Odonatologica I (2) 73-102 June I. 1972
Synopsis of the main cytotaxonomic data
in the order Odonata
B. Kiauta
Institute of Genetics, University of Utrecht, Utrecht, The Netherlands
Received April 13, 1972
So far 371 species and subspecies of 20 families of the three living suborders have studied been cylologically. this figure represents more than 6% of the total number of species described, making dragonflies cylologically by far the
best studied order of insects, A review is given of the main data on the
chromosomal cytology, viz. haploid chromosome number, general morphology
and origin of the karyotype, distinctions, if any, between geographic popula- chromosome tions, peculiarities in behaviour and mode of sex determination
of all In biblio- species and subspecies hitherto studied. addition, a complete graphy related to the subject is presented.
INTRODUCTION
Since several years the author is working on a monograph on the cytotaxono-
and of my cytogenetics the Order, which will include also a detailedannotated
of catalogue the chromosomal cytology of dragonflies. It may take some time before the will therefore useful work appear in print, it seems to give here a
of the data the chromosomal of and synopsis main on cytology the 371 species
far studied and subspecies so to present a complete bibliography related to the subject.
Because of the limited the information listed had be space available, to re- stricted to the most essential data, viz. locality, haploid chromosome number, occurrence of w-chromosomes, general morphology and origin of the karyotype
if (if peculiar), distinctions, any, between geographic populations, peculiarities in
chromosome behaviour, mode of sex determination(if other than the usual XO type) and reference to the source publication.
73 The listed under valid species are taxonomic names, in alphabetic sequence within of the a genus, regardless synonym under which originally cytotaxono- mically described.
For the sake of convenience the system followed is that outlined by FRASER
reclassification (1957, A of the order Odonata, R. zool. Soc. N.S.W., Sydney),
not though this does imply that the author agrees with all of his suggestions.
Several of his families and subfamilies are certainly not tenable, e.g. Macrodiplac-
subdivision of Petaluridae — tidae, Sympecmatinae, etc. to mentionjust some of them.
REPRESENTATIVENESS OF THE CYTOLOGICALLY STUDIED MATERIAL
The available sample covers 20 families of the three living suborders and represents more than 6% of the total number of species described. Though this figure makes dragonflies cytologically by far the best studied insect order, the cytotaxonomic record does not include several phylogenetically important families.
In Fraser’s Figure 1 genealogical tree has been used to present graphically the
affiliation of the material hitherto studied family (in figure family names
first framed). At a glance at least two important lacunae in the cytotaxonomic record are evident:
The (1) superfamily Hemiphlebioidea, with a single family Hemiphlebiidae, con-
taining a monotypic genus Hemiphlebia (species H. mirabilis Selys, from
Gouldburn River, Victoria, Australia) is considered, on the basis of venatio-
nal characters, as a transitional form between the Coenagrionidea on one
hand, and the extinct suborder Protozygoptera on the other. It could, there-
furnish fore, perhaps some evidence on the cytological conditions during the
earliest history of Zygoptera.
(2) In view of the peculiar cytotaxonomic situation found in the only hitherto of studied member the primitive Pseudolestidae, it would he interesting to
information gather some on the karyotypic condition in the Amphipterygi-
which the of dae, occupy a key position at base the higher Calopterygoidea
and in their direct derivatives, viz. Heliocharitidae and Chlorocyphidae.
Aside from the fact that in several families too little material has been examined to of permit any generalisations, we are the opinion that cytotaxono- mic data the and on Hemiphlebiidae Amphipterygidae are urgently needed be- fore more far reaching cytophylogenetic considerations on the Order can be
discussed.
74 Fig. 1. Cytologically studied (framed) and unknown (unframed) odonate families, placed in Fraser’s genealogical tree.
CHROMOSOME NUMBER
The chromosome number has been determined in all species studied; for 20 of these more than one number has been reported. The variation occurs either in
different of different numbers cells the same specimen or chromosome are pecu- liar to different geographic populations. The forms with numeric variation in the complement were omitted from the graphs in Figure 2.
75 Fig. 2. Histograms of haploid chromosome numbers in the order and in the three living suborders (in %of total number of species studied). The width of columns is proportional to the absolute number of cytotaxonomically examined species within the order and sub- orders.
76 Haploid numbers range from 3 to 15 (cf. Fig. 2). In spite of this rather broad range there is, in general, but little numericalvariation in the dragonfly comple- ments. The haploid numbers 12, 13 and 14 are represented in nearly 93.8% of the Odonata of examined; n= 13 can be considered as the type number the
Order. It is found in 55.9% of the species studied cytologically.
On the suborder level the pattern is essentially similar in Anisoptera. The haploid number, n = 13, is represented in 62.2% of the sample. The same type number can be assumed also for Anisozygoptera, since it was found in the single cytologically studied representative of the suborder, that consists of two living species only. The situation seems more complicated in Zygoptera. The haploid number, n = 13, was found only in 39.4%of cases studied, whereas 14 elements occur the of 53.6% of them. would from this in haploid sets It appear, evidence,
14 that n = should be regarded as the type number of the suborder, if it were not for the fact that 14 elements are peculiar only to the familiesProtoneuridaeand
Coenagrionidae and that the latter covers more than 51.5% of the total number of the representatives of the suborder hitherto examined cytologically. This
is being so, it is clear that the studied sample not representative for the suborder and that for the be Zygoptera too, the haploid number, n = 13, can more safely assumed as the true type number.
The family type numbers are as follows: n = 9 (Pseudolestidae [? ], Petaluri-
= dae), n 12 (Polythoridae, Gomphidae), n = 13 (Platystictidae [? ],Platycnemi- didae, Megapodagrionidae, Lestidae, Epallagidae, Hetaerinidae, Calopterygidae,
Epiophlebiidae, Cordulegasteridae, Corduliidae, Macrodiplactidae, Libellulidae),
and = n 14 (.Protoneuridae, Coenagrionidae, Aeshnidae).
Some genera deviate distinctly from the prevailing chromosome number pat- tern ofthe families they belong to, notably Leptagrion (n = 15) in Coenagrioni-
dae, Mecistogaster (n = 3 and 15) in Pseudostigmatidae, Trigomphus (n = 10 and
12) in Gomphidae, Tetragoneuria (n = 11 and 14) in Corduliidaeand Orthemis
= Libellulidae. of these of (n 12) in Some complements, however, are secondary origin and only point to the genetic unstability of the genera involved.
In the cytologically studied material with uniform chromosome numbers
the 79.8% of throughout species range the taxa have a number that is equal to or
higher than the type number of the Order. If the distribution is analyzed from the point of view of the geological age of the families concerned, a significant shift can be observed. A tendency towards lower chromosome numbers is clearly
in in apparent the geologically old families, whereas the younger groups numeri- cally higher complements prevail. Thus, Epallagidae. Gomphidae, Petaluridae,
Aeshnidae and Cordulegasteridae are the only families of our sample the repre- sentatives of which are known already from the Mesozoic beds. In these only
77 some 10.5% of species studied possess chromosome numbers equal to or higher than the present type number of the Order.
Summarizing the above information, it can be concluded that the chromo-
affinities some numbers do reflect, to some extent, the taxonomic and phyloge- netic character of some superspecific taxa in dragonflies.
PRIMARY AND SECONDARY COMPLEMENTS
Karyotypes of dragonflies can be divided roughly in two groups: (l)the
“normal”, high-n complements (n = 9 to 14). and (2) the low-n complements
(n = 3 to 8). Chromosome size in the high-n species is approximately halfthat in
the low-n forms.
In the between the high-n species a parallel is apparent increase in specializa- tion and advancement on one hand, and the increase of the chromosome number
found on the other. The low-n species can be in any systematic group, of any
number. has type It been demonstrated in numerous cases, that the low-n com-
plement arises by fusion of some or all elements of the original complement,
hence the nomenclature “primary” and “secondary” complements.
Aside from of fusions, in several species fragmentation one or more elements of of chromosome the original set has taken place, resulting in an increase the
number. These complements too, are called “secondary” and may occur in single
of species any taxonomic group.
THE m-CHROMOSOMES
The m-chromosomes (= micro-chromosomes) represent a peculiar feature of
dragonfly cytotaxonomy. They can be found in any species of any family,
in though some groups they appear to be scarcer than in others. Thus, they are
all of lacking in species the genera Argia and Ophiogomphus, in all primary
complements of Enallagma and Megapodagrion and in most representatives of
the subfamilies Coenagrioninae, Ischnurinae and Gomphoidinae, whereas in
many genera they were found in all species studied.
Contrary to the chromosome numbers, the distribution of m-chromosomes
does not reflect any taxonomic affinities, save perhaps for a few cases in the
above mentioned taxa.
On the geographic population level, on the other hand, the w-chromosomes
feature; be and may represent a peculiar they may present in one population
78 lacking in the other. This condition has been observed in the following species:
Lestes dryas, Gomphus exilis, Anax junius, Cordulegaster boltoni, Macromia
magnifica, Pachydiplax longipennis, Leucorrhinia hudsonica, L. intacta and
Brechmorrhoga mendax. Also the relative size of the m-elements may be dif- ferent different different forms of in populations or in infraspecific the same species, e.g. in Calopteryx virgo, Libellula quadrimaculata etc.
SEX CHROMOSOMES AND SEX DETERMINATION
The original mode of sex determinationin dragonflies is XO/XX, the male being the heterogametic sex. It is observed in all primary complements, regard- less of the chromosome number viz. the degree of advancement and specializa- tion achieved by the taxa concerned.
In secondary complements a neo XY-sex determination occurs in those cases where the involved fusion original X was in a with an autosome. Its occurrence is related to to neither any taxonomic groups nor phylogeny, though in some families(e.g. Aeshnidae) it is found more frequently than in others.
The original sex chromosome is usually one of the smallest elements of the karyotype, in some cases it is medium-sized, while it is the largest in a few species other than Gomphidae. It has been argued that the present type number of the latter family is of secondary rather than of primary origin and that the
X-es with fusions large met in many gomphidan dragonflies arose in complicated with autosomes (cf. KIAUTA, 1969 a), but the suggestion should be understood as a working hypothesis only.
It is a matter of course that the neo-X element is at least medium sized, ifit is not the largest of the complement. The unfused homologue of the autosomal
of part of the neo-sex element is usually small, it has the function a neo-Yand is
confined to the maleline.
REVIEW OF THE MAIN CYTOTAXONOMIC DATA IN THE ORDER
The following explanations on the tabulation of the data are considered
useful:
(1) As a rule the male haploid numbers are indicated. If a number was derived
from brackets. Where female a mitotic figure, it is given in only comple-
ments were described, the sex is indicated in brackets, preceding the num-
ber. If for a species several numbers are listed, those connected by “+”
and while those appear in one the same specimen, separated by were
reported for different specimens or populations. 79 (2) The presence of m-chromosomes is indicated in brackets behind the chromo-
some number. In the latter, however, the m-chromosomes are also included.
(3) The remarks are derived both from the text and from the illustrations of the
score publication.
(4) If the name of an author in brackets behind the listed appears synonym as under “Remarks” and is printed in italics, it refers to the author of the cyto-
logical publication where the synonym was used, and not to the author of
the synonym.
SYNOPSIS
FAMILY Locality dn (and Remarks References
Subfamily presencepresenceof
Genus, Species. Subspecies m)m )
PLATYSTICTIDAE
Palaemnematinae
PalaemnemaPalaemnemapaulina Costa Rica 1313(m)(m) GUMMING,GUMMING, 1964a
(Drury. 1773)
PROTONEURIDAE
Protoneurinae
Epipleoneurasp. Bolivia 14 GUMMING, 1964a
NeoneuraNeoneurarubriventrisrubriventris Bolivia 14 (m) GUMMING, 1964a
(Selys, 1860)
PLATYCNEMIDIDAE
Platycnemidinae
Copera annulateannulate Japan, India 13 (m) subPlatycnemisPlatycnemis K1CHIJO,K1CHIJO, 1941,1941, 1942a,1942a. (Selys,(Selys. 1863)1863) 1942c;
DASGUPTA, 1957
Platycnemis pennipes Finland, Italy 13 in ddoccasionally OKSOKSALA,ALA, 1945;
(Pallas,(Pallas, 1771) twotwo chiasmata inin KIAUTA,KIAUTA, 1971b
one ofofthe one longer
bivalentsbivalents
COENAGRIONIDAE
AmphicnemininaeAmphicnemininae
Diceratobasismacrogastermacrogaster Jamaica 1414(m) GUMMING, 1964a
(Selys,(Selys, 1875)
PseudagrioninaePseudagrioninae
Ceriagrioncerinorubellumcerinorubellum India 1414(m) DASGUPTA,DASGUPTA. 19571957
(Brauer,(Brauer, 1866)
C. RAY CHAUDHURICHAUDHURI C coromandelianum India 1414((m)m ) (Fabricius,(Fabridus, 1798) && DAS GUPTA. 1949; SRIVASTAVASRIVASTAVA& DAS,
1953;DAS,DAS, 1956
C.C fallaxfallaxRis,Ris, 1914 BanglaBangla DeshDesh 1414((m)m ) DASGUPTA,DASGUPTA. 19571957
C.C. rubiae Laidlaw, 1916 IndiaIndia 1414(m)(m) ASANA & MAKING, 1935;1935; MAKING, 1935;
KIGH1JO,KICH1JO, 1942e
CC. tenellum tenellum ItalyItaly 1414 (m) KIAUTA,KIAUTA. 1971b
(Villers, 1789)
MegalagrionMêgalagrion oahuense HawaiiHawaii 1414 (/n)(m) KIAUTA,KIAUTA. 1969d
(Blackburn, 1884)
Pseudagrion australasiae India 1414 (m) subP.P. bengalensebengalense DASGUPTA,DASGUPTA. 1957 Selys,Selys. 18761876 LaidlawLaidlaw
80 FAMILY Locality tindn (and RemarksRemarks References
SubfamilySubfamily presenceofm)m)
Genus,Genus, Species, Subspecies
P. decorum(Rambur,(Rambur, 1842) India 14 ((m)m ) DASGUPTA, 1957
P.P. microcephalum (Rambur, 1842)India1842)lndia 14 (m) DASGUPTA, 1957
P. rubriceps Selys, 18761876 India 14 (m) DASGUPTA,DASGUPTA. 1957
P. spencei Fraser, 1922 India 14 (m) DASGUPTA, 1957
Coenagrioninae
CercionUndentlindeni(Selys,(Selys, 1840) Italy 14 (m) KIAUTA, 19711971bb
Chromagrion conditumconditurn U.S.A. 14 CRUDEN, 1968 (Hagen,(Hagen, 1876)1876)
armatumturn CoenagrionCoenagrionarma Finland, 14 sub Agrion OKSALA, 19391939a;a;
(Charpentier,(Charpentier, 1840) U.S.S.R. MAKALOWSKAJA,MAKALOWSKAJA, 1940
C. hastulatumhastulatum U.S.S.R. 14 tabsubAgrion MAKALOWSKAJA, 1940 (Charpentier,(Charpentier, 1825)
C.C hieroglyphicum Japan 14 (m) subAgrion KICHIJO,KICHIJO, 1941, (Brauer.(Brauer, 1865) 19421942a.a, 19421942c c
C.C pulchellumpulehelium U.S.S.R. 14 sub.sub Agrion MAKALOWSKAJA,MAKALOWSKAJA. 1940;
(Vander(Vander Linden, 1823) Netherlands (Makalowskaja) KIAUTA,KIAUTA. 19691969aa
C. resolutumresobitum (Hagen, 1876) U.S.A. 14 CRUDEN, 1968
Coenagrion sp. Japan 14 (m) subAgrion KICHIJO,KICHIJO. 1941, 19421942a,a, 19421942c c
Erythromma najas Finland, 14 OKSALA, 19391939a;a;
(Hansemann,(Hansemann, 1823) U.S.S.R., MAKALOWSKAJA,MAKALOWSKAJA. 1940; Netherlands KIAUTA,KIAUTA, 19691969a a
Nehalenniaireneirene (Hagen, 1861) U.S.A. 14 CRUDEN. 1968)
N. speciosa Finland (5)(9l 14 (m ?)? ) OKSALA, 1945 (Charpentier,(Charpentier, 1840)
Pyrrhosoma nymphula Finland (9)14 OKSALA,OKSALA. 1945 (Sulzer,(Sulzer, 1776)
Ischnurinac
Acanthagrion ascendens Bolivia 14 (m) GUMMING, 19641964aa Calvert, 1909
A. chacoenseCalvert, 1909 Bolivia 14 (m) GUMMING, 19641964aa
Aeolagrion foliaceumfolia ceurn Bolivia 14 GUMMING, 19641964aa (Sjostedt,(Sjöstedt, 1918)
Amphiagrion abbreviatum U.S.A. 14 CRUDEN,CRUDEN. 1968 (Selys.(Selys. 1876)
Enallagma aspersum U.S.A. 14 CRUDEN,CRUDEN. 1968 (Hagen,(Hagen, 1861)
E. borealeSelys,Selys, 1875 U.S.A. 14 CRUDEN,GRUDEN, 1968
E. carunculatum U.S.A. 14 CRUDEN.GRUDEN, 1968 Morse,Morse. 1891895 S
E. civile (Hagen, 1861) U.S.A. 14 CRUDEN, 1968
E. 14+15 of cyathigerum Finland, 14; (m); fragmentation one OKSALA, 19391939a,a, 1945; (Charpentier, 1840)1840) U.S.S.R. 15 (m)+l4;(m)+14; 15 ofthethesmaller MAKALOWSKAJA, 1940;
Netherlands, (m) bivalentsin some or VAN BRINK & KIAUTA,
U.S.A. in all cells of some 1964;CRUDEN, 1968;
populations (Netherlands,(Netherlands, KIAUTA,KIAUTA. 19691969a,a, 19691969b b U.S.A.)
81 FAMILY Locality (5ndn(and(and RemarksRemarks References
of Subfamily presence m)m ) Genus, Species, Subspecies
E. ebrium(Hagen, 1861) U.S.A. 14 CRUDEN, 1968
E. praevarum (Hagen, 1861) U.S.A. 1414 CRUDEN, 1968
Ischnura cervula U.S.A. 14 CRUDEN,GRUDEN, 1968
Selys, 1876
I. denticollis U.S.A. 14 CRUDEN,GRUDEN, 1968 (Burmeister, 1839)
I. elegans Finland, 14 at 6ddiakinesis OKSALA,19391939a,a, 1945
(VanderLinden,Linden. 1823) Netherlands X occasionally KIAUTA, 19691969a a
negatively allocyclic
(,(Kiauta)Kiauta)
I. fluviatilis Bolivia 14 CUMMING,GUMMING, 19641964aa
Selys. 1876
I. perparvaperparvaSelys.Selys, 1876 U.S.A. 14 CRUDEN.GRUDEN, 1968
I./. senegalensissenegalensis Japan.Japan, India, 14 (m) KICHIJO, 1941,19421941,1942a.a, (Rambur,(Ram bur, 1842) Ethiopia 19421942c;c; DASGUPTA, 1957;
KIAUTA,KIAUTA. 19691969c c
I. verticalis (Say.(Say, 1839) U.S.A. 14 CRUDEN.GRUDEN. 1968
I./. cf. ultima Ris, 1908 Bolivia 14 CUMMING.GUMMING, 19641964a a
Leptagrion macrurummacrurum Brasil (15)(15) neo-XY KIAUTA,KIAUTA. 1971c,1972
(Burmeister, 1839)
Tigriagrion aurantinigrum Bolivia 14 CUMMING.GUMMING, 19641964aa Calvert, 1909
Zoniagrion exclamationis U.S.A. 14 CRUDEN,CRUDEN. 1968 (Selys, 1876)
Agriocneminae CeraturaCentura capreola Bolivia 14 CUMMING.GUMMING, 19641964aa
(Hagen, 1861)
Mortonagrion selenion Japan 14 (m) subAgriocnemis KICHIJO, 1941,19421941,1942a,a, (Ris, 1916)1916) 1942c,1941942c,1942e2e
Argiinae Argia funebris Mexico (9)(9)(14)(14) KIAUTA,KIAUTA.
(Hagen, 1861) unpublished
A. sedula (Hagen, 1861) U.S.A. 14 CUMMING,GUMMING, 1964a1964a
A. violaceaviolaceaif( Hagen,Hagen, 1861) U.S.A. 14 CRUDEN, 1968
A. vivida Hagen, 1865 U.S.A. 14 CRUDEN, 1968
PSEUDOSTIGMATIDAEPSEUDOST1GMATIDAE
Pseudostigmatinae
MecistogasterMecistogaster sp. 1 Bolivia 15 (m) XO; fragmentation CUMMING,GUMMING, 19641964a;a;
of two bivalents KIAUTA, 19691969a a
Mecistogaster sp.2 Bolivia 6 neo-XY (?);(? ); fusion CUMMING,GUMMING, 19641964a;a;
of all elements KIAUTA, 19691969a a
MEGAPODAGRIONIDAEMEG APODAGRIONI DAE
Megapodagrioninae
13 of Megapodagrion contortumcon torturn Brasil 13(m);13(/n); 13(m)(m) fragmentation ofa KIAUTA,
(Selys, 1862) +l4+14 (m) bivalentin some figures unpublished
82 dn FAMILY Locality <5n (and Remarks References
Subfamily presenceofm)
Genus, Species, Subspecies
M. macropus Bolivia 13 X largest of the GUMMING, 1964a
(Selys, 1862) set a 6d
metaphase I
M. setigerum Bolivia 13 GUMMING, 1964a Selys, 1886
Argiolestinae
Heteragrion flavidorsum Bolivia 13 GUMMING, 1964a Calvert, 1909
H. inca Calvert, 1909 Bolivia 13(m) GUMMING, 1964a
Philogeniacarrilica Costa Rica 13(m) GUMMING, 1964a Calvert, 1907
LEST1DAE
Sympecmatinae
Sympecma fusca Japan 13(m) subSympycna;Sympycrur, KICHIJO, 1941,1942a,
(Vander Linden, 1823) one bivalent extra 1942c
large
Lestinae
Chalcolestesviridis Netherlands 13(m) KIAUTA, 1969a (Vander Linden, 1825)
Lestes congener U.S.A. one bivalent extra 1968 13(m) one CRUDEN,
Hagen, 1861 large; X minute
L. disjunctus U.S.A. 13 one bivalent extra CRUDEN, 1968
Selys, 1862 large
L. 1890 U.S.A. some dryas Kirby, 13; 13(m) m present in some CRUDEN, 1968
populations; one
bivalentextra large
LL. forcipatus U.S.A. 11 two extra large bivalents, CRUDEN, 1968
Rambur, 1842 probably due to fusion; XO
LL. forficula Rambur, 18421842 Jamaica 13(m) GUMMING, 1964a
L.L. rectangularis Say, 1839 U.S.A. 13(m) one bivalentextra CRUDEN,CRUDEN. 1968
large
LL. simulatrix McLachlan, 1895 Madagascar 13(m) bivalents ofgradually KIAUTA, 1968f; decreasing magnitude 1969c
L.L. sponsa (Hansemann, 1823) U.S.S.R., Japan 13; 13(m) in Japanese material one MAKALOWSKAJA, 1940;
extra large bivalent KICHIJO, 1941,1941,1942a,1942a.
and m, both absent 1942c, 1942e
in the Russian population
L.L. stultus 1861 U.S.A. extra stultusHagen, 1861 U.S.A. 13(/n)13(m) one bivalent CRUDEN, 1968 large
L.L. vidua Hagen, 1861 U.S.A. 13(m) GUMMING, 1964a
L.L. virens vestalis Netherlands 13(m) KIAUTA, 1969a1969a
Rambur, 1842
PSEUDOLEST1DAE
Pseudolestinae
Hypolestes clara Jamaica 9 GUMMING, 1964a (Calvert, 1891)1891)
POLYTHORIDAE
Polythorinae Corairene Ris, 1918 BoliviaBolivia 12(m) GUMMING, 1964a
Polythore bolivianaboliviano BoliviaBolivia 1212(m)(m) GUMMING, 1964a
(McLachlan, 1878)
83 FAMILY FAMILY Locality <5n(and<5n (and Remarks References
Subfamily presence of m) Genus. Species. Subspecies
EPALLAG1DAE
Epallaginae
Epallaget'pal läge fatime Greece 13 KIAUTA, 1970b (Charpentier, 1840)
HETAER1N1DAE
Hetaerininae
Hetaerina americana U.S.A. 13(m) in a publishedfigure GUMMING, 1964a; (Fabricius, 1798) two bivalentsextremely CRUDEN, 1968 minute(([Cruden)Cruden)
H. chorcachoreaCalvert.Calvert. 1909 Bolivia 13(m) GUMMING, 1964a
H. rosea 1853 Bolivia Selys, 1414(m)(m) the increase in n GUMMING, 1964a;
probably due to KIAUTA, 1969b fragmentation
H. sanguinea Selys, 1853 Bolivia 13 GUMMING, 1964a
H. titia (Drury.(Drury, 1773) Bolivia 13(m) GUMMING, 1964a
H.H tricolor ((Burmeister,Burmeister, 1839) Mexico 13(m) KIAUTA,KIAUTA. 1970c
H. vulnerata (Selys, 1853 Mexico (9) (13) (m?)(m? ) KIAUTA, 1970c
CALOPTERYGIDAE
CalopterygjnaeCalopteryginae
Anaciagrion corneliaCornelia Japan I3(m)13(m) sub Calopteryx OGUMA, 1930;
(Selys,(Selys. 1853) K1CHIJO.K1CHIJO, 1942e
Calopteryx aequabile U.S.A. 13(m) subAgrion CRUDEN. 1968 Say,1839
C. atrata 1853 C Selys, 1853 Japan 13(m) OGUMA. 1930;
KICHIJO, 1942e;
OMURA, 1957
C.C maculata U.S.A. 13(m) subAgrion maculatum GUMMING.GUMMING, 1964a; (Beauvois.(Beauvois, 1805) ( Cruden)Cruden ) CRUDEN. 1968
C. splendens caprai Italy 13(m) at ddiplotene at KIAUTA. 1971b
Conci, 1956 least four bivalents with
two chiasmata; high re-
combinationindex is in
accordancewith high morphologicalvariability
C.C splendens splendens U.S.S.R., splendens U.S.S.R., 13 MAKALOWSKAJA, 1940; (Harris, 1782) Finland, OKSOKSALA,ALA. 1945;
Germany KIAUTA, 1969a, 1971b
C. virgo japonica Japan 13(m) m minute KICHIJO, 1942e;I942e; SeJys.Selys, !8691869 HIRAI, 1956; OMURA.OMURA, 1957;
KIAUTA. 1968b, 1968c
CC. meridionalis of bivalent 1971a virgo Spain 13(m); 13(m) fragmentation a bivalent KIAUTA,KIAUTA.
1873 Selys, +14(m) in some figures; often precocious segregation of
several bivalents
C. virgo padana Slovenia(Yugo- 13(m) sub C. virgo KIAUTA, 1967a.1967a, 1968b, _ _ Conci, 1956 Conci, 1956 slavia), Austria (1967a) 1968c)
C. 13; 13(m) virgo virgo Belgium, Finland, occasionally precocious CARNOY, 1885(of historic
(Linnaeus, 1758) U.S.S.R., GermanyGermany segregation of some biva- value only); OKSALA, 1939a;
Luxembourg.Luxembourg, lents (Makalowskaja ) MAKALOWSKAJA, 1940; Netherlands KIAUTA, 1968b, 1968c,
unpublished
Matrona basilaris Taiwan 13 KIAUTA,KIAUTA. 1968f
Selys.Selys, 1853
84 FAMILY Locality dn(anddn (and RemarksRemarks ReferencesReferences
Subfamily presence ofm)m) Genus, Species, Subspecies
Mnais costalis Japan 13(m) absolutemagnitudeof all OGUMA, 1930;
Selys, 1869 elements andrelativerelative size KICHIJO,KICH1JO, 1942e
of m smaller thanin m smaller thanin
M.M. strigata
M. strigalastrigata Selys, 1853 Japan 13(m) absolutemagnitudeof all OGUMA, 1930;
elements andrelative size KICHIJO, 1942e;1942c;
of m thanin 1957 m bigger OMURA,
M. costalis
EPIOPHLEBIIDAE
Epiophlebiinae
Epiophlebia superstessuperstes Japan 13 OGUMA, 1951 Selys, 1889
GOMPHIDAE
Gomphinae
Anisogomphus bivittatus India 1212(m)(m) X medium sized at DAS, 1956 (Selys,(Selys. 1854) metaphase II
Davidius nanus Japan 12 sub Gomphuskakiensis;hakiensis; KICHIJO, 1939,
(Selys.(Selys, 1869) no extralarge element; 1942e X smallest at metaphase I
Dromogomphus spinosus U.S.A. 12(m) CRUDEN, 1968
(Selys.(Selys, 1854)
extra D. spoliatus (Hagen, 1857) U.S.A. 12(m)12(m) no large element CRUDEN, 1968
ErpelogomphusErpetogomphus designatusdésignants U.S.A. 12(/n)12(m) GUMMING, 1964a
Hagen,Hagen, 1857
E. diadophisCalvert, 1905 U.S.A. 12 GUMMING, 1964a
E. ophibolus Calvert, 1905 Mexico 12(m) X most voluminous at KIAUTA, 1970c metaphase I1
extra GomphusGomphus confraternus U.S.A. 12(m) no extra large element CRUDEN, 1968
Selys,Selys, 1873
of G.G. exilis Selys, 1854 U.S.A., 12(m); 12 m present in one the CRUDEN, 1968;
Canada twoU.S.A. populations; in KIAUTA, 1969a
some oogonialfiguresfigures of
Canadianmaterial two smaller
elementsattachedto two longest
chromosomes (KiautaKiauta))
at G. graslini France 12(m) X largest at all stages;stages; KIAUTA,KIAUTA. 1968d, 1969a Rambur, 1842 nco-X/neo-nco-Y (? )
G. lentulus U.S.A. 12 CRUDEN,CRUDEN. 1968 Needham, 1902
G. lividus Selys, 1854 U.S.A. 12(m) no extra large elementclement CRUDEN, 1968
G. melaenops Selys, 1854 Japan 12(m) for identityof the species TOYOSHIMA &&. HIRAI,HIRAI. 1953; listed by Toyoshima & HIRAI,HIRAI. 1956;
HiraiHind and Hirai cf. OMURA, 1957
CRUDEN,CRUDEN. 1968 andand KIAUTA, 1969a
G. militarismilitons Hagen, 1857 U.S.A.U.S.A. 12 CRUDEN, 1968
G. palliduspalliJus Rambur,Rambur, 1842 U.S.A.U.S.A- 12 GUMMING, 1964a
G. plagiatusplagiants Selys.Selys, 1854 U.S.A.U.S.A. 12(m) CRUDEN,CRUDEN. 1968
G. postocularis Selys, 1869 JapanJapan 12(m?)12(m? ) OMURA, 1957
G. scudderi Selys, 1873 U.S.A.U.S.A. 12 CRUDEN, 1968
85 (5n FAMILY Locality dn(and(and Remarks References
Subfamily presence ofm)ff»)
Genus,Genus.Species. Subspecies
G. spicatusSelys.Selys, 18541854 U.S.A. 1212(m)(m) CRUDEN, 1968
G. submedianus U.S.A. 12 CRUDEN, 1968 Williamson, 1914
Nihonogomphus viridisviridis Japan 12(m? ) OMURA, 1957
Oguma, 1926
extra Octogomphus specularis U.S.A. 12 nono largelarge element CRUDEN, 1968 (Hagen, 1859)
Onychogomphus forcipatus Finland, 12-1-1312+13 in Austrian material in the OKSALA, 1945;
(Linnaeus, 1758) Austria same specimen two different KIAUTA, 1969a complements; XO
in n= 13; nco-XY
in nco-Xneo-X nn= 12; longest;
X smallest; kinetic behaviour
ofneo-Xneo-Xnormal
Ophiogomphusbisonbison U.S.A. 12+13;12+13;13+1213+12 one element extra large CRUDEN, 1968
1873 in variation in Selys, n= 13;
toto n due fragmentation
extra element O. colubrinusSelys, 1854 U.S.A. 12 no large element CRUDEN,CRUDEN. 1968
O.0. occidentalis Hagen, 1882 U.S.A. 12 no extra large element CRUDEN.CRUDEN, 1968
O. one extra 1968 rupinsulensis (Walsh, 1862) U.S.A. 12 one large element CRUDEN,
O. serpentinus Finland (9)(9)1212 X largest OKSALA,OKSALA. 1945 (Charpentier, 1825)
suzukii sub no extraextra 1930; Stylogomphus Japan 12(m) Gomphus;Gomphus ; no OGUMA,OGUMA. 1930;
(Matsumura, 1926) largelarge element;element;XX second KICHIJO, 1942c
smallest at metaphasc I1
tabei one element TOYOSHIMA & TrigomphusTrigomphus citimuscilimus tabei Japan ll(m) subsub GomphuGomphuss\; element TOYOSHIMA A HIRAI,HIRAI.
extra Asahina,Asahina, 1949 extra large; X marked in 1953; HIRAI, 1956
as one of smallest figure as
elementselementsat metaphasemetaphase II
T.T interruptusinlerruptus JapanJapan KK10(m)m) sub Gomphus melampus OGUMA,OGUMA, 1930; (Selys, 1854) (Oguma){Oguma) and G. melampus TOYOSHIMAA& HIRAI,MIRAI. bifasciatusbifasciatus (other authors); 1953; HIRAI,H1RAI, 1956;
forthetheidentity of the OMURA, 1957 species listed by Toyoshima
& MiraiHirai andandMiraiHirai cf.cf.
CRUDEN,GRUDEN, 19681968and KIAUTA,KIAUTA, 1969a
T.T “unifasciatus”"unifasciatus" JapanJapan 11 sub Gomphus;Gomphus-, the“species"“species" OGUMA,OGUMA. 1930, 19421942
(Oguma, 1926) is in factcomposed of three species
Epigomphinae
Epigomphusllama Bolivia 12 GUMMING, 1964a
Calvert. 1903
Ictinogomphinae1 ctinogomphinae
IctinogpmphusIctinogomphusrapax IndiaIndia I2(m)12(m) subIctinus;Ictinus', ASANA A& MAKING,MAKINO, 1935;1935;
(Rambur, 1942) X largest at all stages MAKING, 1935;
KICHIJO, 1942e; OMURA,OMURA. 1949, 1952,1952, 1953;1953; DASCUPTA,DASGUPTA, 1957
Gomphoidinae
AphyllaedentataSelys, 1869 Bolivia 12 GUMMING, 1964a
A. producta Selys, 1854 Bolivia 12 GUMMING, 1964a
Bolivia 12 1964a Gomphoides sp. GUMMING,
86 FAMILYFAMILY Locality dn(andd n (and Remarks References
of Subfamily presence m) Genus, Species, Subspecies
Bolivia 12 1964a Phyllocyclasp. Bolivia 12 GUMMING,
Progomphus borealis U.S.A. 12 CRUDEN,GRUDEN, 1968
McLachlan, 1873
P. intricatus(Hagen, 1857) Bolivia 12 GUMMING, 1964a
P. obscurus (Rambur, 1842) U.S.A. 12 CRUDEN,GRUDEN, 1968
P. phyllochromusphyUochrontus Ris, 1918 Bolivia 12(m) GUMMING, 1964a
Hageninae
Sieboldiusalbardae Japan 12(m) X largest OMURA,OMURA, 1957 Selys, 1886
PETALURIDAE
PetalurinacPetalurinae
Tachopteryx thoreyithoreyi U.S.A. 10(m) at ddiplotened diplotene multiple GUMMING, 1964a1964a
(Hagen, 1857) chiasmata
Uropetala carovei New Zealand 9(m) XX medium-sized WOLFE, 1953 (White, 1846)
Tanypterictinae
Tanypteryx hahagenigem U.S.A. 9(m)9(m) CRUDEN,GRUDEN, 1968
(Selys, 1879)
T.T pryeri (Selys, 1889) Japan 9(m)9(m) m minute;X second K1CH1JO, 1939, 1942e
smallest at all stages
AESHN1DAE
Gomphacschninae
Basiaeschnajanatajanata U.S.A. 13 CRUDEN,GRUDEN, 1968
(Say. 1839)
armatala Mexico X and 1970c OplonaeshnaOplonaeshna arma Mexico 14(m) X and m nearly equal KIAUTA, (Hagen,(Hagen. 1861) in size and extremely minuteminute
at metaphase II
Brachytrinac
Acanthaeschnaanacantha Australia 14(m) X smallest at KIAUTA, 1968f
(Tillyard, 1908) metaphase I
A. multipunctata Australia 14(m) KIAUTA, 1968f (Martin, 1901)
Boyeria maclachlani Japan 14(m) X secondsmallest, the OMURA,OMURA. 1957
(Selys, 1883) m’s occasionally unpaired
at at metaphase 11
B. vinosa(Say, 1839) U.S.A. 14 CRUDEN,GRUDEN, 1968
Planaeschnamilnei Japan 14(/n)14(m) KIAUTA, 1968f, 1969a
(Selys, 1883)
Aeshninae
Aeshnacanadensis U.S.A. 14 CRUDEN,GRUDEN, 1968
Walker, 19081908
A. clepsydra Say,Say, 1839 U.S.A. 14(m) HUNG, 1971
A. coerulea (Strom,(Ström, 1783) Finland 12 original m fused with the OKSALA,OKSALA, 1943a
original relatively small
X; heterokinesis neo-XY; no
of neo-Xat anaphase II
A. crenata Hagen, 1856 Finland 14(/n) in one figure the m’s OKSALA,1939a,
unpaired at metaphase II 1943a, 1944a, 1952
87 FAMILY Remarks References FAMILY Locality (5n(and<5 n(and Remarks References
Subfamily presence ofofm) Genus, Species, Subspecies
A. cyanea (Mueller, 1764) Finland, 1414(m)(m) mm minute OKSALA, 1943a; Netherlands KIAUTA,KIAUTA. 1969a
to fusion A. diffinis diffinis Bolivia I1l(m)Km) decreasein n due GUMMING, 1964a Rambur,Rambur, 1842 of autosomes;fourextra large
bivalents; XO
A. FUCHS6wNA&FUCHSÖWNA A. grandis (Linnaeus, 1758)1758) U.S.S.R., 13(m)+14(m),13(mH14(m), decreasedecreaseinin nndueto fusionfusion &
Finland, 14(/n)+13(m),14(m)+13(m), ofofXwithanautosome(usu-X with an autosome(usu- SAWCZYNSKA.SAWCZYNSKA, 1928;
Netherlands 13(m) allyally both kindsofofcomplc-comple- OKSALA, 1939a,1943a,1943a,
ments in the same specimen); 1944a,1945;1945;
neo-XYneo-XYin fused comple- MAKALOWSKAJA,MAKALOWSKAJA. 1940;
ments;neo-Xneo-Xlargest KIAUTA, 1967b, 1967c, 1967d, 1967e, 1968a,1968a,
1968d,1969a1968d,1969a
A. intricata 1908 due to fusion intricateMartin, 1908 Bolivia 10(m) decreasedecreaseinin nndue fusion GUMMING, 1964a
ofofautosomes;five extra large
bivalents; XO; X smallestsmallest at
metaphase II
A. to A. juncea (Linnaeus. 1758)1758) Finland, 13(m), 13(m) decreasedecreaseinin nn due fusion OKSALA,OKSALA.1939a, 1943a, of X with U.S.S.R., +14(m) an autosome,pri- 1944a, MAKALOWSKAJA,MAKALOWSKAJA.
Italy marymary complementsin aa few 1940; KIAUTA, 1971b1971b
cells; neo-XY in secondary
complements, no heterokinesisheterokinesis
of neo-Xat anaphase II11
A. mixta Latreille, 18051805 Netherlands 14(m)14(m) KIAUTA,KIAUTA, 1969a
A. palmate Hagen,Hagen, 1856 U.S.A. 14(m) X minute CRUDEN,GRUDEN, 19681968
A. peralta Ris, 1918 Bolivia 14(m)14(m) GUMMING, 1964a
A. senateserrata fennica Finland 13(m)13(m) subsubA.A. osiliensisfennica;fennica; OKSALA,OKSALA. 1943a
1938 element extraextra Valle,Valle, 1938 oneone element large;
neo-XY
A. subarctica elisabethae Finland 14(/n)14(m) X smallest at metaphase 11 OKSALA. 1939a, 1943a, Djakonov, 19221922 1952
A. umbrosaoccidentaloccidentalis U.S.A. 14(m)14(m) absoluteabsolutesize ofofall elements CRUDEN,GRUDEN, 1968
Walker,Walker, 1912 smaller thaninin nominatenominateform
A. umbrosa umbrosa U.S.A. 1414 absolutesize ofofall elements CRUDEN,GRUDEN, 1968 Walker, 1908 larger thanthanin A. umbrosa
occidentalis
U.S.A.U.S.A. 1971 A. verticalisverticalis Hagen, 18611861 I4(m)14(m) m hardly recognizable HUNG,HUNG,
at diakinesis
A. viridisviridis Eversman, 1836 Finland 13(m)13(m) decreaseinin n due to fusion OKSALA,OKSALA,1943a1943a
ofXX with an autosome;neo-XY
A. walkeri Kennedy,Kennedy, 1917 U.S.A. 14(m)14(m) GRUDEN, 1968
A. cf.unicolor Bolivia 1414(m)(m) GUMMING,GUMMING, 1964a1964a
Martin, 1908
Castoraeschnacastor Brasil 14(m) KIAUTA, unpublished
(Brauer, 1865)
CoryphaeschnaCoryphaeschna adnexa Bolivia 1414 GUMMING, 1964a (Hagen, 1861)
Anactinae
Anax imperator Leach, 1815 France 14(m)14(m) KIAUTA,KIAUTA. 1965, 1969a
88 FAMILY <5dn(and Remarks References FAMILY Locality n (and Remarks References
Subfamily presence ofm)
Genus, Species. Subspecies
A. juniusJunius (Drury, 1773) U.S.A. 14(m); 14 oim lacking in one ofthethe McGILL, 1904, 1907;
two populations studied by LEFEVRE & McGILL, 1908;
CrudenCruden KICHIJO, 1942e;CRUDEN,CRUDEN. 1968
A. longipes Hagen, 1861 U.S.A. 14(m) CRUDEN, 1968
A. parthenope julius Japan 14(m) OMURA, 1957
Brauer, 18651865
to Hemianax ephippiger India 7(m)7(m) decreasein ndue fusion; SESHACHAR & BAGGA,
(Burmeister,(Burmeister, 1839) probably neo-XY CKiauta)-,Kiauta); 1962;KIAUTA,KIAUTA. 1969a
m extremely minute
H. papuensis Australia 14(m) KIAUTA, 1968f, 1969a
(Burmeister,(Burmeister, 1839)
GynacanthaginaeGynacanthaginae
Gynacantha japonica Japan 14(m)14(oi) OMURA,OMURA, 1957 Bartenev, 1909
CORDU LEGASTERI DAE
Cordulegasterinae
Anotogaster sieboldiisie boldii Japan 13(m)13(oi) OGUMA, 1930; KICHIJO. (Selys, 1854) 1942e; KIAUTA. 1969a
Cordulegaster boltoni Finland,Finland, 13(m);1313(oi); 13 subsubCC. annulatusr (Oksala, OKSALA, 1939a,
m in (Donovan, 1807) Austria, Kichi/o)\KichijoY, m present 1939b;KICHIJO,KICHIJO, SwedenSweden Fennoscandianpopulations 1942e; KIAUTA, 1968c,
butlackinglacking in Austrian mate- 1968e, 1969a rial;X occasionally negatively
heterocyclic at diakinesis
C. deserticola U.S.A.U.S.A. 13(m?)13(oi?) CRUDEN.CRUDEN, 1969
Cruden,Crudcn, 1969
C. diastatops (Selys, 1854) U.S.A. 13(m)13(oi) CRUDEN. 1968
C.C dorsalis Hagen, 1857 U.S.A. 13(m)13(oi) CRUDEN.GRUDEN, 1968
C. maculamaculatustus Selys, 1854 U.S.A. 13 CORDULI1DAE Corduliinae Cordulia 13 Cordulia aenea (Linnaeus.(Linnaeus, 1758) Finland, OKSALA, 1939a; U.S.S.R., MAKALOWSKAJA. 1940; Netherlands KIAUTA,KIAUTA, 1968d, 1969a C. shurtleffi Scudder, 1866 U.S.A. 13 CRUDEN. 1968 Dorocordulia libera U.S.A. 6+7; 7 decreasein ndue to fusion;fusion; CRUDEN, 1968; (Selys,(Selys. 1871) the two different comple- KIAUTA,KIAUTA, 1969a ments in twodifferentdifferentgeogra- phic populations;allelements extra large;large; neo-XY in 7 assumed (Kiauta) n = ( ) Epicorduliaprinceps U.S.A. 13(m)13(oi) HUNG,HUNG, 1971 (Hagen. 1861) Epitheca canis McLachlan, 1886 U.S.A. 13(m)13(oi) CRUDEN,CRUDEN, 1968 E. cynosuracynosura(Say, 1839) U.S.A. 10+11;11 decreaseinin n duetoto fusion; CRUDEN,CRUDEN, 1968; the two different comple- KIAUTA, 1969a ments ments in differentgeographic in n= 10 populations; n= one elementextra large; mode of sex sex determinationuncertain E. semiaquea(Burmeister, 1839) U.S.A. 13 CRUDEN, 1968 89 FAMILY Locality ddnn (and Remarks References of Subfamily presence m)m ) Genus, Species. Subspecies E. spinigera(Selys, 1871) U.S.A. 1313(m)(m) CRUDEN, 1968 Somatochloraflavomaculata U.S.S.R. 13 MAKALOWSKAJA, 1940 (Vandcr Linden, 1825) S. Finland 1945 S. memetallicatallica Finland (9)13(9) 13 OKSALA, 1945 (Vander Linden, 1825) S. semicircularissemicircularis U.S.A. 13 CRUDEN,CRUDEN. 1968 (Selys, 1871) S. uchidaiuchidaiFocrstcr, 1909 Japan 13(m) m minute and occasionally OGUMA, 1915,1930;1915, 1930; unpaired at diakinesis KICHIJO, 1942d S. viridiaenea Japan 13 OGUMA, 1915, 1930; (Uhler, 1858) KICHIJO. I942d1942d Tetragoneuriapetechialis U.S.A. 11 GUMMING, 1964a Muttkowski. 1911 T. spinigera (Selys, 1871)1871) U.S.A. 14 HUNG, 1971 Fpophthalmiinae Didymops transversa U.S.A. 13(m) CRUDEN,CRUDEN. 1968 (Say. 1839) EpophthalmiaEpophlhalmiafrontalisfrontalisfrontalis India 13(m) DASGUPTA, 1957 Selys, 1871 Macromia magnifica U.S.A. 13(m);1313(m); 13 m present in one ofthe CRUDEN, 19681968 (McLachlan,(MeLachlan, 1874) two geographic populations studied MACROD1PLACTIDAEMACRODIPLACTIDAF. Macrodiplactinae AethriamantaAelhriamantabrevipennis India 13(m) DASGUPTA, 1957 (Rambur, 1842) Urothemis signatasignala signatasignala India 13(m) DAS, 1956; (Rambur.(Rambur, 1842) DASGUPTA, 1957 LIBELLULIDAE TetratheminaeTctratheminae Nannothemisbella U.S.A. 13(m) CRUDEN,CRUDEN. 1968 (Uhler, 1857) Libellulinae Cannaphilavibex (Hagen,(Hagen. 1861) Bolivia 13(m) GUMMING, 1964a Dasythemis esmeralda Ris, 1910 Bolivia 13(m) GUMMING, 1964a D. venosa (Burmeister, 1839) Brasil 13(m) KIAUTA, unpublished Ladonajulia (Uhler, 1857) U.S.A. 13(m) CRUDEN, 1968 LathrecistaiMlhrecista asiatica India 13(m) DASGUPTA,DASGUPTA. 1957 (Fabricius, 1798) LibelluleLibellulaangelina Selys, 1883 Japan 13(m) OGUMA,OGUMA. 1915, 1930; KICHIJO, 1942e L. axilena Westwood, 1837 U.S.A. 12 GUMMING, 1964a L.L compositacomposite (Hagen, 1873) U.S.A. 13(m) CRUDEN, 1968 L. croceipennis Selys, 18681868 U.S.A. 13(m) CRUDEN, 1968 L. 1775 U.S.A. 13 1968 L. cyanea Fabricius, 1775 U.S.A. 13 CRUDEN, 90 FAMILY Locality dn(anddn (and Remarks References Subfamily presenceof m) Genus. Species, SubspeciesSubspecies to L. depressa Linnaeus, 17S81758 Belgium, 12; 12+13(m) increase in ndue CARNOY, 188S1885(of historic of England fragmentation aa bivalent valueonly);only); Austria in Austrian material onlyonly HOGBEN, 1921; (Kiauta)Kiauta) KIAUTA,KIAUTA. 1968c, 1969b L. flavida Rambur,Rambur, 1842 U.S.A. 13(m) CRUDEN, 1968 L. forensis Hagen, 18611861 U.S.A. 13(m) X secondsecond smallest at CRUDEN, 1968 diakinesis L.incesta Hagen, 1861 U.S.A. 13 GUMMING, 1964a; CRUDEN. 1968 L.L luctuosaluctuosa Burmeister, 1839 U.S.A.U.S.A. 13 subL.L. basalis SMITH,SMITH. 1916 L.L pulchellapuleHella Drury,Druiy, 1773 U.S.A.,U.S.A.,CanadaCanada 13(m) CRUDEN, 1968; KIAUTA. 1969a1969a L. quadrimaculataquadrimaculataasahinaiasahinai Japan 13(m)13(m) subsubL. quadrimaculata OGUMA,OGUMA. 1915,1915, 1930;1930; Schmidt.Schmidt, 1957 ((OgumaOguma,, Kichi/o.Kichijo,OmuraOmura)) KICHIJO,KICHIJO. 1942d; OMURA,OMURA. 1955; KIAUTA. 1968b,1968b, 1968c L. quadrimaculata quadrimaculata U.S.S.R., 13(m) FUCHSOWNA && SAWCZYNSKA, Linnaeus, 1758 Finland.Finland, 1928; OKSALA, 1939a, 1939b. Netherlands, 1945; MAKALOWSKAJA, 1940; U.S.A. KIAUTA, 1968b, 1968c; CRUDEN,CRUDEN, 19681968 L.L. saturatasaturate Uhler, 1857 U.S.A. 13(m) CRUDEN. 19681968 L.L. semifasciata U.S.A. 13(m) CRUDEN, 19681968 Burmeister, 18391839 L. vibrans Fabricius,Fabricius, 1793 U.S.A. 13(m) CRUDEN. 19681968 Lyriothemis pachygastra Japan 13(m?) OMURA.OMURA, 1955 (Selys, 1878) Nesciothemisfarinosumfarinosum Kenya (13) (ml(m?)) KIAUTA,KIAUTA. 1969c (Focrster, 1898) Orthemisbiolleyibiolleyi Bolivia 12 XO GUMMING,GUMMING, 1964a Calvert, 1906 O.0.cultiformiscultiformisCalvert. 19061906 Bolivia 12(m) XO GUMMING, 1964a O.0.ferruginea(Fabricius,(Fabricius, 1775) Bolivia,U.S.A.,U.S.A., 5; 12; 12(m) decreasein n due to fusion; GUMMING, 1964a; 1968; Domi- n 5 in Bolivia only; CRUDEN,CRUDEN. 1968; Guatemala,Guatemala, == only; nican neo-XY in KIAUTA,KIAUTA. 1969a,1969a, 1971c Republic,Republic, n = 5; Peru m in some populations onlyonly O. levis 1906 Bolivia 4 decreasein n dueduetoto 1964b; Calvert,Calvert, 3; n fusion; GUMMING, 1964a, 1964b; neo-XY probable ((Kiauta)Kiauta) KIAUTA, 1969a Orthetrumalbistylumalbistylumalbistylum ItalyItaly 13(m) KIAUTA,KIAUTA. 1971b (Selys,(Selys, 1848)1848) O.0. albistylum speciosum Japan 13(m)13(m) OGUMA,OGUMA. 1915, 1917,1930; (Uhler.(Uhler, 1858) KICHIJO,KICHIJO. 1942d; OMURA. 1955 O.0.azureum (Rambur, 1842) Madagascar 13(m)13(m) KIAUTA, 1969b, 1969c O.O. brachiale 11 decreasein n due to 1969c1969c (Beauvois, 1805) Kenya n KIAUTA, 1969b, fusionof autosomes; one bivalentbivalentextra large; in eightororninebivalentsbivalents least two chiasmata at perper bivalent;XO 91 dn FAMILY Locality dn (and Remarks References Subfamily presenceofm) Genus, Species. Subspecies O. brunneum Italy 1313(m)(m) KIAUTA, 1971b (Fonscolombe, 1837) O. cancellatum Finland, India, 13(m) OKSALA, 1939a; (Linnaeus, 1758) Netherlands DASGUPTA, 1957; KIAUTA, 1969a, 1969b in dueto 1971b O.coerulescens Austria, 13(m); 12+13(m); increase n KIAUTA, 1969b, (Fabricius, 1798)1798) Italy 13(m)+14(m) fragmentation ofautosomes; different complements in the same specimen;specimen ; m occasi-occasi- onally unpaired at metaphase II O. glaucum (Brauer, 1865) India 13(m) DASGUPTA, 1957 O.0.japonicum (Uhler, 1858) Japan 13(m) OGUMA, 1917, 1930; KICHIJO, 1942d; OMURA, 1955 O. pruinosum neglectum India, Taiwan 13(m) DASGUPTA, 1957; (Rambur, 1842) KIAUTA,KIAUTA. 1969a, 1969b O. sabina (Drury, 1773) India 13(m) ASANA & MAKING, 1935; MAKING, 1935; KICHIJO. 1942d; RAY CHAUDHURI && DASGUPTA. 1949 O. taeniolatumtaeniobtum Greece 13(m) KIAUTA, unpublished (Schneider, 1845) O. triangulatetrbngulare melaniamebnia Japan 13(m) subO. triangulatetrbngubre OGUMA, 1917; (Selys, 1883) ( Oguma)Oguma ) OMURA,OMURA. 1955 O. triangulatetrbngubre triangulatetriangubre Taiwan 13(m) KIAUTA, 1969a, 1969b (Selys, 1878) PlathemisPbthemis lydia (Drury, 1773) U.S.A. 13(m)(m) McCILL,McGILL, 1907; CRUDLN, 1968 Potamarcha obscura India 13(m) ASANA & MAKING,MAKINO, 1935; (Rambur, 1842) MAKING, 1935; KICHIJO, 1942d; DASGUPTA, 1957 Diastatopodinae intensa DiastatopsDbstatops intense Bolivia 13(m) subD. intense RisRis GUMMING, 1964a Montgomery, 1940 D. obscura(Fabricius, 1775) Bolivia I3(m)13(m) GUMMING, 1964a Perithemis corneliacornelb Ris, 1910 Bolivia 13 GUMMING, 1964a P. domitiadomitb(Drury,(Drury, 1773) Jamaica 13(m) GUMMING, 1964a P. electraelectro Ris, 1928 Bolivia 13 GUMMING, 1964a in due to GUMMING, 1964a P.P laisbis (Perty, 1834) Bolivia 9 decrease n fusion ofautosomes; five extra large bivalents; XO BoliviaBolivia 1964a P. moomamoo maKirby,Kirby, 1889 13(m) GUMMING, P. seminole Calvert, 1907 U.S.A. 13(m) GUMMING, 1964a Perithemis Bolivia 13 1964a sp. GUMMING, ZenithopteraZenithopteraviolaviob Ris, 1910 Bolivia 13(m) GUMMING, 1964a 92 FAMILY Locality dn (and Remarks References Subfamily presence ofm)m ) Genus,Genus. Species, Subspecies Brachydiplactinae Brachydiplax chalybea India 13(m) DASGUPTA, 1957 Brauer, 1868 B. farinosa Krueger, 1902 India 13(m)13(/n) X occasionally fused DASGUPTA,DASGUPTA. 1957 with a dividing autosome, to givingrise to neo-XY B.B.sobrina sobrina (Rambur, 1842) India 13(m) RAY CHAUDHURI & DASGUPTA.DASGUPTA, 1949 Micrathyria atra Bolivia 13(m)13(/n) GUMMING, 1964a (Martin, 1897) M. didyma (Selys,(Selys. 1857) Jamaica 13(m) GUMMING, 1964a M. hageniKirby,Kirby. 1890 Jamaica 13(/n)13(m) GUMMING, 1964a M. iheringi Dos Santos.Santos, 1946 Bolivia 12(m) GUMMING,GUMMING. 1964a M. laevigata Calvert, 1909 Bolivia 13(m) GUMMING, 1964a M. ocellatadentiens Bolivia 13(m) GUMMING, 1964a Calvert, 1909 M. spuria (Selys.( Selys, 1900) Bolivia 13(m) GUMMING. 1964a M. cf. eximia Kirby, 1897 Bolivia 11 GUMMING.GUMMING, 1964a Micrathyria sp. (ungulata(ungulata Bolivia 12 GUMMING, 1964a Foerster, 1907-group) Sympetrinae Acisomapanorpoides panorpoides Bangla Desh,Dcsh, 13(m) DASGUPTA,DASGUPTA. 1957 Rambur, 1842 India Brachythemis contaminatacontaminata India 13(m) ASASANAAN A&&. MAKING, 1935; (Fabricius, 1793) MAKING, 1935; KICH1JO,KICHIJO, 1942d; DASGUPTA, 1957 Bradinopyga geminata India 13(m) DASGUPTA,DASGUPTA. 1957 (Rambur, 1842) two Crocothemis erythraea India, Kenya, 13(m) chiasmata in a DASGUPTA, 1957;1957; (Brullé, 1832) medium-sized pair Italy pair KIAUTA,KI AUTA, 1969c,1969c, 1971b occasionally in Italian material;twotwo Italian populations cytotaxonomically identic,inin Indian material a larger X and slightly higher chiasmachiasma frequency C.C serviliasen'ilia (Drury, 1773) India, Japan 13(m);12(/n?)13(m);12(m?) in Japanese material ASANA & MAKING, 1935; (Omura) permanent fusion MAKING, 1935; ofX with anan autosome KICHIJO, 1942d;1942d; (save in a single cell), RAY CHAUDHURI & DASGUPTA, thereforeneo-XY 1949; OMURA,OMURA. 1955 to Diplacodes bipunctata Australia 13(m)+15(/n)13(m)+15(m) increase in nndue KIAUTA, 1969b (Brauer, 1866) fragmentation of a bivalent D. haematodes Australia 13(m>+12 increase in ndue to KIAUTA,KIAUTA. 1969b (Burmeister, 1839)1839) fragmentation of a bivalent / D. lefebvrei (Rambur, 1842) Madagascar 13(m) KIAUTA. 1968f, 1969c D. nebulosa(Fabricius, 1793) India 13(m) DASGUPTA. 1957 93 FAMILY Locality 6<5n(andn (and Remarks References Subfamily presence ofm)m ) Genus. Species, Subspecies trivialis D.D. trivialis (Rambur, 1842) India, Australia 13(m) ASANA & MAKING, 1935; MAKING, 1935;1935; DASGUPTA, 1957; K1AUTA, 1969b Erythemis attain (Selys, 1857) Bolivia 13 GUMMING, 1964a E. collocata(Hagen,(Hagen, 1861) U.S.A. U.S.A. 13(m) CRUDEN, 1968 E. plebeja (Burmeister, plebeja (Burmeister, 1839) Bolivia 13 GUMMING, 1964a E. simplicicollis (Say,{Say. 1839) U.S.A. 13(m) CRUDEN,CRUDEN. 1968 Erythrodiplax basalis basalis basalts basalis Bolivia 13 GUMMINGGUMMING, 1964a (Kirby. 1897) E. berenice (Drury, 1770) U.S.A. 13; 13+14(m)13+14(m) CRUDEN,CRUDEN. 1968;HUNG, 1971 castanea E. castanea (Burmeister, 1839) Bolivia 12(?12(7),), 13(?)13(? ) probably due to printer’s GUMMING, 1964a error differentnumbersnumbers are given onon different pages E. connata connataconnala Chile 13(m) Chile 13(m) KIAUTA,KIAUTA, unpublished (Burmeister, 1839) connata E. connata fusca Bolivia. 13 GUMMING, 1964a; (Rambur, 1842) Guatemala CRUDEN, 1968 E. fervida (Erichson, 1848) Jamaica 13(m) GUMMING,GUMMING. 1964a E. justiniana (Selys, 1857) Jamaica 13(m) GUMMING, 1964a E. mediaBorror, 1942 Bolivia ll(m) GUMMING, 1964a E. melanorubraBorror, 1942 Bolivia 13{m)13(m) GUMMING, 1964a E. paraguayensis Bolivia I2(m)12(m) GUMMING, 1964a (Focrster, 1904) E. umbrataumbrala (Linnaeus, 1758) Bolivia, Domi- 13(m) GUMMING, 1964a; nican Republic CRUDEN, 19681968 E. unimaculata (De Geer, 1773) Bolivia 13(m) GUMMING, 1964a Lepthemis vesiculosa Bolivia 13 GUMMING, 1964a (Fabricius, 1775) NesogoniaNesogpniablackburniblackburnt Hawaii 1313(m)(m) KIAUTA,KIAUTA. 1969d (McLachlan, 1883) NeurothemisNeurothemistullia tullia to tullia India 1414(m)(m) increasein n due to RAY CHAUDHURI & (Drury,(Drury. 1773) fragmentation oftwo DASGUPTA, 1949; autosomalpairs and KIAUTA,KIAUTA. 1969a subsequent fusion ofthe originalXX with a fragment; neo-XY((Kiauta);Kiauta); delayed pairing of neo-sexneo-sex elements in primary spermatocytes U.S.A. 13 m Pachydiplax longipennis U.S.A. 13(m); 13 m present in three GUMMING, 1964a; (Burmeister, 1839) Californianpopulations CRUDEN, 1968 butlacking in Florida and W. Virginia material Rhodopygia cardinalis Bolivia 13(m) GUMMING, 1964a (Erichson, 1848) Sympetrum costiferum U.S.A. 13(m) CRUDEN, 19681968 (Hagen, 1861) 94 FAMILY <5n6 Locality n (and Remarks References Subfamily presence ofm) Genus.Genus, Species, Subspecies S. danaedame (Sulzer, 1776) U.S.S.R.,U.S.S.R., Finland, 13(m) sub S. scoticum MAKALOWSKAJA,MAKALOWSKAJA. 1940; U.S.A.U.S.A. (Makalowskaja)CMakalowskaja) OKSALA 1945; CRUDEN, 1968 S. eroticum eroticum Japan 11 one pair (bivalent)(bivalent) KICHIJO,K1CHIJO, 1942b, 1942d; (Selys.(Selys, 1883) extra large;large, XO; at HIRAI, 1956;KIAUTA, 1969b least three bivalents multiple chiasmata S. flaveolum (Linnaeus, 1758) U.S.S.R. 13(m) MAKALOWSKAJA, 1940 S. 12 frequens frequens Japan 12 no elementextra large OGUMA, 1917,1930; (Selys.(Selys, 1883) KICHIJO,K1CHIJO, 1942d, 1942e; KIAUTA. 1969b S. madidum (Hagen,(Hagen. 1861) U.S.A. 13(m) CRUDEN,CRUDEN. 1968 S. meridionaleméridionale(Selys, 1841) Switzerland 13(m) KIAUTA, unpublished S. obtrusum(Hagen, 1867) U.S.A. 13(m) CRUDEN. 1968 S. parvulum Bartenev, 1912 Japan 13(m) KIAUTA. 1968f S. pedemontanum elatumdatum Japan 13(/n) subS. pedemontanum;pedemontanum ; OGUMA,OGUMA. 1917, 1930; (Selys,(Selys. 1872) m extremely minute KICHIJO, 1942d at metaphase II S. rubicundulumrubicundulum (Say, 1839) U.S.A. 13(m) CRUDEN, 1968 indoftentwo6 S.S. sanguineum Italy 1313(m)(m) in often two KIAUTA. 1971b (Mueller, 1764) chiasmata at leastin one ofthe small bivalents S. semicinctum (Say, 1839) U.S.A. 13(m) SMITH. 1916; CRUDEN. 1968 S. striolatum Luxembourg 13 KIAUTA,KIAUTA. 1966.1966, 1969a. (Charpentier, 1840) 1969b S. vicinum (Hagen, 1861) U.S.A. 13(m) CRUDEN, 1968 S. vulgalumvulgatum (Linnaeus, 1758) Netherlands 13(m) KIAUTA, unpublished Tarnetrum corruptumcorruplum U.S.A. 13(/n) CRUDEN,GRUDEN, 1968; (Hagen, 1861) KIAUTA. 1969a, 1969b TT. illotum (Hagen, 1861) Jamaica 13(m) GUMMING, 1964a; U.S.A. CRUDEN. 1968 Leucorrhiniinae CannacriaCanmcria gravida U.S.A. I3(m) CRUDEN.GRUDEN, 1968 (Calvert. 1890) CC. herbida (Gundlach,(Gundlach. 1889) Jamaica 13(m) GUMMING, 1964a) Celithemiselisa U.S.A. I3(m) CRUDEN, 1968 (Hagen, 1861) C. fasciolafasciata Kirby, 1889 U.S.A.U.S.A. 13(m) CRUDEN, 1968 Leucorrhiniaalbifrons U.S.S.R.U.S.S.R. 13(m) MAKALOWSKAJA,MAKALOWSKAJA. 1940 (Burmeister, 1839) L. dubia (Vander Linden, 1825) FinlandFinland (9)13(9) 13 OKSALA. 1939a, 1945 L.L frigida Hagen, 1890 U.S.A.U.S.A. ll+12(m)ll+12(m) decreasein ndue to CRUDEN, 1968 fusion of autosomes; both complements in the extra same specimen; one extra large bivalentin 12;XO n = 12; 95 FAMILY Remarks References Locality 6dnn (and Subfamily presenceofm)m) Genus, Species, Subspecies L. glacialisHagen. 18901890 U.S.A.U.S.A. 13(m)13(m) CRUDEN, 1968 L. hudsonica (Selys, 1850) U.S.A.U.S.A. 13(m); 13 m present inin Californian CRUDEN, 1968 and Oregon populations, but lacking in Wisconsin material mtacta m Californian 19681968 L. intacta(Hagen, 1861) U.S.A.U.S.A. 13(m); 13 m present inin CRUDEN,CRUDEN. and Wisconsin populations, but lacking in Ohio material L. pectoralispectoralis Finland (9)(9) 13(m?)13(m? ) OKSALA, 1945 (Charpentier, 1825)1825) L. proximaproximo Calvert, 1890 U.S.A. 13(m) CRUDEN, 1968 L.L rubicunda (Linnaeus, 1758) Finland, 13 OKSALA,OKSALA. 1939a; U.S.S.R. MAKALOWSKAJA, 1940 Planiplaxsanguiniventris Guatemala 13(m) CRUDEN, 1968 (Calvert, 1907) Tritheminae Pseudothemiszonatazonala Japan 12 X end-to-endattached OMURA, 1955 to (Burmeister, 1839) toan autosome;neo-XY; neo-Xlargest 1932 Trithemis aurora India 13(m) OGUMA && ASANA, 1932 (Burmeister, 1839) T. pallidinervis India 13(m)13(m) ASANA & MAKING, 1935; (Kirby, 1889) MAKING, 1935; KICH1JO,KICHIJO, 1942d; DASGUPTA, 1957 Dytheminae Brechmorrhoga mendax U.S.A. 13(m);1313(m); 13 of thetwotwo Californian CRUDEN, 1968 (Hagen, 1861) populations studied m present inin materialfrom Nevadaco, but lacking in in that of Humboldt Co. B. nubecula (Rambur, 1842) Bolivia 13(m) CUMMING,GUMMING, 1964a B. pertinax peruviana Ris, 1913 Bolivia 13 CUMMING.GUMMING, 1964a Dythemis cannacroides Bolivia 12(m) CUMMING.GUMMING, 1964a Calvert. 1906 D. fugax Hagen, 1861 U.S.A. 13(w)13(m) CRUDEN, 1968 D. rufinervisrufinervis Jamaica 13(m)13(m) CUMMING,GUMMING, 1964a1964a (Burmeister, 1839) D. velox Hagen, 1861 Bolivia 13(m)13(m) sub,subD.D. multipunctata CUMMING,GUMMING, 1964a to Macrothemis declivata Brasil 12(m) decreasein ndue KIAUTA, unpublishedunpublished Calvert,Calvert. 19091909 fusion of twoautosomes; one bivalent extra large; XO to M. hemichlora Bolivia 3 decreasein n due to CUMMING,GUMMING, 1964a (Burmeister, 1839)1839) fusion; two extra large and one smaller bivalent; one of these two chiasmata; no sex-element recognizable at neo-XY at any stage; probableprobable minuteat KIAUTA, M. imitansimitons imitansimitons Brasil 13(m) m extremely KIAUTA, unpublishedunpubUshed Karsch, 18901890 metaphase II 96 FAMILY Locality <5ndn (and Remarks References Subfamily presenceofm) Genus, Species, Subspecies M.M. mortoni Ris, 19131913 Bolivia 1313(m)(m) GUMMING, 1964a M.M. musiva, Calvert, 1898 Bolivia 13(m) GUMMING, 1964a Scapanea frontalis Jamaica 13(m) GUMMING, 1964a (Burmeister, 1861)1861) Rhyotheminae RhyothemisRhyothemis fuliginosa Japan I2(m);12(m); 13(m) subSympetrum (Toyoshima TOYOSHIMA& HIRAI.HIRAI, Selys, 1883 && Mini.Hirai, Hirai);Hirai ); 1953; OMURA, 1955;1955; decreasein ndue to in n due HIRAI, 1956; K1AUTA,KIAUTA, 1969 fusion; one extra largelarge bivalent in n= 12; different n in two different geographic populations; of sex mode of sex determinationdetermination unclear R.R. variegatavariegata India 13(m) RAY CHAUDHURI & DASGUPTA, (Linnaeus & Johansson, 1763) 1949 Pantaliinac MiathyriaMiathyria marcella Bolivia 13(m) GUMMING, 1964a (Selys, 1857) PantalaPantala flavescens India, 13(m) ASANA& MAKING, 1935;1935; (Fabricius, 1798) Bolivia,Bolivia. MAKING,MAKINO, 1935;KICHIJO,KICHIJO, Madagascar; 1942d;DASGUPTA, 1957; Hawaii SESHACHAR& BAGGA, 1963;1963; GUMMING, 1964a; KIAUTA, 1969c1969c P. hymenaea (Say,(Say, 1836)1836) Bolivia, 13(m) GUMMING, 1964a; U.S.A. CRUDEN,GRUDEN, 1968 Tauriphila australis Bolivia 13(m) GUMMING, 1964a (Hagen, 1867) T. azteca Calvert.Calvert, 1906 Mexico 13(/n)13(m) CRUDEN,GRUDEN, 1968 Trapezostigma abdominalis Bolivia 13 sub Tramea GUMMING, 1964a (Rambur, 1842)1842) T.T. basilare burmeisteri India 13(m) sub Tramea basilaris DAS,DAS. 1956; (Kirby, 1889) DASGUPTA, 1957 TT. carolinaCarolina U.S.A. 13 subTramea GUMMING, 1964a; (Linnaeus(Linnaeus & Johansson, 1763) CRUDEN,GRUDEN, 1968 T.T. cophysa (Hagen, 1867) Bolivia 13(m) subTramea GUMMING, 1964a TT. lacerata (Hagen, 1861) U.S.A. 13 subTramea CRUDEN,GRUDEN, 1968 TT. limbata (Desjardins, 1832) India I3(m)13(m) subTramea ASANA & MAKING,MAKINO, 1935; MAKING,MAKINO, 1935; KICHIJO, 1942d T. Virginiavirginia (Rambur, 1842) India 13(/n)13(m) subTramea chinensis OGUMA &.& ASANA,ASANA. (Oguma &Asana,& Asana, 1932; KICHIJO, 1942d; Kichijo) DASGUPTA, 1957 ACKNOWLEDGEMENTS The author is obliged to Dr. JANNY M. VAN BRINK (Utrecht) for valuable suggestions on the tabulation of the cytotaxonomic data, and to Dr. M.A. L1EFTINCK (Rhenen, Netherlands) for help with the synonymy. The author’s laboratory assistant. Miss M. BRINK, was helpful with some calculations, while his secretary, Miss E. VAN DER GRIJN typed the manuscript. 97 BIBLIOGRAPHY ASANA, J.J. & S. MAKING, 1935. A comparative study of the chromosomes in the Indian dragonflies. J. Fac. Sci. Hokkaido. Univ., VI, 4 (2): 67-86. BATTAGLIA, E. & J.W. BOYES, 1955. Postreductional meiosis: its mechanism and causes. Caryologia 8; 87-134. BRINK, B. J.M.VAN & KIAUTA, 1964. Notes on chromosome behaviour in the spermato- genesis of the damselfly Enallagma cyathigerum (Charp.) (Odonata: Coenagrionidae). Genetica 35: 171-174. 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