Checklist of Kansas Damselflies
by Roy J. Beckemeyer and Donald G. Huggins
ABOUT THIS ISSUE
Published by EMPORIA STATE UNIVERSITY
Prepared and Issued by THE DIVISION OF BIOLOGICAL SCIENCES
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Current knowledge of Kansas Odonata is based largely on the extensive work of the Kansas Biological Survey. Personnel of the Kansas Department of Wildlife and Parks, Wichita Parks Department, and the Quivira and Flint Hills National Wildlife Refuges kindly gave their permission to collect. Thanks are due to the following individuals for allowing the use of their excellent damselfly photographs: Dr. Sidney W. Dunkle of Collin Co. Community College, Plano, TX (Dr. Dunkle is preparing a field guide to the Odonata of North America), Blair Nikula of Cape Cod, MA, and Dr. Forest Mitchell, James Lasswell, and Nathan McConal of the Texas Agricultural Experiment Station, Stephenville, TX. Mr. Nikula and Dr. Mitchell both have web sites containing color images of live Odonata. Links to their web sites may be found at: http://www2.southwind.net/~royb/odonata.html. Dr. Ralph Charlton of Kansas State University quite generously shared his records for the species Lestes eurinus and Erythemis vesiculosa, both of which were new for Kansas.
Cover Photo: No. 5. Lestes disjunctus australis, Common Spreadwing. Pair in tandem, ovipositing into a grass stem at the edge of a pond. Photo by Roy Beckemeyer, Boone Co., AR, May, 1997.
Publication and distribution of this issue was made possible in part by grants from The Price and Flora Reid Foundation Trust and Central States Entomological Society.
Kansas School Naturalist is indexed in Wildlife Review/Fisheries Review; the text of this issue and of other KSNs is available at http://www.emporia.edu/ksn/.
ABOUT THE AUTHORS
Roy J. Beckemeyer (957 Perry, Wichita, KS 67203-3141) has studied the taxonomy, distribution, and biology of dragonflies and damselflies for a number of years. He is also interested in the mechanics of flight in insects and birds.
Donald G. Huggins (The University of Kansas, 2041 Constant Ave., Lawrence, KS 66047-2906) is Senior Scientist and Director of the Ecotoxicological Program of the Kansas Biological Survey. He studies the taxonomy and ecology of aquatic invertebrates, with a focus on the Odonata. Much of the knowledge of damselflies in Kansas is based on his 20 years of work in the state.
CHECKLIST OF KANSAS DAMSELFLIES
by Roy J. Beckemeyer and Donald G. Huggins
As their name implies, damselflies are delicate insects. They have a gentle, fluttering flight style, so they often fly amid vegetation rather than in the open. But, like their larger relatives the dragonflies, they are predatory insects with voracious appetites; they take a toll of midges, mosquitoes, and other insect prey as they feed.
In size the Kansas species range from tiny pond damsels, an inch or so in length, to the Great Spreadwing, almost two and a half inches long. Their colors vary widely. The spectacular Ebony Jewelwing has black, shining wings that look as if they were carved from a semi-precious stone. It has a metallic green body with an iridescent glow that brightens the shady woods it frequents. On the other hand, some of the drab female pond damsels are shades of tan and muted pastel colors. They are so inconspicuous that we have to search carefully through the sedges and grasses to find them.
While their retiring habits require more care on our part to bring them to our notice, once we find them, their habits also make them easy to study, since they carry out their life's work in confined areas. Some of the most thorough studies of damselfly behavior were done by George and Juanda Bick, who spent many summers observing pond and stream species in Oklahoma. Their articles (Ref. 6-10) are worth looking for in the library, as they document the lives of many of the damselflies commonly found in Kansas.
Damselflies and dragonflies are placed into the Order Odonata, damselflies in Suborder Zygoptera, dragonflies in Suborder Anisoptera. The dragonflies of Kansas were the subject of an earlier issue of the Kansas School Naturalist (Ref. 5). Both suborders are aquatic in their immature stages, and are usually found near the water as adults.
Damselflies lay their eggs into plant tissue. The immature damselfly, called a nymph or larva, lives in the water for one or more years, undergoing as many as 15 molts as it grows. The larva possesses a hinged and hooked lower lip which folds beneath the head and can be extended out to grasp and pull back its prey. Damselfly larvae have three leaf-like external gills at the ends of their abdomens which are also used like fins to help them swim.
Eventually the damselfly larva climbs from the water to undergo a final molt in which the adult insect emerges from the larval skin. The newly emerged insect is relatively soft, and is quite vulnerable to predation. It flies away from the water to mature, returning later to mate.
Damselflies have a variety of approaches to reproduction. Members of the family Calopterygidae are quite territorial and the males sometimes use a courtship display to woo potential mates. In the families Coenagrionidae and Lestidae there are no courtship displays, but the female usually oviposits while being held in tandem by the male. In some cases, the male guards the female as she oviposits to protect her from other males.
It is fascinating and instructive to watch damselflies at the pond or stream and to learn from observation how they go about their lives. This issue will help you to identify living adult damselflies; Ref's. 27 & 28 cover larval identification. Use these resources to help you learn more about damselflies in Kansas.
Key to the Genera of Live Damselflies (Odonata: Zygoptera) of Kansas
This key requires the damselflies to be netted and inspected in hand, although it does include some behavioral characteristics in addition to morphological ones. Its use to distinguish the Pond Damselfly genera should be used carefully outside of Kansas, as the characters used will not always yield the correct genus for some of the species and genera from outside the area. Male and female damselflies can be distinguished by the bump containing the accessory genitalia under the male second abdominal segment, and by the prominent ovipositor under the last abdominal segments on females.
|1a. Heavy-bodied; eyes closer together than one diameter (touching in most species); wings always held wide open when perched; hind wings much broader near base than fore wings; usually strong fliers; usually oviposit into water (one family oviposits into plant tissue).||Suborder ANISOPTERA, (The Dragonflies). Not covered in this publication. (81 KS species. See Ref. 5 and p. 15 of this issue.)|
|1b. Slender-bodied; eyes farther apart than one diameter; wings held together over abdomen when perched (open slightly in one family); front and hind wings similar in shape; usually weak fliers; oviposit in plant tissue.||(2) Suborder ZYGOPTERA, (The Damselflies) - 40 KS species|
|2a. Wings not petiolate; numerous antenodal and quadrangular cross veins; wings pigmented with some black, brown or red in males, black, brown or non-pigmented in females; found along streams or in riparian habitats; perch horizontally near water on twigs or vegetation; males quite territorial, guarding females during oviposition; do not oviposit in tandem.||(3) Calopterygidae (The Broadwinged Damselflies) - 3 KS species|
|3a. Wings of male entirely black, of female dark brown with a striking white stigma; no cross veins in space proximal to arculus; body metallic green in reflected light (blue when back-lit); shaded streams and wooded riparian areas; females usually oviposit with tip of abdomen under water.||Calopteryx maculata (Ebony Jewelwing)|
|3b. Wings with brown or red markings; several cross veins in space proximal to arculus; body with metallic sheen that is red in males, green in females; often in vegetation along more open areas of streams; males very territorial; females descend underwater completely to oviposit.||Hetaerina sp. (Ruby spots) - 2 KS species|
|4a. Vein M3 separates from M1-2 nearer the arculus than the nodus; stigma as long as diameter of eye; long legs with tibial spurs longer than the space between them; often perches with wings partially spread, usually clinging to vertical stems with abdomen vertical or at an oblique angle; usually oviposit in tandem into upright stems; seem to prefer marshes or grassy ponds.||(5) Lestidae (The Spreadwing Damselflies) - 5 KS species|
|4b. Vein M3 separates from M1-2 nearer the nodus than the arculus; stigma not as wide as diameter of eye; perches with wings held together over back; often from vertical stems but with body usually held horizontally.||(6) Coenagrionidae (The Pond Damselflies) - 32 KS species|
|5a. Vein M2 branches from M1 one cell distal to the nodus; diagonal yellow stripe on side of thorax.||Archilestes grandis (Great Spreadwing)|
|5b. Vein M2 branches from M1 more than two cells distal to the nodus; typically dull colors, dorsum of abdomen usually dark; rear of head and thorax and basal and terminal abdominal segments often become pruinose with age.||Lestes sp. (Spreadwings) - 4 KS species|
|6a. Spurs on the tibia at least twice as long as the distance between them; oviposit in tandem; often perch horizontally on the ground.||Argia sp. (Dancers) - 10 KS species|
|6b. Spurs on the tibia barely longer than the distance between them. Oviposit in tandem or alone; males may guard females during oviposition.||(7)|
|7a. No post-ocular spots on dorsum of head.||(8)|
|7b. With round, oval or triangular postocular spots on dorsum of head.||(9)|
|8a. Male with black thorax, brick red abdomen with black on dorsum of abdominal segments 8-10; female mostly brown to reddish-brown and with prominent vulvar spine under segment 8 just forward of the ovipositor; short (about 1 inch long) with a "stout" appearance compared to most other Pond Damsels; forehead rounded in lateral view.||Amphiagrion sp. (Red Damsel) - 1 KS species|
|8b. Male with red abdomen and red-brown thorax with black stripes; abdomen almost entirely red; female similar to male but paler; females without a vulvar spine; in lateral view there is a sharp angle between top and front surfaces of the forehead.||Telebasis salva (Desert Firetail)|
|9a. Vein M2 arises nearest the 3rd postnodal vein in the hind wing and the 4th postnodal vein in the fore wing.||(10)|
|9b.Vein M2 arises near the 4th postnodal cross vein in the hind wing and the 5th postnodal cross vein in the fore wing.||(11)|
|10. Male superior appendages not strongly directed downward; males of most species blue with black markings, but 3 species are yellow or orange with black and blue/green markings; females usually similar in color to males, but usually lighter and often tan where males are blue or green.||Enallagma sp. (Bluets) - 12 KS species|
|11a. Male blue with blue mid-dorsal carina dividing black dorsal thoracic stripe and two black and two blue stripes on sides of thorax; female similar in color but paler, often more of a tan color.||Enallagma basidens (Doublestriped Bluet)|
|11b. Black dorsal thoracic stripe and mid-dorsal carina; small to medium sized; males with a forked projection on the dorsum of the tip of the abdomen; male superior appendages strongly directed downward and inward; males of most species with black and green head and thorax, black dorsal abdomen, with yellow or blue markings (but one species yellow-orange and black); some females have form similar to male, some have more than one color form (see photos of No. 3, Ischnura verticalis).||Ischnura sp. (Forktails) - 8 KS species|
A NOTE ON THE EVOLUTION OF DRAGONFLIES AND DAMSELFLIES
AND THEIR PLACE IN THE INSECT WORLD
Over ninety-nine percent of the insect species that have been described by scientists are placed in a group (Infraclass) called the Neoptera. These insects (including flies, bees, true bugs and beetles, for example) have the ability to fold their wings down flat against their abdomens.
Damselflies and dragonflies belong to a smaller group, the Palaeoptera, which lack the added hinge which allows the Neoptera to fold their wings flat. The group Palaeoptera includes one other order of extant insects, the Ephermeroptera, or mayflies, and several orders of extinct insects known only from their fossils.
Dragonflies hold their wings out to the side when resting, somewhat like the pages of an open book. Damselflies and mayflies hold their wings together perpendicular to their thorax like little sails.
The ability of the Neoptera to fold their wings flat allowed them to radiate into many different habitats that the Palaeoptera were unable to use because of their protruding wings. This advantage may account for the fact that the Neoptera so greatly outnumber the Paleaoptera today.
Fossil Palaeoptera, including the extinct order Protodonata (which shares many of the characteristics of modern Odonata), have been found from Upper Carboniferous strata of 300 million years ago. Some of these insects had astounding wingspans of 700 mm (27 inches). Order Odonata first appeared in the Triassic period, but became much more diverse in the Jurassic (145-210 million years ago).
Because they lived near water, many specimens of Odonata have been preserved as fossils, and there is a long and rich fossil history. Interestingly, many important fossil Odonata have been taken from the Permian beds of Kansas and Oklahoma.
In addition to the suborders Anisoptera and Zygoptera, many entomologists recognize a third suborder, the Anisozygoptera. This was a very diverse suborder of Odonata during the Jurassic. While it contains many extinct species, there are only two living species in the suborder, one from Japan and a second from the Himalayas. These strange insects have bodies like dragonflies, with wings that have many characteristics in common with damselflies.
Reference 11 contains information and pictures of fossil insects and is a good starting place for those who want to learn more about the fossil record and the evolution and diversity of damselflies and dragonflies.
No. 1. CALOPTERYGIDAE: Calopteryx maculata, Ebony Jewelwing (Male).
No. 2 CALOPTERYGIDAE: Hetaerina americana, American Rubyspot (Pair in tendem).
No. 4. LESTIDAE. Archilestes grandis, Great Spreadwing (Male).
No. 7. LESTIDAE: Lestes rectangularis, Slender Spreadwing (Male).
No. 11. COENAGRIONIDAE: Argia Apicalis, Blue-fronted Dancer (Male).
No. 15. COENAGRIONIDAE: Argia nahuana, Aztec Dancer (Pair in copula).
No. 16. COENAGRIONIDAE: Argia plana, Springwater Dancer (Female).
No. 16. COENAGRIONIDAE: Argia plana, Springwater Dance (Male).
No. 17. COENAGRIONIDAE: Argia sedula, Blue-ringed Dancer (Male).
No. 19. COENAGRIONIDAE: Argia translata, Dusky Dancer (Pair in tandem).
No. 21. COENAGRIONIDAE: Enallagma aspersum, Azure Bluet (Pair in copula).
No. 22. COENAGRIONIDAE: Enallagma basidens, Double-striped Bluet (Pair in copula).
No. 24. COENAGRIONIDAE: Enallagma civile, Familiar Bluet (Male).
No. 25. COENAGRIONIDAE: Enallagma divagans, Turquoise Bluet (Male).
No. 26. COENAGRIONIDAE: Enallagma exsulans, Stream Bluet (Pair in copula).
No. 29. COENAGRIONIDAE: Enallagma signatum, Orange Bluet (Male).
No. 31. COENAGRIONIDAE: Enallagma vesperum, Vesper Bluet (Male).
No. 32. COENAGRIONIDAE: Ischnura barberi, Desert Forktail (Male).
No. 36. COENAGRIONIDAE: Ischnura hastat, Citrine Forktail (Male).
No. 38. COENAGRIONIDAE: Ischnura posita, Fragile Forktail (Male).
No. 39. COENAGRIONIDAE: Ischnura verticalis, Eastern Forktail (Male).
No. 39. COENAGRIONIDAE: Ischnura verticalis, Eastern Forktail (Female, blue form).
No. 39. COENAGRIONIDAE: Ischnura verticalis, Eastern Forktail (Female, orange form).
No. 40. COENAGRIONIDAE: Telebasis salva, Desert Firetail (Pair in copula).
ANNOTATED CHECKLIST OF KANSAS DAMSELFLIES
This list includes only those species for which the authors know of an extant voucher specimen. Entries are comprised of: Genus species Author, Date species was described - Common Name (Ref. 34) / Abundance & Distribution / Probable Flight Dates / Habitat / [References, bold if ref. contains pictures or sketches.] The author's name is in parentheses if the species has been assigned to a new genus since the original description. Key to abbreviations: C = common; UC = uncommon; VC = very common; R = rare; E = east; W = west; N = north; S = south; KS = Kansas.
ODONATA (ZYGOPTERA) - DAMSELFLIES - 40 SPECIES
CALOPTERYGIDAE - BROAD-WINGED DAMSELFLIES - 3 SPECIES
1. Calopteryx maculata (Beauvois), 1805 - Ebony Jewelwing / C throughout state. / May to Sept. / Shaded streams. / [12, 16, 27, 31, 33, 37, 39]
2. Hetaerina americana (Fabricius), 1798 - American Rubyspot / C throughout state. / May to Oct. / Rivers & streams. / [10, 16, 20, 27, 30, 31, 33, 37, 39]
3. Hetaerina titia (Drury), 1773 - Smoky Rubyspot / R in E KS. / July to Oct. / Rivers & streams. / [16, 20, 27, 30, 31, 33, 39]
LESTIDAE - SPREADWING DAMSELFLIES - 5 SPECIES
4. Archilestes grandis (Rambur), 1842 - Great Spreadwing / UC across state. / July to Oct. / Temporary pools and slow streams. Oviposits in woody plants overhanging the water. / [9, 27, 30, 33, 39]
5. Lestes disjunctus australis Walker, 1952 - Common Spreadwing / C but not numerous across state. / May to Sept. / Ponds with emergent vegetation. / [16, 27, 30, 31, 33, 36, 37, 39]
6. Lestes eurinus Say, 1839 - Amber-winged Spreadwing / R one record in Pottawatomie Co. / June / Vegetated ponds / [12, 16, 27, 31, 33, 37, 39]
7. Lestes rectangularis Say, 1839 - Slender Spreadwing / R / May to Sept. / Shaded to partly open waters. / [12, 16, 27, 31, 33, 37, 39]
8. Lestes unguiculatus Hagen, 1861 - Lyre-tipped Spreadwing / C across state. / May to Sept. / Semi-permanent and temporary water with emergent vegetation. / [33, 37, 39]
COENAGRIONIDAE - POND DAMSELS - 32 SPECIES
9. Amphiagrion sp. Selys, 1876 - Red Damsel / C in appropriate habitat across state. / May to Aug. / Spring seeps & springy marshes in shelter of grasses & sedges. / [27, 39]*
10. Argia alberta Kennedy, 1918 - Paiute Dancer / UC across state. / July to Aug. / Spring runs, seeps, small spring-fed streams. / [21, 33, 39]
11. Argia apicalis (Say), 1839 - Blue-fronted Dancer / VC across state. / May to Oct. / Banks of streams, rivers, and sometimes ponds & lakes. / [8, 16, 21, 27, 30, 31, 33, 37, 39]
12. Argia bipunctulata (Hagen), 1861 - Seepage Dancer / R Chautauqua Co. / July / Small spring seeps. / [16, 21, 30, 31, 33, 39]
13. Argia fumipennis violacea (Hagen), 1861 - Variable Dancer / C across state, more so in E. / May to Sept. / Shallow streams, woodland ponds, small impoundments. / [12, 21, 22, 30, 31, 37, 39]
14. Argia moesta (Hagen), 1861 - Powdered Dancer / VC across state. / May to Oct. / Medium to large and swift streams & rivers. / [16, 21, 27, 30, 31, 33, 37, 39]
15. Argia nahuana Calvert, 1902 - Aztec Dancer / UC across state. / June to Aug. / Springs, spring runs with moderate current. / [15, 30, 39]
16. Argia plana Calvert, 1902 - Springwater Dancer / UC across state. / June to Aug. / Seeps and springs. / [21, 30, 39]
17. Argia sedula (Hagen), 1861 - Blue-ringed Dancer / UC across state. / June to Sept. / Small to medium streams & rivers. / [16, 21, 27, 31, 33, 37, 39]
18. Argia tibialis (Rambur), 1842 - Blue-tipped Dancer / C in E KS. / June to Aug. / Small to medium streams & rivers. / [16, 21, 27, 30, 31, 33, 37, 39]
19. Argia translata Hagen in Selys, 1865 - Dusky Dancer / Locally C in E third of state. / July to Sept. / Clear water streams with rocky substrate. / [21, 30, 33, 37, 39]
20. Enallagma antennatum (Say), 1839 - Rainbow Bluet / C / May to Sept. / Streams. / [37, 39]
21. Enallagma aspersum (Hagen), 1861 - Azure Bluet / R Woodson & Labette Co's. / June to Aug. / Small ponds. / [12, 27, 33, 37, 39]
22. Enallagma basidens Calvert, 1902 - Double-striped Bluet / VC / May to Sept. / Ponds, still waters. / [16, 18, 19, 25, 27, 30, 31, 33, 39]
23. Enallagma carunculatum Morse, 1895 - Tule Bluet / UC, more numerous in W KS. / June to Aug. / Ponds & lakes. / [18, 19, 30, 33, 37, 39]
24. Enallagma civile (Hagen), 1861 - Familiar Bluet / VC across state. / May to Oct. / Virtually any slow water habitat. Tolerates low oxygen levels and organic pollution. / [7, 12, 16, 18, 19, 27, 30, 31, 33, 37, 39]
25. Enallagma divagans Selys, 1876 - Turquoise Bluet / UC E third of KS. / Apr. to July / Shaded areas of ponds, lakes, slow streams. / [16, 24, 27, 30, 31, 33, 39]
26. Enallagma exsulans (Hagen), 1861 - Stream Bluet / C E half of KS. / May to Sept. / Streams & rivers. / [27, 30, 33, 37, 39]
27. Enallagma geminatum Kellicott, 1895 - Skimming Bluet / R and local. / May to Sept. / Vegetated areas of ponds and lakes. / [12, 16, 30, 31, 37, 39]
28. Enallagma praevarum (Hagen), 1861 - Arroyo Bluet / R Cheyenne Co. / June to Aug. / Spring Creek, sand pit. / [18, 19, 30, 33, 39]
29. Enallagma signatum (Hagen), 1861 - Orange Bluet / C E 2/3 of KS. / May to Sept. / Lentic habitats. / [12, 16, 27, 30, 31, 33, 37, 39]
30. Enallagma traviatum westfalli Donnelly, 1964 - Slender Bluet / R Chautauqua, Labette, Montgomery CO's / Apr. to July. / Small shady streams. / [8, 9, 14, 15, 30, 31, 39]
31. Enallagma vesperum Calvert, 1919 - Vesper Bluet / UC E KS. / June to Sept. / Ponds & lakes. Crepuscular, so rarely seen. / [12, 16, 27, 30, 31, 33, 37, 39]
32. Ischnura barberi Currie, 1903 - Desert Forktail / Locally C in Pratt & Stafford CO's / June to Sept. / Marsh N of Cairo, KS & Big Salt Marsh, Quivira NWR. / [30, 33, 39]
33. Ischnura damula Calvert, 1902 - Plains Forktail / R Sherman & Cheyenne CO's, NW KS. / May to Aug. / Heavily vegetated ponds. / [33, 37, 39]
34. Ischnura demorsa (Hagen), 1861 - Mexican Forktail / UC in SW KS. / May to Aug. / Wide range of habitats including spring runs, irrigation pools, river backwaters. / [30, 33, 39]
35. Ischnura denticollis (Burmeister), 1839 - Black-fronted Forktail / UC across state. / May to Sept. / Wide range, from drainage ditches to springs and streams. / [30, 33, 39]
36. Ischnura hastata (Say), 1839 - Citrine Forktail / C E KS. / May to Oct. / Tolerates low oxygen levels & organic pollution. / [12, 16, 30, 31, 33, 37, 39]
37. Ischnura perparva Selys, 1876 - Western Forktail / R N part of KS, Brown, Cheyenne, Lincoln, Rawlins CO's / March to June. / Pond & lake outflows, spring creeks. / [33, 37, 39]
38. Ischnura posita (Hagen), 1861 - Fragile Forktail / UC E KS. / May to Sept. / Vegetated ponds, tolerant of low oxygen levels. / [12, 16, 27, 30, 31, 33, 37, 39]
39. Ischnura verticalis (Say), 1839 - Eastern Forktail / VC across state. / Apr. to Sept. / Virtually all still or slow waters. / [12, 27, 30, 33, 37, 39]
40. Telebasis salva (Hagen), 1861 - Desert Firetail / R Historic record from Sumner Co.** / Sept. / Ponds, slow streams. / [30, 33, 39]
* [Two species of Amphiagrion have been described for the USA: A. saucium (Eastern) and A. abbreviatum (Western). The late Mrs. L.K. Gloyd studied Amphiagrion for many years and at one time was planning on describing the mid-American form (AZ to the Midwestern states) as a third species. Some workers today consider Amphiagrion as a single species with gradual clinal variation from east to west. The problem remains to be closed, but in their recent master work on the Zygoptera of North America, Westfall and May (Ref. 39) state: "There is evidence that populations from the midwest, southwest into Arizona, probably represent a third species, but their status is as yet unclear. Amphiagrion not clearly assignable to either A. abbreviatum or A. saucium are known from: AZ, CO, IL, IN, IA, KS, MI, NE, NM, ND, OK, SD, WI." We have chosen to continue to list the Kansas specimens as Amphiagrion sp.]
** [A series of specimens of T. salva were collected by Eldon Kile in Sumner County, KS from Hunters Mill Pond in September of 1936. Ten specimens dated 1 Sept. 1936 and 4 dated 11 Sept. 1936 were determined by L.K. Gloyd and deposited in the Univ. of Michigan Museum of Zoology (UMMZ) collection. One male and one female specimen of the group now reside in the Florida State Collection of Arthropods, Gainesville, Florida. A second pair have been donated to the Kansas Biological Survey Research Collection in Lawrence, KS. These are the only records for this species we have found for Kansas. Thanks to M.L. May of Rutgers Univ., W.L. Mauffray of the International Odonatological Reserach Inst., and M.F. O'Brien of the UMMZ for help in locating these specimens, and for making them available to us. This species should be searched for in the southern counties of Kansas.]
1. Allison, V.C., 1921, "Some dragonflies of southeastern Kansas", Kansas Acad. Sci., 30:45-58
2. Banks, N., 1894, "On a collection of neuropteroid insects from Kansas", Ent. News, 4:178-180
3. Beckemeyer, R.J., 1995, "Some county records for Kansas and Oklahoma", Argia: The news journal of the Dragonfly Society of the Americas, 7(3):28-29
4. Beckemeyer, R.J., & R. Todd, 1996, "Additions to Kansas Odonata Records for 1996", Argia: The news journal of the Dragonfly Society of the Americas, 8(4):13-14
5. Beckemeyer, R.J., & D.G. Huggins, 1997, Checklist of Kansas Dragonflies, Kansas School Naturalist, 43(2):1-16
6. Bick, G.H. & J.C. Bick, 1961, "An adult population of Lestes disjunctus australis Walker (Odonata: Lestidae)", Southwestern Naturalist, 6(3-4):111-137
7. Bick, G.H. & J.C. Bick, 1963, "Behavior and population structure of the damselfly, Enallagma civile (Hagen) (Odonata: Coenagrionidae)", Southwestern Naturalist, 8(2):57-84
8. Bick, G.H. & J.C. Bick, 1965, "Demography and behavior of the damselfly Agria apicalis (Say), (Odonata: Coenagrionidae)", Ecology, 46(4):461-472
9. Bick, G.H. & J.C. Bick, 1970, "Oviposition in Archilestes grandis (Rambur) (Odonata: Lestidae)", Ent. News, 81-157-163
10. Bick, G.H. & D. Sulzbach, 1966, "Reproductive behaviour of the damselfly, Hetaerina Americana (Fabricius) (Odonata: Calopterygidae)", Animal Behaviour, 14(1):156-158
11. Carpenter, F.M., 1992, Treatise on Invertebrate Paleontology, Part R, Arthropoda 4, Volume 3: Superclass Hexapoda, The Geological Soc. of Am. & The Univ. of Kansas, Boulder CO and Lawrence, KS
12. Carpenter, V., 1991, Dragonflies and damselflies of Cape Cod, Cape Cod Mus. Nat. Hist. Series, No. 4, Brewster, MA
13. Cringan, M.S., 1978, "Dragonflies and damselflies of McKinney Marsh", The Emporia State University Research Studies, XXVII(3):1-28
14. Donnelly, T.W., 1964, "Enallagma westfalli, a new damselfly from eastern Texas, with remarks on the genus Teleallagma Kennedy", Proc. Ent. Soc. Wash., 66(2):103-109
15. Donnelly, T.W., 1973, "The status of Enallagma traviatum and westfalli (Odonata: Coenagrionidae)", Proc. Ent. Soc. Wash., 75(3):297-302
16. Dunkle, S.W., 1990, Damselflies of Florida, Bermuda, and the Bahamas, SCI Publ. Nat. Guide No. 3, Gainesville, FL
17. Edmonds, W.T., 1976, Collecting and preserving Kansas invertebrates, Tech. Pub. No. 14 of the State Biol. Surv. of Kansas
18. Garrison, R.W., 1979, Population dynamics and systematics of the damselfly genus Enallagma of the Western United States, PhD. Dissertation, Univ. Calif., Berkeley, CA.
19. Garrison, R.W., 1984, Revision of the genus Enallagma of the United States west of the Rocky Mountains and identification of certain larvae by discriminant analysis, Univ. Cal. Publ. Entomol. No. 105, Berkeley, CA.
20. Garrison, R.W., 1990, "A synopsis of the genus Hetaerina, with descriptions of four new species", Trans. Am. Ent. Soc., 116(1):175-279.
21. Garrison, R.W., 1990, "A synopsis of the genus Argia of the United States, with keys and descriptions of new species Argia sabina, A. leonorae, and A. pima (Odonata: Coenagrionidae)", Trans. Am. Ent. Coc., 120(4):287-368 (Excellent coverage of a very difficult genus. Highly recommended.)
22. Gloyd, L.K., 1968, "The union of Argia fummmipennis (Burmeister, 1839) with Argia violacea (Hagen, 1861) and the recognition of three subspecies (Odonata)", Occ. Pap. Mus. Zool., Univ. Mich., No. 658, 6 pp.
23. Huggins, D.G., 1978, "Additional Records of Kansas Odonata", in New Records of the Fauna and Flora of Kansas for 1977, Tech. Publ. Of the State Biol. Surv. Of Kansas, 6:1-35
24. Huggins, D.G., 1978, "Description of the nymph of Enallagma divagans Selys (Odonata: Coenagrionidae)", J. Kansas Ent. Soc., 51(1):140-143
25. Huggins, D.G., 1978, "Redescription of the nymph of Eanallagma basidens Calvert (Odonata: Coenagrionidae)", J. Kansas Ent. Soc., 51(2):222-227
26. Huggins, D.G., 1983, "New Records of Odonata", in New Records of the Fauna and Flora of Kansas for 1982, Tech. Publ. Of the State Biol. Surv. Of Kansas, 13:24-25
27. Huggins, D.G., & W.U. Brigham, 1982, "Odonata", Chapter 14 in Aquatic Insects and Oligochaetes of North and South Carolina, ed. by W.U. Brigham & A. Gnilka, Midwest Aquatic Enterprises, Mahomet, IL
28. Huggins, D.G., P.M. Liechti, & L.C. Ferrington, 1985, Guide to the freshwater invertebrates of the Midwest, 2nd Edition, Tech. Pub. No. 11 of the State Biol. Surv. Of Kansas.
29. Huggins, D.G., P.M. Liechti, and D.W. Roubik, 1976, "Species Accounts for Certain Aquatic Macroinvertebrates from Kansas (Odonata, Hemiptera, Coleoptera and Sphaeriidae)", in New Records of the Fauna and Flora of Kansas for 1975, Tech. Publ. Of the State Biol. Surv. Of Kansas, 1:13-77
30. Johnson, C., 1972, "The Damselflies (Zygoptera) of Texas", Bulletin of the Florida Sate Museum of Biological Sciences, 16(2):55-128
31. Johnson, C., & M.J. Westfall, Jr., 1970, "Diagnostic keys and notes on the damselflies (Zygoptera) of Florida", Bulletin of the Florida State Museum of Biological Sciences, 15(2):45-89
32. Kennedy, C.H. 1917, "The Dragonflies of Kansas", Univ. KS Dept. Ent. Bull. 11:129-145
33. Needham, J.G. & H.B. Heywood, 1929, A handbook of the dragonflies of North America, Charles C. Thomas, Springfield, IL
34. Paulson, D.R. & SW Dunkle, 1996, "Common names of North American dragonflies and damselflies, adopted by the Dragonfly Society of the Americas", ARGIA, 8(2): (Supplement)
35. Tucker, 1907, "Some results of desultatory collections of insects of Colorado and Kansas", Univ. Ks. SCI Bull. (Odonata), 4:78-79
36. Walker, E.M., 1952, "The Lestes disjunctus and forcipatus complex (Odonata: Lestidae)", Trans. Am. Eng. Soc., LXXVII:59-74
37. Walker, E.M., 1953, The Odonata of Canada and Alaska. Part 1, General and Part 2, The Zygoptera - damselflies, Univ. Toronto Press, Vol. 1
38. Westfall, M.J., Jr., 1984, "Odonata", in Aquatic Insects of North America, ed by Merritt & Cummins, Kendall/Hunt Publ. Co., Dubuque, Iowa, pp. 126-176. (Keys to genus).
39. Westfall, M.J., Jr., & ML May, 1996, Damselflies of North America, Scientific Publishers, Gainesville, FL (Definitive work, up-to-date nomenclature and keys to species with references to primary literature. Highly recommended.)
ADDENDUM TO THE"CHECKLIST OF KANSAS DRAGONFLIES" (REF. 5)
The 81st Anisoptera species recorded for Kansas was Erythemis vesiculosa (Fabricius) 1775 - Great Pondhawk (Family Libelludlidae), collected by Dr. Ralph Charlton of Kansas State University in Geary Co., near the Kansas River. This brings the total number of Odonata species recorded for Kansas to 121 (40 Zygoptera and 81 Anisoptera).
No. 14. COENAGRIONIDAE: Argia moesta, Powdered Dancer (Male).
No 2. CALOPTERYGIDAE: Hetaerina americana, American Rubyspot (Male).
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