Odonatologica I (2) 73-102 June I. 1972

Synopsis of the main cytotaxonomic data

in the order

B. Kiauta

Institute of Genetics, University of Utrecht, Utrecht, The Netherlands

Received April 13, 1972

So far 371 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 , 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 , 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 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, , 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.

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102