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A checklist of chromosome numbers and a review of karyotype variation in Odonata of the world
expand article infoValentina G. Kuznetsova, Natalia V. Golub
‡ Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia
Open Access

Abstract

The ancient insect order Odonata is divided into three suborders: Anisoptera and Zygoptera with approximately 3000 species worldwide each, and Anisozygoptera with only four extant species in the relict family Epiophlebiidae. An updated list of Odonata species studied regarding chromosome number, sex chromosome mechanism and the occurrence of m-chromosomes (= microchromosomes) is given. Karyotypes of 607 species (198 genera, 23 families), covering approximately 10% of described species, are reported: 423 species (125 genera, 8 families) of the Anisoptera, 184 species (72 genera, 14 families) of the Zygoptera, and one species of the Anisozygoptera. Among the Odonata, sex determination mechanisms in males can be of X(0), XY and X1X2Y types, and diploid chromosome numbers can vary from 6 to 41, with a clear mode at 2n = 25(60%) and two more local modes at 2n = 27(21%) and 2n = 23(13%). The karyotype 2n = 25(24A + X) is found in each of the three suborders and is the most typical (modal) in many families, including the best-covered Libellulidae, Corduliidae (Anisoptera), Lestidae, Calopterygidae, and Platycnemididae (Zygoptera). This chromosome set is considered ancestral for the Odonata in general. Chromosome rearrangements, among which fusions and fissions most likely predominated, led to independent origins of similar karyotypes within different phylogenetic lineages of the order. The karyotype 2n = 27(26A + X) prevails in Aeshnidae and Coenagrionidae, whereas the karyotype 2n = 23(22A + X) is modal in Gomphidae and Chlorocyphidae, in both pairs of families one being from the Anisoptera while the other from the Zygoptera.

Keywords

Chromosome numbers, damseldragons, damselflies, dragonflies, m-chromosomes, sex chromosome mechanisms

Introduction

The order Odonata, which comprises slightly more than 6,000 described species worldwide, is one of the most ancient among winged insects (Pterygota), dating from the Permian (Grimaldi and Engel 2005). Extant Odonata include two main suborders with approximately 3,000 species each, the Zygoptera or damselflies with about 308 genera and the Anisoptera or true dragonflies with about 344 genera. Within these suborders, up to 21 and 11 families (and sometimes more), respectively, are currently recognized. The third suborder, the Anisozygoptera or damseldragons, includes only one genus Epiophlebia Calvert, 1903 with four extant species in the relict family Epiophlebiidae. A substantial body of evidence indicates that Anisoptera and Zygoptera are each monophyletic, and Zygoptera are sister to Epiophlebia plus Anisoptera (Rehn 2003; Kalkman et al. 2008; Dijkstra et al. 2013, 2014; Schorr and Paulson 2020).

The field of Odonata cytogenetics was heavily influenced by Bastiaan Kiauta, who has published dozens of papers and analyzed karyotypes of about 260 species and subspecies of this group (see References and Table 1). During the years that have passed since the publication of chromosome number checklist of Odonata (Kiauta 1972c), approximately 90 chromosome papers have been published. The number of examined species has since increased by more than 2.3 times, and now it seems appropriate to publish an updated list. In this review article, all data available today are presented in two tables and one figure. Table 1 includes all species studied so far cytogenetically and compiles data on their chromosome numbers, sex chromosome mechanisms and the occurrence of the so-called m-chromosomes (= microchromosomes). Table 2 summarizes data presented in Table 1 and shows the family-level variability of the above-mentioned traits (except m-chromosomes, since data on their presence or absence in specific species are often questionable) together with the most characteristic (modal) karyotypes for each of the families explored. On the Fig. 1, the modal karyotypes are mapped onto phylogenetic tree of Odonata families taken from Bybee et al. (2016) who in turn redrawn and synthesized it from Dijkstra et al. (2014) and Carle et al. (2015). In the final section of the review, the main characteristics of Odonata karyotypes are briefly discussed and prospects for future research are outlined.

Figure 1.

Mapping of modal karyotypes onto phylogenetic tree of Odonata families. The phylogenetic tree is taken from Bybee et al. (2016) who synthesized it based on trees from Dijkstra et al. (2014) and Carle et al. (2015). Plesiomorphic karyotype state is indicated by a black solid square (■), apomorphic karyotype states are indicated by black solid circles (●).

Concluding remarks and future directions

In total, karyotypes of 607 species (198 genera, 23 families) of Odonata are studied up to now. Table 1, presented in our work, includes 423 species (125 genera, 8 families) of the Anisoptera, 184 species (72 genera, 14 families) of the Zygoptera, and one species of the Anisozygoptera. Thus, the presently available karyotype data cover about 10% of the world species diversity of the order in general.

Table 1.

Cytogenetically analyzed species of Odonata and their main karyotype characteristics (chromosome numbers, sex chromosomes, m-chromosomes).

Taxon Karyotype formula 2n ♂ m-chromo somes Country References
Anisozygoptera
Epiophlebioidea
Epiophlebiidae
1. Epiophlebia superstes Selys, 1889 25(24A+X) Japan Oguma 1951
Anisoptera
Aeshnoidea
Aeshnidae
2. Aeshna caerulea (Ström, 1783) 24(22A+neo-XY) Finland Oksala 1943
3. A. canadiensis Walker, 1908 27(26A+X) + USA Cruden 1968
4. A. clepsydra Say, 1839 27(26A+X) + USA Hung 1971
5. A. crenata Hagen, 1856 27(26A+X) + Finland Oksala 1939a, 1943, 1944, 1952
– » – Russia Perepelov and Bugrov 2002
6. A. cyanea (Müller, 1764) 27(26A+X) + Finland Oksala 1943
– » – + Netherlands Kiauta 1969a
7. A. grandis (Linnaeus, 1758) 27(26A+X) + Former USSR Fuchsówna and Sawczyńska 1928
25(24A+X) + Former USSR Makalowskaja 1940
26(24A+neo-XY) + Finland Oksala 1939a, 1943, 1944, 1945
– » – + Netherlands Kiauta 1967ad 1968a, b, 1969a
– » – + Russia Perepelov and Bugrov 2002
25(24A+X) Finland Nokkala et al. 2002
8. A. isoceles (Müller, 1767) 27(26A+X) USA Kiauta 1978 as Anaciaeschna isosceles (Müller, 1767)
25(24A + X) + Russia Kuznetsova et al. 2020b
9. A. juncea (Linnaeus, 1758) 26(24A+neo-XY) + Finland Oksala 1939a, 1943, 1944
– » – + Former USSR Makalowskaja 1940
27(26A+X) + Italy Kiauta 1971a
26(24A+neo-XY) + Russia Perepelov and Bugrov 2002
10. A. mixta Latreille, 1805 27(26A+X) + Netherlands Kiauta 1969a
25(24A+X) + India Sandhu and Malhotra 1994a
– » – + India Sharma and Durani 1995
27(26A+X) + Russia Perepelov and Bugrov 2001b
11. A. nigroflava Martin, 1909 27(26A+X) + Japan Katatani 1987
– » – Russia Perepelov and Bugrov 2002
12. A. palmata Hagen, 1856 27(26A+X) + USA Cruden 1968
13. A. serrata Hagen, 1856 26(24A+neo-XY) + Finland Oksala 1943 as A. osiliensis Mierzejewski, 1913 and A. s. fennica Valle, 1938
14. A. subarctica Walker, 1908 27(26A+X) + USA Oksala 1939a, 1943, 1952 as A. s. elisabethae Djakonov, 1922
– » – + Switzerland Kiauta and Kiauta 1980a as A. s. elisabethae
15. A. umbrosa Walker, 1908 27(26A+X) + USA Cruden 1968 as A. u. occidentalis Walker, 1908 and A. u. umbrosa Walker, 1908
16. A. verticalis Hagen, 1861 27(26A+X) + USA Hung 1971
17. A. viridis Eversmann, 1836 26(24A+neo-XY) + Finland Oksala 1943
– » – + Russia Perepelov et al. 1998
18. A. walkeri Kennedy, 1917 27(26A+X) + USA Cruden 1968
19. Anaciaeschna jaspidea (Burmeister, 1839) 27(26A+X) + India Walia and Sandhu 1999
20. Anax amazili (Burmeister, 1839) 27(26A+X) Argentina Capitulo et al. 1991
– » – + Argentina Mola et al. 1999
21. A. concolor Brauer, 1865 27(26A+X) + Surinam Kiauta 1979a
22. A. ephippiger (Burmeister, 1839) 13(12A+X) + India Seshachar and Bagga 1962 as Hemianax ephippiger (Burmeister, 1839)
14(12A+neo-XY) + India Kiauta 1969a as H. ephippiger
23. A. guttatus (Burmeister, 1839) 15(14A+X) + Nepal Kiauta and Kiauta 1982
24. A. immaculiformis Rambur, 1842 27(26A+X) + India Sangal and Tyagi 1982
– » – + India Walia et al. 2018
25. A. imperator Leach, 1815 27(26A+X) + France Kiauta 1965, 1969a
– » – Kenya Wasschner 1985
– » – + Russia Perepelov and Bugrov 2002
26. A. junius (Drury, 1773) 27(26A+X) + USA McGill 1904, 1907
– » – + USA Lefevre and McGill 1908
– » – Japan Kichijo 1942a
– » – + USA Cruden 1968
– » –
27. A. longipes Hagen, 1861 27(26A+X) + USA Cruden 1968
28. A. nigrofasciatus Oguma, 1915 27(26A+X) + Nepal Kiauta 1974, 1975 (A. n. nigrolineatus Fraser, 1935)
25(24A+X) + India Sandhu and Malhotra 1994a (A. n. nigrolineatus)
27(26A+X) + India Walia and Sandhu 1999 (A. n. nigrolineatus)
– » – + India Walia et al. 2018 (A. n. nigrolineatus)
29. A. papuensis (Burmeister, 1839) 27(26A+X) + Australia Kiauta 1968c, 1969a as Hemianax papuensis (Burmeister, 1839)
30. A. parthenope (Selys, 1839) 27(26A+X) + Japan Omura 1957 as A. parthenope julius Brauer, 1865
– » – + India Thomas and Prasad 1986
– » – + China Zhu and Wu 1986 as A. p. julius
25(24A+X) + Japan Suzuki and Saitoh 1990 as A. p. julius
27(26A+X) + India Sandhu and Malhotra 1994a
31. Andaeschna unicolor (Martin, 1908) 27(26A+X) + Bolivia Cumming 1964 as Aeshna cf. unicolor Martin, 1908
32. Austroaeschna anacantha Tillyard, 1908 27(26A+X) + Australia Kiauta 1968c as Acanthaeschna anacantha (Tillyard, 1908)
33. A. multipunctata (Martin, 1901) 27(26A+X) + Australia Kiauta 1968c as Acanthaeschna multipunctata (Martin, 1901)
34. Basiaeschna janata (Say, 1939) 25(24A+X) USA Cruden 1968
35. Boyeria maclachlani (Selys, 1883) 27(26A+X) + Japan Omura 1957
36. B. vinosa (Say, 1839) 27(26A+X) USA Cruden 1968
37. Caliaeschna microstigma (Schneider, 1845) 16(14A+neo-XY) + Greece Kiauta 1972a
38. Castoraeschna castor (Brauer, 1865) 27(26A+X) + Brazil Kiauta 1972b
39. Cephalaeschna orbifrons Selys, 1883 25(24A+X) + Nepal Kiauta 1975
40. Cephalaeschna sp. 25(24A+X) + India Sandhu and Malhotra 1994a
41. Coryphaeschna adnexa (Hagen, 1961) 27(26A+X) Bolivia Cumming 1964
42. C. perrensi (McLachlan, 1887) 25(24A+X) Argentina Capitulo et al. 1991
27(26A+X) + Argentina Mola et al. 1999
– » – + Argentina De Gennaro et al. 2008
43. C. viriditas Calvert, 1952 23(22A+X) + Surinam Kiauta 1979a
44. Gynacantha bayadera Selys, 1891 25(24A+X) + India Walia 2007 as G. milliardi Fraser, 1936
27(26A+X) +
45. G. hyalina Selys, 1882 28(26A+XX)* + India Tyagi 1978a, b
46. G. interioris Williamson, 1923 26(24A+neo-XY) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
47. G. japonica Bartenev, 1909 27(26A+X) + Japan Omura 1957
48. Gynacanthaeschna sikkima (Karsch, 1891) 27(26A+X) + India Walia et al. 2016
49. Oplonaeschna armata (Hagen, 1861) 27(26A+X) + Mexico Kiauta 1970a
50. Planaeschna milnei (Selys, 1883) 27(26A+X) + Japan Kiauta 1968c, 1969a
51. Remartinia luteipennis (Burmeister, 1839) 25(24A+X) + Surinam Kiauta 1979a as Coryphaeschna l. luteipennis Burmeister, 1839
27(26A+X) + Brazil Ferreira et al. 1979 as C. l. luteipennis
52. Rhionaeschna bonariensis (Rambur, 1842) 26(24A+neo-XY) + Argentina, Uruguay Mola and Papeschi 1994 as Aeschna bonariensis Rambur, 1842
52. Rhionaeschna bonariensis (Rambur, 1842) – » – + Argentina, Uruguay Mola 1995 as A. bonariensis
53. Rh. californica (Calvert, 1895) 27(26A+X) + Canada Kiauta 1973a as Aeshna californica Calvert, 1895
54. Rh. confusa (Rambur, 1842) 27(26A+X) + Argentina, Uruguay Mola and Papeschi 1994 as Aeshna confuse Rambur, 1842
– » – + Argentina, Uruguay Mola 1995 as A. confuse
55. Rh. diffinis (Rambur, 1842) 21(20A+X) + Bolivia Cumming 1964 as Aeshna d. diffinis Rambur, 1842
56. Rh. intricata (Martin, 1908) 19(18A+X) + Bolivia Cumming 1964 as Aeshna intricata Martin, 1908
57. Rh. peralta (Ris, 1918) 27(26A+X) + Bolivia Cumming 1964 as Aeshna peralta Ris, 1918
58. Rh. planaltica (Calvert, 1845) 16(14A+neo-XY) + Argentina Mola and Papeschi 1994 as Aeschna cornigera planaltica Calvert, 1952
– » – + Argentina Mola 1995 as A. c. planaltica
59. Staurophlebia reticulata (Burmeister, 1839) 27(26A+X) + Brazil Souza Bueno 1982 (S. r. reticulata (Burmeister, 1839))
Petaluroidea
Petaluridae
60. Tachopteryx thoreyi (Hagen, 1857) 19(18A+X) + USA Cumming 1964
61. Tanypteryx hageni (Selys, 1879) 17(16A+X) + USA Cruden 1968
62. T. pryeri (Selys, 1889) 17(16A+X) + Japan Kichijo 1939, 1942a
63. Uropetala carovei (White, 1846) 17(16A+X)** + New Zealand Wolfe 1953
25(24A+X) + New Zealand Jensen and Mahanty 1978
– » – + New Zealand Jensen 1980
Gomphoidea
Gomphidae
64. Anisogomphus bivittatus (Selys, 1854) 23(22A+X) + India Das 1956
– » – + India Walia and Chahal 2020
65. A. occipitalis (Selys, 1854) 23(22A+X) Nepal Kiauta 1974, 1975
66. Aphylla edentata Selys, 1869 23(22A+X) Bolivia Cumming 1964
67. A. producta Selys, 1854 23(22A+X) Bolivia Cumming 1964
68. A. theodorina (Navas, 1933) 23(22A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
69. A. williamsoni (Gloyd, 1936) 23(22A+X) + USA Kiauta and Brink 1978
70. Aphylla sp. 23(22A+X) + Argentina Mola 2007
71. Arigomphus lentulus (Needham, 1902) 23(22A+X) USA Cruden 1968 as Gomphus lentulus Needham, 1902
72. A. pallidus (Rambur, 1842) 23(22A+X) USA Cumming 1964 as Gomphus pallidus Rambur, 1842
73. A. submedianus (Williamson, 1914) 23(22A+X) USA Cruden 1968 as Gomphus submedianus Williamson, 1914
74. Asiagomphus melaenops (Selys, 1854) 23(22A+X) + Japan Toyoshima and Hirai 1953 as Gomphus melaenops Selys, 1854
– » – + Japan Hirai 1956 as G. melaenops
– » – + USA Cruden 1968 as G. melaenops
75. Burmagomphus pyramidalis Laidlaw, 1922 23(22A+X) + India Tyagi 1977
76. Davidius nanus (Selys, 1869) 23(22A+X) Japan Kichijo 1939, 1942a
77. Dromogomphus spinosus (Selys, 1854) 23(22A+X) + USA Cruden 1968
78. D. spoliatus (Hagen, 1857) 23(22A+X) + USA Cruden 1968
79. Epigomphus llama Calvert, 1903 23(22A+X) Bolivia Cumming 1964
80. Erpetogomphus designatus Hagen, 1857 23(22A+X) + USA Cumming 1964
81. E. diadophis Calvert, 1905 23(22A+X) USA Cumming 1964
82. E. ophibolus Calvert, 1905 23(22A+X) + Mexico Kiauta 1970a
83. Gomphoides sp. 23(22A+X) Bolivia Cumming 1964
84. Gomphus confraternus Selys, 1873 23(22A+X) + USA Cruden 1968
85. G. exilis Selys, 1854 23(22A+X) + USA Cruden 1968
– » – + Canada Kiauta 1969a
86. G. graslini Rambur, 1842 12(10A+neo-neo-XY) + France Kiauta 1968d, 1969a
87. G. pulchellus Selys, 1840 23(22A+X) + France Kiauta 1973b
88. G. vulgatissimus (Linnaeus, 1758) 23(22A+X) Russia Perepelov et al. 2001
89. Ictinogomphus rapax (Rambur, 1942) 23(22A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942a
– » – + India Omura 1949, 1952, 1953
– » – + India Dasgupta 1957
90. Nepogomphus modestus (Selys, 1878) 23(22A+X) India Walia et al. 2006
– » – India Walia and Chahal 2014
91. Nihonogomphus ruptus (Selys, 1858) 23(22A+X) Russia Perepelov et al. 2001
92. N. viridis Oguma, 1926 23(22A+X) + Japan Omura 1957
93. Nychogomphus duaricus (Fraser, 1924) 22(20A+neo-XY) + India Tyagi 1977
94. Octogomphus specularis (Hagen, 1859) 23(22A+X) USA Cruden 1968
95. Onychogomphus forcipatus (Linnaeus, 1758) 25(24A+X) Finland Oksala 1945
22(20A+neo-XY) Austria Kiauta 1969a
25(24A+X)
96. O. saundersii Selys, 1854 22(20A+neo-XY) + India Tyagi 1977 (O. s. duaricus Fraser, 1924)
97. Ophiogomphus bison Selys, 1873 23(22A+X) USA Cruden 1968
25(24A+X)
98. O. cecilia (Fourcroy, 1785) 24(22A+XX)* Finland Oksala 1945
23(22A+X) Russia Perepelov et al. 1998
– » – Russia Perepelov and Bugrov 2001a
99. O. colubrinus Selys, 1854 23(22A+X) USA Cruden 1968
100. O. obscurus Bartenev, 1909 23(22A+X) Russia Perepelov and Bugrov 2001b
101. O. occidentalis Hagen, 1882 23(22A+X) USA Cruden 1968
102. O. rupinsulensis (Walsh, 1862) 23(22A+X) USA Cruden 1968
103. Phanogomphus lividus (Selys, 1854) 23(22A+X) + USA Cruden 1968 as Gomphus lividus Selys, 1854
104. Ph. militaris (Hagen, 1858) 23(22A+X) USA Cruden 1968 as Gomphus militaris Hagen, 1858
105. Ph. spicatus (Selys, 1854) 23(22A+X) + USA Cruden 1968 as Gomphus spicatus Selys, 1854
106. Paragomphus lineatus (Selys, 1850) 23(22A+X) Nepal Kiauta 1974, 1975
– » – India Walia and Chahal 2014
107. P. capricornis (Förster, 1914) 23(22A+X) Thailand Kiauta and Kiauta 1983
108. Phyllocycla propinqua Belle, 1972 21(20A+X) Argentina De Gennaro 2004
109. Phyllocycla sp. 23(22A+X) Bolivia Cumming 1964
110. Phyllocycla sp. 1 23(22A+X) + Argentina Mola 2007
111. Phyllocycla sp. 2 23(22A+X) Argentina Mola 2007
112. Phyllogomphoides undulatus (Needham, 1944) 23(22A+X) + Surinam Kiauta 1979a
113. Progomphus borealis McLachlan, 1873 23(22A+X) USA Cruden 1968
114. P. intricatus (Hagen, 1857) 23(22A+X) Bolivia Cumming 1964
115. P. obscurus (Rambur, 1842) 23(22A+X) USA Cruden 1968
116. P. phyllochromus Ris, 1918 23(22A+X) + Bolivia Cumming 1964
117. Scalmogomphus bistrigatus (Hagen, 1854) 23(22A+X) Nepal Kiauta 1974, 1975 as Onychogomphus bistrigatus (Hagen, 1854)
118. Shaogomphus postocularis (Selys, 1869) 23(22A+X) + Japan Omura 1957 as Gomphus postocularis Selys, 1869
– » – Russia Perepelov et al. 2001 as Gomphus epophtalmus Selys, 1872
119. Sieboldius albardae Selys, 1886 23(22A+X) + Japan Omura 1957
120. Stylogomphus suzukii (Matsumura, 1926) 23(22A+X) + Japan Oguma 1930
– » – + Japan Kichijo 1942a
121. Stylurus flavipes (Charpentier, 1825) 23(22A+X) + Russia Perepelov and Bugrov 2001b
122. S. plagiatus (Selys, 1854) 23(22A+X) + USA Cruden 1968 as Gomphus plagiatus Selys, 1854
123. S. scudderi (Selys, 1873) 23(22A+X) USA Cruden 1968 as Gomphus scudderi Selys, 1873
124. S. townesi Gloyd, 1936 22(20A+neo-XY) USA Kiauta and Brink 1978 as Gomphus townesi Gloyd, 1936
125. Temnogomphus bivittatus (Selys, 1854) 23(22A+X) + Nepal Kiauta 1975
126. Trigomphus citimus (Needham, 1931) 21(20A+X) + Japan Toyoshima and Hirai 1953 (T. c. tabei Asahina, 1949)
– » – + Japan Hirai 1956 (T. c. tabei)
127. T. interruptus (Selys, 1854) 19(18A+X) + Japan Oguma 1930
– » – + Japan Toyoshima and Hirai 1953
– » – + Japan Hirai 1956
– » – + Japan Omura 1957
128. T. melampus (Selys, 1869) 21(20A+X) Japan Oguma 1930, 1942 as T. unifasciatus (Oguma 1926)
129. Zonophora callipus Selys, 1869 23(22A+X) + Surinam Kiauta 1979a
Libelluloidea
Macromiidae
130. Didymops transversa (Say, 1839) 25(24A+X) + USA Cruden 1968
131. Epophthalmia frontalis (Selys, 1871) 25(24A+X) + India Dasgupta 1957 (E. f. frontalis (Selys, 1871))
132. Macromia daimoji Okumura, 1949 25(24A+X) Japan Katatani 1987
133. M. amphigenia Selys, 1871 25(24A+X) Russia Perepelov and Bugrov 2001b (M. a. fraenata Martin, 1906)
134. M. magnifica (McLachlan, 1874) 25(24A+X) + USA Cruden 1968
– » –
135. M. moorei Selys, 1874 25(24A+X) + Nepal Kiauta 1977
– » – + India Walia and Chahal 2018
Corduliidae
136. Cordulia aenea (Linnaeus, 1758) 25(24A+X) Finland Oksala 1939a
– » – Former USSR Makalowskaja 1940
– » – Netherlands Kiauta 1968b, 1969a
– » – Russia Perepelov et al. 1998
– » – Bulgaria Grozeva and Marinov 2007
– » – Russia Kuznetsova et al. 2018
137. C. shurtleffi Scudder, 1866 25(24A+X) + USA Cruden 1968
– » – + Canada Kiauta 1973a
138. Dorocordulia libera (Selys, 1871) 11(10A+X) USA Cruden 1968
13(12A+X)
14(12A+neo-XY) USA Kiauta 1969a
13(12A+X)
139. Epicordulia princeps (Hagen, 1861) 25(24A+X) + USA Hung 1971
140. Epitheca bimaculata (Charpentier, 1825) 25(24A+X) Russia Perepelov 2003
– » – Russia Kuznetsova et al. 2018
141. E. canis McLachlan, 1886 25(24A+X) + USA Cruden 1968
142. E. cynosura (Say, 1839) 19(18A+X) USA Cruden 1968
21(20A+X)
143. E. petechialis (Muttkowski, 1911) 21(20A+X) USA Cumming 1964 as Tetragoneuria petechialis Muttkowski, 1911
144. E. semiaquea (Burmeister, 1839) 25(24A+X) USA Cruden 1968
145. E. spinigera (Selys, 1871) 25(24A+X) + USA Cruden 1968
27(26A+X) USA Hung 1971 as Tetragoneuria spinigera (Selys, 1871)
146. Procordulia grayi (Selys, 1871) 25(24A+X) + New Zealand Jensen 1980
147. P. smithii (White, 1846) 25(24A+X) + New Zealand Jensen 1980
148. Rialla villosa Rambur, 1842 25(24A+X) + Argentina De Gennaro 2004
149. Somatochlora alpestris (Selys, 1840) 25(24A+X) Switzerland Kiauta and Kiauta 1980a
27(26A+X) +
150. S. arctica (Zetterstedt, 1840) 25(24A+X) + Russia Perepelov 2003
151. S. borisi Marinov, 2001 20(18A+XY) Bulgaria Grozeva and Marinov 2007
152. S. flavomaculata (Van der Linden, 1825) 25(24A+X) Former USSR Makalowskaja 1940
– » – Russia Perepelov 2003
– » – + Russia Kuznetsova et al. 2020b
153. S. graeseri Selys, 1887 25(24A+X) Russia Perepelov et al. 2001
154. S. meridionalis Nielsen, 1935 25(24A+X) Slovenia Kiauta and Kiauta 1995
– » – Bulgaria Grozeva and Marinov 2007
155. S. metallica (Van der Linden, 1825) 26(24A+XX)* Finland Oksala 1945
25(24A+X) Finland Nokkala et al. 2002
– » – Finland Grozeva and Marinov 2007
– » – Russia Perepelov and Bugrov 2001b
156. S. semicircularis (Selys, 1871) 25(24A+X) USA Cruden 1968
157. S. uchidai Fürster, 1909 25(24A+X) + Japan Oguma 1915, 1930
– » – + Japan Kichijo 1942b
158. S. viridiaenea (Uhler, 1858) 25(24A+X) Japan Oguma 1915, 1930
– » – Japan Kichijo 1942b
Libellulidae
159. Acisoma panorpoides Rambur, 1842 25(24A+X) + Bangladesh, India Dasgupta 1957 (A. p. panorpoides Rambur, 1842)
– » – + Nepal Kiauta 1975 (A. p. panorpoides)
– » – + Thailand Kiauta and Kiauta 1983 (A. p. panorpoides)
– » – + India Tyagi 1982
160. Aethriamanta brevipennis (Rambur, 1842) 25(24A+X) + India Dasgupta 1957
161. Anatya guttata (Erichson, 1848) 25(24A+X) Surinam Kiauta 1979a
162. Atoconeura biordinata Karsch, 1899 21(20A+X) + Sudan Wasscher 1985
163. Brachydiplax chalybea Brauer, 1868 25(24A+X) + India Dasgupta 1957
– » – + India Taygi 1982
– » – + Thailand Kiauta and Kiauta 1983
– » – + India Prasad and Thomas 1992
164. B. farinosa Krueger, 1902 25(24A+X) + India Dasgupta 1957
– » – + India Taygi 1982
– » – Thailand Kiauta and Kiauta 1983
165. B. sobrina (Rambur, 1842) 25(24A+X) + India Ray Chaudhuri and Dasgupta 1949
– » – + India Taygi 1982
– » – + Nepal Kiauta and Kiauta 1982
166. Brachvmesia furcata (Hagen, 1861) 25(24A+X) + Surinam Kiauta 1979a
– » – + Argentina Agopian and Mola 1988
– » – Brazil Ferreira et al. 1979
– » – Brazil Souza Bueno 1982
167. B. gravida (Calvert, 1890) 25(24A+X) + USA Cruden 1968 as Cannacria gravida (Calvert, 1890)
168. B. herbida (Gundlach, 1889) 25(24A+X) + Jamaica Cumming 1964 as Cannacria herbida (Gundlach, 1889)
169. Brachythemis contaminata (Fabricius, 1793) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942b
– » – + India Dasgupta 1957
– » – + Nepal Kiauta 1975
– » – + India Tyagi 1982
– » – + Thailand Kiauta and Kiauta 1983
170. B. lacustris (Kirby, 1899) 25(24A+X) + Sudan Wasscher 1985
171. Bradinopyga cornuta Ris, 1911 25(24A+X) + Republic of South Africa Boyes et al. 1980
172. B. geminata (Rambur, 1842) 25(24A+X) + India Dasgupta 1957
– » – + India Tyagi 1982
173. Brechmorhoga mendax (Hagen, 1861) 25(24A+X) + USA Cruden 1968
– » –
174. B. nubecula (Rambur, 1842) 25(24A+X) + Bolivia Cumming 1964
175. B. pertinax (Hagen, 1861) 25(24A+X) Bolivia Cumming 1964 (B. p. peruviana Ris, 1913)
176. Cannaphila vibex (Hagen, 1861) 25(24A+X) + Bolivia Cumming 1964
177. Celithemis amanda (Hagen, 1861) 25(24A+X) + USA Kiauta and Brink 1978
178. C. elisa (Hagen, 1861) 25(24A+X) + USA Cruden 1968
179. C. fasciata Kirby, 1889 25(24A+X) + USA Cruden 1968
180. C. ornata (Rambur, 1842) 25(24A+X) + USA Kiauta and Brink 1978
181. Crocothemis erythraea (Brulle, 1832) 25(24A+X) + India Dasgupta 1957
– » – + Kenya Kiauta 1969b
– » – + Italy Kiauta 1971a
– » – + India Prasad and Thomas 1992
– » – + Republic of South Africa Boyes et al. 1980
– » – + India Tyagi 1982
182. C. sanguinolenta (Burmeister, 1839) 25(24A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
183. C. servilia (Drury, 1773) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942b
– » – + India Ray Chaudhuri and Dasgupta 1949
– » – + Nepal Kiauta 1975
– » – + Philippines Kiauta and Kiauta 1980b
– » – + Nepal Kiauta and Kiauta 1982
– » – + India Tyagi 1982
– » – + Thailand Kiauta and Kiauta 1983
– » – + Japan Katatani 1987
– » – + Japan Higashi and Kayano 1993
– » – + Japan, Taiwan Higashi et al. 2001
24(22A+neo-XY) + Japan Omura 1955 (C. s. mariannae Kiauta, 983)
– » – Japan Kiauta 1983 (C. s. mariannae)
– » – Japan Katatani 1987 (C. s. mariannae)
– » – Japan Higashi et al. 2001 (C. s. mariannae)
184. Dasythemis esmeralda Ris, 1910 25(24A+X) + Bolivia Cumming 1964
185. D. mincki (Karsch, 1890) 25(24A+X) + Brazil Souza Bueno 1982
186. D. venosa (Burmeister, 1839) 25(24A+X) + Brazil Kiauta and Boyes 1972
187. Diastatops intensa Montgomery, 1940 25(24A+X) + Bolivia Cumming 1964
188. D. obscura (Fabricius, 1775) 25(24A+X) + Bolivia Cumming 1964
189. D. pullata (Burmeister, 1839) 23(22A+X) + Surinam Kiauta 1979a
190. Diplacodes bipunctata (Brauer, 1865) 25(24A+X) + Australia Kiauta 1969b
29(28A+X) +
191. D. haematodes (Burmeister, 1839) 25(24A+X) + Australia Kiauta 1969b
23(22A+X)
192. D. lefebvrei (Rambur, 1842) 25(24A+X) + Madagascar Kiauta 1968c, 1969b
193. D. nebulosa (Fabricius, 1793) 25(24A+X) + India Dasgupta 1957
– » – + India Kiauta and Kiauta 1982
– » – + India Tyagi 1982
194. D. trivialis (Rambur, 1842) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Dasgupta 1957
– » – + Australia Kiauta 1969c
– » – + Nepal Kiauta 1975
– » – + India Tyagi 1982
195. Dythemis fugax Hagen, 1861 25(24A+X) + USA Cruden 1968
196. D. multipunctata Kirby, 1894 25(24A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
197. D. rufinefris (Burmeister, 1839) 25(24A+X) + Jamaica Cumming 1964
198. D. velox Hagen, 1861 25(24A+X) + Bolivia Cumming 1964
– » – + Peru Kiauta and Boyes 1972
199. Elasmothemis cannacrioides (Calvert, 1906) 21(20A+X) Bolivia Cumming 1964 as Dythemis cannacrioides Calvert, 1906
23(22A+X) + Surinam Kiauta 1979a as D. cannacrioides
– » – + Brazil Ferreira et al. 1979
200. E. williamsoni (Ris, 1919) 22(20A+neo-XY) Surinam Kiauta 1979a as Dythemis williamsoni (Ris, 1919)
25(24A+X)
201. Erythemis attala (Selys, 1857) 25(24A+X) Bolivia Cumming 1964
– » – + Argentina Agopian and Mola 1988
202. E. collocata (Hagen, 1861) 25(24A+X) + USA Cruden 1968
203. E. credula (Hagen, 1861) 25(24A+X) + Surinam Kiauta 1979a
204. E. haematogastra (Burmeister, 1839) 25(24A+X) Surinam Kiauta 1979a
205. E. peruviana (Rambur, 1842) 25(24A+X) Surinam Kiauta 1979a
206. E. plebeja (Burmeister, 1839) 25(24A+X) Bolivia Cumming 1964
207. E. simplicicollis (Say, 1839) 25(24A+X) + USA Cruden 1968
208. E. vesiculosa (Fabricius, 1775) 25(24A+X) Bolivia Cumming 1964 as Lepthemis vesiculosa (Fabricius, 1775)
– » – Surinam Kiauta 1979a as L. vesiculosa
– » – + Brasil Ferreira et al. 1979 as L. vesiculosa
209. Erythrodiplax anomala (Brauer, 1865) 25(24A+X) + Brazil Souza Bueno 1982
210. E. atroterminala Ris, 1911 25(24A+X) + Uruguay Goni and Abenante 1982
– » – + Argentina Mola 1996
211. E. attenuata (Kirby, 1889) 25(24A+X) + Surinam Kiauta 1979a
– » – + Brasil Ferreira et al. 1979
212. E. basalis (Kirby, 1897) 25(24A+X) Bolivia Cumming 1964
– » – + Surinam Kiauta 1979a (E. b. basalis (Kirby, 1897))
– » – + Brasil Ferreira et al. 1979 (E. b. basalis)
213. E. berenice (Drury, 1770) 25(24A+X) USA Cruden 1968
27(26A+X) + USA Hung 1971
25(24A+X) +
214. E. castanea (Burmeister, 1839) 25(24A+X) Bolivia Cumming 1964
215. E. chromoptera Borror, 1942 23(22A+X) + Uruguay Goni and Abenante 1982
216. E. cleopatra Ris, 1911 25(24A+X) + Peru Kiauta and Boyes 1972
217. E. connata (Burmeister, 1839) 25(24A+X) + Chile Kiauta and Boyes 1972 (E. c. connata (Burmeister, 1839))
– » – + USA Kiauta and Brink 1978 (E. c. minuscula (Rambur, 1842))
218. E. coralline (Brauer, 1865) 25(24A+X) + Argentina Mola 1996
219. E. famula (Erichson, 1848) 25(24A+X) + Brazil Souza Bueno 1982
220. E. fusca (Rambur, 1842) 25(24A+X) Bolivia Cumming 1964 as E. connata fusca (Rambur, 1842)
– » – Guatemala Cruden 1968 as E. c. fusca
– » – + Surinam Kiauta 1979a as E. c. fusca
– » – + Brazil Ferreira et al. 1979 as E. c. fusca
– » – + Brazil Souza Bueno 1982
– » – + Argentina Mola 1996
221. E. fervida (Erichson, 1848) 25(24A+X) + Jamaica Cumming 1964
222. E. justiniana (Selys, 1857) 25(24A+X) + Jamaica Cumming 1964
223. E. juliana Ris, 1911 25(24A+X) + Brazil Souza Bueno 1982
224. E. latimaculata Ris, 1911 25(24A+X) + Surinam Kiauta 1979a
– » – + Brasil Ferreira et al. 1979
225. E. lygaea Ris, 1911 25(24A+X) + Argentina Capitulo et al. 1991
– » – + Argentina Mola 1996
226. E. media Borror, 1942 21(20A+X) + Bolivia Cumming 1964
22(20A+XX)* + Brazil Kiauta and Boyes 1972
21(20A+X) + Surinam Kiauta 1979a
– » – + Brasil Ferreira et al. 1979
22(20A+neo-XY) + Argentina Mola 1996
227. E. melanorubra Borror, 1942 25(24A+X) + Bolivia Cumming 1964
– » – + Venezuela Kiauta and Boyes 1972
– » – + Argentina Capitulo et al. 1991
– » – + Argentina Mola 1996
228. E. minuscula (Rambur, 1842) 25(24A+X) + USA Kiauta and Brink 1978
22(20A+neo-XY) + Argentina Mola and Agopian 1985
229. E. nigricans (Rambur, 1842) 25(24A+X) + Uruguay Goni and Abenante 1982
229. E. nigricans (Rambur, 1842) – » – + Argentina Mola 1996
– » – Argentina De Gennaro 2004
– » – + Argentina De Gennaro et al. 2008
230. E. ochracea (Burmeister, 1839) 25(24A+X) + Argentina Mola 1996
231. E. paraguayensis (Foerster, 1904) 23(22A+X) + Bolivia Cumming 1964
– » – + Surinam Kiauta 1979a
232. E. umbrata (Linnaeus, 1758) 25(24A+X) + Bolivia Cumming 1964
– » – + Dominica Cruden 1968
– » – + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al 1979
– » – + Argentina Mola 1996
233. E. unimaculata (DeGeer, 1773) 25(24A+X) + Bolivia Cumming 1964
– » – + Surinam Kiauta 1979a
234. Hydrobasileus croceus (Brauer, 1867) 25(24A+X) + India Prasad and Thomas 1992
235. Ladona julia (Uhler, 1857) 25(24A+X) + USA Cruden 1968
236. Lathrecista asiatica (Fabricius, 1798) 25(24A+X) + India Dasgupta 1957
– » – + India Tyagi 1982
237. Leucorrhinia albifrons (Burmeister, 1839) 25(24A+X) + Former USSR Makalowskaja 1940
238. L. dubia (Van der Linden, 1825) 26(24A+XX)* Finland Oksala 1939a, 1945
25(24A+X) + Russia Kuznetsova et al. 2020b
239. L. frigida Hagen, 1890 21(20A+X) USA Cruden 1968
23(22A+X) +
240. L. glacialis Hagen, 1890 25(24A+X) + USA Cruden 1968
241. L. hudsonica (Selys, 1850) 25(24A+X) + USA Cruden 1968
– » –
242. L. intacta (Hagen, 1861) 25(24A+X) + USA Cruden 1968
– » –
243. L. pectoralis (Charpentier, 1825) 26(24A+XX)* Finland Oksala 1945
244. L. proxima Calvert, 1890 25(24A+X) + USA Cruden 1968
245. L. rubicunda (Linnaeus, 1857) 25(24A+X) Finland Oksala 1939a
– » – Former USSR Makalowskaja 1940
– » – Russia Kuznetsova et al. 2018
246. Libellula angelina Selys, 1883 25(24A+X) + Japan Oguma 1915, 1930
– » – + Japan Kichijo 1942a
247. L. auripennis Burmeister, 1839 25(24A+X) + USA Kiauta and Brink 1978
248. L. axilena Westwood, 1837 23(22A+X) USA Cumming 1964
249. L. basalis (Say, 1840) 25(24A+X) USA Smith 1916
250. L. composita (Hagen, 1873) 25(24A+X) + USA Cruden 1968
251. L. croceipennis Selys, 1868 25(24A+X) + USA Cruden 1968
252. L. cyanea Fabricius, 1775 25(24A+X) USA Cruden 1968
253. L. depressa Linnaeus, 1758 23(22A+X) Belgium Carnoy 1885
– » – England Hogben 1921
25(24A+X) + Austria Kiauta 1968c, 1969b
23(22A+X)
25(24A+X) + France Kiauta 1973b
– » – + Croatia Francovič and Jurečic 1986, 1989
– » – + Russia Perepelov et al. 1998
– » – + Russia Kuznetsova et al. 2018
254. L. flavida Rambur, 1842 25(24A+X) + USA Cruden 1968
255. L. forensis Hagen, 1861 25(24A+X) + USA Cruden 1968
256. L. fulva Muller, 1764 25(24A+X) + Switzerland Kiauta and Kiauta 1979
27(26A+X) + Croatia Francovič and Jurečic 1986, 1989
257. L. insecta Hagen, 1861 25(24A+X) USA Cumming 1964
– » – USA Cruden 1968
258. L. luctuosa Burmeister, 1839 25(24A+X) USA Smith 1916
259. L. pulchella Drury, 1773 25(24A+X) + USA Cruden 1968
– » – + Canada Kiauta 1969a
260. L. quadrimaculata Linnaeus, 1758 25(24A+X) + Japan Oguma 1915, 1930 (L. q. asahinai Schmidt, 1957)
260. L. quadrimaculata Linnaeus, 1758 25(24A+X) + Japan Kichijo 1942d (L. q. asahinai)
– » – + Japan Omura 1955 (L. q. asahinai)
– » – + Japan Kiauta 1968b, c (L. q. asahinai)
– » – + Former USSR Fuchsówna and Sawczyńska 1928 (L. q. quadrimaculata Linnaeus, 1758)
– » – + Finland Oksala 1939a, b, 1945 (L. q. quadrimaculata)
– » – + Former USSR Makalowskaja 1940 (L. q. quadrimaculata)
– » – + Netherlands Kiauta 1968b, c (L. q. quadrimaculata)
– » – + USA Cruden 1968 (L. q. quadrimaculata)
– » – + Russia Perepelov et al. 1998 (L. q. quadrimaculata)
– » – + Russia Kuznetsova et al. 2018 (L. q. quadrimaculata)
261. L. saturata Uhler, 1857 25(24A+X) + USA Cruden 1968
262. L. semifasciata Burmeister, 1839 25(24A+X) + USA Cruden 1968
263. L. vibrans Fabricius, 1793 25(24A+X) + USA Cruden 1968
264. Lyriothemis pachygastra (Selys, 1878) 25(24A+X) Japan Omura 1955
265. Macrothemis declivata Calvert, 1909 23(22A+X) + Brazil Kiauta and Boyes 1972
266. M. hemichlora (Burmeister, 1839) 6(4A+neo-XY) Bolivia Cumming 1964
267. M. imitans Karsch, 1890 25(24A+X) + Brazil Kiauta and Boyes 1972 (M. i. imitans Karsch, 1890)
268. M. mortoni Ris, 1913 25(24A+X) + Bolivia Cumming 1964
269. M. musiva Calvert, 1898 25(24A+X) + Bolivia Cumming 1964
270. Macrothemis sp. 25(24A+X) + Argentina Mola 2007
271. Miathyria artemis (Selys, 1857) 25(24A+X) + Surinam Kiauta 1979a
272. M. marcella (Selys, 1857) 25(24A+X) + Bolivia Cumming 1964
– » – + Surinam Kiauta 1979a
– » – + Argentina Mola and Agopian 1985
– » – + Brazil Ferreira et al. 1979
273. Micrathyria artemis Ris, 1911 25(24A+X) + Brazil Ferreira et al. 1979
– » – + Brazil Souza Bueno 1982
274. M. atra (Martin, 1897) 25(24A+X) + Bolivia Cumming 1964
275. M. catenata Calvert, 1909 25(24A+X) + Brazil Souza Bueno 1982
– » – + Argentina Mola 2007
276. M. didyma (Selys, 1857) 25(24A+X) + Jamaica Cumming 1964
277. M. exima Kirby, 1897 25(24A+X) + Surinam Kiauta 1979a
278. M. hagenii Kirby, 1890 25(24A+X) + Jamaica Cumming 1964
279. M. hesperis Ris, 1911 25(24A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
– » – + Argentina Mola et al. 1999
280. M. hypodydima Calvert 1906 23(22A+X) + Brazil Souza Bueno 1982
25(24A+X) + Argentina Agopian and Mola 1988
281. M. iheringi Santos, 1946 23(22A+X) + Bolivia Cumming 1964
282. M. laevigata Calvert, 1909 25(24A+X) + Bolivia Cumming 1964
– » – + Brazil Kiauta and Boyes 1972
283. M. longifasciata Calvert, 1909 24(22A+neo-XY) Argentina Agopian and Mola 1988
284. M. ocellata (Martin, 1897) 25(24A+X) + Bolivia Cumming 1964 (M. o. dentiens Calvert, 1909)
285. M. spuria (Selys, 1900) 25(24A+X) + Bolivia Cumming 1964
– » – + Argentina Mola et al. 1999
286. M. stawiarskii Santos, 1953 25(24A+X) + Brazil Souza Bueno 1982
287. M. ungulata Foerster, 1907 23(20A+X1X2Y) Argentina Mola et al. 1999
288. M. cf. eximia Kirby, 1879 21(20A+X) Bolivia Cumming 1964
289. M. sp. (ungulata Foerster, 1907-group) 23(22A+X) Bolivia Cumming 1964
290. Nannothemis bella (Uhler, 1857) 25(24A+X) + USA Cruden 1968
291. Nesciothemis farinosa (Foerster, 1898) 25(24A+X) + Kenya Kiauta 1969c
– » – + Kenya Wasscher 1985
292. Nesogonia blackburni (McLachlan, 1883) 25(24A+X) + Hawaii Kiauta 1969d
293. Neurothemis fulvia (Drury, 1773) 25(24A+X) + Nepal Kiauta 1974, 1975
294. N. intermedia (Rambur, 1842) 25(24A+X) + Nepal Kiauta 1974, 1975 (N. i. intermedia (Rambur, 1842))
– » – + Nepal Kiauta and Kiauta 1982 (N. i. degener (Sel, 1842))
295. N. terminata Ris, 1911 25(24A+X) + Philippines Kiauta and Kiauta 1980b
296. N. tullia (Drury, 1773) 28(26A+neo-XY) + India Ray Chaudhuri and Dasgupta 1949
– » – + India Kiauta 1969a (N. t. tullia (Drury, 1773))
– » – + India Tyagi 1982 (N. t. tullia)
25(24A+X) + Thailand Kiauta and Kiauta 1983
297. Oligoclada amphinome Ris, 1919 25(24A+X) + Surinam Kiauta 1979a
298. O. laetitia Ris, 1911 23(22A+X) + Argentina Mola and Agopian 1985
21(20A+X) Brazil Souza Bueno 1982
299. O. monosticha Borror, 1931 23(22A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
300. O. pachystigma Karsch, 1890 23(22A+X) + Brazil Souza Bueno 1982
301. Orthemis aequilibris Calvert, 1909 12(10A+neo-XY) Surinam Kiauta 1979a
302. O. ambinigra Calvert, 1909 12(10A+neo-XY) Argentina Agopian and Mola 1984
303. O. biolleyi Calvert, 1906 23(22A+X) + Bolivia Cumming 1964
304. O. cultiformis Calvert, 1906 23(22A+X) + Bolivia Cumming 1964
– » – + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
305. O. discolor Burmeister, 1839 23(22A+X) + Argentina Mola 2007
306. O. ferruginea (Fabricius, 1775) 10(8A+neo-XY)*** Bolivia Cumming 1964
23(22A+X) USA
– » – + Guatemala, Dominica Cruden 1968
– » – + Peru Kiauta 1969a, 1971c
– » – + Peru Kiauta and Boyes 1972
23(22A+X) + Surinam Kiauta 1979a
25(24A+X) +
23(22A+X) + Brazil Ferreira et al. 1979
23(22A+X) Brazil, Argentina Mola and Agopian 1985
24(22A+XX)* +
307. O. levis Calvert, 1906 6(4A+neo-XY)*** Bolivia Cumming 1964
8(6A+neo-XY)***
308. O. nodiplaga Karsch, 1891 41(40A+X) Argentina Agopian and Mola 1984
309. Orthetrum abbotti Calvert, 1892 25(24A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
310. O. albistylum (Selys, 1848) 25(24A+X) + Italy Kiauta 1971a (O. a. albistylum (Selys, 1848))
– » – + Russia Perepelov et al. 1998
– » – + Japan Oguma 1915, 1917, 1930 (O. a. speciosum (Uhler, 1858))
– » – + India Kichijo 1942b (O. a. speciosum)
– » – + Japan Omura 1955 (O. a. speciosum)
311. O. azureum (Rambur, 1842) 25(24A+X) + Madagascar Kiauta 1969b, c
312. O. brachiale (Beauvois, 1805) 21(20A+X) Kenya Kiauta 1969b, c
25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979 (O. b. brachiale (Beauvois, 1805))
313. O. brunneum (Fonscolombe, 1837) 25(24A+X) + Italy Kiauta 1971a
– » – + Russia Perepelov et al. 1998
314. O. cancellatum (Linnaeus, 1758) 25(24A+X) + Finland Oksala 1939a
– » – + India Dasgupta 1957
– » – + Netherlands Kiauta 1969a, b
– » – + India Tyagi 1982
– » – + Russia Kuznetsova et al. 2018
315. O. chrysostigma (Burmeister, 1839) 25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979
– » – + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
– » – + Kenya Wasscher 1985
316. O. coerulescens (Fabricius, 1798) 25(24A+X) + Austria Kiauta 1969c
23(22A+X)
25(24A+X) + Italy Kiauta 1971a
27(26A+X) +
317. O. glaucum (Brauer, 1865) 25(24A+X) + India Dasgupta 1957
– » – + India Tyagi 1978a, b
– » – + India Handa and Batra 1980
– » – + India Tyagi 1982
– » – + India Handa et al. 1984
– » – + India Walia and Sandhu 2002
– » – + India Kumari and Gautam 2017
318. O. guineese (Ris, 1909) 25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979
319. O. japonicum (Uhler, 1858) 25(24A+X) + Japan Oguma 1917, 1930 (O. j. internum McLachlan, 1894)
– » – + Japan Kichijo 1942b (O. j. internum)
– » – + Japan Omura 1955 (O. j. internum)
– » – + Nepal Kiauta 1975 (O. j. internum)
– » – + Nepal Kiauta and Kiauta 1976 (O. j. internum)
320. O. julia Kirby, 1900 25(24A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980 (O. j. falsum (Longfeild, 1955))
– » – + Kenya Wasscher 1985 (O. j. falsum)
321. O. luzonicum (Brauer, 1868) 25(24A+X) + Nepal Kiauta 1975
– » – + Nepal Kiauta and Kiauta 1982
– » – + India Thomas and Prasad 1981
– » – + India Prasad and Thomas 1992
322. O. melania (Selys, 1883) 25(24A+X) + Japan Oguma 1917
– » – + Japan Omura 1955
– » – + Russia Perepelov 2003
323. O. monardi (Schmidt, 1951) 25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979
324. O. poecilops (Ris, 1916) 25(24A+X) + Japan Suzuki et al. 1991 (O. p. miyajimaensis Yuki et Doi, 1938)
325. O. pruinosum (Burmeister, 1839) 25(24A+X) + India Dasgupta 1957 (O. p. neglectum (Rambur, 1842))
– » – + Taiwan Kiauta 1969a, c (O. p. neglectum)
– » – + India Tyagi 1982 (O. p. neglectum)
– » – + India Prasad and Thomas 1992 (O. p. neglectum)
– » – + India Tyagi 1978a, b (O. p. neglectum)
– » – + Nepal Kiauta and Kiauta 1982 (O. p. neglectum)
– » – + India Walia and Sandhu 2002 (O. p. neglectum)
– » – + India Kumari and Gautam 2017 (O. p. neglectum)
326. O. sabina (Drury, 1773) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942b
– » – + India Ray Chaudhuri and Dasgupta 1949
– » – + Nepal Kiauta 1975
326. O. sabina (Drury, 1773) – » – + India Tyagi 1982
– » – + India Prasad and Thomas 1992
– » – + India Walia and Sandhu 2002 (O. s. sabina (Drury, 1773))
327. O. taeniolatum (Schneider, 1845) 25(24A+X) + Greece Kiauta 1972a
– » – + Nepal Kiauta 1975
– » – + India Tyagi 1978a, b
– » – + India Handa and Batra 1980
– » – + India Tyagi 1982
– » – + India Handa et al. 1984
– » – + India Thomas and Prasad 1986
– » – + India Walia and Sandhu 2002a
– » – + India Walia et al. 2015
328. O. testaceum (Burmeister, 1839) 25(24A+X) + Nepal Kiauta and Kiauta 1982
329. O. triangulare (Selys, 1878) 25(24A+X) + Japan Omura 1955 (O. t. melania (Selys, 1883))
– » – + Taiwan Kiauta 1969a, b (O. t. triangulare (Selys, 1878))
– » – + Nepal Kiauta 1975 (O. t. triangulare)
– » – + India Tyagi 1978a, b (O. t. triangulare)
– » – + India Handa and Batra 1980 (O. t. triangulare)
– » – + India Tyagi 1982 (O. t. triangulare)
– » – + India Walia and Sandhu 2002 (O. t. triangulare)
330. Pachydiplax longipennis (Burmeister, 1839) 25(24A+X) USA Cumming 1964
– » – + USA Cruden 1968
– » – + USA Kiauta and Brink 1978
331. Palpopleura jucunda Rambur, 1842 25(24A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
332. P. lucia (Drury, 1773) 25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979 (P. l. portia (Drury, 1773))
– » – + Kenya Wasscher 1985 (P. l. portia)
333. P. sexmaculata (Fabricius, 1787) 25(24A+X) + Nepal Kiauta 1974, 1975
– » – + India Tyagi 1982 (P. s. sexmaculata (Fabricius, 1787))
334. Pantala flavescens (Fabricius, 1798) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942b
– » – + India Dasgupta 1957
– » – + India Seshachar and Bagga 1963
– » – + Bolivia Cumming 1964
– » – + Madagascar Kiauta 1969b
– » – + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
– » – + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
– » – + Brazil Souza Bueno 1982
– » – + Argentina Agopian and Mola 1988
– » – + India Prasad and Thomas 1992
– » – + Russia Perepelov and Bugrov 2001b
23(22A+X) + India Walia et al. 2011
335. P. hymenaea (Say, 1836) 25(24A+X) + Bolivia Cumming 1964
– » – + USA Cruden 1968
336. Perithemis cornelia Ris, 1910 25(24A+X) Bolivia Cumming 1964
337. P. domitia (Drury, 1773) 25(24A+X) + Jamaica Cumming 1964
338. P. electra Ris, 1928 25(24A+X) Bolivia Cumming 1964
339. P. icteroptera (Selys in Sagra, 1857) 25(24A+X) + Argentina Mola and Agopian 1985
340. P. lais (Petry, 1834) 17(16A+X) Bolivia Cumming 1964
– » – Surinam Kiauta 1979a
– » – Brazil Ferreira et al. 1979
341. P. mooma Kirby, 1889 25(24A+X) + Bolivia Cumming 1964
– » – + Surinam Kiauta 1979a
– » – Brazil Ferreira et al. 1979
– » – + Argentina Mola and Agopian 1985
342. P. tenera (Say, 1839) 25(24A+X) + USA Kiauta and Brink 1978
343. P. seminole Calvert, 1907 25(24A+X) + USA Cumming 1964
344. Perithemis sp. 25(24A+X) Bolivia Cumming 1964
345. Planiplax erythropyga (Karsch, 1891) 25(24A+X) + Argentina Mola et al. 1999
– » – + – » – De Gennaro 2004
346. P. sanguiniventris (Calvert, 1907) 25(24A+X) + USA Cruden 1968
347. Plathemis lydia (Drury, 1773) 25(24A+X) + USA McGill 1907
– » – + USA Cruden 1968
348. Potamarcha congener (Rambur, 1842) 25(24A+X) + India Asana and Makino 1935 as P. obscura (Rambur, 1842)
– » – + India Makino 1935 as P. obscura
– » – + India Kichijo 1942b as P. obscura
– » – + India Dasgupta 1957 as P. obscura
– » – + India Tyagi 1982 as P. obscura
– » – + India Prasad and Thomas 1992
– » – + India Sandhu and Walia 1995
349. Pseudothemis zonata (Burmeister, 1839) 24(22A+neo-XY) Japan Omura 1955
350. Pseudotramea prateri Fraser, 1920 25(24A+X) + Nepal Kiauta 1974, 1975
351. Rhodopygia cardinalis (Erichson, 1848) 25(24A+X) + Bolivia Cumming 1964
352. R. geijskesi Belle, 1964 25(24A+X) + Surinam Kiauta 1979a
353. Rhodothemis rufa (Rambur, 1842) 25(24A+X) + India Prasad and Thomas 1992
354. Rhyothemis fuliginosa Selys, 1883 25(24A+X) + Japan Toyoshima and Hirai 1953
– » – + Japan Omura 1955
– » – + Japan Hirai 1956
25(24A+X) + Japan Kiauta 1969c
23(22A+X) +
355. R. variegata (Linnaeus et Johansson, 1763) 25(24A+X) + India Ray Chaudhuri and Dasgupta 1949
– » – + Nepal Kiauta 1975
356. Scapanea frontalis (Burmeister, 1839) 25(24A+X) + Jamaica Cumming 1964
357. Sympetrum commixtum (Selys, 1884) 25(24A+X) India Tyagi 1978a, b, 1982
358. S. corruptum (Hagen, 1861 25(24A+X) + USA Cruden 1968 as Tarnetrum corruptum (Hagen, 1861)
– » – + USA Kiauta 1969a, c as T. corruptum
359. S. costiferum (Hagen, 1861) 25(24A+X) + USA Cruden 1968
360. S. croceolum (Selys, 1840) 25(24A+X) + Russia Perepelov 2003
361. S. danae (Sulzer, 1776) 25(24A+X) + Former USSR Makalowskaja 1940
– » – + Finland Oksala 1945
– » – + USA Cruden 1968
– » – + Russia Perepelov 2003
– » – + Russia Kuznetsova et al. 2018
362. S. eroticum (Selys, 1883) 21(20A+X) Japan Kichijo 1942b, c
– » – Japan Hirai 1956
– » – Japan Kiauta 1969c
363. S. flaveolum (Linnaeus, 1758) 25(24A+X) + Former USSR Makalowskaja 1940
– » – + Russia Perepelov 2003
364. S. fonscolombii (Selys, 1840) 25(24A+X) + Russia Perepelov 2003
365. S. frequens (Selys, 1883) 23(22A+X) Japan Oguma 1917, 1930
– » – Japan Kichijo 1942a, b
– » – Japan Kiauta 1969c
366. S. infuscatum (Selys, 1883) 25(24A+X) + Russia Perepelov 2003
367. S. internum Montgomery, 1943 27(26A+X) + Canada Kiauta 1973a
368. S. madidum (Hagen, 1861) 25(24A+X) + USA Cruden 1968
368. S. madidum (Hagen, 1861) – » – + Canada Kiauta 1973a
369. S. meridionale (Selys, 1841) 25(24A+X) + Switzerland Kiauta 1966
370. S. obtrusum (Hagen, 1867) 25(24A+X) + USA Cruden 1968
371. S. parvulum Bartenev, 1912 25(24A+X) + Japan Kiauta 1968c
372. S. pedemontanum Müller in Allioni, 1766 25(24A+X) + Japan Oguma 1917, 1930 (S. p. elatum (Selys, 1872))
– » – + Japan Kichijo 1942b (S. p. elatum)
– » – + Japan Kiauta and Brink 1975 (S. p. elatum)
– » – + Switzerland Kiauta and Brink 1975 (S. p. pedemontanum (Müller, 1766))
– » – + Russia Perepelov et al. 1998 (S. p. pedemontanum)
– » – + Russia Perepelov and Bugrov 2001b
373. S. rubicundulum (Say, 1839) 25(24A+X) + USA Cruden 1968
374. S. sanguineum (Müller, 1764) 25(24A+X) + Italy Kiauta 1971a
– » – + Russia Perepelov and Bugrov 2001b
375. S. semicinctum (Say, 1839) 25(24A+X) + USA Smith 1916
– » – + USA Cruden 1968
376. S. striolatum (Charpentier, 1840) 25(24A+X) Luxembourg Kiauta 1966
377. S. vicinum (Hagen, 1861) 25(24A+X) + USA Cruden 1968
378. S. vulgatum (Linnaeus, 1758) 25(24A+X) + Netherland Kiauta 1972c
– » – + Russia Perepelov 2003
– » – + Russia Kuznetsova et al. 2018
379. Tarnetrum illotum (Hagen, 1861) 25(24A+X) + Jamaica Cumming 1964
– » – + USA Cruden 1968
380. Tauriphila australis (Hagen, 1867) 25(24A+X) + Bolivia Cumming 1964
381. T. azteca Calvert, 1906 25(24A+X) + Mexico Cruden 1968
382. T. risi Martin 1896 25(24A+X) + Argentina, Uruguay Mola and Agopian 1985
383. Tholymis citrina Hagen, 1867 25(24A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
384. Th. tillagra (Fabricius, 1798) 25(24A+X) + India Prasad and Thomas 1992
– » – + Nepal Kiauta and Kiauta 1982
– » – + Thailand Kiauta and Kiauta 1983
385. Tramea abdominalis (Rambur, 1842) 25(24A+X) Bolivia Cumming 1964
386. T. basilaris (Palisot de Beauvois, 1817) 25(24A+X) + India Das 1956 (T. b. burmeisteri (Kirby, 1889))
– » – + India Dasgupta 1957 (T. b. burmeisteri)
– » – + Nepal Kiauta and Kiauta 1982 (T. b. burmeisteri)
– » – + India Prasad and Thomas 1992 (T. b. burmeisteri)
387. T. binotata (Rambur, 1842) 25(24A+X) + Surinam Kiauta 1979a
– » – Brazil Ferreira et al. 1979
388. T. carolina (Linnaeus, 1763) 25(24A+X) USA Cumming 1964
– » – USA Cruden 1968
389. T. cophysa (Hagen, 1867) 25(24A+X) + Bolivia Cumming 1964
390. T. lacerata (Hagen, 1861) 25(24A+X) USA Cruden 1968
391. T. limbata (Desjardins, 1832) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942b
392. T. virginia (Rambur, 1842) 25(24A+X) + India Oguma and Asana 1932
– » – + India Kichijo 1942b
– » – + India Dasgupta 1957
393. Trithemis annulata (Palisot de Beauvois, 1805) 25(24A+X) Republic of South Africa Boyes et al. 1980
– » – + Kenya Wasscher 1985
394. T. arteriosa (Burmeister, 1839) 25(24A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
395. T. atra Pinhey, 1961 25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979
396. T. aurora (Burmeister, 1839) 25(24A+X) + India Oguma and Asana 1932
– » – + Nepal Kiauta 1975
– » – + India Tyagi 1982
397. T. dorsalis (Rambur, 1842) 25(24A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
398. T. festiva (Rambur, 1842) 25(24A+X) + Nepal Kiauta 1974, 1975
– » – + India Tyagi 1982
– » – + India Prasad and Thomas 1992
399. T. furva Karsch, 1899 25(24A+X) + Sudan Wasscher 1985
400. T. imiata Pinhey, 1961 25(24A+X) Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979
401. T. kirbyi Selys, 1891 25(24A+X) Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979 (T. k. ardens Gerstaecker, 1891)
– » – + Kenya Wasscher 1985 (T. k. ardens)
402. T. pallidinervis (Kirby, 1889) 25(24A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942b
– » – + India Dasgupta 1957
– » – + Philippines Kiauta and Kiauta 1980b
403. T. werneri Ris, 1912 25(24A+X) + Kenya Wasscher 1985
404. Uracis imbuta (Burmeister, 1839) 25(24A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
405. U. ovipositrix Calvert, 1909 25(24A+X) + Surinam Kiauta 1979a
– » – Brazil Ferreira et al. 1979
406. Urothemis edwardsi (Selys, 1849) 25(24A+X) + Sudan Wasscher 1985
407. U. signata (Rambur, 1842) 25(24A+X) + India Das 1956 (U. s. signata (Rambur, 1842))
– » – + India Dasgupta 1957 (U. s. signata)
– » – + Nepal Kiauta 1975
– » – + India Prasad and Thomas 1992
408. Zenithoptera fasciata (Linnaeus, 1758) 25(24A+X) + Surinam Kiauta 1979a
409. Z. lanei Santos, 1941 25(24A+X) + Surinam Kiauta 1979a
– » – + Brazil Ferreira et al. 1979
410. Z. viola Ris, 1910 25(24A+X) + Bolivia Cumming 1964
411. Zygonyx iris Kirby, 1900 23(22A+X) + Thailand Kiauta and Kiauta 1983 (Z. i. malayanus (Laidlaw, 1902))
412. Z. torrida (Kirby, 1889) 25(24A+X) + India Tyagi 1978a, b
413. Zyxomma petiolatum (Rambur, 1842) 25(24A+X) + India Prasad and Thomas 1992
Cordulegastroidea
Chlorogomphidae
414. Watanabeopetalia atkinsoni (Selys, 1878) 25(24A+X) + India Walia and Chahal 2019
Cordulegastridae
415. Anotogaster basalis Selys, 1854 23(22A+X) India Sandhu and Malhotra 1994b
416. A. kuchenbeiseri (Förster, 1899) 25(24A+X) + China Zhu and Wu 1986
417. A. sieboldii (Selis, 1854) 25(24A+X) + Japan Oguma 1930
– » – + Japan Kichijo 1942a
– » – + Japan Kiauta 1969a
– » – + Russia Perepelov et al. 2001
418. Cordulegaster boltoni (Donovan, 1807) 25(24A+X) + Finland Oksala 1939a, b
– » – Austria Kichijo 1942a
– » – + Sweden Kiauta 1968d, e, 1969a
419. C. brevistigma Selys, 1854 25(24A+X) + India Walia and Chahal 2019
420. C. diastatops (Selys, 1854) 25(24A+X) + USA Cruden 1968
421. C. dorsalis Hagen, 1857 25(24A+X) + USA Cruden 1968
422. C. maculata Selys, 1854 25(24A+X) + USA Cruden 1968
423. Neallogaster hermionae (Fraser, 1927) 25(24A+X) + Nepal Kiauta and Kiauta 1976
Zygoptera
Lestoidea
Lestidae
424. Austrolestes colensonis (White, 1846) 25(24A+X) + New Zealand Jensen 1980
425. Chalcolestes viridis (Van der Linden, 1825) 25(24A+X) + Netherlands Kiauta 1969a
426. Indolestes cyaneus (Selys, 1862) 25(24A+X) + Nepal Kiauta and Kiauta 1976 as I. cyanea (Selys, 1862)
427. Lestes barbarus (Fabricius, 1798) 25(24A+X) + Former Yugoslavia Kiauta 1972a
428. L. congener Hagen, 1861 25(24A+X) + USA Cruden 1968
429. L. disjunctus Selys, 1862 25(24A+X) USA Cruden 1968
430. L. dorothea Fraser, 1924 25(24A+X) + Nepal Kiauta 1974, 1975
431. L. dryas Kirby, 1890 25(24A+X) USA Cruden 1968
– » – + Russia Perepelov and Bugrov 2001b
432. L. forcipatus Rambur, 1842 21(20A+X) USA Cruden 1968
433. L. forficula Rambur, 1842 25(24A+X) + Jamaica Cumming 1964
434. L. paulistus Calvert, 1909 25(24A+X) + Brazil Souza Bueno 1982
435. L. rectangularis Say, 1839 25(24A+X) + USA Cruden 1968
436. L. similatrix McLachlan, 1895 25(24A+X) + Madagascar Kiauta 1969b
437. L. sponsa (Hansemann, 1823) 25(24A+X) Former USSR Makalowskaja 1940
– » – + Japan Kichijo 1941, 1942a, d, e
– » – + Russia Perepelov and Bugrov 2001b
438. L. stultus Hagen, 1861 25(24A+X) + USA Cruden 1968
439. L. vidua Hagen, 1861 25(24A+X) + USA Cumming 1964
440. L. vigilax Selys, 1862 19(18A+X) USA Kiauta and Brink 1978
441. L. virens Charpentier, 1825 25(24A+X) + Netherlands Kiauta 1969a (L. v. vestalis Rambur, 1842)
442. Sympecma fusca (Van der Linden, 1823) 25(24A+X) + Japan Kichijo 1941, 1942d, e
443. S. paedisca (Brauer, 1877) 25(24A+X) + Netherlands Kiauta and Kiauta-Brink 1975 (S. annulata braueri (Bianchi, 1904))
– » – + Russia Perepelov 2003 (S. a. braueri)
Synlestidae
444. Megalestes major Selys, 1862 25(24A+X) Nepal Kiauta 1974, 1975
Platystictoidea
Platystictidae
445. Drepanosticta sp. 25(24A+X) Nepal Kiauta and Kiauta 1976
446. Drepanosticta sp. 25(24A+X) India Tyagi 1978a, b
447. Palaemnema paulina (Drury, 1773) 25(24A+X) + Costa Rica Cumming 1964
448. Protosticta sp. 25(24A+X) Tailand Kiauta and Kiauta 1983
Calopterygoidea
Calopterygidae
449. Atrocalopteryx atrata (Selys, 1853) 25(24A+X) + Japan Oguma 1930 as Calopteryx atrata Selys, 1853
– » – + Japan Kichijo 1942d as C. atrata
– » – + Japan Omura 1957 as C. atrata
450. Calopteryx aequabilis Say, 1839 25(24A+X) + USA Cruden 1968
451. C. cornelia (Selys, 1853) 25(24A+X) + Japan Oguma 1930 as Anaciagrion cornelia (Selys, 1853)
– » – + Japan Kichijo 1942a as A. cornelia
452. C. dimidiata Burmeister, 1839 25(24A+X) + USA Kiauta and Brink 1978
453. C. japonica Selys, 1869 25(24A+X) + Japan Kichijo 1942a
– » – + Japan Hirai 1956
– » – + Japan Omura 1957
– » – + Japan Kiauta 1968e, f
454. C. maculata (Beauvois, 1805) 25(24A+X) + USA Cumming 1964a
– » – + USA Cruden 1968
455. C. splendens (Harris, 1780) 25(24A+X) + Turkey Kiauta 1972a (C. s. amasina Bartenev, 1912)
455. C. splendens (Harris, 1780) – » – + Italy Kiauta 1971a (C. s. caprai Conci, 1956)
– » – Former USSR Makalowskaja 1940 (C. s. splendens (Harris, 1782))
– » – Finland Oksala 1945 (C. s. splendens)
– » – Germany Kiauta 1969a, 1971b (C. s. splendens)
– » – France Kiauta 1973b (C. s. splendens)
– » – Russia Perepelov et al. 1998 (C. s. splendens)
– » – + Russia Kuznetsova et al. 2020b
456. C. virgo (Linnaeus, 1758) 25(24A+X) + Spain Kiauta 1971b (C. v. meridionalis Selys, 1873)
27(26A+X) +
25(24A+X) + Slovenija Kiauta 1967a, 1968b, c (C. v. padana Conci, 1956)
– » – + Austria Kiauta 1967a, 1968b, c (C. v. padana)
– » – Belgium Carnoy 1885 (C. v. virgo (Linnaeus, 1758))
– » – + Finland Oksala 1939 (C. v. virgo)
– » – + Former USSR Makalowskaja 1940 (C. v. virgo)
– » – + Germany, Luxembourg Kiauta 1968e, f (C. v. virgo)
– » – + Netherlands Kiauta 1972c (C. v. virgo)
– » – + Russia Kuznetsova et al. 2020b
457. Hetaerina americana (Fabricius, 1798) 25(24A+X) + USA Cumming 1964
– » – USA Cruden 1968
458. H. charca Calvert, 1909 25(24A+X) + Bolivia Cumming 1964
459. H. longipes (Hagen in Selys, 1853) 25(24A+X) + Brazil Souza Bueno 1982 as H. carnifex Hagen in Selys, 1853
– » – + Brazil Agopian and Mola 1984 as H. carnifex
460. H. rosea Selys, 1853 27(26A+X) + Bolivia Cumming 1964
– » – + Bolivia Kiauta 1969c
25(24A+X) Brazil Ferreira et al. 1979
27(26A+X) +
461. H. sanguinea Selys, 1853 25(24A+X) Bolivia Cumming 1964
462. H. titia (Drury, 1773) 25(24A+X) + USA Cumming 1964
– » – + Mexico Kiauta 1970a as H. tricolor (Burmeister, 1839)
463. H. vulnerata (Selys, 1853) 25(24A+X) + Mexico Kiauta 1970a
464. Matrona basilaris Selys, 1853 25(24A+X) Taiwan Kiauta 1968c
465. Mnais costalis Selys, 1869 25(24A+X) + Japan Oguma 1930
– » – + Japan Kichijo 1942a
466. M. pruinosa Selys, 1853 25(24A+X) + Japan Oguma 1930 as M. strigata Selys, 1853
– » – + Japan Kichijo 1942a as M. strigata
– » – + Japan Omura 1957 as M. strigata
467. Neurobasis chinensis (Linnaeus, 1758) 23(22A+X) Nepal Kiauta 1975 (N. c. chinensis (Linnaeus, 1758))
25(24A+X)
23(22A+X) India Tyagi 1978b (N. c. chinensis)
– » – + Nepal Kiauta and Kiauta 1982 (N. c. chinensis)
– » – Thailand Kiauta and Kiauta 1983 (N. c. chinensis)
– » – + India Walia and Sandhu 2002 (N. c. chinensis)
– » – India Walia et al. 2016 (N. c. chinensis)
– » – India Walia and Katnoria 2018 (N. c. chinensis)
468. Phaon iridipennis (Burmeister, 1839) 25(24A+X) + Republic of South Africa Boyes et al. 1980
Chlorocyphidae
469. Aristocypha fenestrella Rambur, 1842 23(22A+X) Thailand Kiauta and Kiauta 1983 as Rhinocypha fenestrella Rambur, 1842
470. A. quadrimaculata (Selys, 1853) 23(22A+X) + India Chatterjee and Kiauta 1973 as Rhinocypha quadrimaculata Selys, 1853
– » – + Nepal Kiauta and Kiauta 1982 as Rh. quadrimaculata
471. A. trifasciata (Selys, 1853) 23(22A+X) India Tyagi 1978a, b as Rhinocypha trifasciata Selys, 1853
– » – + Nepal Kiauta and Kiauta 1982 as Rh. trifasciata
472. Heliocypha biforata (Selys, 1859) 23(22A+X) India Tyagi 1978a, b as Rhinocypha biforata beesoni Selys, 1859
473. H. biseriata (Selys, 1859) 23(22A+X) Thailand Kiauta and Kiauta 1983 as Rhinocypha b. biforata Selys, 1859
474. Libellago lineata (Burmeister, 1839) 23(22A+X) India Walia et al. 2018 (L. l. lineata (Burmeister, 1839))
25(24A+X)
475. Paracypha unimaculata (Selys, 1879) 23(22A+X) + Nepal Kiauta 1974, 1975 as Rhinocypha unimaculata Selys, 1879
– » – + Nepal Kiauta and Kiauta 1982 as Rh. unimaculata
476. Rhinocypha colorata Selys, 1869 23(22A+X) Philippines Kiauta and Kiauta 1980b
25(24A+X)
477. Vestalis gracilis (Rambur, 1842) 25(24A+X) + Thailand Kiauta and Kiauta 1983
Polythoridae
478. Cora irene Ris, 1918 23(22A+X) Bolivia Cumming 1964
479. Polythore boliviana (McLachlan, 1878) 23(22A+X) Bolivia Cumming 1964
Euphaeidae
480. Anisopleura comes Hagen, 1880 25(24A+X) + Nepal Kiauta and Kiauta 1976, 1982
481. Bayadera indica (Selys, 1853) 25(24A+X) + Nepal Chatterjee and Kiauta 1973
– » – + Nepal Kiauta 1975
482. Euphaea guerini Rambur, 1842 25(24A+X) Thailand Kiauta and Kiauta 1983
483. Epallage fatime (Charpentier, 1840) 25(24A+X) Greece Kiauta 1970b
– » – Greece Chatterjee and Kiauta 1973
Megapodagrionidae
484. Allopodagrion contortum (Selys, 1862) 25(24A+X) + Brazil Kiauta 1972b as Megapodagrion contortum (Selys, 1862)
485. Teinopodagrion macropus (Selys, 1862) 25(24A+X) Bolivia Cumming 1964 as Megapodagrion macropus (Selys, 1862)
486. T. setigerum (Selys, 1886) 25(24A+X) Bolivia Cumming 1964 as Megapodagrion setigerum Selys, 1886
Heteragrionidae
487. Heteragrion flavidorsum Calvert, 1909 25(24A+X) Bolivia Cumming 1964
488. H. inca Calvert, 1909 25(24A+X) + Bolivia Cumming 1964
Philogeniidae
489. Philogenia carrillica Calvert, 1907 25(24A+X) + Costa Rica Cumming 1964
Hypolestidae
490. Hypolestes clara (Calvert, 1891) l7(16A+X) Jamaica Cumming 1964
Coenagrionoidea
Platycnemididae
491. Calicnemia miniata (Selys, 1886) 25(24A+X) + Nepal Kiauta and Kiauta 1982
492. C. pulverulans (Selys, 1886) 25(24A+X) Nepal Kiauta 1975
493. Calicnemia sp. 25(24A+X) Nepal Kiauta 1975
494. Calicnemia sp. 25(24A+X) India Tyagi 1978b
495. Coeliccia chromothorax (Selys, 1891) 25(24A+X) India Walia and Devi 2020b
496. C. bimaculata (Laidlaw, 1914) 25(24A+X) India Walia and Devi 2020b
497. C. didyma (Selys, 1863) 25(24A+X) India Walia and Devi 2020b
498. C. fraseri (Laidlaw, 1932) 25(24A+X) India Walia and Devi 2020b
499. C. renifera (Selys, 1886) 25(24A+X) Nepal Kiauta 1974, 1975
– » – India Walia and Devi 2020b
500. Copera annulata (Selys, 1863) 25(24A+X) + Japan Kichijo 1941, 1942a, c
– » – + India Dasgupta 1957
– » – Thailand Kiauta and Kiauta 1983
– » – + India Walia and Devi 2018
501. C. marginipes (Rambur, 1842) 25(24A+X) India Tyagi 1978a, b
– » – Thailand Kiauta and Kiauta 1983
– » – + India Walia and Devi 2018
502. C. vittata (Selys, 1863) 25(24A+X) + India Walia and Devi 2018
– » – + India Walia and Devi 2018 (C. v. assamensis (Laidlaw, 1914))
503. Disparoneura quadrimaculata (Rambur, 1842) 25(24A+X) India Walia and Devi 2020a
504. Esme cyaneovittata Fraser, 1922 25(24A+X) India Walia and Devi 2020a
505. E. longistyla Fraser, 1931 25(24A+X) India Walia and Devi 2020a
506. Onychargia atrocyana (Selys, 1865) 25(24A+X) Thailand Kiauta and Kiauta 1983
507. Platycnemis pennipes (Pallas, 1771) 25(24A+X) Finland Oksala 1945
– » – Italy Kiauta 1971a
– » – Russia Perepelov and Bugrov 2001b
508. Prodasineura autumnalis (Fraser, 1922) 25(24A+X) + Thailand Kiauta and Kiauta 1983
509. P. nigra (Fraser, 1922) 25(24A+X) India Walia and Devi 2020a
510. P. verticalis (Selys, 1860) 25(24A+X) India Walia and Devi 2020a
511. Prodasineura sp.1 25(24A+X) Thailand Kiauta and Kiauta 1983
512. Prodasineura sp.2 25(24A+X) Thailand Kiauta and Kiauta 1983
Coenagrionidae
513. Acanthagrion ascendens Calvert, 1909 27(26A+X) + Bolivia Cumming 1964
514. A. chacoense Calvert, 1909 27(26A+X) + Bolivia Cumming 1964
515. A. gracile (Rambur, 1842) 27(26A+X) Surinam Kiauta 1979a (A. g. minarum Selys, 1876)
– » – Brazil Ferreira et al. 1979 (A. g. minarum Selys, 1876)
516. Aeolagrion inca Selys, 1876 27(26A+X) Bolivia Cumming 1964 as A. foliaceum (Sjöstedt, 1918)
517. Agriocnemis clauseni Fraser, 1922 27(26A+X) + India Tyagi 1978a, b
518. A. femina (Brauer, 1868) 27(26A+X) Philippines Kiauta and Kiauta 1980b
– » – + Thailand Kiauta and Kiauta 1983
519. A. pygmaea (Rambur, 1842) 27(26A+X) India Tyagi 1978b
– » – + Thailand Kiauta and Kiauta 1983
520. Amphiagrion abbreviatum (Selys, 1876) 27(26A+X) USA Cruden 1968
521. Amphiallagma parvum (Selys, 1876) 27(26A+X) + India Handa and Kochhar 1985 as Enallagma parvum Selys, 1876
522. Argia apicalis (Say, 1839) 37(36A+X) USA Kiauta and Kiauta 1980b
523. A. fumipennis (Burmeister, 1839) 27(26A+X) USA Kiauta and Kiauta 1980c (A. f. atra Gloyd, 1968)
– » – USA Kiauta and Brink 1978 (A. f. fumipennis (Burmeister, 1839))
– » – USA Kiauta and Kiauta 1980c (A. f. fumipennis)
– » – + Canada Kiauta and Kiauta 1980c (A. f. violacea (Hagen, 1861))
524. A. funebris (Hagen, 1861) 27(26A+X) USA Kiauta 1972b
28(26A+XX)* Mexico Kiauta and Kiauta 1980c
525. A. immunda (Hagen, 1861) 27(26A+X) USA Kiauta and Kiauta 1980c
526. A. moesta (Hagen, 1861) 25(24A+X) Canada Kiauta 1978
– » – USA Kiauta and Kiauta 1980c
527. A. nahuana Calvert, 1902 25(24A+X) USA Kiauta and Kiauta 1980c
528. A. sedula (Hagen, 1861) 27(26A+X) USA Cruden 1968
– » – USA Kiauta and Kiauta 1980c
529. A. tibialis (Rambur, 1842) 37(36A+X) USA Kiauta and Kiauta 1980c
530. A. translata Hagen, 1865 25(24A+X) + USA Kiauta and Kiauta 1980c
531. A. violacea (Hagen, 1861) 27(26A+X) USA Cruden 1968
532. A. vivida (Hagen, 1861) 27(26A+X) USA Cruden 1968
533. Ceriagrion auranticum Fraser, 1922 27(26A+X) + Thailand Kiauta and Kiauta 1983 as C. latericium Lieftinck, 1951
534. C. azureum (Selys, 1891) 27(26A+X) Nepal Kiauta 1974, 1975
535. C. cerinomelas Lieftinck, 1927 27(26A+X) Nepal Kiauta 1974, 1975
536. C. cerinorubellum (Brauer, 1866) 27(26A+X) + India Dasgupta 1957
– » – + India Prasad and Thomas 1992
537. C. coromandelianum (Fabricius, 1798) 27(26A+X) + India Ray Chaudhuri and Dasgupta 1949
– » – + India Srivastava and Das 1953
– » – + India Das 1956
– » – + Nepal Kiauta and Kiauta 1982
– » – + India Prasad and Thomas 1992
538. C. fallax Ris, 1914 27(26A+X) + Republic of South Africa Dasgupta 1957
539. C. glabrum (Burmeister, 1839) 27(26A+X) Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
540. C. rubiae Laidlaw, 1916 27(26A+X) + India Asana and Makino 1935
– » – + India Makino 1935
– » – + India Kichijo 1942a
541. C. tenellum (Villers, 1789) 27(26A+X) + Italy Kiauta 1971a (C. t. tenellum (Villers, 1789))
542. Chromagrion conditum (Hagen, 1876) 27(26A+X) USA Cruden 1968
543. Coenagrion armatum (Charpentier, 1840) 27(26A+X) Finland Oksala 1939a
– » – Former USSR Makalowskaja 1940
544. C. hastulatum (Charpentier, 1825) 27(26A+X) Former USSR Makalowskaja 1940
– » – Russia Perepelov and Bugrov 2001b
545. C. hylas (Trybom, 1889) 27(26A+X) Austria Kiauta and Kiauta 1991 (C. h. freyi (Bilek, 1954))
546. C. lunulatum (Charpentier, 1840) 27(26A+X) + Russia Perepelov and Bugrov 2001b
547. C. pulchellum (Vander Linden, 1823) 27(26A+X) Former USSR Makalowskaja 1940
– » – Netherlands Kiauta 1969c
– » – + Russia Kuznetsova et al. 2020b
548. C. puella (Linnaeus, 1758) 27(26A+X) + Russia Kuznetsova et al. 2020b
549. C. resolutum (Hagen, 1876) 27(26A+X) USA Cruden 1968
550. Coenagrion sp. 27(26A+X) + Japan Kichijo 1941, 1942d, e
551. Diceratobasis macrogaster (Selys, 1875) 27(26A+X) + Jamaica Cumming 1964
552. Enallagma aspersum (Hagen, 1861) 27(26A+X) USA Cruden 1968
553. E. boreale Selys, 1875 27(26A+X) USA Cruden 1968
554. E. carunculatum Morse, 1895 27(26A+X) USA Cruden 1968
555. E. circulatum Selys, 1883 27(26A+X) + Russia Perepelov and Bugrov 2001b
556. E. civile (Hagen, 1861) 27(26A+X) USA Cruden 1968
557. E. cyathigerum (Charpentier, 1840) 27(26A+X) Finland Oksala 1939a, 1945
– » – Former USSR Makalowskaja 1940
– » – + USA Brink and Kiauta 1964
27(26A+X), USA Cruden 1968
29(28A+X)
27(26A+X) + Netherlands Kiauta 1969a, c
29(28A+X) +
558. E. ebrium (Hagen, 1861) 27(26A+X) USA Cruden 1968
559. E. praevarum (Hagen, 1861) 27(26A+X) USA Cruden 1968
560. Erythromma lindeni (Selys, 1840) 27(26A+X) + Italy Kiauta 1971a
561. E. najas (Hansemann, 1823) 27(26A+X) Finland Oksala 1939a
– » – Former USSR Makalowskaja 1940
– » – Netherlands Kiauta 1969a
– » – Russia Perepelov and Bugrov 2001b
– » – + Russia Kuznetsova et al. 2020b
562. Homeoura chelifera (Selys, 1876) 27(26A+X) + Surinam Kiauta 1979a as Enallagma cheliferum (Selys, 1876)
– » – + Brazil Ferreira et al. 1979 as E. cheliferum
563. Ischnura aurora (Brauer, 1865) 27(26A+X) Nepal Kiauta 1974, 1975
– » – India Handa and Kochhar 1985
564. I. capreola (Hagen, 1861) 27(26A+X) Bolivia Cumming 1964 as Ceratura capreola (Hagen, 1861)
565. I. cervula Selys, 1876 27(26A+X) USA Cruden 1968
566. I. denticollis (Burmeister, 1839) 27(26A+X) USA Cruden 1968
567. I. elegans (Van der Linden, 1823) 27(26A+X) Finland Oksala 1939a, 1945
– » – Netherlands Kiauta 1969a
– » – Russia Perepelov 2003
568. I. fluviatilis Selys, 1876 27(26A+X) Bolivia Cumming 1964
569. I. forcipata Morton, 1907 27(26A+X) Nepal Kiauta 1974, 1975
570. I. nursei (Morton, 1907) 25(24A+X) + India Tyagi 1978b as Rhodischnura nursei (Morton, 1907)
571. I. pumilio (Charpentier, 1825) 27(26A+X) + Netherlands Kiauta 1979b
572. I. perparva Selys, 1876 27(26A+X) USA Cruden 1968
573. I. ramburii (Selys, 1850) 27(26A+X) + USA Kiauta and Brink 1978
574. I. rufostigma Selys, 1876 27(26A+X) Nepal Kiauta 1974, 1975 (I. r. annandalei Laidlaw, 1919)
575. I. senegalensis (Rambur, 1842) 27(26A+X) + Japan Kichijo 1941, 1942d, e
– » – + India Dasgupta 1957
– » – + Ethiopia Kiauta 1969b
– » – + Philippines Kiauta and Kiauta 1980b
– » – Thailand Kiauta and Kiauta 1983
– » – + India Prasad and Thomas 1992
576. I. verticalis (Say, 1839) 27(26A+X) USA Cruden 1968
577. I. ultima Ris, 1908 27(26A+X) Bolivia Cumming 1964
578. Leptagrion macrurum (Burmeister, 1839) 30(28A+neo-XY) Brazil Kiauta 1971c, 1972d
579. Mecistogaster . sp. 1 29(28A+X) + Bolivia Cumming 1964
580. Mecistogaster sp. 2 12(10A+neo-XY) Bolivia Cumming 1964
581. Megalagrion oahuense (Blackburn, 1884) 27(26A+X) + Hawaii Kiauta 1969b
582. Mortonagrion selenion (Ris, 1916) 27(26A+X) + Japan Kichijo 1941, 1942a, d, e
583. Nehalennia irene (Hagen, 1861) 27(26A+X) USA Cruden 1968
584. N. speciosa (Charpentier, 1840) 28(26A+XX)* Finland Oksala 1945
585. Oxyagrion hempeli Calvert, 1909 27(26A+X) Brazil Souza Bueno 1982
586. O. terminale Selys, 1876 27(26A+X) Surinam Kiauta 1979a
– » – Brazil Ferreira et al. 1979
587. Paracercion hieroglyphicum (Brauer, 1865) 27(26A+X) + Japan Kichijo 1941, 1942d, e as Coenagrion hieroglyphicum (Brauer, 1865)
588. P. malayanum (Selys, 1876) 27(26A+X) + Nepal Kiauta 1974, 1975
589. Proischnura subfurcata (Selys, 1876) 27(26A+X) Kenya Wasscher 1985 as Enallagma subfurcatum Selys, 1876
590. Pseudagrion acaciae Förster, 1906 27(26A+X) + Republic of South Africa Boyes et al. 1980
591. P. australasiae Selys, 1876 27(26A+X) + India Dasgupta 1957
592. P. decorum (Rambur, 1842) 27(26A+X) + India Dasgupta 1957
593. P. kersteni (Gerstaker, 1869) 27(26A+X) Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
594. P. microcephalum (Rambur, 1842) 27(26A+X) + India Dasgupta 1957
– » – + Philippines Kiauta and Kiauta 1980b
595. P. pruinosum (Burmeister, 1839) 27(26A+X) + Thailand Kiauta and Kiauta 1983
596. P. rubripes (Selys, 1876) 27(26A+X) + India Dasgupta 1957
– » – + Philippines Kiauta and Kiauta 1980b
– » – + Thailand Kiauta and Kiauta 1983
597. P. salisburyense Ris, 1921 27(26A+X) + Kingdom of Eswatini (Former Swaziland) Boyes et al. 1980
598. P. spencei Fraser, 1922 27(26A+X) + India Dasgupta 1957
599. P. whellani Pinhey, 1956 25(24A+X) + Burkina Faso (Former Voltiac Republic) Kiauta and Ochssée 1979
600. Pyrrhosoma nymphula (Sutzer, 1776) 28(26A+XX)* Finland Oksala 1945
601. Telebasis carmesina Calvert, 1909 27(26A+X) Surinam Kiauta 1979a
– » – Brazil Ferreira et al. 1979
602. Tigriagrion aurantinigrum Calvert, 1909 27(26A+X) Bolivia Cumming 1964
603. Xanthocnemis zealandica (McLachlan, 1873) 27(26A+X) New Zealand Jensen 1980 as X. zelandica (McLachlan, 1873)
604. Zoniagrion exclamationis (Selys, 1876) 27(26A+X) USA Cruden 1968
Protoneuridae
605. Caconeura autumnalis Fraser, 1922 25(24A+X) + India Tyagi 1978b
606. Epipleoneura sp. 27(26A+X) Bolivia Cumming 1964
607. Protoneura rubriventris (Selys, 1860) 27(26A+X) + Bolivia Cumming 1964 as Neoneura rubriventris Selys, 1860

Within Odonata, chromosome numbers in males vary over a relatively wide range, from 2n = 6 in Macrothemis hemichlora and Orthemis levis to 2n = 41 in O. nodiplaga. Both low chromosome number species are suggested to have an evolutionarily secondary neo-XY system (Cumming 1964; Kiauta 1972c) that could have arisen through an X-autosome fusion from an X(0) system. All three of the above species belong to the largest dragonfly family Libellulidae, in which nearly 89% of studied species (255 in total) have the karyotype 2n = 25(24A + X). The last one is the most common in Odonata in general: it occurs in each of the three suborders, Zygoptera, Anisoptera and Anisozygoptera, and in all families with the exception of two damselfly families, the Polythoridae with only two studied species sharing 2n = 23(22A + X) and a monotypic family Hypolestidae with 2n = 17(16A + X) in male Hypolestes clara. Besides Libellulidae, the karyotype 2n = 25(24A + X) is currently the presumed modal one in 14 other families, such being the case at least in six better covered (at species and/or generic level) families, i.e. the dragonfly families Corduliidae, Cordulegastridae, and Macromiidae, and the damselfly families Lestidae, Calopterygidae, and Platycnemididae (Table 2, Fig. 1). This chromosome set is suggested to be an ancestral one for the order Odonata in general (Oguma 1930; Kuznetsova et al. 2020b) although this suggestion remains questionable at this stage.

Table 2.

The diversity of chromosome numbers and sex chromosome mechanisms, and modal karyotypes in 23 families of Odonata: a summary.

Taxa N of species/ genera studied Male karyotypes Modal karyotype N of species/genera with modal karyotype (occurrence in percent)
(N of species/genera described*)
Anisozygoptera
Epiophlebioidea Epiophlebiidae (4/1) 1/1 25, X0 24A + X 1 (100) / 1 (100)
Anisoptera
Aeshnoidea Aeshnidae (456/51) 58/18 13, X0; 14, neo-XY; 15, X0; 16, neo-XY; 19, X0; 21, X0; 24, neo-XY; 25, X0; 26, neo-XY; 27, X0 26A + X 44 (76) / 14 (78)
Petaluroidea Petaluridae (10/5) 4/3 17, X0; 19, X0; 25, X0 16A + X 3 (75) / 2 (67)
Gomphoidea Gomphidae (980/87) 66/31 12, neo-neo-XY; 21, X0; 22, neo-XY; 23, X0; 24, neo-XY; 25, X0 22A + X 57 (86) / 28 (90)
Libelluloidea Macromiidae (125/4) 6/3 25, X0 24A + X 6 (100) / 3 (100)
Corduliidae (154/20) 23/7 10, neo-XY; 11, X0; 13, X0; 14, neo-XY, 20, XY; 21, X0; 25, X0; 26, neo-XY; 27, X0 24A + X 19 (83) / 6 (86)
Libellulidae (1037/142) 255/59 6, neo-XY; 6 neo-XY; 8, neo-XY; 10, neo-XY; 12, neo-XY; 17, X0; 21, X0; 22, neo-XY; 23, X0; 23, X1X2Y; 24, neo-XY; 25, X0; 27, X0; 28, neo-XY; 29, X0; 41, X0 24A + X 227 (89) / 57 (97)
Cordulegastroidea Cordulegastridae (46/3) 9/3 23, X0; 25, X0 24A + X 8 (89) / 3 (100)
Chlorogomphidae (47/3) 1/1 25, X0 24A + X 1 (100) / 1 (100)
Zygoptera
Lestoidea Lestidae (151/9) 20/5 19, X0; 21, X0; 25, X0 24A + X 18 (90) / 5 (100)
Synlestidae (39/9) 1/1 25, X0 24A + X 1 (100) / 1 (100)
Platystictoidea Platystictidae (224/6) 4/3 25, X0 24A + X 4 (100) / 3 (100)
Calopterygoidea Calopterygidae (185/21) 20/8 23, X0; 25, X0; 27, X0 24A + X 20 (100) / 8 (100)
Chlorocyphidae (144/19) 9/6 23, X0; 25, X0 22A + X 8 (89) / 5 (84)
Polythoridae (59/7) 2/2 23, X0 22A + X 2 (100) / 2 (100)
Euphaeidae (68/12) 4/4 25, X0 24A + X 4 (100) / 4 (100)
Megapodagrionidae (296/42) 3/2 25, X0 24A + X 3 (100) / 2 (100)
Heteragrionidae (57/2) 2/1 25, X0 24A + X 2 (100) / 1 (100)
Philogeniidae (40/2) 1/1 25, X0 24A + X 1 (100) / 1 (100)
Hypolestidae (6/4) 1/1 17, X0 16A + X 1 (100) / 1 (100)
Coenagrionoidea Platycnemididae (404/40) 22/8 25, X0 24A + X 19 (100) / 7 (100)
Coenagrionidae (1267/114) 92/28 12, neo-XY; 25, X0; 27, X0; 29, X0; 30, neo-XY; 37, X0 26A + X 81 (89) / 26 (90)
Protoneuridae (260 / 25) 3/3 25, X0; 27, X0 26A + X 2 (70) / 2 (70)

Chromosomal rearrangements, among which fission and fusions apparently predominated (Kiauta 1969c, 1972c), led to the appearance of divergent karyotypes in the evolution of Odonata. As a result, in many dragonfly and damselfly families, other karyotypes, when occurring, are of secondary origin as indicated by either a diverged number of autosomes or a secondary sex chromosome system of an XY-type or both (e.g. Cumming 1964; Kiauta 1969a, c; Agopian and Mola 1984, 1988; Mola et al. 1999; Perepelov and Bugrov 2002). Some interesting examples of this kind can be found in the family Libellulidae, in which 2n = 25(24A + X) is most likely an evolutionarily initial karyotype (e.g. Agopian and Mola 1988). These examples are as follows (see Table 1): Orthemis nodiplaga and O. ambinigra with 2n = 41(40A + X) and 2n = 12(10A + neo-XY), respectively; Erythrodiplax media and E. minuscula, both with 2n = 22(20A + neo-XY); Micrathyria longifasciata and M. ungulata with 2n = 24(22A + neo-XY) and 2n = 23(20A + X1X2Y), respectively. In some families, any of these presumably derived karyotypes not only occurs but also prevails and may be considered modal (see Table 2 and Fig. 1). Within Anisoptera, such families are Aeshnidae (2n = 26A + X) and Gomphidae (2n = 22A + X), whereas within Zygoptera, these are Chlorocyphidae (2n = 22A + X) and Coenagrionidae (2n = 26A + X). Thus, Odonata, despite the fact that they have holokinetic chromosomes (Nokkala et al. 2002), demonstrate rather high karyotypic stability, with most species showing 2n = 25 (found in 60% of studied species), 2n = 27(21%) and 2n = 23(13%) which may point to some selective constraints acting to stabilize chromosome number in their evolution (Kuznetsova et al. 2020b).

There are the species for which different authors give various karyotypes that are sometimes difficult to interpret (see Table 1). In some cases, this might be due to misidentifications of a particular species or an error in determining the karyotype. For example, Wolfe (1953) reported 2n = 17(16A + X) for males of Uropetala carovei (Petaluridae, Anisoptera) from New Zealand. However, according to later studies of this species in the same locality (Jensen and Mahanty 1978; Jensen 1980), it has 2n = 25(24A + X), and Jensen (1980) therefore considers the Wolfe data as erroneous. We cannot exclude, however, that the above authors studied different U. carovei subspecies, U. c. carovei White, 1846 and U. c. chiltoni Tillyard, 1921, that may indeed have different karyotypes. In other cases, the chromosome number difference between geographic populations might be indicative of the inter-population variation within the bounds of one taxonomic species or even the existence of a species complex with several morphologically cryptic species. For example, 4 of the 17 studied species of the dragonfly genus Aeshna Fabricius, 1775 were reported to have different karyotypes in different populations. These are: Aeshna grandis – 2n = 26A + X (former USSR), 2n = 24A + X (former USSR, Finland), and 2n = 24A + neo-XY (Netherlands, Finland); A. isoceles – 2n = 26A + X (USA) and 2n = 24A + X (Russia); A. juncea – 2n = 26A + X (Italy) and 2n = 24A + neo-XY (Finland, former USSR, Italy); A. mixta – 2n = 26A + X (Netherlands) and 2n = 24A + X (India) (Table 1). In all such cases, special studies involving a combined analysis of karyotypes, morphology, distribution patterns and molecular markers are needed.

Approximately 80% of Odonata species have a pair of very small chromosomes, i.e. microchromosomes or m-chromosomes (Mola 2007, Table 1). A number of speculations have been forwarded to explain the origin of these chromosomes in Odonata. Kiauta (1968e) suggested m-chromosomes to be fragments of “normal” chromosomes, whereas Oguma (1930) considered them the remnants of an autosome pair in the process of its elimination by progressive loss of chromatin. The size of the smaller chromosome pair was shown to be variable within different species (Kiauta 1968e; see Mola 2007 for other references) which is consistent with both hypotheses. Closely related species and different populations of the same species often differ from each other in the presence/absence of m-chromosomes (Table 1). This is most likely due to the lack of clear criteria for the identification of a small chromosome pair as m-chromosomes in a particular karyotype (Mola 2007; Kuznetsova et al. 2020b).

Most cytogenetic studies of Odonata have been made only to determine the chromosome number and sex chromosome mechanism for which the routine staining was used. Although a considerable amount of such data was obtained (Table 1, 2), standard karyotypes of many Odonata taxa remain totally unknown (Fig. 1). Lack of data on more “primitive” families of Zygoptera (e.g. Hemiphlebiidae) and Anisoptera (e.g. Austropetaliidae and Neopetaliidae) makes difficult understanding karyotype evolution of the order in general.

During the last decades, karyotypes of a few dozen Odonata species were studied using various techniques of differential staining of chromosomes such as C-banding, AgNOR-staining and DNA specific fluorochrome banding visualiszing constitutive heterochromatin, nucleolus organizing regions (NORs) and AT- and GC-rich chromosome segments, respectively. Such data can be found in the following publications: Thomas and Prasad (1986), Prasad and Thomas (1992), Perepelov et al. (1998), Perepelov and Bugrov (2001a, b, 2002), Grozeva and Marinov (2007), De Gennaro et al. (2008), Walia et al. (2011, 2018), Walia and Chahal (2014, 2018), Walia and Devi (2018), Walia and Katnoria (2018), Walia and Devi (2020a, b). Unfortunately, these data alone did not shed much light on the karyotypic evolution of Odonata.

Although the classical cytological techniques remain necessary starting points for cytogenetic studies of Odonata to get an overview of their genomes, the future of Odonata cytogenetics must be coupled with the application of new cytogenetic molecular techniques that enable the localization of specific DNA sequences in chromosomes and the identification of individual chromosomes in karyotypes. In the article by Frydrychová et al. (2004) and, on a larger scale, in two of our recent publications (Kuznetsova et al. 2018, 2020b), the fluorescence in situ hybridization (FISH) technique was used for the first time for analyzing Odonata karyotypes. Several species belonging to the Anisoptera (from the families Aeshnidae, Libellulidae, and Corduliidae) and the Zygoptera (from the families Coenagrionidae and Calopterygidae) were studied regarding the occurrence of the TTAGG telomeric repeats and the distribution of the 18S rRNA genes in their karyotypes. The TTAGG repeats proved to be the canonical motif of telomeres in the class Insecta in general, which, however, was repeatedly lost in the evolution of different phylogenetic lineages (Kuznetsova et al. 2020a). It was shown in the listed Odonata publications that the (TTAGG)n motif does not occur in all but one (Sympetrum vulgatum) species, and the 18S is located on one of the largest pairs of autosomes in all studied dragonfly species but on m-chromosomes in all studied damselfly species (Kuznetsova et al. 2020b).

The results obtained showed great promise of the combined use of FISH and classical and banding cytogenetics in order to identify new chromosomal markers, reveal differences between species, particularly when they share the same or very close karyotypes, and speculate about the mechanisms involved in the karyotype evolution of Odonata (Kuznetsova et al. 2020b). Another promising line of future research could be to test hypotheses (Mola and Papeschi 1994; Ardila-Garcia and Gregory 2009) about whether there is a relationship between karyotype evolution and genome size diversity in the Odonata or there is no such relationship.

Acknowledgements

The present study was performed within the research project no. AAAA-A19-119020790106-0. The authors deeply grateful to the late Evgeny A. Perepelov whose thesis “Karyotype evolution of Odonata (Insecta) of Northern Palearctics” (2003; Novosibirsk State University, Novosibirsk, Russia) served as a starting point for preparing Table 1 of our paper. We thank two reviewers, Dr. M. Marinov and Dr. S. Grozeva, for their useful remarks to a draft of this MS. Special thanks to Dr. M. Marinov for his nomenclatural and taxonomic corrections and updates.

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