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International Journal of Odonatology 9(1) 2006: 1-44 1
Voltinism of Odonata: a review
Philip S. Corbet 1, Frank Suhling 2& Dagmar Soendgerath2
1I.C.A.P.B., University of Edinburgh, Scotland, UK;
present address: Crean Mill, St. Buryan, Cornwall TR19 6HA, UK.
<mail@pcorbet.vispa.com>
2Institut für Geoökologie, Technische Universität Braunschweig,
Langer Kamp 19c, 38106 Braunschweig, Germany.
<f.suhling@tu-bs.de>, <d.soendgerath@tu-bs.de>
Key words: Odonata, dragonfly, voltinism, life cycle, latitude, growth rate, seasonal regulation.
Abstract
We classified 542 records of voltinism for 275 species and subspecies of Odonata
according to three variables: geographical latitude, systematic position and habitat
type. We sorted records according to voltinism – categories being three or more
generations per year, two generations per year, one generation per year, one gene-
ration in two years and one generation in three or more years. We sought to cor-
relate the voltinism of each record with latitude of the study site, thus demonstrating
an overall negative correlation between voltinism and latitude. After allowing for
phylogenetic similarity a negative correlation remains, although it decreases in
strength after removal of taxonomic correlates, mainly between family and genus
levels. A negative correlation exists at the species level within most families, with
the exception of Lestidae. In genera for which we lacked data for latitudes
0-31°N/S no significant correlation between latitude and voltinism exists. In tem-
porary waters most species complete at least one generation per year; most species
in lentic perennial waters complete one generation or fewer; and the majority of
species in lotic waters complete half a generation or less. We discuss the roles of
exogenous and endogenous factors in influencing voltinism and identify those that
may be affecting the correlation that the data reveal. We suggest projects that could
improve understanding of voltinism in the context of seasonal regulation and the
main types of odonate life cycle so far recognised.
Introduction
Knowledge of voltinism, i.e. the number of generations completed within one year
in the field, is needed to understand how life cycles have become appropriate to
environmental conditions in different regions especially with regard to latitude,
and consequently how seasonal regulation has been achieved. We assume that the
Odonata are primarily tropical in origin and that, while colonising temperate lati-
tudes, they have retained components of warm adaptation. Thus, unlike the
Received 06 July 2005; revised and accepted 08 December 2005
Ephemeroptera and Plecoptera in temperate latitudes, many species of Odonata
have evolved one or more diapause stages that confine cold-intolerant stages – i.e.,
the reproductively active adult, larvae in early stadia, and sometimes also the
egg – to the warmer times of year (Pritchard 1982). Such diapause stages, commonly
a feature of one or more larval stadia, occasionally occur also in the egg and/or the
pre-reproductive adult. The ways in which the incidence and completion of dia-
pause, be it obligate or facultative, are regulated by responses to temperature and
photoperiod are manifold and complex (see review in Corbet 1999a: 230-237) and
provide an essential template against which to interpret patterns of voltinism in
Odonata. For example, at the highest latitudes, the constraints imposed by the
brief, cool, variable summer may result in mechanisms of seasonal regulation that
significantly extend (or shorten) the time needed to complete a generation as a
result of cohort-splitting (Norling 1984a). This phenomenon can enable some ‘out-
lier’ members of a hatching cohort to emerge either one year sooner (Corbet
1957c) or one year later (Norling 1984a) than the rest of the cohort. An estimate
of voltinism for a given species in a given latitude (or habitat) will be subject to
variation which will reflect lack of uniformity in the temporal status of members
of each population, itself an expression of such variables as times of emergence,
oviposition and hatching from the egg, and larval growth rate and the impact of
predation. Voltinism in Odonata depends mainly on regulating mechanisms and on
growth rates, which usually increase with increasing temperature up to a species-
specific maximum (Krishnaraj & Pritchard 1995). We can therefore expect that, if
climate does not require a diapause, ambient temperature will affect voltinism
directly, and therefore at low latitudes development should be more rapid. In-
creased development rate at lower latitudes has, for instance, been shown for
Onychogomphus uncatus (Ferreras-Romero et al. 1999).
Latitude and its physical correlates probably have a major influence on voltinism.
Our primary purpose in this review is to try to characterise such an influence, while
making allowance for certain other variables, namely phylogeny and habitat. For
tropical species in seasonal-rainfall areas, constraints imposed by obligate migration
will obviously affect the voltinism of populations, though to an extent that may be
almost impossible to measure. Our hypothesis in this review is that voltinism of
Odonata correlates inversely with latitude.
Comparative analysis between species should take phylogeny into account
because the life-history traits of a taxon may be ecologically constrained by the
characters of its ancestors (Felsenstein 1985; Harvey & Pagel 1991). It has been
shown, for instance, that growth rates and temperature optima are characteristic
of groups of species (e.g. Krishnaraj & Pritchard 1995; Pritchard et al. 2000).
Recent research indicates also that species-specific differences in ingestion rates are
responsible for differences in growth rate (McPeek et al. 2001). In this article we
therefore correct statistically for variation associated with phylogenetic affinity (cf.
Harvey & Pagel 1991) before analysing for latitude effects.
A preliminary, broad analysis of voltinism in Odonata was presented by one of
us (Corbet 1999a: table 7.2) without specifying the source data, and with the inten-
tion of publishing those data, together with a more searching analysis, in due course.
The present review fulfils that intention. A brief synopsis, foreshadowing, but not
duplicating, parts of this review, was presented later (Corbet 1999b). Here we pre-
sent an overview and analysis based on more than 250 publications on 275 species
International Journal of Odonatology 9(1) 2006: 1-44
2
Corbet, Suhling & Soendgerath
International Journal of Odonatology 9(1) 2006: 1-44 3
Voltinism of Odonata
and subspecies of Odonata. For this review we use the derivative terms ‘univoltine’
and ‘bivoltine’ to mean completing, respectively, one and two generations per year,
‘multivoltine’ to mean completing three or more generations per year, ‘semivoltine’
to mean completing one generation in two years and ‘partivoltine’ to mean com-
pleting one generation in more than two years.
Historic outline
The earliest published record of voltinism in Odonata known to us is that of Jan
Swammerdam. Referring to a species that was almost certainly Gomphus vulgatis-
simus, occupying a river in The Netherlands, he surmised that the species was semi-
voltine (Swammerdam 1758: 99, although the observation was made before 1669).
Almost 100 years later John Bartram reported that the (exophytic) odonates he
observed in Pennsylvania were univoltine: “The Eggs are soon hatch’d and the
young Reptiles creep amongst the Stones and Weeds etc. and so continue [as]
Water-Animals the greatest Part of the Year, until the Season comes round for their
Appearance in the beautiful Fly ...” (Collinson 1750). Bartram did not (could not)
assign a scientific name to the species he observed but if it was a species of Sym-
petrum his assertion about its univoltinism would have been correct. The first
systematic study of the topic was by Wesenberg-Lund (1913) who investigated vol-
tinism in Denmark by making field observations, including the inspection of two-
weekly samples of larvae, and concluded that within a single population voltinism
could vary from year to year. His conclusions were general, however, and he did
not assign values for voltinism to any particular species, apart from correctly inter-
preting the univoltine life cycles of Lestes dryas and L. sponsa (Wesenberg-Lund
1913: 377). Tillyard used qualitative field observations to infer voltinism of some
Australian Odonata, including Austrolestes leda (Selys) (Tillyard 1906), Petalura
gigantea (Tillyard 1911) and Anax papuensis (Tillyard 1916). Portmann (1921)
likewise used field observations to infer that Anax imperator in Switzerland was
univoltine but did not present quantitative data to support this inference. Calvert,
in his classic papers on larval development in Odonata, first (1929) considered
growth rates only in the laboratory, but later (1934) used phenological data for
several species of Anax to infer voltinism in the field. The first large-scale, syste-
matic study of voltinism was launched by Münchberg in northern Germany in the
late 1920s. He sampled larvae in nature at regular intervals throughout the year,
recording their dimensions and state of development and placing these findings in
the context of the flying season to infer the voltinism of several species of
Sympetrum (1930a), Aeshnidae (1930b, 1936), Gomphidae (1932a), Corduliidae
(1932b), and Lestidae (1933). Prominence was given to the topic of voltinism by
the inclusion of findings by Münchberg (mainly) and Portmann in a popular book
on German Odonata by Schiemenz (1953). Records of voltinism were included
also in the book by Robert (1958), who reared several European species of Odo-
nata. A new development occurred in the 1950s when information about growth
rates and voltinism in the field was combined with knowledge of the temporal pat-
tern of emergence to construct hypotheses regarding the responses controlling such
phenomena as the synchronisation and seasonal placement of emergence and the
contribution to voltinism of cohort splitting (Corbet 1957c). This led to an ecolo-
gical classification of Odonata based on the means they employ for achieving sea-
sonal regulation (Corbet 1954; Corbet & Corbet 1958) and thus to a template for
classifying their life cycles (Corbet 1960, 1999a: table 7.3).
Such templates for Palaearctic Odonata have been proposed by Corbet (1960: 143)
and Norling (1975). Corbet recognised two main categories: spring and summer
species, distinguished according to the presence or absence of a diapause in the
final larval stadium. Norling (1975) likewise recognised two categories, distinguis-
hed according to the overwintering stage and voltinism. Both authors stressed the
distinction between the obligatorily univoltine life cycle and the rest. Later, in a
searching review of the relationship between voltinism and latitude, Norling
(1984a) confirmed his recognition of two basic types of life cycle, venturing to
explain the relationship between latitude and generation length in the context
of the northern expansion of Odonata, with special reference to the induction of
larval diapause by photoperiod. Corbet (2003), in proposing an hypothesis that
would rationalise the existence of univoltinism at high latitudes, recognised
three broad types of life cycle: Type 1: spring species; Type 2: summer species; and
Type 3: obligatorily univoltine species.
Light was thrown on voltinism of tropical Odonata by Gambles (1960), Corbet
(1962) and Kumar (e.g. 1972, 1976, 1979). Using the analogy of locust migration,
elucidated by Rainey (1951), Corbet (1962: 195) hypothesised that several species
of temporary-pool-breeding odonates in seasonal-rainfall zones in the Tropics (e.g.
Pantala flavescens) were likewise travelling with the Inter-Tropical Convergence
Zone (ITCZ), the rain-bearing frontal system, and were thereby being delivered to
a succession of localities where rain was falling or about to fall. In this way it was
hypothesised that such populations would be able to complete several generations
in rapid succession, perhaps as many as five within a year, though in different
localities (see Corbet 1999a: 219). This hypothesis has an important bearing on
estimates of voltinism among tropical Odonata. So far this hypothesis has been
sustained, evidence in support of it being persuasive (e.g. Corbet 1984) though
unavoidably circumstantial. Here we assume that the hypothesis has not been
falsified. Studies of non-migratory Odonata in wet/dry climates in the Tropics have
also been exceptionally informative. Gambles (1960) showed that some species in
Nigeria (at 9.68°N), e.g. Lestes virgatus, Gynacantha vesiculata and Crocothemis
divisa, maintain a (regulated) univoltine life cycle which features a long-lived,
siccatating adult and a drought-resistant egg for which the hatching stimulus is
apparently wetting. Kumar (1972, 1976, 1979a), studying tropical-centred species
at about 30°N, revealed similarly regulated univoltine life cycles in Platylestes
praemorsus and Bradinopyga geminata and showed also that certain other non-
migratory species, e.g. Crocothemis servilia, Orthetrum sabina, (apparently un-
regulated) exhibit facultative bivoltinism if suitable aquatic habitats are available
during the dry season and also that the duration of larval development depends on
ambient temperature, a phenomenon detected also by Jödicke (2003) in subtropical
Tunisia at about 33°N. These observations by Kumar and Jödicke have provided
valuable insights into the origins of seasonal regulation in low temperate latitudes,
where temperature rather than rainfall serves as the dominant environmental
variable. A study by Schnapauff et al. (2000) from rice fields in Greece reveals that
bi- and multi-voltinism associated with habitat change from dry to wet season, e.g.
in Crocothemis erythraea and Sympetrum fonscolombii, may be a general pheno-
menon at low temperate latitudes.
International Journal of Odonatology 9(1) 2006: 1-44
4
Corbet, Suhling & Soendgerath
Patterns of voltinism in relation to life cycle and latitude on a global scale were
discussed by Corbet (1999a: 217), citing 172 records representing 27 families.
A crude analysis (making no allowance for life cycle) revealed that (as expected)
tropical-centred species exhibit higher levels of voltinism than do temperate-
centred species, but no attempt was made to seek a regression of latitude against
voltinism among temperate-centred species, an omission we correct in this review.
It is already known, from discrete observations, that several extrinsic factors, both
abiotic (including latitude) and biotic, can modify voltinism within a species
(Corbet 1999a: table A.7.2), but no correlation with latitude has previously been
sought on a global scale.
Methods
Data base
In the Appendix we present records of odonate voltinism, published and unpub-
lished, available to us at the time of writing and from many parts of the world
(Fig. 1). We included only those records that appear to us secure, but they vary
widely in rigour and quality between two extremes: from the detailed, quantitati-
ve analysis of many successive samples of larvae taken during more than one year
to a strong inference based on a single sample or on temporal patterns of emer-
gence or flight. For example, the observation of emergence from a habitat known
by the observer to have been available for oviposition no more than a year pre-
viously would in our view constitute reliable evidence of univoltinism. Similarly,
the developmental stage of larvae remaining in a water body when annual emer-
gence has just ended can yield useful information about voltinism in that popula-
tion. There are several other considerations that affect the strength of generalisati-
ons about voltinism drawn from field data:
(1) Voltinism is most readily determined for species that are consistently uni-
voltine, such as most species of Lestes and Sympetrum, because the identity
of size-groups is not blurred by cohort splitting.
(2) When emergence or oviposition continues without interruption throughout
the year, as in many tropical species, it may be difficult or impossible to infer
voltinism in the field, except by elaborate analytical methods (see Yule 1996).
(3) The practical difficulty of continuing a study for several consecutive years
means that records for partivoltine species with long life cycles tend to be
under-represented.
(4) Most publishable studies are conducted at habitats with large populations
because they are liable to yield clear-cut results; this leads to populations in
secondary and latency habitats (Sternberg 1995; Corbet 1999a: 11) being
under-represented.
(5) An investigator’s personal circumstances seldom permit a study of voltinism to
continue beyond one or two years; accordingly temporal variation in voltinism
will be understudied, especially in partivoltine species.
(6) Evidence for voltinism may differ widely in quality from one study to another,
presenting the challenge of assigning greater weight to some records than to
others when interpreting data.
International Journal of Odonatology 9(1) 2006: 1-44 5
Voltinism of Odonata
Having regard to the last consideration, and though confident in the security of the
data we have cited, we advise any investigator intending to use information in the
Appendix for comparative purposes to consult the original sources. In some cases,
e.g. when citing findings by Aguesse (1955) and Montes et al. (1982), we accepted
the authors’ unequivocal statements about voltinism without being provided with
evidence. Some other reports, though consistent with one type of voltinism, do not
explicitly exclude alternatives. For example, observations by some authors describe
adults of Uracis imbuta leaving rain forest en masse and rapidly attaining repro-
ductive maturity at the onset of the rains, a pattern of behaviour consistent with
that of species having a regulated, univoltine life cycle of the A.2.1.2 type (Corbet
1999a: 220). However, such observations do not exclude the (unlikely) possibility
that such populations might complete more than one generation in a year were
larval habitats to be available. Accordingly we omitted observations of this kind
from the Appendix.
On the assumption that climatic regimes are likely to result in major differences
between species in different latitudes, we used degrees of latitude for the analyses
(see below). Where possible we derived the latitude from the original article or
from the author directly. Failing this, we tried to reconstruct the latitude from the
locality information provided, e.g. the nearest town, using an atlas or the Internet.
Therefore the information is variably precise; we recorded latitude to the nearest
0.01 degree, thus adopting the normal format for geographical information
systems. Altitude data, though obviously relevant, could not be obtained with
acceptable accuracy. This we regret, because (as expected) altitude can affect volti-
nism (Deacon 1979).
Beyond the primary segregation according to latitude, we classified individual
records according to two other variables that we predict will impose constraints
that prevent voltinism being a simple resultant of the main physical correlates of
latitude. These variables are (1) systematic position and (2) larval habitat. We gave
weight to systematic position (i.e. phylogenetic affinity, see below) to allow for pat-
terns compellingly exemplified by taxa such as Lestes and Sympetrum, many mem-
bers of which exhibit a univoltine life cycle (type B.2.1.2), usually maintained
across a wide latitudinal range by an obligate diapause in the egg and/or the adult,
International Journal of Odonatology 9(1) 2006: 1-44
6
Figure 1: Distribution of localities from which data of Odonata voltinism (Table 1) were derived.
Corbet, Suhling & Soendgerath
Arctic circle
Tropic of Cancer
Equator
Tropic of Capricorn
even in representatives that occur north of the Arctic Circle. Because any one
species can be opportunistic in choice of habitat, we used broad habitat categories,
each of which included the habitat type most representative of the one in the
original account. We have identified three categories: perennial lotic, perennial len-
tic and temporary lentic.
Analysis of the influence of latitude
For analysing the influence of latitude on voltinism we used Spearman rank corre-
lations because we predicted that our data are dependent and not normally distri-
buted, particularly in respect of phylogenetic relationships, and the fact that many
species have more than one entry in the database (Appendix). We used Stearns’
method (1983) of directly subtracting the variation associated with phylogenetic
similarity from the species’ data. Because no consensual phylogenetic tree was avail-
able for all species of Odonata included in our database, we used taxonomic simi-
larity to stand in for phylogeny. For higher taxonomy we referred to the work of
Misof (2002) and Rehn (2003). The species were classified by taxonomic related-
ness into suborder, family and genus. The families Corduliidae and Calopterygidae
were used in the widest sense, i.e. to include Macromiidae and Synthemistidae in
Corduliidae and Hetaerinidae in Calopterygidae. The study of Rehn (2003) implies
that the number of zygopteran families may be combined into some recent ‘super-
families’, e.g. Coenagrionoidea. However, because the zygopteran taxonomy is not
yet resolved, we referred here to the families in current use. Taxa were given cate-
gorical codings to produce dummy variables for membership of suborder, family
and genus. Each categorical code was used as an independent variable in a multiple
regression to remove the taxonomic correlates, resulting in a set of residuals that
are not correlated with taxonomy (for details on methods see Harvey & Pagel
1991: 130, box 5.1). For this kind of analysis we had to remove from the database
all taxa for which no replicates were available. For example, in the Euphaeidae
three genera, Anisopleura, Bayadera and Epallage, were each represented only
once in the data set, and therefore, when removing generic relatedness, only one
replicate was available for each of these genera, resulting in singularities in the
regression analysis. Then, after having removed these three genera, only one genus
with replicates was left within the Euphaeidae, so that the whole family had to be
removed from the analysis. The same occurred with the Chlorocyphidae, Poly-
thoridae, Platystictidae, Protoneuridae, Epiophlebiidae, Petaluridae, and Cordule-
gastridae. Likewise, all genera with only one entry had to be removed. Due to these
statistical requirements eventually only 481 from a total of 542 entries in the data-
base could be used.
Were phylogenetic affinity to have influenced our results, we expected the corre-
lations based on the taxonomy-independent residuals to be weaker than those
obtained using the original voltinism data. Therefore, as a first step, we correlated
voltinism data, expressed as number of generations completed per year (multivol-
tine = 3, bivoltine = 2, univoltine = 1, semivoltine = 0.5, partivoltine = 0.3) against
latitude without removing the effect of taxonomy. For the next step, to determine
at which taxonomic level voltinism was most influenced, we first removed mem-
bership of suborder and family and thereafter, in another analysis, membership of
suborder, family and genus.
International Journal of Odonatology 9(1) 2006: 1-44 7
Voltinism of Odonata
Seven families, Calopterygidae, Lestidae, Coenagrionidae, Aeshnidae, Gomphidae,
Corduliidae and Libellulidae, are represented by sufficient numbers of entries in
the database to allow correlations between voltinism and latitude at the family
level to be carried out. The residuals from the removal of genus membership, as
well as the voltinism data without removal of other taxonomic correlates, were
used (see above). Additionally, we performed correlations at the genus level for
some genera. For the genus level we used the voltinism data without removing the
effect of taxonomy. We also did so for two species that have enough entries in our
database: Ischnura elegans and Gomphus vulgatissimus.
In the Appendix we scored for records, and not solely for species (or subspecies).
A frequency table for voltinism derived directly from entries in the Appendix
would be misleading, mainly because some species, e.g. G. vulgatissimus, have
several entries, resulting in the overall voltinism frequencies for the Gomphidae
being correspondingly weighted by the voltinism of G. vulgatissimus. Another, dif-
ferent, distortion resulted from the fact that entries for some species feature popu-
lations exhibiting different types of voltinism, e.g. Pyrrhosoma nymphula and
Anax imperator. Hence, to avoid misinterpretations we did not summarise data at
the family level.
Analysis of habitat effects
The larval habitat may constrain the duration of larval development. Temporary
waters usually require rapid growth to permit successful development (cf. Wellborn
et al. 1996; Suhling et al. 2005). Except for those species that siccatate, most tem-
porary-water species may therefore be able to produce more than one generation
per year. By contrast, species occurring in perennial waters, particularly riverine
calopterygids and gomphids (Rüppell 2005; Suhling & Müller 1996), often have a
‘slow’ life style (sensu Johnson 1991) leading to low growth rates (e.g. Johansson
2000; Suhling 2001). Major differences in voltinism should therefore appear bet-
ween each of (1) temporary waters, (2) perennial lentic waters and (3) perennial
lotic waters. We recognised that some species exhibit opportunistic habitat occu-
pancy within and between seasons, choices that may influence their voltinism (see
Kumar 1976). We assigned such species to one of the three types of habitat by
choosing the one which probably most influenced their voltinism. We tested for the
effect of larval habitat by linking each habitat type to each species entry in the
database (Appendix). Because of multiple entries for species, the data set was not
independent. Therefore we used the Kruskal-Wallis rank test to evaluate differences
in voltinism between habitat types. This operation was conducted for the raw data
as in the Appendix, as well as for data corrected for taxonomic relationship at the
family and genus levels (see above).
International Journal of Odonatology 9(1) 2006: 1-44
8
Figure 2: Scattergrams of Odonata voltinism vs latitude — (a) Correlation of numbers of
generations per year with latitude. In (b) and (c) phylogenetic correlations are removed using
the method of Stearns (1983). (b) Correlation of residuals of voltinism from a multiple regres-
sion with suborder and family relatedness. (c) Correlation of residuals of voltinism from a
multiple regression with suborder, family and genus relatedness (see Methods). Because
some cases had to be removed from the original data set due to singularities, the number of
replicates was only 484 in all diagrams. Negative (southern) latitudes are multiplied by -1.
Corbet, Suhling & Soendgerath
International Journal of Odonatology 9(1) 2006: 1-44 9
3
2
1
0
2
1
0
-1
-2
2
1
0
-1
-2
010203040506070
Latitude [°]
a
b
c
Generations per year
Residuals of generations per year Residuals of generations per year
Voltinism of Odonata
Results
Relationship between voltinism and latitude or phylogenetic affinity
In the Appendix we compiled voltinism data for 269 species and six additional
subspecies, many of which have several entries. The types of voltinism were not
equally distributed over all families. For instance, multivoltine species were recor-
ded more often from Coenagrionidae and Libellulidae, whereas partivoltinism
occurred mainly in the Anisoptera, for instance in the Cordulegastridae, though
rarely in the Zygoptera.
Number of generations per year including all entries was significantly negatively
correlated with latitude (Spearman rank correlations, Fig. 2). This was true for
voltinism data without removal of phylogenetic correlates (rho = -0.505, p< 0.001,
Fig. 2a) as well as for the residuals from such phylogenetic membership removals
(Figs 2b, 2c). After removing suborder and family membership the correlation
was somewhat reduced compared with that without such removal (rho = -0.427,
p< 0.001, Fig. 2b). However, after also removing genus membership the correlation
coefficient dropped distinctly, although still being significant (rho = -0.203,
p< 0.001). This indicates that most variation attributed to phylogeny occurs bet-
ween the family and the genus levels. When completely eliminating from the data
set the family Lestidae, most species of which are obligatorily univoltine because
the life cycle is regulated, the correlation slightly strengthened (rho = -0.527,
p< 0.001). In the Calopterygidae, Coenagrionidae, Aeshnidae, Gomphidae,
Corduliidae, and Libellulidae the number of generations per year was significantly
negatively correlated with latitude, whereas no such correlation was found in the
Lestidae (Table 1, Fig. 3). After removal of taxonomic correlates at the genus level
the correlations were reduced in strength, implying effects of phylogenetic inertia
at the genus level. However, the correlations were still negative and significant for
the Coenagrionidae, Aeshnidae and Libellulidae, and there was a trend in the
Gomphidae (Table 1).
Some genera contained enough entries in the Appendix to allow correlations of
voltinism with latitude at the genus level (Table 2). Spearman rank correlations
indicated negative correlations between numbers of generations per year and lati-
tude for most genera tested, these being significant only in Coenagrion, Ischnura,
Anax and Orthetrum (Table 2). However, for most of the non-significant genera
we did not have data between 30°S and 30°N, which may have influenced the results.
In Lestes there was no good evidence for variation in voltinism, all species being
univoltine (cf. Appendix). For several species there were records for at least two
different generation times within a single species, which sometimes revealed a high
within-species variability of the duration of a generation (Appendix). The clearest
example was the widespread Palaearctic species Ischnura elegans, which had been
observed to be multivoltine (in Spain, southern France and Greece), bivoltine (in
Belgium, France and Germany), mainly univoltine (in northern England, Italy and
The Netherlands) and semivoltine (in Scotland) (Appendix). In this species the data
revealed a pronounced inverse relation between latitude and voltinism (Spearman
International Journal of Odonatology 9(1) 2006: 1-44
10
Corbet, Suhling & Soendgerath
International Journal of Odonatology 9(1) 2006: 1-44 11
rank correlation: rho = -0.867, p= 0.004, n= 12). By contrast, voltinism of
Gomphus vulgatissimus was positively correlated with latitude, although not
significantly so (rho = 0.237, p= 0.344, n= 17).
Influence of larval habitat
Most entries (56%) for species of temporary waters were multi- or bivoltine,
whereas for species in lentic perennial and lotic waters most entries were uni- or
semi-voltine (65%) and semi- or parti-voltine (68%), respectively (Fig. 4). Hence,
the median numbers of generations per year were highest in the temporary-water
Figure 3: Scattergrams of Odonata voltinism vs latitude segregated according to family — in
the Calopterygidae the multivoltine entry represents Hetaerina capitalis in Costa Rica; in the
Lestidae the single partivoltine entry represents an upland population (579 m a.s.l.; 43.05°S)
of Austrolestes colensonis in New Zealand; in the Gomphidae bi- and multi-voltine entries
represent the predominantly tropical species Paragomphus genei in Namibia and Tunisia, re-
spectively.
Calopterygidae Aeshnidae
GomphidaeLestidae
Coenagrionidae Libellulidae
010203040506070
010203040506070
3
2
1
0
3
2
1
0
3
2
1
0
Generations per year
Latidude [°]
Voltinism of Odonata
International Journal of Odonatology 9(1) 2006: 1-44
12
species (2 generations per year) followed by perennial lentic species (1) and lotic
species (0.5). Voltinism differed significantly between the types of habitat, which
was true for voltinism data without removal of phylogenetic correlates (Kruskal-
Wallis test H= 104.56, p< 0.001) as well as for the residuals after removals of the
suborder and family memberships (H= 43.9, p< 0.001) and genus membership
(H= 6.4, p= 0.041).
Discussion
Our analyses reveal an overall negative correlation between voltinism and latitude.
Certain departures from this correlation can be interpreted in terms of the factors,
both exogenous and endogenous, that are known or inferred to influence voltinism
and that therefore act as sources of variation in our analysis, as depicted for
instance in Figure 2.
Exogenous factors
Major factors influencing development and therefore also voltinism are ambient
temperature and photoperiod (Corbet 1999a: 228). Latitude is correlated with both
these factors and may therefore be used as a surrogate variable, as in this study.
Our results reveal that the general pattern supports our hypothesis: the number of
generations per year decreases with latitude. In low latitudes (i.e. in the Tropics)
most species are multi- or bi-voltine, whereas in high latitudes most species are
semi- or parti-voltine. In mid-latitudes and the Subtropics the frequencies are inter-
mediate (Fig. 5). This pattern can also be observed at the levels of most families
and of some genera. However, our analysis reveals also that voltinism is influenced
by phylogenetic constraints. On the one hand, this can be explained by life-cycle
strategies (i.e. endogenous factors; see below) of certain taxonomic groups. For in-
Corbet, Suhling & Soendgerath
Table 1. Latitude vs Odonata voltinism — Spearman rank correlations for higher taxonomic
groups between latitude and voltinism and between latitude and residuals of a multiple re-
gression between voltinism and taxonomic relationship at the genus level. ngives the num-
bers of entries which could be used for analysis after removal of taxa without replicates (see
Methods).
Family nLatitude vs voltinism Latitude vs residuals
rho p rho p
Aeshnidae 88 –0.462 < 0.001 –0.222 0.038
Corduliidae 30 –0.634 < 0.001 0.066 0.945
Gomphidae 69 –0.610 < 0.001 –0.218 0.072
Libellulidae 124 –0.528 < 0.001 –0.235 0.009
Calopterygidae 19 –0.524 0.026 0.021 0.929
Coenagrionidae 99 –0.694 < 0.001 –0.309 0.002
Lestidae 38 0.113 0.491 –0.171 0.297
International Journal of Odonatology 9(1) 2006: 1-44 13
stance in the Lestidae these lead to obligate univoltinism in most species (Appendix).
On the other hand, the uneven representation of entries in our database has surely
influenced the results. Many genera, for instance, have very few entries often deri-
ving from a very small latitudinal range. Therefore they show little or no variation
with latitude. Entries for the genera Ischnura, Anax and Orthetrum, of which we
have examples from low to high latitudes, reveal that in genera in which the species
have unregulated life cycles (sensu Corbet 1999a: 220) distinct correlations bet-
ween voltinism and latitude occur. Thus, increasing temperatures and decreasing
winter periods in lower latitudes often lead to increased growth rates and therefore
allow completion of more than one generation per year. This is evident in Ischnura
elegans (Table 2), which shows the clearest correlation between latitude and
voltinism.
Voltinism differs significantly among perennial lotic, perennial lentic and tem-
porary lentic habitats (Fig. 4). This is to be expected, particularly due to the life-
cycle constraints caused by drying out of temporary waters. Temporary waters
usually require rapid growth to permit successful development (cf. Wellborn et al.
1996). This pattern is especially evident in arid countries, such as Namibia
(Johansson & Suhling 2004; Suhling et al. 2004, 2005). Species occupying tem-
porary waters there develop more rapidly than those occupying permanent waters,
and those occupying lentic waters there develop more rapidly than those occupy-
ing lotic waters. This correlation is not independent of latitude – seasonal rainfall
and ephemeral pools are commoner at lower latitudes – or phylogenetic affinity
because certain taxa, e.g. Pantala flavescens, oviposit by preference in ephemeral
Temporary Perennial lentic Perennial lotic
n= 96 n= 271 n= 109
100
80
60
40
20
0
Percentage of records
Figure 4: Percent occurrence of the different types of Odonata voltinism listed in the Appendix
to habitat-type — the semivoltine entry for temporary habitats is represented by Libellula depressa
from France (Blois 1985). — Black: multivoltine; grey: bivoltine; white: univoltine; vertical-
hatching: semivoltine; horizontal-hatching: partivoltine.
Voltinism of Odonata
International Journal of Odonatology 9(1) 2006: 1-44
14
pools lacking surface vegetation (Weir 1974), which on that account can be expected
to be warmer. With the exception of those species, e.g. Lestidae, that hibernate in
the egg or adult stage or aestivate or siccatate as adults, most temporary-water
species may therefore be able to be bi- or multi-voltine (cf. Suhling et al. 2003). On
the other hand, species occurring in perennial waters, particularly lotic habitats,
such as Gomphidae and Cordulegastridae, often have a slow life style (sensu
Johnson 1991) associated with low growth rates and therefore longer generation
times (e.g. Johansson 2000; Suhling 2001).
The effect of type of larval habitat on voltinism influences the relationship bet-
ween latitude, taxonomy and voltinism in two major ways. First, in lower latitu-
des the frequency of temporary waters increases – as does the frequency of entries
in our database for temporary-water species. In genera that have unregulated life
cycles, such as Ischnura, Anax and Orthetrum, growth rate depends mainly on
temperature, enabling them to develop rapidly in such temporary waters, which
more often occur in lower latitudes. Hence, the finding that in these three genera,
for instance, voltinism correlates well with latitude (see above) also reflects their
ability to use these habitats at higher ambient temperatures. Second, the distribu-
tions shown in the Appendix allow the special status of several families to be high-
lighted as occupants of habitats that are either solely lotic (Calopterygidae, Chloro-
cyphidae, Euphaeidae, Platystictidae, Polythoridae, Cordulegastridae) or mainly
lotic (Platycnemididae, Gomphidae). Wheras in other families several species occur
in habitats that sometimes or always feature drying, such as temporary pools
(Lestidae, Coenagrionidae and Libellulidae). Hence, taxonomy and habitat are
often correlated, which explains some of the obvious variation caused by taxo-
nomic relatedness in the correlation between latitude and voltinism. Besides the
known differences between the habitat types chosen in our study, small-scale
differences in habitat may matter also. For instance, in Gomphus vulgatissimus,
which shows no significant correlation between voltinism and latitude, informative
conclusions can be drawn from the larval habitat: G. vulgatissmus is semivoltine
in large rivers, which can feature high summer water temperatures (> 30°C in shal-
low water near the banks) even at about 50°N, whereas it is usually partivoltine,
requiring up to four years per generation, in smaller streams as well as in cold
lakes, regardless of latitude (Müller et al. 2000).
Corbet, Suhling & Soendgerath
Table 2. Latitude vs Odonata voltinism — Spearman rank correlations for selected genera.
Genus nrho p
Coenagrion 17 –0.504 0.044
Enallagma 15 –0.332 0.214
Ischnura 30 –0.657 < 0.001
Aeshna 44 –0.240 0.116
Anax 25 –0.516 0.004
Gomphus 25 –0.056 0.785
Orthetrum 16 –0.560 0.030
Sympetrum 32 –0.168 0.349
International Journal of Odonatology 9(1) 2006: 1-44 15
Endogenous factors
We have considered endogenous factors only by allowing that phylogenetic affinity
(sometimes reflected in life-cycle regulation) may impose constraints, examples
being the obligatorily univoltine life cycles exhibited by many species of Lestes and
Sympetrum under almost all circumstances, and the low voltinism exhibited by the
predominantly lotic Cordulegastridae. We now consider other endogenous factors
(sometimes reflecting phylogeny) that can influence interspecific variability. Many
species of Odonata in temperate latitudes possess an elaborate suite of responses
to temperature and daylength that result in the emergence period being placed at a
time of season favourable for reproduction (Norling 1984a), a process known as
seasonal regulation. Such species are described as having a ‘regulated’ life cycle.
The better-known examples of seasonal regulation entail extending generation
time, as when (in spring species, with the Type-1 life cycle) development of the
Polar Temperate Tropical
n= 12 n= 131 n= 66
100
80
60
40
20
0
Percentage of records
Figure 5: Percent occurrence of the different types of Odonata voltinism in the major climatic
zones of the earth — tropical (latitude 0° - 23.45°N/S), subtropical (23.45° - 35°N/S), tem-
perate (35° - 58°) and polar (> 58°) derived from all entries in the Appendix. We allow for
departures from the conventional (geographical) definitions of climatic zones: in the Dehra
Dun valley in India the tropics reach 30.5°; in western Europe (Mediterranean France only),
reflecting local climate determined by the Gulf Stream, we consider the subtropics to extend
to about 44°N; the polar zone is conventionally regarded as falling within latitudes excee-
ding 66.5°N/S but here, to reflect the realities of the climatic environment within which the
northernmost Odonata occur, we place the southern limit of this zone at 58°N. — Black:
multivoltine; grey: bivoltine; white: univoltine; vertical-hatching: semivoltine; horizontal-
hatching: partivoltine.
Subtropical
n= 273
Voltinism of Odonata
International Journal of Odonatology 9(1) 2006: 1-44
16
most precocious individuals is arrested by the intercalation of a final-stadium dia-
pause, which allows retarded individuals to ‘catch up’ in the autumn preceding a
spring emergence. Likewise, onset of this late season diapause may oblige a larva
in summer to postpone emergence for an extra year. Or, in summer species (with
the Type-2 life cycle), a rising series of temperature thresholds for entry to succes-
sive late stadia in spring (Corbet 1957b; Lutz 1968) can enable laggards to catch
up with the most precocious individuals that are thereby held back. Another
mechanism causing univoltinism is postponed reproductive maturation of species
that emerge in spring and oviposit in autumn to avoid the drought of their habi-
tats in summer (cf. Samraoui et al. 1998). Such species, for instance Aeshna mixta,
Sympetrum meridionale and S. s. striolatum in Algeria, spend the hot summer as
immature adults away from their reproductive habitats, as does S. sinaiticum in
Tunisia (Jödicke 2003). Such a resting period, representing a summer diapause is
known as aestivation, as distinct from the siccatation-resting period exhibited by
some tropical species during the dry season.
There are other, less well-known, responses that can shorten generation time and,
by serving thereby to offset the effects of latitude, may obscure the direct correla-
tion we postulate. Such responses are found among most species of Lestes and help
to maintain the obligatorily univoltine life cycle typical of the genus. Most tempe-
rate species of Lestes feature a diapause in the egg (see Jödicke 1997), which pre-
vents small larvae being exposed to low (winter) temperatures and synchronises
egg hatching in spring (Corbet 1956). Thereafter the larvae, which have an intrinsi-
cally high thermal coefficient for growth (Krishnaraj & Pritchard 1995) and a high
intrinsic attack coefficient (Pickup & Thompson 1990), grow unusually rapidly and
emerge promptly in early to mid summer – early enough to complete the repro-
ductive period before the advent of autumn, whereupon the adults die. This suite
of responses alone would normally be able to maintain the univoltine life cycle of
Lestes, but in L. congener (Johansson and Rowe 1999) and L. sponsa (Johansson
et al. 2001) a mechanism exists that enables retarded larvae in spring and early
summer to grow more rapidly (under the progressively longer photoperiods that
characterise the advancing season up to the summer solstice) and thus to ‘make up
for lost time’. In this case seasonal regulation is not extending generation time but
abbreviating it. This response, the so-called ‘light-growth effect’ (see Saunders
2002), is well known in insects and has been detected in at least ten other species
of Odonata (Danks 1987: 204). A far-reaching consequence of this photoperiod
response is that it could provide a mechanism for continuously compensating for
the retarding effects of latitude between the spring equinox and the summer solstice,
because of the relationship between latitude and daylength (Corbet 2003; see also
Corbet 1999a: 229). Any mechanism that compensates for latitude in such a way
will weaken the regression of latitude on voltinism and so weaken the correlation
we have revealed in this paper.
Another respect in which responses of odonate larvae may help to compensate
for latitude is their possession of genetic heterogeneity in regard to a photoperiod
that is critical for the induction of larval diapause. In Leucorrhinia dubia, for
example, populations in Sweden at different latitudes have different critical values
for the photoperiod that induces larval diapause. In this way the seasonal regula-
tion of larvae can be made appropriate to latitude (Norling 1984b). Unlike the
light-growth effect, however, such a response is likely to prolong, not abbreviate,
larval duration.
Corbet, Suhling & Soendgerath
International Journal of Odonatology 9(1) 2006: 1-44 17
The factors discussed above achieve seasonal regulation and find expression in the
different kinds of life cycles shown by Odonata. On a global scale, twelve types of
odonate life cycle have been identified (Corbet 1999a: 220), falling under two
major dichotomies: tropical-centred vs temperate-centred species and regulated vs
unregulated life cycles. From what has been said, we conclude that a meaningful
correlation between voltinism and latitude can be expected only among species
that occupy temperate latitudes and whose life cycle is not regulated so as to be
invariably univoltine. Species appropriate for such an analysis are those with one
of three main types of life cycle as defined by Corbet (1999a: 220), namely B.1
(unregulated; facultatively multivoltine), B.2.2.1 (regulated; facultative diapause in
larva) and B.2.2.2 (regulated; facultative diapause in larva and obligate diapause
in egg). Species in the second and third of these categories correspond broadly to
Types 2 and 3 respectively as defined by Corbet (2003) and are appropriate for
such analysis because the regulatory processes (as far as we know) relate mainly to
the arrest or acceleration of larval development connected with the seasonal pla-
cement and synchronisation of emergence, and not to the temperature-dependent
rate of growth during the growing season. These processes occur in two episodes:
before the summer solstice, when they result in acceleration of growth before emer-
gence; and after the summer solstice, when they result in growth being arrested in
an overwintering stage (see Corbet 2003).
Having regard to the inferred effects on odonate distribution of global warming
(e.g. J. Ott 1996, 2000; Hickling et al. 2005), we anticipate that in future the dis-
tributions illustrated in Figures 2 and 3 will undergo displacement towards the
right.
Conclusions
We have given examples of mechanisms underlying seasonal regulation. These show
that voltinism is often correlated with, but never likely to be a simple resultant of,
latitude, or indeed of any other exogenous factor. Voltinism is a resultant of the
interaction of exogenous and endogenous factors that, acting in concert, maintain
a viable, seasonally appropriate life cycle under varying environmental conditions.
Our review points up six lines of enquiry that are likely to be fruitful in clarifying
the underlying causes of voltinism in Odonata. They focus especially on odonate
populations in the Tropics and the far north.
(1) To determine voltinism in the Tropics, especially in perennial waters. Most
existing records derive from temporary waters, which, by their nature, do
not provide information about the number of generations that may be
completed in other habitats.
(2) To try to obtain information that would bear on the current hypothesis
regarding the migratory behaviour and voltinism of ITCZ migrants. A
productive approach might be to make co-ordinated observations of adult
arrivals and larval development in a region (such as India) in which the
track and approximate timing of the passage of the annual monsoon rains
are known. To conduct such an investigation would provide an opportunity
to obtain an estimate of the voltinism of a (suspected) ITCZ migrant (such
as Pantala flavescens) and also to test the hypothesis advanced to explain
its seasonal appearance and disappearance. Odonatologists in western India
are exceptionally well placed to undertake such a project because the path
followed by the annual monsoon there is well known.
Voltinism of Odonata
International Journal of Odonatology 9(1) 2006: 1-44
18
(3) To determine voltinism of species from habitats which are poorly represented
in the Appendix – e.g. perennial waters in the Tropics and perennial water
courses in a desert or semi-desert environment.
(4) To explore the effects on voltinism of habitat change. In the Mediterranean
Region, multivoltinism is associated with habitat change: one generation
survives the winter in the larval stage in natural ponds and swamps, whereas
one or more summer generations develop in rice fields (Schnapauff et al.
2000). In central Europe such a summer generation may occur facultatively
in shallow, warm ponds (cf. Inden-Lohmar 1997). Facultative bivoltinism
(even sometimes multivoltinism) can occur among tropical-centred species
in India when perennial habitats are used to bridge the interval between
monsoons (Kumar 1976).
(5) To determine (in the laboratory) the light-growth response of larvae of the
species of Lestes and Sympetrum that occur north of the Arctic Circle, namely
L. disjunctus, L. dryas and S. danae.
(6) To determine (in the field) the voltinism of as many species as possible close
to the northern limit of their distribution. Now that Cannings et al. (1991)
have identified specific habitats for 24 species, representing three main types
of life cycle, north of the Arctic Circle, an opportunity exists to obtain this
information. Such a study could use strong inference based on the size-
distribution of larvae remaining at, or near, the end of the annual emergence
period. Such larvae form the cohort due to emerge one or more years sub-
sequently. They could be distinguished from the current year’s emergence
group because the latter would be present as F-0 larvae, typically showing
signs of metamorphosis.
Acknowledgements
We are greatly indebted to the many, careful investigators whose effort and dedi-
cation provided the data in the Appendix – the backbone of this paper. We thank
those who provided data for co-ordinates of habitats where this information was
unavailable from a published source. We also thank Sally Corbet for valued com-
ments on a late draft and Ulf Norling and Gordon Pritchard for critical and help-
ful reviews. We acknowledge with pleasure the stimulus provided by Ulf Norling
in his seminal paper on seasonal regulation published in 1984 (Norling 1984a).
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Appendix: Records of voltinism of 269 species plus six additional subspecies of dragonflies,
varying from detailed, long-term analyses of cohorts and their growth rates to informed infe-
rences based on knowledge of the flying season, ambient temperatures and growth rate deri-
ved from various studies. Information on latitudes and longitudes refer to the study sites and
are in decimal degrees (accuracy: 0.01 degree); negative values stand for S and W respec-
tively. Multiple entries for a single species or subspecies are listed in order of increasing vol-
tinism, namely partivoltine (P), semivoltine (S), univoltine (U), bivoltine (B), and multivoltine
(M). The habitat type for the study site is classified as temporary (TEM), perennial lentic (PER),
and perennial lotic (LOT).
Taxon Voltinism Lat Long Habitat Reference
ZYGOPTERA
CALOPTERYGIDAE
Atrocalopteryx atrata Selys U 36.02 140.13 LOT Takamura (1996)
Calopteryx dimidiata Burmeister U 34.18 -81.12 LOT Smock (1988)
haemorrhoidalis (Vander Linden) S 37.93 -4.87 LOT Ferreras-Romero et al. (2000)
japonica Selys U 36.50 137.87 LOT Watanabe et al. (1998)
splendens (Harris) S 52.40 12.53 LOT Göcking (1999)
U52.40 12.53 LOT Göcking (1999)
U52.32 10.45 LOT C. Ott (1996)
U52.32 10.45 LOT Schütte & Schrimpf (2002)
virgo (Linnaeus) S 52.32 10.45 LOT C. Ott (1996)
S52.32 10.45 LOT Schütte & Schrimpf (2002)
S50.82 -1.58 LOT Corbet (1957a)
S50.38 4.17 LOT Lambert (1994)
S47.00 8.00 LOT Robert (1958)
S37.37 -4.75 LOT Ferreras-R. & García-R. (1995)
U37.37 -4.75 LOT Ferreras-R. & García-R. (1995)
International Journal of Odonatology 9(1) 2006: 1-44
32
Corbet, Suhling & Soendgerath
Taxon Voltinism Lat Long Habitat Reference
Hetaerina americana (Fabricius) U 10.50 -85.25 LOT Pritchard (1991)
capitalis Selys M 10.50 -85.25 LOT Pritchard (1991)
rosea Selys U 20.77 -42.88 LOT De Marco & Cardoso P. (2004)
Mnais andersoni McLachlan in Selys U 22.42 114.18 LOT Dudgeon (1987)
pruinosa costalis Selys U 35.50 139.48 LOT Taguchi & Watanabe (1993)
Neurobasis chinensis (Linnaeus) B 30.50 78.00 LOT Kumar (1976)
CHLOROCYPHIDAE
Platycypha caligata (Selys) B -15.00 35.30 LOT Parr (1984)
Rhinocypha perforata (Percheron) U 22.42 114.18 LOT Dudgeon (1987)
quadrimaculata Selys U 30.50 78.00 LOT Kumar (1976)
EUPHAEIDAE
Anisopleura lestoides Selys U 30.50 78.00 LOT Kumar (1976)
Bayadera indica (Selys) U 30.50 78.00 LOT Kumar (1972)
Epallage fatime (Charpentier) U 36.17 28.00 LOT Norling (1982)
Euphaea decorata Hagen in Selys U 22.42 114.18 LOT Dudgeon (1989b)
formosa Hagen U 23.50 121.00 LOT Hayashi (1990)
yayeyamana Oguma U 24.33 123.83 LOT Hayashi (1990)
POLYTHORIDAE
Cora marina Selys U 10.50 -85.25 LOT Pritchard (1996)
LESTIDAE
Austrolestes colensonis (White) P -43.05 171.78 PER Deacon (1979)
U-36.87 174.77 PER Rowe (1987)
U-37.87 174.77 PER Winstanley (1979)
U-43.45 172.63 PER Crumpton (1979)
U-43.47 172.53 PER Deacon (1979)
Lestes congener Hagen U 52.25 -106.50 PER Sawchyn & Gillott (1974a)
U43.78 -72.26 PER Stoks & McPeek (2003)
disjunctus Selys U 53.18 -115.42 PER Baker & Clifford (1981)
U52.25 -106.50 PER Sawchyn & Gillott (1974b)
U51.08 -114.12 PER Krishnaraj & Pritchard (1995)
U43.78 -72.26 TEM Stoks & McPeek (2003)
U35.08 -83.17 PER Ingram (1976)
dryas Kirby U 55.93 12.32 TEM Wesenberg-Lund (1913)
U52.25 -106.50 PER Sawchyn & Gillott (1974b)
U43.78 -72.26 TEM Stoks & McPeek (2003)
eurinus Say U 46.00 -74.00 PER Pilon et al. (1993)
U43.78 -72.26 PER Stoks & McPeek (2003)
U36.07 -79.78 PER Lutz (1968)
inaequalis Walsh U 43.78 -72.26 PER Stoks & McPeek (2003)
rectangularis Say U 43.78 -72.26 PER Stoks & McPeek (2003)
U41.65 -80.42 PER Gower & Kormondy (1963)
sponsa (Hansemann) U 55.93 12.32 TEM Wesenberg-Lund (1913)
U52.50 15.00 TEM Münchberg (1933)
U51.47 -0.98 PER Corbet (1956)
tenuatus Rambur U 10.00 -84.00 TEM Paulson (1983)
unguiculatus Hagen U 52.25 -106.50 PER Sawchyn & Gillott (1974b)
U43.78 -72.26 PER Stoks & McPeek (2003)
vigilax Selys U 43.78 -72.26 PER Stoks & McPeek (2003)
U35.08 -83.17 PER Ingram (1976)
International Journal of Odonatology 9(1) 2006: 1-44 33
Voltinism of Odonata
Taxon Voltinism Lat Long Habitat Reference
Lestes virgatus (Burmeister) U 9.68 8.70 TEM Gambles (1960)
viridis (Vander Linden) U 52.50 15.00 PER Münchberg (1933)
U47.00 8.00 PER Robert (1958)
U37.37 -4.75 TEM Ferreras-R. & García R. (1995)
U37.55 -5.11 TEM Agüero-Pelegrin et al. (1999)
U37.53 -5.08 TEM Agüero-Pelegrin et al. (1999)
Platylestes praemorsus (Hagen) U 30.50 78.00 TEM Kumar (1976)
Sympecma fusca (Vander Linden) U 52.50 15.00 PER Münchberg (1933)
U47.00 8.00 PER Robert (1958)
paedisca (Brauer) U 52.25 5.40 PER Prenn (1928)
HEMIPHLEBIIDAE
Hemiphlebia mirabilis Selys U -38.92 146.28 PER Sant & New (1988)
COENAGRIONIDAE
Aciagrion migratum (Selys) B 35.00 129.00 PER Asahina (1991)
B34.78 135.45 Kansai Research Group (1977)
Agriocnemis dabreui Fraser B 13.87 100.53 PER Asahina et al. (1972)
f. femina Brauer B 13.87 100.53 TEM Asahina et al. (1972)
femina oryzae Lieftinck B 34.17 131.58 TEM Ikeda & Sawano (1965)
Agriocnemis pygmaea (Rambur) B 13.87 100.53 PER Asahina et al. (1972)
B30.50 78.00 TEM Kumar (1972)
M30.50 78.00 TEM Kumar (1976)
Amphiagrion abbreviatum (Selys) U 51.17 -115.57 LOT Pritchard et al. (2000)
Amphiallagma parvum (Selys) M 23.83 78.72 PER Srivastava & Suri Babu (1994)
Argia vivida Hagen P 50.83 -117.92 LOT Pritchard (1989)
S42.40 -112.73 LOT Pritchard (1989)
U50.60 -117.87 LOT Pritchard (1989)
Ceriagrion auranticum Fraser B 13.87 100.53 PER Asahina et al. (1972)
coromandelianum (Fabricius) M 30.50 78.00 TEM Kumar (1972)
M23.83 78.72 PER Suri Babu & Srivastava (1994)
tenellum (de Villers) S 50.82 -1.58 PER Corbet (1957b)
Coenagrion angulatum Walker U 52.25 -106.50 PER Sawchyn & Gillott (1975)
armatum (Charpentier) S 63.83 20.25 PER Johansson & Norling (1994)
caerulescens (Fonscolombe) U 37.37 -4.75 LOT Ferreras-R. & García-R. (1995)
hastulatum (Charpentier) P 67.83 21.67 PER Norling (1984c)
S63.83 20.25 PER Johansson & Norling (1994)
S58.70 16.52 PER Norling (1984c)
U63.83 20.25 PER Johansson & Norling (1994)
U58.70 16.52 PER Norling (1984c)
U47.00 8.00 PER Robert (1958)
mercuriale (Charpentier) S 50.82 -1.58 LOT Corbet (1957b)
puella (Linnaeus) S 53.47 -2.38 PER Parr (1970)
U53.47 -2.38 PER Parr (1970)
U48.27 15.70 PER Waringer & Humpesch (1984)
U47.00 8.00 PER Robert (1958)
U41.73 12.40 PER Nicolai & Carchini (1985)
pulchellum (Vander Linden) U 47.00 8.00 PER Robert (1958)
resolutum (Selys) S 53.18 -115.42 PER Baker & Clifford (1981)
U53.18 -115.42 PER Baker (1982)
U52.25 -106.50 PER Sawchyn & Gillott (1975)
U51.08 -114.12 PER Krishnaraj & Pritchard (1995)
International Journal of Odonatology 9(1) 2006: 1-44
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Corbet, Suhling & Soendgerath
Taxon Voltinism Lat Long Habitat Reference
Enallagma aspersum (Hagen) U 43.58 -71.67 PER McPeek et al. (2001)
U35.08 -83.17 PER Ingram & Jenner (1976)
boreale Selys S 53.18 -115.42 PER Baker & Clifford (1982)
S49.32 -122.55 PER Pearlstone (1973)
U49.32 -122.55 PER Pearlstone (1973)
carunculatum Morse S 44.73 -63.67 PER Paterson (1994)
cyathigerum (Charpentier) S 56.53 -3.53 PER Corbet & Chowdhury (2002)
S53.40 -2.35 PER Parr (1976)
U53.40 -2.35 PER Parr (1976)
U52.32 10.45 PER Steiner et al. (2000)
U47.48 19.17 PER Steinmann (1961)
B52.32 10.45 PER Steiner et al. (2000)
B48.68 11.18 PER Burbach (2000)
divagans Selys U 34.18 -81.12 LOT Smock (1988)
hageni (Walsh) U 35.08 -83.17 PER Ingram & Jenner (1976)
Erythromma lindenii (Selys) U 41.73 12.40 PER Nicolai & Carchini (1985)
U37.77 -5.17 PER Ferreras-Romero (1991)
B37.77 -5.17 PER Ferreras-Romero (1991)
najas (Hansemann) U 47.00 8.00 PER Robert (1958)
Ischnura asiatica Brauer B 34.78 135.45 Kansai Research Group (1977)
B34.50 135.50 PER Naraoka (1976)
aurora Brauer M 30.50 78.00 TEM Kumar (1979a)
damula Calvert B 34.00 -106.33 PER Johnson (1964)
elegans (Vander Linden) S 57.43 -7.35 PER Parr (1969)
S53.88 -0.72 PER Thompson (1978)
S53.43 -2.37 PER Parr (1973)
U53.88 -0.72 PER Thompson (1978)
U53.43 -2.37 PER Parr (1973)
U41.73 12.40 PER Nicolai & Carchini (1985)
B50.88 4.07 PER Dumont (1971)
B50.58 7.10 PER Inden-Lohmar (1997)
B43.60 3.88 PER Cassagne-Méjean (1963)
M43.57 4.57 TEM Aguesse (1955)
M43.52 4.67 TEM Aguesse (1961)
M40.68 24.73 TEM Schnapauff et al. (2000)
fountaineae Morton M 33.34 9.00 PER Jödicke (2003)
graellsii (Rambur) B 42.26 -8.38 PER Cordero-Rivera (1987)
M37.77 -5.17 TEM Montes et al. (1982)
M34.03 -6.84 TEM Azzouz & Aguesse (1990)
posita (Hagen) U 36.07 -79.78 PER Patrick (1969)
pumilio (Charpentier) U 52.13 -0.48 PER Cham (1993)
B50.58 7.10 PER Inden-Lohmar (1997)
B48.68 11.18 PER Burbach (2000)
B37.77 -5.17 TEM Aguesse (1955)
M40.68 24.73 TEM Schnapauff et al. (2000)
saharensis Aguesse M 33.45 10.13 PER Jödicke (2003)
senegalensis (Rambur) B 13.87 100.53 PER Asahina et al. (1972)
M13.87 100.53 PER Asahina et al. (1972)
verticalis (Say) U 43.58 -78.33 PER Baker & Feltmate (1987)
International Journal of Odonatology 9(1) 2006: 1-44 35
Voltinism of Odonata
Taxon Voltinism Lat Long Habitat Reference
Mortonagrion selenion (Ris) U 39.72 140.67 PER Sonehara (1994)
Paracercion calamorum (Ris) B 32.50 123.50 PER Naraoka (1976)
hieroglyphicum (Brauer) B 34.78 135.45 Kansai Research Group (1977)
sieboldii (Selys) U 40.75 140.33 PER Naraoka (1987)
U40.50 140.50 PER Naraoka (1976)
B40.75 140.33 PER Naraoka (1987)
B40.50 140.50 PER Naraoka (1976)
Pericnemis triangularis Laidlaw M 0.50 114.00 TEM Orr (1994)
Pseudagrion rubriceps Selys M 30.50 78.00 TEM Kumar (1979b)
salisburyense Ris B 26.12 28.12 LOT Chutter (1961)
Pyrrhosoma nymphula (Sulzer) S 56.73 -3.02 PER Corbet & Harvey (1989)
S54.75 -1.75 PER Lawton (1970)
S54.43 -2.97 PER Macan (1964)
S53.82 -1.00 PER Bennett & Mill (1993)
S50.82 -1.58 PER Corbet (1957a)
U56.73 -3.02 PER Corbet & Harvey (1989)
U47.00 8.00 PER Robert (1958)
Xanthocnemis zealandica (McLachlan) P -43.05 171.78 PER Deacon (1979)
S-43.47 172.53 PER Deacon (1979)
U-43.45 172.63 PER Crumpton (1979)
PLATYCNEMIDIDAE
Copera annulata (Selys) B 34.78 135.45 LOT Inoue (1979)
tokyoensis Asahina B 34.78 135.45 LOT Inoue (1979)
Lieftinckia kimminsi Lieftinck U -6.00 155.00 LOT Yule & Pearson (1995)
B-6.00 155.00 LOT Yule & Pearson (1995)
Platycnemis echigoana Asahina B 34.78 135.45 LOT Inoue (1979)
foliacea Selys B 34.78 135.45 LOT Inoue (1979)
latipes Rambur U 37.93 -4.87 PER Agüero-P. & Ferreras-R. (1992)
pennipes (Pallas) U 52.40 12.53 LOT Göcking (1999)
U52.32 10.45 PER Steiner et al. (2000)
PLATYSTICTIDAE
Drepanosticta carmichaeli (Laidlaw) U 30.50 78.00 LOT Kumar (1976)
Protosticta taipokuensis Asahina & U 22.42 114.18 LOT Asahina & Dudgeon (1987)
Dudgeon
PROTONEURIDAE
Elattoneura campioni (Fraser) U 30.50 78.00 LOT Kumar (1972)
Roppaneura beckeri Santos S -19.91 -43.93 PER Machado (1984)
PSEUDOSTIGMATIDAE
Mecistogaster linearis (Fabricius) U 8.95 -79.57 TEM Fincke (1992a)
ornata Rambur U 8.95 -79.57 TEM Fincke (1992b)
Megaloprepus caerulatus (Drury) U 8.95 -79.57 TEM Fincke (1992b)
B8.95 -79.57 TEM Fincke (1992b)
M8.95 -79.57 TEM Fincke (1992b)
ANISOZYGOPTERA
EPIOPHLEBIIDAE
Epiophlebia laidlawi Tillyard P 28.00 85.50 LOT Asahina (1982)
superstes (Selys) P 32.22 130.75 LOT Tabaru (1984)
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Corbet, Suhling & Soendgerath
Taxon Voltinism Lat Long Habitat Reference
ANISOPTERA
PETALURIDAE
Petalura gigantea Leach P -33.35 150.28 PER Tillyard (1911)
Tanypteryx pryeri (Selys) P 35.33 139.63 PER Taketo (1971)
Uropetala chiltoni Tillyard P -42.95 171.57 PER Wolfe (1953)
AESHNIDAE
Adversaeschna brevistyla (Rambur)* U -33.83 151.17 LOT Tillyard (1916)
U-37.87 174.77 PER Winstanley (1979)
Aeshna affinis Vander Linden U 52.45 16.50 TEM Bernard & Samolag (1997)
caerulea (Ström) P 55.13 -4.42 PER Clarke (1994)
P57.67 -5.43 PER Smith et al. (2000)
P47.90 8.10 PER Sternberg (1990)
californica Calvert U 47.00 -119.50 PER Kime (1974)
cyanea (Müller) P 58.70 16.52 PER Norling (1984a)
P51.15 -0.93 PER Corbet (1959)
S55.67 13.21 PER Norling (1984a)
S51.15 -0.93 PER Corbet (1959)
S48.58 7.75 PER Schaller (1960)
S37.93 -4.87 PER Ferreras-R. & Puchol-C. (1995)
U52.83 7.83 PER Jödicke (1999)
U48.58 7.75 PER Schaller (1960)
U48.08 -1.68 PER Blois (1985)
U47.32 8.78 PER Wildermuth (1994)
U47.00 8.00 PER Robert (1958)
grandis (Linnaeus) S 52.50 15.00 PER Münchberg (1930b)
isoceles (Müller) S 52.50 15.00 PER Münchberg (1930b)
juncea (Linnaeus) P 67.83 21.67 PER Norling (1984a)
P54.43 -2.97 PER Macan (1964)
P47.90 8.10 PER Sternberg (1990)
P48.17 -88.50 PER Van Buskirk (1992)
P48.17 -88.50 PER Van Buskirk (1993)
P47.00 8.00 PER Robert (1958)
P36.70 137.80 PER Kurata (1980)
P35.98 139.08 PER Arai & Murabayashi (1983)
S58.70 16.52 PER Norling (1984a)
S54.43 -2.97 PER Macan (1964)
S48.17 -88.50 PER Van Buskirk (1993)
mixta Latreille U 52.25 5.40 PER Münchberg (1930a)
U47.00 8.00 PER Robert (1958)
U37.88 -4.77 PER Munoz-P. & Ferreras-R. (1996)
U36.85 8.38 PER Cheriak (1993)
U36.85 8.38 PER Samraoui et al. (1998)
U34.50 -6.00 PER Jacquemin (1987)
multicolor Hagen U 47.00 -119.50 PER Kime (1974)
nigroflava Martin P 36.70 137.80 PER Kurata (1980)
subarctica elisabethae Djakonov P 47.90 8.10 PER Sternberg (1995)
tuberculifera Walker S 38.57 -79.03 PER Halverson (1984)
umbrosa Walker S 38.57 -79.03 PER Halverson (1984)
S36.52 -82.62 PER Johnson et al. (1980)
International Journal of Odonatology 9(1) 2006: 1-44 37
Voltinism of Odonata
*Syn. Aeshna b.
Taxon Voltinism Lat Long Habitat Reference
Aeshna viridis Eversmann P 55.67 13.18 PER Norling (1971)
S55.67 13.18 PER Norling (1971)
S52.50 15.00 PER Münchberg (1930b)
Anax ephippiger (Burmeister) B 47.67 7.52 TEM Hunger & Schiel (1999)
B43.52 4.67 TEM Katzur (1998)
B40.68 24.73 TEM Schnapauff et al. (2000)
B33.54 8.09 PER Jödicke (2003)
B9.68 8.70 TEM Gambles (1960)
immaculifrons Rambur U 30.50 78.00 TEM Kumar (1976)
imperator Leach S 51.47 -0.98 PER Corbet (1957c)
U52.30 10.78 PER Martens (1986)
U51.47 -0.98 PER Corbet (1957c)
U51.45 -2.58 PER Holmes & Randolph (1994)
U48.08 -1.68 TEM Blois (1985)
U47.55 7.58 PER Portmann (1921)
U47.00 8.00 PER Robert (1958)
U36.85 8.38 PER Cheriak (1993)
B41,10 0,92 PER R. Jödicke (pers. comm.)
julius Brauer B 34.78 135.45 Kansai Research Group (1975)
junius (Drury) U 47.00 -119.50 PER Kime (1974)
U43.87 -80.87 PER Trottier (1971)
U40.33 -86.10 PER Wissinger (1988)
nigrofasciatus Oguma U 35.97 139.08 TEM Arai (1991)
papuensis (Burmeister) U -36.25 147.00 LOT Hawking & New (1996)
U-31.93 115.83 TEM Hodgkin & Watson (1958)
B-31.93 115.83 TEM Hodgkin & Watson (1958)
parthenope Selys S 52.50 15.00 TEM Münchberg (1936)
S47.00 8.00 TEM Robert (1958)
S49.00 11.50 TEM Werzinger & Werzinger (2001)
B47.67 7.52 TEM Hunger & Schiel (1999)
S47.17 8.07 TEM Wuest-Graf (2003)
B43.52 4.67 TEM Katzur (1998)
B40.68 24.73 TEM Schnapauff et al. (2000)
B33.52 9.56 PER Jödicke (2003)
tristis Hagen B 9.68 8.70 TEM Gambles (1960)
Austroaeschna unicornis (Martin) U -36.25 147.00 LOT Hawking & New (1996)
Basiaeschna janata (Say) S 36.52 -82.62 PER Johnson et al. (1980)
Boyeria irene (Fonscolombe) S 37.93 -4.87 LOT Ferreras-Romero (1997)
vinosa (Say) S 34.18 -81.12 LOT Smock (1988)
Brachytron pratense (Müller) P 52.50 15.00 PER Münchberg (1930b)
S52.50 15.00 PER Münchberg (1930b)
U51.45 -2.58 PER Holmes (1984)
Coryphaeshna perrensi (McLachlan) U -22.93 -42.83 PER Carvalho (1992)
Dendroaeschna conspersa (Tillyard) U -33.83 151.17 LOT Tillyard (1916)
Gynacantha membranalis Karsch U 8.95 -79.57 TEM Fincke (1992b)
B8.95 -79.57 TEM Fincke (1992b)
vesiculata Karsch U 9.68 8.70 TEM Gambles (1960)
Indaeschna grubaueri (Förster) B 0.50 114.00 TEM Orr (1994)
Limnetron debile (Karsch) S -22.45 -42.93 LOT Assis et al. (2000)
International Journal of Odonatology 9(1) 2006: 1-44
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Corbet, Suhling & Soendgerath
Taxon Voltinism Lat Long Habitat Reference
Nasiaeschna pentacantha (Rambur) U 29.90 -82.32 LOT Dunkle (1985)
Oplonaeschna armata (Hagen) P 34.05 -106.88 LOT Johnson (1968)
Planaeschna milnei (Selys) P 36.00 139.50 LOT Arai (1988)
P41.70 140.45 LOT Yokohama (2001)
GOMPHIDAE
Anisogomphus maacki (Selys) P 34.80 137.78 LOT Fukui (1982)
S34.80 137.78 LOT Fukui (1982)
Asiagomphus melaenops (Selys) S 36.50 137.87 LOT Kurata (1971)
S34.80 137.78 LOT Fukui (1982)
pryeri (Selys) P 35.02 135.17 LOT Aoki (1999)
P34.68 135.17 LOT Aoki (1993)
Austrogomphus cornutus Watson S -36.25 147.00 LOT Hawking & New (1996)
ochraceus (Selys) S -36.25 147.00 LOT Hawking & New (1996)
Davidius nanus (Selys) P 34.80 137.78 LOT Fukui (1982)
S34.80 137.78 LOT Fukui (1982)
Dromogomphus spinosus Selys S 36.52 -82.62 PER Mahato & Johnson (1991)
U36.52 -82.62 PER Mahato & Johnson (1991)
Gomphus flavipes (Charpentier) P 52.83 14.78 LOT Müller (1995)
P52.50 15.00 LOT Münchberg (1932a)
P51.93 46.00 LOT Popowa (1923)
S52.83 14.78 LOT Müller (1995)
lividus Selys S 34.18 -81.12 LOT Smock (1988)
pulchellus Selys P 52.27 10.52 LOT Suhling (1994)
S52.27 10.52 LOT Suhling (1994)
S37.37 -4.75 LOT Ferreras-R.& García-R. (1995)
vulgatissimus (Linnaeus) P 53.47 13.10 LOT Müller et al. (2000)
P53.43 12.92 LOT Müller et al. (2000)
P52.83 14.78 LOT Müller (1995)
P52.62 8.88 LOT Kern (1999)
P52.62 8.88 LOT Müller et al. (2000)
P52.50 15.00 LOT Münchberg (1932a)
P52.47 10.93 LOT Müller et al. (2000)
P52.43 10.38 LOT Müller et al. (2000)
P52.11 7.54 LOT Artmeyer (1997)
P52.10 7.68 LOT Müller et al. (2000)
P49.73 10.95 LOT Müller et al. (2000)
P48.52 12.08 LOT Müller et al. (2000)
P48.50 7.84 LOT Foidl et al. (1993)
P45.80 4.98 LOT Müller et al. (2000)
S52.83 14.78 LOT Müller (1995)
S52.68 14.78 LOT Müller et al. (2000)
S52.25 7.47 LOT Müller et al. (2000)
Hagenius brevistylus Selys P 32.17 -91.88 LOT Wright (1944)
Heliogomphus scorpio (Ris) S 22.42 114.18 LOT Dudgeon (1989a)
U22.42 114.18 LOT Dudgeon (1989a)
Hemigomphus gouldii (Selys) S -36.25 147.00 LOT Hawking & New (1996)
Ictinogomphus decoratus melaenops (Selys) U 1.36 103.48 PER Lieftinck (1978)
pertinax (Selys) S 34.68 135.17 PER Aoki (1997)
U33.55 133.55 PER Ishida et al. (1988)
cited in Aoki (1997)
International Journal of Odonatology 9(1) 2006: 1-44 39
Voltinism of Odonata
Taxon Voltinism Lat Long Habitat Reference
Ictinogomphus rapax (Rambur) S 30.50 78.00 PER Kumar (1985)
U30.50 78.00 PER Kumar (1985)
Lanthus vernalis Carle P 39.28 -89.10 LOT Folsom & Manuel (1983)
S39.28 -89.10 LOT Folsom & Manuel (1983)
Nihonogomphus viridis Oguma S 34.80 137.78 LOT Fukui (1982)
Onychogomphus f. forcipatus (Linnaeus) P 49.04 7.77 LOT Herden (1990)
f. unguiculatus (Vander Linden) P 43.57 4.83 LOT Suhling (2001)
S43.57 4.83 LOT Suhling (2001)
S37.37 -4.75 LOT Ferreras-R. & García-R. (1995)
modestus Selys* U 30.50 78.00 LOT Kumar (1976)
uncatus (Charpentier) P 43.57 4.83 LOT Schütte et al. (1998)
P37.93 -4.87 LOT Ferreras-Romero et al. (1999)
S43.57 4.83 LOT Schütte et al. (1998)
S37.93 -4.87 LOT Ferreras-Romero et al. (1999)
viridicostus (Oguma) S 34.80 137.78 LOT Fukui (1982)
Ophiogomphus australis Carle S 30.60 -92.05 LOT Carle (1992)
cecilia (Fourcroy) P 52.83 14.78 LOT Müller (1995)
S52.83 14.78 LOT Müller (1995)
S52.50 15.00 LOT Münchberg (1932a)
colubrinus Selys S 46.08 -87.50 LOT Cornelius & Burton (1987)
howei Bromley S 36.65 -80.98 LOT Kennedy & White (1979)
sinicus (Chao) S 22.42 114.18 LOT Dudgeon (1989a)
U22.42 114.18 LOT Dudgeon (1989a)
Paragomphus genei (Selys) U 36.78 -6.37 LOT Testard (1975)
B35.00 10.63 PER Jödicke (2001)
M33.52 7.52 PER Jödicke (2003)
M-22.42 15.73 TEM Suhling et al. (2004)
lineatus (Selys) U 30.50 78.00 LOT Kumar (1976)
Phyllogomphoides duodentatus Donnelly P 32.42 -115.08 LOT Novelo-Gutiérrez (1993)
Progomphus obscurus (Rambur) U 30.00 -97.00 LOT Phillips (2001)
Sieboldius albardae Selys S 34.80 137.78 LOT Fukui (1982)
Stylogomphus suzukii (Oguma) S 34.80 137.78 LOT Fukui (1982)
CORDULEGASTRIDAE
Cordulegaster bidentata Selys P 51.53 8.08 LOT Dombrowski (1989)
P47.97 7.95 LOT Salowsky (1989)
b. boltonii (Donovan) P 51.67 14.00 LOT Donath (1987)
P46.17 14.30 LOT Kiauta (1964)
boltonii immaculifrons Selys S 43.60 4.83 LOT Schütte (1997)
S37.93 -4.87 LOT Ferreras-R. & Corbet (1999)
dorsalis Hagen P 37.33 -121.93 LOT Kennedy (1917)
CORDULIIDAE s.l.
Apocordulia macrops Watson S -36.25 147.00 LOT Hawking & New (1996)
Cordulia aenea (Linnaeus) P 60.17 24.97 PER Halkka (1980)
P52.50 15.00 PER Münchberg (1932b)
P47.00 8.00 PER Robert (1958)
S52.50 15.00 PER Münchberg (1932b)
S47.00 8.00 PER Robert (1958)
amurensis (Selys) P 43.05 141.35 PER Ubukata (1981)
shurtleffii Scudder P 46.00 -74.00 PER Caron & Pilon (1992)
International Journal of Odonatology 9(1) 2006: 1-44
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Corbet, Suhling & Soendgerath
*Syn. Nepogomphus m.
International Journal of Odonatology 9(1) 2006: 1-44 41
Taxon Voltinism Lat Long Habitat Reference
Epitheca bimaculata sibirica Selys S 35.98 138.38 PER Sonehara (1967)
cynosura (Say) S 36.52 -82.62 PER Johnson (1986)
U36.52 -82.62 PER Johnson (1986)
marginata Selys S 35.98 138.38 PER Sonehara (1967)
princeps Hagen S 40.33 -86.10 PER Wissinger (1988)
U40.33 -86.10 PER Wissinger (1988)
Hemicordulia australiae (Rambur) S -36.87 174.77 PER Rowe (1987)
tau Selys U -31.93 115.83 TEM Hodgkin & Watson (1958)
U-36.08 148.75 PER Faragher (1980)
Macromia amphigena Selys P 34.80 137.78 LOT Fukui (1982)
S34.80 137.78 LOT Fukui (1982)
daimoji Okumura S 34.80 137.78 LOT Fukui (1982)
illinoiensis georgina (Selys) S 34.18 -81.12 LOT Smock (1988)
moorei Selys U 30.50 78.00 LOT Kumar (1976)
splendens (Pictet) S 44.12 3.93 LOT Leipelt & Suhling (2005)
Procordulia smithii (White) P -42.67 171.33 PER Deacon (1979)
Somatochlora alpestris (Selys) P 64.42 19.50 PER Johansson & Nilsson (1991)
P47.90 8.10 PER Sternberg (1990)
S47.90 8.10 PER Sternberg (1990)
arctica (Zetterstedt) P 47.90 8.10 PER Sternberg (1990)
flavomaculata (Vander Linden) P 52.50 15.00 PER Münchberg (1932b)
metallica (Vander Linden) P 47.00 8.00 PER Robert (1958)
S47.00 8.00 PER Robert (1958)
sahlbergi Trybom P 67.10 -137.00 PER Cannings & Cannings (1985)
Synthemis leachi Selys S -31.93 115.83 LOT Watson (1967)
LIBELLULIDAE
Acisoma p. ascalaphoides Rambur U 36.85 8.38 PER Cheriak (1993)
p. panorpoides Rambur B 13.87 100.53 PER Asahina et al. (1972)
M30.50 78.00 PER Kumar (1976)
Brachythemis contaminata (Fabricius) B 13.87 100.53 PER Asahina et al. (1972)
leucosticta (Burmeister) B 34.50 -6.00 PER Jacquemin (1987)
Bradinopyga geminata (Rambur) U 30.50 78.00 TEM Kumar (1973)
Celithemis elisa (Hagen) U 40.33 -86.10 PER Wissinger (1988)
fasciata Kirby U 33.00 -82.00 PER Benke & Benke (1975)
ornata (Rambur) U 33.00 -82.00 PER Benke & Benke (1975)
Crocothemis divisa Karsch U 34.80 137.78 TEM Fukui (1982)
erythraea (Brullé) B 44.00 5.00 TEM Aguesse (1960)
B43.52 4.67 TEM Aguesse (1961)
B43.52 4.67 TEM Katzur (1998)
B37.00 -6.00 TEM Montes et al. (1982)
B36.85 8.38 PER Cheriak (1993)
B33.54 8.09 PER Jödicke (2003)
B32.00 -5.00 Jacquemin & Boudot (1999)
M-22.42 15.73 TEM Suhling et al. (2004)
servilia (Drury) B 34.35 45.37 PER Sage (1960)
B13.87 100.53 PER Asahina et al. (1972)
M30.50 78.00 TEM Kumar (1976)
Diplacodes bipunctata (Brauer) U -31.93 115.83 TEM Hodgkin & Watson (1958)
haematodes (Burmeister) U -36.25 147.00 LOT Hawking & New (1996)
Voltinism of Odonata
Taxon Voltinism Lat Long Habitat Reference
Diplacodes lefebvrii (Rambur) B 33.54 8.09 PER Jödicke (2003)
nebulosa (Fabricius) B 13.87 100.53 PER Asahina et al. (1972)
trivialis (Rambur) B 13.87 100.53 PER Asahina et al. (1972)
M30.50 78.00 PER Kumar (1976)
Erythemis simplicicollis (Say) U 40.33 -86.10 PER Wissinger (1988)
B40.33 -86.10 PER Wissinger (1988)
B33.67 -117.85 PER Morin (1984)
Erythrodiplax berenice (Drury) B 43.08 -70.08 PER Kelts (1979)
Leucorrhinia caudalis (Charpentier) U 53.23 13.58 PER Mikolajewski et al. (2004)
dubia (Vander Linden) P 67.83 21.67 PER Norling (1984b)
P58.70 16.52 PER Norling (1984b)
P47.90 8.10 PER Sternberg (1990)
S63.83 20.25 PER Johansson (2000)
S36.70 137.80 PER Kurata (1980)
intacta (Hagen) U -43.53 80.02 PER Deacon (1975)
U40.33 -86.10 PER Wissinger (1988)
pectoralis (Charpentier) P 47.32 8.78 PER Wildermuth (1994)
S47.32 8.78 PER Wildermuth (1994)
Libellula deplanata Rambur S 33.00 -82.00 PER Benke & Benke (1975)
depressa Linnaeus S 48.08 -1.68 TEM Blois (1985)
U52.83 14.78 PER Weisheit (1995)
U52.30 10.78 PER Martens (1986)
U51.45 -2.58 PER Holmes & Randolph (1994)
U47.33 8.23 PER Winsland (1991)
U47.00 8.00 PER Robert (1958)
fulva Müller S 44.00 6.00 PER Aguesse (1960)
incesta Hagen S 33.00 -82.00 PER Benke & Benke (1975)
luctuosa Burmeister U 40.33 -86.10 PER Wissinger (1988)
pulchella Drury U 40.33 -86.10 PER Wissinger (1988)
quadrimaculata Linnaeus U 52.30 10.78 PER Martens (1986)
U47.32 8.78 PER Wildermuth (1994)
Lyriothemis cleis (Brauer) M 0.50 114.00 TEM Orr (1994)
Malgassophlebia aequatoris Legrand B 0.56 12.86 LOT Legrand (1979)
Neurothemis intermedia atalanta (Ris) B 13.87 100.53 PER Asahina et al. (1972)
tullia feralis (Burmeister) B 13.87 100.53 PER Asahina et al. (1972)
t. tullia (Drury) U 30.50 78.00 TEM Kumar (1988)
M5.13 100.50 TEM Che Salmah et al. (1999)
Orthetrum albistylum (Selys) U 47.33 8.23 TEM Wildermuth et al. (1986)
B40.68 24.73 TEM Schnapauff et al. (2000)
brunneum (Fonscolombe) U 30.50 78.00 LOT Kumar (1976)
B30.50 78.00 LOT Kumar (1976)
caledonicum (Brauer) U -36.25 147.00 LOT Hawking & New (1996)
cancellatum (Linnaeus) S 52.50 13.42 PER Schmidt (1982)
B43.52 4.67 TEM Katzur (1998)
chrysostigma (Burmeister) B 34.23 7.55 PER Jödicke (2003)
M-22.42 15.73 TEM Suhling et al. (2004)
coerulescens anceps (Schneider) U 37.37 -4.75 LOT Ferreras-R. & García-R. (1995)
B34.23 7.55 PER Jödicke (2003)
japonicum (Uhler) U 36.72 136.52 PER Watanabe (1986)
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Taxon Voltinism Lat Long Habitat Reference
Orthetrum pruinosum (Burmeister) M 30.50 78.00 PER Kumar (1988)
sabina (Drury) B 30.50 78.00 TEM Kumar (1976)
B13.87 100.53 PER Asahina et al. (1972)
trinacria (Selys) M 34.50 -6.00 PER Jacquemin (1987)
Pachydiplax longipennis (Burmeister) S 35.90 -79.05 PER Eller (1963)
U35.90 -79.05 PER Eller (1963)
B40.33 -86.10 PER Wissinger (1988)
B33.67 -117.85 PER Morin (1984)
Palpopleura lucia (Drury) M 7.28 30.90 PER Hassan (1976)
Pantala flavescens (Fabricius) U 40.33 -86.10 PER Wissinger (1988)
U-37.23 145.92 PER Hawking & Ingram (1994)
M30.50 78.00 TEM Kumar (1984)
M-22.42 15.73 TEM Suhling et al. (2004)
Perithemis tenera (Say) U 40.33 -86.10 PER Wissinger (1988)
B40.33 -86.10 PER Wissinger (1988)
B33.67 -117.85 PER Morin (1984)
Plathemis lydia (Drury) U 40.33 -86.10 PER Wissinger (1988)
Sympetrum commixtum (Selys) B 30.50 78.00 TEM Kumar (1979a)
danae (Sulzer) U 48.23 15.68 PER Waringer (1983)
depressiusculum (Selys) U 47.00 8.00 TEM Robert (1958)
U43.52 4.67 TEM Katzur (1998)
flaveolum (Linnaeus) B 47.83 16.75 TEM Schmidt (1982)
fonscolombii (Selys) B 43.52 4.67 TEM Aguesse (1961)
B32.00 -5.00 Jacquemin & Boudot (1999)
B34.50 -6.00 TEM Jacquemin (1987)
B33.25 8.48 PER Jödicke (2003)
B47.67 7.52 TEM Hunger & Schiel (1999)
B40.68 24.73 TEM Schnapauff et al. (2000)
M43.52 4.67 TEM Katzur (1998)
M37.00 -6.00 PER Montes et al. (1982)
internum Montgomery U 43.38 -80.48 TEM Peterson (1975)
meridionale Selys U 36.85 8.38 PER Cheriak (1993)
U36.85 8.38 PER Samraoui et al. (1998)
pedemontanum (Müller in Allioni) U 52.50 15.00 PER Münchberg (1930a)
rubicundulum (Say) U 43.38 -80.48 TEM Peterson (1975)
sanguineum (Müller) U 51.47 -0.98 TEM Clausnitzer (1974)
U47.00 8.00 TEM Robert (1958)
U36.85 8.38 PER Cheriak (1993)
sinaiticum Dumont U 33.54 8.09 PER Jödicke (2003)
striolatum imitoides Bartenef U 35.08 135.75 TEM Matsura et al. (1995)
s. striolatum (Charpentier) U 50.47 -1.45 PER Corbet (1956)
U47.00 8.00 TEM Robert (1958)
U36.85 8.38 PER Samraoui et al. (1998)
B51.30 6.27 TEM Jödicke & Thomas (1993)
vicinum (Hagen) U 40.33 -86.10 TEM Wissinger (1988)
U35.90 -79.05 TEM Boehms (1971)
International Journal of Odonatology 9(1) 2006: 1-44 43
Voltinism of Odonata
Taxon Voltinism Lat Long Habitat Reference
Sympetrum vulgatum (Linnaeus) U 47.48 19.17 TEM Steinmann (1961)
Tramea lacerata Hagen U 40.33 -86.10 PER Wissinger (1988)
B33.67 -117.85 PER Morin (1984)
virginia (Rambur) U 30.50 78.00 TEM Kumar (1989)
Trithemis annulata (Palisot de Beauvois) B 37.27 -6.95 PER Hartung (1985)
B37.37 -4.75 PER Agüero-P. & Ferreras-R. (1992)
M33.54 8.09 PER Jödicke (2003)
kirbyi ardens (Gerstäcker) M 33.52 7.52 PER Jödicke (2003)
M-22.42 15.73 TEM Suhling et al. (2004)
pallidinervis (Kirby) B 13.87 100.53 TEM Asahina et al. (1972)
Urothemis assignata (Selys) M 7.28 30.90 PER Hassan (1977)
Zygonyx iris insignis (Kirby) U 22.33 114.17 LOT Dudgeon & Wat (1986)
Zyxomma petiolatum Rambur U 30.50 78.00 PER Kumar (1972)
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