Odonaiolugica9 (I): 5-18 March I. 1980

PROCEEDINGS OF THE FIFTH INTERNATIONAL

SYMPOSIUM OF ODONATOLOGY

Montreal, August 5-11, 1979

Part I

A bibliography of reproductive behavior of

Zygoptera of Canada and conterminous United

States

G.H. Bick and J.C. Bick

1928 SW 48th Avenue, Gainesville, Florida 32608, United States

Received November 26, 1979

A bibliography of 170 references for the post 1892 literature on reproductive

is of Canadaand conterminous U.S. behavior presented for 119 spp. Zygoptera in

References and are presented by species reproductive event, includingspecies/sex

recognition, courtship, 9 refusal, intra d sperm translocation, copula,oviposition,

and those morphological aspects directly related to reproductive behavior. There

for 6 and is no information representatives of Protoneuridae,nonefor genera none

47 14 there but is for spp. For spp. is onereference. Information on oviposition

most frequent, on copula and sperm translocation less so. Seldom has a single

detailed 3 of for needs for: author all these events onesp. Outstanding are (I) data

the 47 for there data the 14 but on spp. which is none, (2) additional for with one

reference, (3) studies supported by continuous observations, timing and

descriptions of all activity from seizure until the 9 leaves the water.

INTRODUCTION

1892 literature searched for information The post was on reproduction, primarily behavorial rather than morphological, of the 119 taxa in 20 genera and four families present in Canada and conterminous United States. The

which ended Nov. 1,1979, included in which survey, only accounts species are named, not those on family or generic levels. Events considered were: species and sex recognition (Sp./sex recog.); courtship; female refusal; intra male sperm translocation (Sp.tr.), sometimes given in the literature only by supposition; copula (Cop.); oviposition (Ovi.).

three Only morphological aspects (Rep.morph.) were considered: (1) male

of female in grasp the thorax tandem, (2) male-malecontact during intramale

translocation, male-female sperm (3) genitalic contact during copula. 6 G.H. Bick & J.C. Bick

Descriptions of genitalia and of abdominal appendages primarily for taxonomic purposes and all considerations of internal anatomy were excluded.

The following data include all descriptions of reproductive events, based on apparently original information however meager or possibly erroneous,

but exclude notes stating only that mating pairs were collected or oviposition

observed without further information. Because it was impossible to check the original determinations, species are usually listed as originally given even though some determinationsmay be erroneous. A few old specific names and

generic designations are replaced by more modern opinions when the newer

nomenclature is well established.

encountered with the ofcertain it is Difficulty was use terms. Forexample, often impossible to decide if an author’s terms ’’mating” or ’’pairing” refer to copula or merely to tandem. In such doubtful cases we arbitrarily interpret

these terms as copula. Also, we use ’’courtship” when the description seems to

warrant it, even though the author did not use the term. Conversely, an author’s ’’courtship” designation is always used even though we consider it

inappropriate.

The information is presented by species and by reproductive events, the

latter abbreviated as above. Genera are given alphabetically in the family as

in the Within each references listed are species genus. reproductive event, are

alphabetically by numbers which relate to the 170 references detailed later.

It is hoped that this bibliography will make it less likely for futureworkers

to overlook items in the very scattered store of informationas we, regrettably,

sometimes have done. It should also point out those species and those aspects

of reproductive activity which need study.

SPECIES AND NUMBERED REFERENCES TO REPRODUCTIVE

ACTIVITY

PROTONEURIDAE

Neoneura

aaroni Calvert, 1903 Nil.

Protoneura

cara Calvert, 1903 Nil.

COEN AGR lON 1 DAE

Amphiagrion

abbreviatum (Selys, 1876) Ovi.: 116.

saucium (Burmeister, 1839) Nil. Bibliography of Zygoptera reproductive behavior 7

Anomalagrion

hastatum (Say, 1839) Rep.morph.: 141; Ovi.: 165.

Apanisagrion

- lais (Selys, 1876) Nil.

Argia

agrioides Calvert, 1895 — Nil.

alberta Kennedy, 1918 — Ovi.: 116.

apicalis (Say, 1839) — Sp./sex recog.: 14; Rep.morph.: 141, 161; Sp.tr.; 10,

16, 56; Cop.: 8, 10. 16; Ovi.: 7, 8, 10. 16, 44, 52, 56. 104, 166.

barretti Calvert, 1902 — Nil.

bipunctulata (Hagen, 1861) — Nil.

— emma Kennedy, 1915 Sp./sex recog.: 111; 9refusal: 75; Sp.tr.: 75; Cop.:

75; Ovi.: 75, 116, 148, 153.

extranea (Hagen, 1861) — Nil.

— fumipennis atra Gloyd, 1968 Nil.

fumipennis fumipennis (Burmeister, 1839) — Nil.

fumipennis violacea (Hagen, 1861) — Rep.morph.: 56; Sp.tr.: 56; Ovi.: 29,

56, 73, 74, 102, 104, 147.

hinei Kennedy, 1918 — Nil.

immunda(Hagen, 1861) — Rep.morph.; 56; Sp.tr.; 10, 15, 56; Cop. 56;

Ovi.; 56.

lugens (Hagen, 1861) — Ovi.: 77.

moesta (Hagen, 1861) Rep.morph.: 56, 68, 141, 161; Sp.tr.: 10, 15, 18,

158; Cop.; 15, 18,56; Ovi.; 2, 10,15, 18,26,36,51,52,56,74,78,81,142,

147, 157, 158, 164, 167.

munda Calvert, 1902 — Nil.

— nahuanaCalvert, 1902 Cop.: 56; Ovi.: 11, 56.

oenea Hagen, 1865 — Nil.

pallens Calvert, 1902 — Nil.

plana Calvert, 1902 Rep.morph.: 56; Sp.tr.: 10, 15, 18, 56; Cop.: 18,56;

Ovi.: 10, 11, 18, 56.

rhoadsi Calvert, 1902 — Nil.

sedula (Hagen. 1861) — Ovi.: 56.

tezpi Calvert, 1902 — Nil.

tibialis (Rambur, 1842) — Nil.

tonto Calvert, 1902 — Nil.

translata Hagen, 1865 — Ovi.: 36, 145, 147.

vivida Hagen, 1865 — Sp./sex recog.: Ill; Cop.: 75; Ovi.: 46, 116. Argiallagma

pallidulum Calvert, 1913 — Nil.

Chromagrion

condit — um (Hagen, 1876) Sp.tr.: 19; Cop.: 19;OvL: 19,83, 115, 121, 147. 8 G.H. Bick & J.C. Bick

Coenagrion

angulatum Walker, 1912 — Sp.tr.: 126; Cop.: 126; Ovi.: 126.

interrogatum (Hagen. 1876) — Cop.: 152; Ovi.; 121.

resolutum (Hagen, 1876) — Ovi.: 126, 147.

Enallagma

anna Williamson, 1900 — Ovi.: 106, 116, 123.

antennatum (Say, 1839) — Ovi.: 44.

— Ovi.: aspersum (Hagen, 1861) Sp.tr.: 21; Cop.: 21; 10, 21, 45, 62, 102.

basidens Calvert, 1902 — Rep.morph.; 56; Cop.: 56; Ovi.: 24, 56.

boreale — (Selys, 1875) Sp./sex recog.: Ill; Sp.tr.; 85; Cop.: 40, 82. 85;

Ovi.: 82, 83, 85, 116, 117, 121, 147.

cardenium Selys, 1876 — Nil.

carunculatum Morse, 1895 — Sp./sex recog.: Ill; Sp.tr.; 85; Cop.: 81,

85; Ovi.: 25, 81, 85, 116. 121, 147.

civile(Hagen, 1861) — Rep.morph.; 56; Sp.tr.: 10, 13. 15, 56; Cop.; 13,52,

56; Ovi.: 7, 10, 13, 24, 39, 56, 95, 167, 169, 170.

clausum Morse, 1895 — Cop.: 77; Ovi.: 32.

concisum Williamson, 1922 — Nil.

— cyathigerum (Charpentier, 1840) Sp./sex recog.: Ill; Rep.morph.; 70,

71; Sp.tr.: 70,71; Cop.: 47,72,122,127; Ovi.: 5, 35, 46, 87. 93, 94. 97,

122, 127, 128, 129, 130, 133, 149.

daeckii (Calvert, 1903) — Nil.

davisi Westfall, 1943 — Nil.

divagans Selys, 1876 — Ovi.: 73.

doubledayi Selys, 1850 — Nil.

dubium Root, 1924 Cop.; 48; Ovi.: 150.

durum (Hagen, 1861) — Nil.

ebrium (Hagen, 1861) — Rep.morph.; 56; Sp.tr.: 56,81; Cop.; 81, Ovi.: 32,

74. 92, 147.

exsulans (Hagen, 1861) — Rep,morph.: 161; Sp.tr.; 21, 158; Cop.; 21,

81; Ovi.: 10, 21, 29, 38, 147, 168.

geminatum Kellicott, 1895 Rep.morph.: 56; Ovi.: 74.

hageni (Walsh, 1863) — Ovi.: 147, 151, 167.

laterale Morse, 1895 — Nil.

minusculum Morse, 1895 — Nil.

novaehispaniae Calvert, 1907 — Nil.

pallidum Root, 1923 — Nil.

pictum Morse, 1895 — Nil.

— pollutum (Hagen, 1861) Sp./sex recog.; 134; 9 refusal: 134;

Rep.morph: 134. 162; Sp.tr.; 134; Cop.: 134; Ovi.: 134.

praevarum (Hagen, 1861) — Sp./sex recog.: Ill; Ovi.; 65.

recurvatum Davis, 1913 — Nil. Bibliography of Zygoptera reproductive behavior 9

signatum — (Hagen, 1861) Sp./sex recog.: 134; 9 refusal: 134;

Rep.morph.: 134; Sp.tr.: 134; Cop.; 134. 156; Ovi.: 24, 25, 44. 79, 134,

156, 159.

sulcatum Williamson, 1922 — Nil.

traviatum Selys, 1876 — Nil.

vernale Gloyd, 1943 — Nil.

vesperum Calvert, 1919 — Ovi.: 118.

— weewa Byers, 1927 Ovi.: 4.

Hesperagrion

heterodoxum (Selys, 1868) — Nil.

Ischnura

barberi Currie, 1903 — Nil.

cervula 1876 — Selys, Sp./sex recog.: Ill; Rep.morph.: 75. Cop.: 75;

Ovi.: 75, 116, 123.

damula Calvert, 1902 — Cop.: 66; Ovi.: 66, 67, 116.

demorsa (Hagen, 1861) — Cop,; 66; Ovi.: 66, 67.

denticollis (Burmeister, — 1839) Sp./sex recog.: Ill; Cop.: 77; Ovi.: 77,

116.

erratica Calvert, 1895 — Cop.: 31; Ovi.: 31, 32.

gemina Kennedy, 1917 — Nil.

kellicotti Williamson, 1898 Ovi.; 69.

perparva Selys, 1876 — Sp./sex recog.: Ill; Cop.: 75; Ovi.: 32, 75, 116.

posita (Hagen, 1861) — Courtship: 110; Ovi.: 6, 11, 155, 170.

prognatha (Hagen, 1861) — Nil.

ramburi (Selys, 1850) — Cop.; 41; Ovi.: 165.

verticalis — (Say, 1839) Courtship: 50; 9 refusal; 9, 10, 84; Rep.morph.: Ovi.: 56; Sp.tr.: 56; Cop.: 40, 50, 81, 84; 10, 29, 40, 44, 50, 74, 81, 84,

92, 98, 101, 147.

Nehalennia

gracilis Morse, 1895 — Ovi.: 120.

integricollis Calvert, 1913 — Nil.

irene (Hagen, 1861) — Cop.: 81; Ovi.: 58, 73, 144, 147.

Neoerythromma

cultellatum (Hagen, 1876) — Ovi.: 41, 105, 154.

Telebasis

byersi Westfall, 1957 — Nil.

salva — (Hagen, 1861) Sp./sex recog.; Ill; Ovi.: 106, 132. Zoniagrion

exclamationis (Selys, 1876) — Ovi.: 77, 132. 10 G.H. Bick & J.C. Bick

LESTIDAE

Archilestes

— californica McLachlan, 1895 Cop.: 75; Ovi,: 75, 76.

— grandis (Rambur, 1842) Sp.tr.: 15, 17; Cop.: 17. 56; Ovi.: 1, 10, 17.43,

163.

Lestes

alacer Hagen, 1861 — Ovi.; 24.

congener Hagen, 1861 — Cop.: 124; Ovi.: 4. 27, 28, 32, 51, 60. 75,80,99,

116, 119, 120, 124, 145, 147.

disjunctus Selys, 1862 — Sp.tr.: 125; Cop.: 12, 125; Ovi.: 7, 10, 12. 80,

81, 83, 116, 120, 121, 125, 147.

dryas Kirby, 1890 — Sp.tr.: 86, 149; Cop.: 86, 153; Ovi.: 32, 42, 53, 72.

86, 87, 103, 104, 108, 113, 121, 122, 125, 127, 135, 145, 147, 149.

eurinus Say, 1839 — Cop.: 83; Ovi.; 83, 89, 90, 91, 112, 120, 147.

forcipatus Rambur, 1842 — Rep.morph.: 30, 161; Cop.: 81; Ovi.; 24. 36,

80, 81, 100, 106, 147.

- forficula Rambur, 1842 Ovi.; 41.

inaequalis Walsh, 1862 Ovi.: 73.

rectangularis Say, 1839 - 9 refusal: 37, 52; Cop.: 49; Ovi.: 49, 121, 147.

sigma Calvert, 1901 — Nil.

simplex Hagen, 1861 — Nil.

stultus Hagen, 1861 — Nil.

— tenuatus Rambur, 1842 Nil.

unguiculatus Hagen, 1861 — Rep.morph.: 56, 161; Sp.tr.: 20; Cop.: 20;

Ovi.: 10, 20, 33, 34, 80, 88, 103, 104. 121, 125, 145, 147.

vidua Hagen, 1861 — Nil.

vigilax Hagen, 1862 — Cop.: 109; Ovi.: 73, 109.

CALOPTERYGIDAE

Calopteryx

aequabilis Say, 1839 — Sp./sex recog.: 136, 138; Courtship: 136, 137,

147; Cop.: 136; Ovi.: 96. 116, 136, 137, 138, 143, 147.

amata Hagen, 1889 — Courtship: 55; Ovi.: 3, 59.

angustipennis (Selys, 1853) — Sp.tr.: 158.

dimidiata Burmeister, 1839 — Nil.

maculata (P. de Beauvois, 1805) — Sp./sex recog,: 136, 138; Courtship;

64, 73, 74, 136, 137, 143, 146, 147, 160; 9 refusal: 64. 136, 137;

Rep.morph.: 140; Sp.tr.: 15, 64, 136, 137, 140, 158; Cop.; 54. 64, 121,

136, 137, 140, 147; Ovi.: 10, 21, 57, 61, 64, 74,79,81,96, 102, 104, 106,

107, 120. 121, 131, 136, 137, 138, 139, 143, 147; Sperm removal: 140. Bibliography of Zygoptera reproductive behavior II

Hetaerina

americana (Fabricius, 1798) — $ refusal: 63; Rep.morph.: 56; Sp.tr.: 10,

15, 22, 23, 56, 63; Cop.: 22, 23, 56, 63; Ovi.: 10, 22, 23, 39, 56, 63, 73.

titia (Drury, 1773) — 9 refusal: 63; Sp.tr.: 63; Cop.: 63; Ovi.: 63, 114.

vulnerata Hagen, 1853 — Nil.

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DISCUSSION

four families in Among the the region, there is no information for

of Protoneuridae; the 20 for representatives among genera, none Neoneura,

Protoneura, Apanisagrion, Argiallagma and Hesperagrion, all of which 18 G.H. Bick & J.C. Bick

barely reach the southern limits of the U.S. Of the 119 taxa, there is no information for 47. In the larger genera there is none for: five of 16 taxa of

Lestes, 14 of 26 Argia, 15 of 35 Enallagma and three of 13 Ischnura. Many of limits of U.S. is these scarcely reach the southern the and the majority habitat both. southwestern. Most are restricted geographically or by or by

But, surprisingly, Argia tibialis, widespread and often abundant in the east,

has not been studied.

which deals with For 14 species there is but a single reference nearly always

is transcontinental Argia sedula and oviposition. Notable among these the

Lestes such widespread eastern species as: inaequalis, Enallagma

antennatum, E. divagans, E. vesperum, Ischnura kellicotti, So, even though

is in there the best chance to work with strictly unstudied species the southwest,

is still opportunity in the east.

The data show that information on oviposition is most frequent, on copula

intra male translocation This is in relatively scarce and on sperm very scarce.

direct relation to the duration and ease of seeing these events. Considering

these three reproductive events, the data show that seldom has a particular

author given information for all three events for a particular species, i.e.. the

complete reproductive sequence was not observed. Hence, for many species, from different information must be gathered from different authors and

This is for ofthe populations to obtain the complete sequence. true even some

studied more frequently species.

Among the 119 taxa, information on the three reproductive events (sp.tr.,

which must occur in all odonates, has been published for only 24 cop., ovi.),

species. Even for the four most frequently studied ( Calopteryx maculata. of Argia moesta, Lestes dryas and Enallagma cyathigerum), many the studies of Palaearctic references relate to work done in Europe. Comparative

and Nearctic populations of the last two taxa should be significant.

of has led to This survey of the literature on reproduction Zygoptera us

conclude that there are three outstanding needs: (1) data for the 47 species for

of which which there is none, (2) additionaldata for the 14 species for each

with there is only a single reference, nearly always dealing only oviposition,

(3) studies supported by continuous observation, timing and description of all This is activity from male seizing the female until the latter leaves water.

particularly important in genera such as Ischnura where the female oviposits

unaccompanied and where, apparently because of the prolonged durationof

has been recorded copula, the complete reproductive sequence not by one

author for a particular species.