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.