously in the sternum of abdominal segment 2 and the anterior part of segment 3; in addition, the abdominal appendages, a superior and inferior pair, occur just posterior to segment 10. These appendages are illustrated for most species. Females lack a genital fossa and possess an oviposi- tor at the terminal end of the abdomen ( Fig. 13J,K). Use of morpho- 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 47 Table 1. bIE DAMSELFLY FAUNA oF FLORIDA LESTIDAE Lestes dis/unctus australis Walker 1952 Lestes tenuatus Rambur 1842 Lestes in€qualis Walsh 1862 Lestes vidua Hagen 1861 Lestes rectangularis Say 1839 Lestes vigilax Hagen 1862 CALOPTERYCIDAE Calopteryx dimidiata Burmeister 1839 Hetaerina titia ( Drury) 1773 Calopterux maculata ( Beauvois) 1805 Hetaerina tricolor (Burmeister) 18391 Hetaerina americana (Fabricius) 1798 COENAGRIONIDAE Anomatagrion hastatum ( Say) 1839 Enallag,na geminatum Kellicott 1895 Argia apicalis (Say) 1839 Enalhgma pallidum Root 1923 Argia bipunctulata ( Hagen) 1861 Enallagma poltutum (Hagen) 1861 Argia fumipennis atra, Gloyd 1'968 Enallagma.signatum ( Hagen) 1861 Argia f. fumipennis ( Burmeister) 1839 Enallagma sulcatum Williamson 1922 Argia .moesta ( Hagen) 1861 Enallagma traviatum Selys 1876 Argia sedula ( Hagen) 1861 Enallagma vespertini Calvert 1919 Argia tibialis C Rambur) 1842 Endllagma weewa Byers 1927 Argiatiagnia pallidulum C Calvert) 1913 Ischnura kenicotti Williamson 1898 ' Enallagma basidens Calvert 1902 Ischnura posita ( Hagen) 1861 Enallagma cardenium Selys 1876 Ischnura prognatha (Hagen) 1861 Enallag,im citile C Hagen) 1861 Ischnura famburi (Selys) 1850 Enallagma concisum Williamson 1922 Nehatennia gracilis Morse 1895 Enallagma davisi Westfall 1943 Nehalennia integricollis Calvert 1913 Enallagma divagans Selys 1876 Neoerythromma culteltatum ( Hagen) Enallagma doubtedayi C Selys) 1850 1876 Enallagma dubium Root 1924 Teleallagma daecki ( Calvert) 1903 Enaltagma dumm C Hagen) 1861 Telebasis byersi Westfall 1957 1Status questionable. See text. logical terms has been kept 'to, a minimum, but familiarity with nu- merous structures and Veins is required. The line drawings are of structural traits and do not include hairs or color patterns. Ex- perience has shown that these illustrations properly orient the user without a lengthy description of odonate morphology. Wing venation is shown in Figure 1 and identified as follows: Longitudinal veins are the costal, C; cubital veins land 2, Cui and Cus; combined 1st and 2nd branches of the median vein, Mi_2; four medial branches, Mi to M*; radial vein, R; and the radial sector, Rs, Specialized 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 48 'BULLETIN FLORIDA STATE MUSEUM Vol. 15 An N A Pn MS Q B. pn -Sc N -CAn MS -R *M i -"- M2 ...-M3 Ac Q M,-2 Pn C N An Sn Ms A - M3 -M4 Q - CU2 ANC -PM M:-2 FIGURE 1. Basal portion of fore wings· in: A, Calopterygidae, Hetaerina ameri- cana; B, L~tidae, Lestes vigilax; C. Coenagrionidae, Argia tibialis. Veins identified in text. Pn. Postnodal cross veins are counted from the first cross vein distal to the nodus ( Fig. 1) outward to the cross vein just proximal to the 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 49 brace vein, b, under the stigma, st, ( Fig. 7C). Specific cells are the antenodal cells, ANC; median space, MS; and quadrangle, Q. Body structures shown in Figure 2 are identified as follows: Pro- MC 62 MPL HPL ~P H LS 2. Abd.Seg. 6 Pc G. n t. -(l L, rli CE 1 -1,\ 9- pe - PSLo FIGURE 2. A, lateral view of thorax ( prothorax disarticulated); B, head; C, dorsal view of head. Structures identified in text. thorax, Tl; mesothorax, T2; metathorax, Ta; pterothorax, T2+Ta; middle and hind prothoracic lobes, MPL and HPL; mesostigmal plates, MP; median carina, MC; humeral suture, H; metapleural suture, LS; ab- dominal segments 1 and 2, abd. seg. 1 and 2; compound eyes, CE; 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 do not include appendages. Greatest width across the compound eyes refers to a line drawn over the width of the head connecting points x' and x'' dhown in Figure 2. 50 BULLETIN FLORIDA STATE MUSEUM Vol. 15 Color patterns of the b-ody consist of alternating dark and pale stripes on the pterothorax. The middle abdominal segments may be predominantly pale or dark with narrow pale or dark transverse rings or bands located over the base or apex of each segment. The termi- nal abdominal segments often havemore extensive pale dorsal sur- faces. The thoracic pattern typically consists of a dark middorsal stripe bordered laterally on either side by a pale antehumeral and dark humeral stripe in that order. The basic pattern is shown by Argia sedula in Figure lOG. An additional dark stripe may occur on the metapleural suture. The head pattern typically consists 6f a pair of pale postocular spots, PS, often a pale postoccipital bar, PB, and a variable facial pattern ( Fig. 2). Pale areas occur in a wide range of colors· while the dark stripes, rings, or bahds are usually brown, black or metallic bronze. Pale areas are largely reduced in some species and the dark pattern then consists of metallic greens, blues, or bronze. Much variation exists on these basic patterns and additional ext)lana- tion, where needed, is provided with the individual keys. See Walker ( 1953) for an introduction to bdonate morphology. Species and sex can usually be recognizdd for adults in the field through a combination of color characters and behavior; consequently a guide to the more useful field traits has been given following the respective key. Use of such characters naturally requires a prior study of defermined museutn .specimens, but specific recognition in the habitat is one of the assets damselflies offer for ecological,investiga- tion. N. W. Moore ( 1960) was correct in referring to odonates as the "bird watchers' insects." Distributional records are certainly not complete, but they pro- vide a general picture of faunal dispersion and are also summarized following the keys. Distributions occurring in north and northwest Florida and south into the peninsula are given as southward to the southernmost county record. Distribution limited to south Florida is given as south from the northernmost county record. Statewide dis- tributions are given as such, and individual counties are listed for species recorded in only one to three counties. A further observation on distribution is given in the Discussion. Description and diagnosis of larval forms are less complete; how- ever, Gloyd and Wright ( 1959) contains an excellent key for the U. S. genera ( except Neoerythromma and Argiallagma). References to larval descriptions are given following species keys. Larva is used rather than nymph in terminology of immature stages following Snod- grass ( 1954). 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 51 KEY TO THE FAMILIES la) Numetous antenodal ahd several quadrangle cross veins: wings not petiolate (Fig. lA), 'Wings pigmented with sume black, browh or red in males, black, brown or nonpigmented in females ____._. Calopterygidae b) Two antenodal and no quadrangle cross veins; wings petiolate (Fig. 1B, C); nonpigmented or with translucent brown -- 2 2 a) V6in Ma separating from Ml-2 nearer the arculus than nodus (Fig. 1B ); stigma more 'than twice as long as wide ___-____.........--_- -_-____- Lestidae b) Vein Ma separating from Ml-2 nearef the nodus than the arculus ( Fig. 1C); stigma not twice as long as wide ___....__.--»_--_- Coenagrionidae LESTIDAE One lestid genus, Lestes, occurring in both eastern and western hemispheres, is represented in Florida by six species. All Florida species have the wings colorless or slightly edged with brown, and the thoracic and abdominal dorsum in metallic green, bronze, or brown colors. Male colors are brighter than females, but they may develop a grayish pruinosity, an external exudate that obscures their patterns in old age. Males of disiunctus and vidua are structurally close and may con- stitute a source of confusion. The thoracic dorsum of females may occasionally possess both green and bronze metallic colors. Such fe- males should be scored as bronze at couplet 1 (b). The stripe pat- tern in couplet 3 of the female key is clarified by the following note. The pale antehumeral stripe of lestids is more lateral than in most damselflies and may overlie part of the humeral suture. The dark humeral stripe is therefore largely posterior to the humeral suture. KEY TO THE MALES 1 a) Inferior abdominal appendages longer than superior abdominal appendages ( Fig. 3A); thoracic dorsum with metallic green .....___»---=- in€quatis b) Inferior abdominal appendages shorter than superior abdominal appendages ( Fig. 3B-F); thoracic dorsum with or without metallic green ___-_ -2 2 a) Median border of superior abdominal appendages in dorsal view with a distinct basal and apical tobth separated by an undulating or serrated margin ( Fig. 3B-D) 3 b) Median border of superior abdominal appendages in dorsal view with dis- tinct basal tooth and lobed margin to apical end, or with flattened basal projection and apical cluster of numerous small bristle-like teeth ( Fig. 3E,F) 5 3 a) Apical tooth of superior abdominal appendage in dorsal view distinctly longer than basal tooth ( Fig. 3B); metapleural suture lacking black stripe or elongated spot; thoracic dorsum metallic bronze .._.._ rectanguZaris 52 BULLETIN FLORIDA STATE MUSEUM Vol. 15 ABC C<4 6l INEQUALIS RECTANGULARIS DISJUNCTUS D E F ( h t 3 VIDUA VIGILAX TENUATUS FIGURE 8. Dorsal view of left superior and inferior male abdominal appendages in Lestes species following sequence appearing in key. b) Apical tuoth of superior abdominal appendage in dorsaI view not longer than basal tooth ( Fig. 3C, D); metapleural suture with blackish, often irregular, stripe 4 4 a) Inferior abdominal appendage in dorsal view extends rearward ( only slightly in some individuals) beyond apical tooth of superior appendage ( Fig. 3C); degree of serration variable between teeth on medial border of superior abdominal appendage; labrum pale bluish __... disiunctus australia b) Inferior abdominal appendage in dorsal view extends rearward no further than apical tooth of superior appendage ( Fig. 3D); labrum dark purple ( blackish in some dried museum specimens) ...______--_.__...__.._--__ vidua 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 58 5 a) Inferior abdominal appendages long and slender, well over half length of superior appendages ( Fig. BE); distance across compound eyes 5 mm or greater; body length 45 mm or greater; legs predominantly black; thoracic dorsum with metallic green on dark ground color __._......__.. vigilax b) Inferior abdominal appendages short, one half 6r less length of superior appendages ( Fig. 3F); distance across compound eyes. less than 5 mm; body length less than 45 mm; legs mottled brown; thoraci6 dorsum with metallic green on tan ground color tenuatus KEY TO THE FEMALES 1 a) Dorsum of thorax with green markings, often metallic ........__--__- 2 b) Dorsum of thorax with bronze or brown markings, often metallic _ 4 2 a) A pair of brownish-black spots on ventrolateral surface of metathorax posterior to 3rd pair of legs; external face of tibia pale brown; stigma less than 2 mm in length tenuatus b) Ventrolateral surface of metathorax pale without dark spots; external face of tibia predominantly black; stigma 2 mm or greater in length ___._ 3 3 a) Antehumeral pale stripe reddish brown, one-third or greater the width of green area on either side of niedian carina; hind wing length 26 mm or less vigilax b) Antehumeral pale stripe yellow, linear and less than one-fourth width of green area on either side of median carina; hind wing length 27 mm or greater inequaNs 4 a) Abdominal segment 7 less than twice as long as ovipositor; abdominal length 30 mm or less disfunctus australia b) Abdominal segment 7 twice as long as ovipositor; abdominal length 30 inm or greater 5 5 a) Ventral surface of metathorax entirely pale, upper end of metapleural suture pale or with very reduced spot; external face of tibia yellow __________ rectangularis b) Ventral surface of metathorax with dark elongated spots and distinct spot on upper end of metapleural suture; external face of tibia predominantly dark brown to black ©idua All species inhabit marshy lake margins and ponds in frequently isolated colonies, fly slowly, and perch with half-spread wings on emergent vegetation. Perching with half-spread wings is character- istic of lestids; other perched damselilies usually fold the wings to- gether over the body if not flexing them back and forth in some be- havioral display. Clear petiolated wings separate lestids from calop- terygids, while large body size and spread-wing perching habit dis- tinguish the group from most coenagrionids. The long slender coena- grionid, Teleallagma daecki, has nonlestid perching habits and a pale bluish or tan body color. Body size and color pattern will allow spe- cies identification of males in the field after a study of museum speci- mens ( excepting perhaps distunctus and uidua). The same traits 54 BULLETIN FLORIDA STATE MUSEUM - Vol.. 15 should prove useful for females, but require closer discrimination. Several studies of lestids have proved their adaptability to problems in population biology, see Bick and Bick ( 1961). Distribution dita are: L. disiunctus australia: sou* to Orange County. L. inequalis: Alachua County. L. rectangularis: Alachua and Clay Counties. L. tenuatus: Dade County. L. uidua: south to Highlands County. L. vigilax: south to Highlands County. Larval descriptions of L. dis/unctus australia, rectangularia, and vigilax were given by Carman ( 1917 as L. forcipatus), Needham ( 1903) and Walker ( 1914) respectively and later supplemented by Walker ( 1953). Larvae of inequalis, tenuatus, and vidua are still un- described. CALOPTERYGIDAE The large body size and conspicuous colors together with terri- torial and often complex courtship behaviors have combined to make these damselflies a favorite tool for several types of field investigation. The calopterygids are represented in Florida by two genera, both ecologically associated with streams. Calopteryx includes a group of species distributed generally in the northern hemisphere; Hetaerina consists of an array of species restricted to North and South America that reaches its highest diversity in the southern lati- tudes. Individuals are readily recognized by their broad, nonpetio- lated wings, and body colors of brown or metallic greens, blue, and bronze with pale areas largely reduced. Red, brown, or black wing pigments, separately or in combination, distinguish sex, with males showing the brighter colors. Females usually have much less or no wing pigment. KEY TO THE GENERA 1 a) No cross veins in median space; little or no pale area in body pattern, coloration metallic green or bluish; wing pigments blackish, red colors absent, stigma absent in males, present and distinctly white in females .-___ Calopteryx b) Several cross veins in median space; distinct pale and metallic dark areas or brownish body patterns; male fore wings have basal red area with or without adjacent brownish pigments, stigma in both sexes -_-_ Hetaerina 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 55 \44146-_-~.-- ---- /. 1:5»-35.«« 4 ~- --- A -",~c j'-- B FIGURE 4. Selected patterns in variation of fore and hind wings of males in Iletaerinu tilia complex . Type A characterizes Hetaerina tricolor. - ®1 • _ I-: 14 tj B t -7- 2- , I ... D FIGURE 5. Wing patterns in Calopterux dimidiata: A, male ; B and C females ; and D, wing pattern of male Hetaerina aniericatia. 56 BULLETIN FLORIDA STATE MUSEUM VoI. 15 Calopteryx KEY TO THE MALES 1 a) Wings translucint brown to opaque black for full length ( Fig. 6A); ventral surface of abdominal segments 9 and 10 whitish __-_-_-_____-_-__.- maculata b) Wings with basal five-sixths to three-fourths area clear, apical portion dis- tinctly marked with brown or black pigment ( Fig. 5A); ventral surface of abdominal segments 9 and 10 black _.--__- -_-__-_-___--_-_-__ dimidiata KEY TO THE FEMALES 1 a) Wings with brownish-black pigment over full length, often with greater intensity in apical fourth ( Fig. 6B); ventrolateral surface of pterothorax blackish. maculata -~ - A B .. C #D TITIA TRICOLOR FIGURE 6. Wing patterns in Calopteryx maculatah: A, male; B, female; C, D, dorsal views of thorneic patterns in females of the Hetaerina titia complex, b) Wings devoid of brownish-black pigment or, if present, restricted to apical fourth or less ( Fig. 5B,C); ventrolateral surface of pterothorax pale dimidiata 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 57 Field identification follows the color pattern differences given in the keys supplemented by the following notes. Female maculata have fully pigmented wings but are usually darker in the apical fourth. Teneral maculata have only smoky-gray wing colors. Female dimidiata may have wings, with little to no pigment, or the apical fourth may be pigmented similar to the male pattern. Females of both species have distinctive white stigmas. Calopteryx species present several intriguing ecological questions. C. maculata occurs along a wide variety of stream-riverine conditions and often disperses well inland. C. dimidiata apparently requires more specific conditions and often exists in isolated colonies. Dis- persal inland from the breeding sites is much more limited in dimid- iata. Where assemblages of the two species occur, niche segregation or differences.in courtship reproductively isolating them have yet to be studied, though the courtship sequence in maculata has been ana- lyzed ( Johnson, 1962). Habitat selection, courtship, and territorial behaviors have been well studied in European species (Buchholtz, 1951; Pajunen, 1966), indicating their suitability for such work. The Florida populations of muculata have been treated as a sub- species, C. m. #oridana, by Huggins ( 1927). The problem is dis- cussed by Byers ( 1930). Distribution data are: C. dimidiata: south to De Soto Co. C. maculata: south to Highlands Co. The larva of C. nmculata was described originally by Needham ( 1903) and in greater detail by Walker ( 1953). Wright ( 1946) de- scribed the larva of C. dimidiata. Hetaerina KEY TO THE MALES: 1 a) Basal red spot on fore wing bordered distally by no pigment or brownish black color extending apically for variable distance, hind wing pigment brown and varying from basal spot to entire wing ( Fig. 4A-D); pigmen- tation paler in teneral specimens ....________.-___________.____.-_ titia complex b) Basal red spot in fore and hind wings, may reach distally to nodus; apical wing areas nonpigmented ( Fig. 5D); red color limited to fore wing in teneral specimens with hind wing spot brown ____________.______-__ americana KEY TO THE FEMALES 1 a) Ventrolateral surface of abdomen brown, slightly lighter, if any, in color 58 BULLETIN FLORIDA STATE MUSEUM Vol. 15 than dorsum, dorsum of head and abdomeh dark br6wn; thorax brown with broad metallic green band on either side of median carina, or band isolated into two eloagate spots (Fig. 6C,D ) _. ____-__.__... titia complex b) Ventrolateral surface of abdomen pale-colored and distinctly contrasting 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 band on either side of median carina ____ americana Hetaerina titia as recognized in the above key may include two species, titia and tricolor. The latter species was originally described in 1839 and synonymized with titia in 1912; Johnson ( 1963) discusses the problem. The two-species interpretation .is based on distinctive behavioral differences, two thoracic color patterns in females ( Fig. 6C,D) and differences, though less distinct, in male wing patterns (Johnson, 1963). The latter studies were made with populations in central Texas where no female intergrades were found. In addition to the above criteria, flight season differences were noted, color differ- ences were found not environmentally modified, and little if any gene exchange occurred between the two forms. Subsequent study of the Florida collections ( unpublished) haye revealed infrequent female intergrades for thoracic pattern and an apparent lack of different flight seasons. Consequently these species are identmed.here under the taxon titia complex, awaiting a more comprehensive study. The only caution required in identifying hetaerinas involves the sequential development of wing patterns during the teneral stages. In titia forms, the basic patterns in pale color may be determined soon after emergence, but in americana,the hind wing spot in males is initially brown, and both fore and hind wings become deep red with age. The hind wing spot in male titia remains brown; however, the veins within the spot may be red. Female titia wings range from clear to variable amounts of brown, and female americana may exist with no wing pigment or with diffuse brown to orange basal spots. The female condition is not an age effect and the variation in americana is similar to that mentioned for Calopteryx dimidiata, possibly representing a sex-limited dimorphism as occurs in Ischnura and suggested in Enallagma. This type of variation is identified more fully below. Distribution data are: H. amen'cana: Jackson County. H. titia complex: south to DeSoto County. The larva of H. americana was described originally by Needham 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 59 ( 1903) and in greater detail by Walker ( 1953). Byers ( 1930) de- scribed the larva of titia, but it is impossible to associate the descrip- tion in the titia complex. COENAGRIONIDAE Most damselflies at any suitable habitat are coenagrionids, and the Florida fauna is no exception with nine genera represented; 34 spe- cies have been collected in the state. Enallagmas and ischnurans have essentially cosmopolitan damselfly distributions, although reach- ing their highest species diversities in North America. Anomala- grion occurs in much of South America and adjacent islands, and Nehalennia occurs in the Palearctic as well as North America. The remaining five genera are limited to smaller nearctic and neotropi- cal distributions. These species, usually with small body size, charac- teristically have brightly colored bodies in males, rather cryptically colored females, and with one notable exception ( Argia fumipennis), clear colorless wings. Anonmlagrion and Teleallagma are mono- typic genera; Argiallagma, Neoerythroinma, and Telebasis are rep- resented by single species in the state while Nehatennia, Ischnura, Argia, and Enallagma have 2,4,6 and 17 species respectively in the state. Taxonomists have largely used adult male characters for recog- nizing genera. Females have fewer and less specific 'structural traits; in addition their color patterns are frequently less distinctive, often obscured with age, or exist in dimorphic phases. Diagnostic keys for nearctic genera have consequently identified only males for Teleal- lagma, Anomalagrion, Ischnura, and Enallagma C Needham and Hey- wood, 1929; Smith and Pritchard, 1956). The following key avoids a unisexual treatment through additional couplets and alternate routes for determination. Color characters involve stripe patterns of the thorax and to a less extent the external faces of the tibiae and shapes of postocular spots. These qualities are considered with thoracic, head, and abdominal colors and their change with age. Where the above characters are not self explanatory, they are explained in the key and referred to necessary figures. The variable color patterns required of female de- termination constitutes the most likely source of confusion. The ex- ternal exudate, pruinescence, may partially obscure patterns in Anomalagrion and Ischnum. A small drop of acetone will usually 60 BULLETIN FLORIDA STATE MUSEUM Vol IS expose the underlying pattern temporarily. Pruinescence is devel- oped to greater degree in males of other genera. KEY TO THE GENERA 1 a) Majority of spines on 2nd and 3rd tibiae long, distance between spines ap- proximately one-half of spine length (Fig. 78); if thoracic dorsum metallic green, a distinct pale antehumeral stripe exists and females with a vulvar spine ( such as Fig. 13K) 2 b) Spines of tibiae short, distance between spines approximately equal to spine length ( Fig. 7A); if thoracic dorsum metallic green, it is never »-1 1-44 - ' 1 RAMBURI A /~ B TIBIALIS 4-Tz _/IA C MOESTA -fz== 3-7 -/E HASTATUM %SEDULA FIGURE 7. Variation in tibial spines and stigma structure in selected coena- grionids. Structures identified in text. Arrows indicate key charac- ters. bordered laterally by pale antehumeral area and such females never with vulvar spine ( such as Fig. 13J ) - 3 2 a) Ms separating from.Ml-„ near the 5th and 4th postnodals on the fore and hind wings respectively; postdodals 8 or less; sternum of abdominal seg- ment 8 with a vulvar spine in females; abdominal appendages of male without distinct arms in lateral view ( Fig. 18D); avg body length 25-27 mm Argiallagma pallidulum b) Mz separating from Ml-2 near the Gth and Sth postnodals or beyohd on the fore and hind wings respectively; postnodals 9 or greater; sternum of abominal segment 8 without a vulvar spine in females; male appendages and body lengths variable Argia 3 a) Thoracic dorsum solid metallic green to bronze; abdominal dorsum pre- 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 61 dominantly greenish-black; sternum of abdominal segment 8 without vulvar spine in females; avg body length 24-27 mm Nehalennia b) No such combination of characters 4 4 a) Thoracic dorsum with black dorsal stripe finely divided by pale-colored carina and distinctly notched laterally in posterior half; a small elongated dark spot midway along each humeral suture; antehumeral area and thoracic sides feddish brown in males, brown in females; abdominal dorsum red in males, brown in females; sternum of abdominal segment 8 without vulvar spine in females; inferior abdominal appendage of male ex= tends rearward beyond apical level of superi6r appendage (.Fig. 18E ); avg body length 28-31 mm Telebasis bversi b) No such combination of characters „· 5 5 a) Abdominal length 34. mm or greater; Cu:! terminating near midway point between nodus and origin of M.; petiole of wing usually extends distally to anal crossing; dorsal thoracic stripe in males narrow and humeral stripes reduced to elongated spots, stripe pattern further reduced in females; su- perior abdominal appendage of males in lateral View with distinct ventrally directed lobe (Fig. 18A ); sternum 6f abdominal segment 8 with vulvar 5pine in females -- - .-„ Teleallagma daecki b) Abdominal length less than 34 mm;. Cua. terminating n6ar origin of M: or beyond, petiole of wing usually terminated by hind margin of wing slightly proximal to anal crossing; variable color patterns, male append- ages and presence of vulvar spine in females --__.__-_-____--_......._-_-_-_- 6 6 a) Males - 7 b) Females _ _ . 11 7 a) Stigma of fore wing removed from wing margin ( Fig. 7E ); abdominal segment 10 elevated into spine, superior abdominal appendage with dis- tinct dorsoposteriorly-directed arm in lateral view ( Fig. 18B); avg body length 23-27 mm Anomatagrion hastatum b) Stigma of fore wing not removed from wing margin; abdominal segment 10 and appendages not as in Fig. 18B 8 8 a) Entire front of head between compound eyes yellowish-white distinctly bordered posteriorly at approximate level of anterior ocellus by black of vertex and rear of head; elongated postocular spots present; abdominal segment 8 and 9 without black dorsal 1narkings, segment 10 dorsally black and not elevated on rear margin; M2 usually separating from Ml-2 near the 4th and 3rd postnodals in the fore and hind wings respectively; superior abdominal appendages longer than segment 10 and with a ventrally di- rected denticle in lateral view ( Fig. 18C); unlikely beyond south Florida .. _-__-___-_-_..._-- Neoeruthromma cultenatum b) Front of head anterior to ocelli with black markings, pale colors ranging from oranges, tans, blues, greens and purples; if black absent from face, abdomi mal segment 8 and 9 with black dorsal markings; M„ separation variable; with or without elevated -rear margin of abdominal segment 10; abdominal appendages never with structure shown in Fig. 18C __...__ 9 9 a) M, separating from Mi-2 near Sth' and 4th postnodals or beyond in the fore and hind wihgs respectively; abdominal segment .10'not elevated on rear dorsal margin; stigma similar in fore a-nd hind wings _..._ Enallagma b) M. separating from MI-2 near the 4th and 3rd' postnodals in the fore and 62 BULLETIN FLORIDA STATE MUSEUM Vol. 15 hind wings respectively; abdominal segment 10 may Or may hot have ele- vation on rear dorsaI margin into spine-like process; stigma color may differ between fore and hind wings in well-matured individuals -- 10 10 a) Abdominal segment 10 elevated on rear margin into spinelike process (Fig. 186-I); or if not, inferior abdominaI appendage consists of dorsally- directed tooth and ventroposteriorly directed arm extending rearward be- yond apical level of superior appendage in lateral view ( Fig. 18F); stigma may differ in »fore and hind wings ( except I. rosita) _- Ischnura b) Abdominal segment 10 not elevated and inferior abdominal appendage never in lateral view as Fig. 18F-I; stigma similar iii fore and hind wings -- _ Enallagina 11 a) Sternum of abdominal sigment 8 without a vulvar spine ( Fig. 13J) _ 12 b) Sternum of ubdominal segment 8 with a vulvar spihe ( Fig. 13K) 13 12 a) Dorsal and humeral dark thoracic stripes bordering distinct pale antehu- - meral stripes ________.__............_ ............ Neoefuthromma cutteltatum b) Thoracic dorsum with dorsal black stripe and orange laterally ( no humeraI stripe) becoming dark brown or greenish with age and d6rsal stripe Ob- scured in older individuals, or each pale antehumeral area exists as two separate spots, the anterior one elongated, whole dorsum becoming ob- scured with age into a solid bluish-gray ( spot pattern revealed by acetone treatment ) - - - - 19 18 a) Distinct dorsal and humeral blackish thoracic stripes bordefing pale ante- humeral stripes; antehumeral colors include light taa, blues, and green __ 14 b) Distinct dorsal thoracic stripe present as solid band 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 ) -- 17 14 a) M" sepapating from Mi-2 near the 5th and 4th postnodals or beyond on the fore and hind wings respectively Enallagma b) M. separating from M·1-2 near the 4th and 3rd ( or 2nd) postnodals on the fore and hind wings respectively , - - 15 15 a) Postocular spots obscured; body length 80 mm or less; dorsum or abdominal segments 3-7 bluish-gray or black; no dark stripe on metapleural suture Anomalagrion hastatum b) Postocular spots distinct or, if obscured, associated With body length of 30 mm or greater; d6rsum of abd6minal segments 8-7 black (often with metallic green tint), brown 6r pale orange; with or without dark stripe on metapleural suture 16 16 a) Postocular spots exist as large triangles occupying majority of postocular area with apex directed anteriorly, or as small circles and associated with a blue abdominal segment 8 - Ischnura b ) Postocular spots are elongated transversely with or without connections to pale postocular bar; if spots exist as circles, abdoininal segment 8 with black markings Enaltagma 17 a) M" separating from Mi-2 near the Sth and 4th postnodals or beyond on the fore and hind wings respectively · 18 b) M2 separating from Mi-2 near the 4th and 3rd ( or End) postnodals on the fore and hind wings respectively . 19 , 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 68 18 a) Dorsum of thorax with dorsal black stripe and orange laterally becoming dark brown or greenish with age ( dorsal stripe obscured in older indi- viduals); 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 bIack stripe and pale blue or bIuish-brown laterally; distinct bluish postocular spots with well defined bbrders, often with pale postocular bar, these patterns not obscured with age Enallagma 19 a) Body length less than 30 mm; dark middorsal thoracic stripe; humeral stripe narrow, faint or absent; pale antehumeral stripe continuous; abdominal seg- ments 8, 9, and 10 bIack Anomalagrion liastatum b) Body length 80 Aim or greater, or antehumeral area dividedinto 2 spots ( re- quires close examination), or abdominal segments 8, 9, and 10 largely pale._. Ischnura Argia The following key to females requires examination of mesostigmal plates and anterior end of the median thoracic carina. These 'struc- tures occurring on the anterior dorsal edge of the mesothorax must be viewed dorsally; therefore, the head and prothorax must often be re- laxed and articulated forward to provide a clear view. The male abdominal appendages are small and their diagnostic features often require study of both dorsal and lateral aspects. KEY TO THE MALES 1 a) Dorsum of abdominal segments 3-6 pale blue with black apical rings; body length usually less than 30 mm; fore wing length usually less than 17 mm; inferior abdominal appendage in lateral view not bifid ( Fig. 8B) bipunctulata b) Dorsum of abdominal segments 3-6 violet with black apical rings, or brownish-black with pale-colored basal rings, or brownish-black without such basal rings; body length usually greater than 30 mm- fore wing langth usually greater than 17 mm; inferior abdominal appendage in lateral View bifid (Fig: BA,C,E,F ); or if not (Fig. BD), stigma surmounts 1.5 to 2 cells ( Fig. 7C) and body length always.greater than 34 mm -- 2 2 a) Stigma surmounts 1.5. to 2 cells (Fig. 7(J ); inferior abdominal appendage in lateral View not bifid (Fig. BD); distinct bluish-white pruinescence on dorsum of head and thorax in nonteneral individuals ...__-__........ moesta b) Stigma surmounts 1 cell ( rarely 1.5 cells) ( Fig. 7D); inferior abdominal appendage in lateral view bifid or notched ( Fig. 8A,C,E,F); little or no pruinescence on dorsum of head and thorax 3 3 a) Abdominal segment 8 black, 9 and 10 bluish on dorsum; ventral arm of inferior appendage in lateral view not extending posteriorly beyond dorsal arm (Fig. BF) -- tibialis 64 BULLETIN FLORIDA STATE MUSEUM Vol. 15 11 APICALIS BIPUNCTULATA -D 1 i /10 t__5 FUMIPENNIS MOESTA 7rR E - 1/ - 6 SEDULA TIBIALIS FIGURE 8. Lateral view of male abdominal appendages in Argia species. b) Dorsum of abdominal segments 8, 9, and 10 blue 6r bluish-purple; ventral arm or denticle of inferior appendage in lateral view extends posteriorly well beyond dorsal arm (:Fig. BA,E); or if not (Fig. 80) wings are dis- tinctly brown 4 4 a) In dorsal view, ventral arms of the inferior abdominal appendages with in- ward curved apical tips (Fig. 9(Da); pale antehumeral area dark blue __ sedula b) In dorsal view, ventral arms of the inferior abdominal appendages with distinct laterally divergent curvature from medial base to apical tips ( Fig. 9A,B); pale antehumeral area light blue to deep violet - 5 5 a) Superior abdominal appendage in lateral view with a distinct apical tooth directed ventrally, a second more basal tooth present but often obscured in lateral view depending on articulation of the appendage ( Fig. 8A); wings clear ( rarely clouded with white or pale brown); middorsal dark thoracic 1970 JOHNSON AND WESTFALL: FLORIDA DAMSELFLIES 65 1 1-- 1 APICALIS FUMIPENNIS SEDULA D E -1J - APICALIS ~ FUMIPENNIS ~ F G S-~73644/*.C ) C--------C+Ii)