The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae)
The life cycle types of mayflies of the subfamily Baetinae in Eastern Ukraine are determined. The scheme for identification of larval age structure is given. The age structure changes of Baetis braaschi, B. nexus and Alainites muticus larvae during the year are described in detail. The flight period...
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Інститут зоології ім. І.І. Шмальгаузена НАН України
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irk-123456789-1096682016-12-07T03:02:10Z The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) Martynov, A.V. Фауна и систематика The life cycle types of mayflies of the subfamily Baetinae in Eastern Ukraine are determined. The scheme for identification of larval age structure is given. The age structure changes of Baetis braaschi, B. nexus and Alainites muticus larvae during the year are described in detail. The flight periods of imagoes throughout the studied region are delimited for almost all species of the subfamily. B. braaschi is shown to have the bivoltine life cycle, with its populations overwintering as the larval stage (MBws) independently on temperature conditions in the water body. The peculiarities of oviposition are described for this species. The following aspects of life cycles were revealed: nymphs, subimagoes and imagoes of different generations of B. braaschi differ considerably in size; the voltinity of B. vernus in the region can vary (Us and/or MBss), mainly due to temperature conditions of the water bodies, where its development takes place; geographically isolated populations of A. muticus in the Eastern Ukraine reproduce only by thelytoky. 2013 Article The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) / A.V. Martynov // Вестник зоологии. — 2013. — Т. 47, № 1. — С. 39–48. — Бібліогр.: 33 назв. — англ. 0084-5604 DOI 10.2478/vzoo-2013-0003 http://dspace.nbuv.gov.ua/handle/123456789/109668 595.734(477.52/6) en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Фауна и систематика Фауна и систематика Martynov, A.V. The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) Вестник зоологии |
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The life cycle types of mayflies of the subfamily Baetinae in Eastern Ukraine are determined. The scheme for identification of larval age structure is given. The age structure changes of Baetis braaschi, B. nexus and Alainites muticus larvae during the year are described in detail. The flight periods of imagoes throughout the studied region are delimited for almost all species of the subfamily. B. braaschi is shown to have the bivoltine life cycle, with its populations overwintering as the larval stage (MBws) independently on temperature conditions in the water body. The peculiarities of oviposition are described for this species. The following aspects of life cycles were revealed: nymphs, subimagoes and imagoes of different generations of B. braaschi differ considerably in size; the voltinity of B. vernus in the region can vary (Us and/or MBss), mainly due to temperature conditions of the water bodies, where its development takes place; geographically isolated populations of A. muticus in the Eastern Ukraine reproduce only by thelytoky. |
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Martynov, A.V. |
| author_facet |
Martynov, A.V. |
| author_sort |
Martynov, A.V. |
| title |
The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) |
| title_short |
The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) |
| title_full |
The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) |
| title_fullStr |
The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) |
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The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) |
| title_sort |
life cycles of mayflies of the eastern ukraine. subfamily baetinae (ephemeroptera, baetidae) |
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Інститут зоології ім. І.І. Шмальгаузена НАН України |
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2013 |
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Фауна и систематика |
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http://dspace.nbuv.gov.ua/handle/123456789/109668 |
| citation_txt |
The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae) / A.V. Martynov // Вестник зоологии. — 2013. — Т. 47, № 1. — С. 39–48. — Бібліогр.: 33 назв. — англ. |
| series |
Вестник зоологии |
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AT martynovav thelifecyclesofmayfliesoftheeasternukrainesubfamilybaetinaeephemeropterabaetidae AT martynovav lifecyclesofmayfliesoftheeasternukrainesubfamilybaetinaeephemeropterabaetidae |
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2025-07-07T23:27:59Z |
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2025-07-07T23:27:59Z |
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1837032674829533184 |
| fulltext |
UDC 595.734(477.52/6)
THE LIFE CYCLES OF MAYFLIES OF THE EASTERN UKRAINE.
SUBFAMILY BAETINAE (EPHEMEROPTERA, BAETIDAE)
A. V. Martynov
Schmalhausen Institute of Zoology, NAS of Ukraine,
vul. B. Khmelnytskogo, 15, Kyiv, 01601 Ukraine
E-mail: martynov_av@ukr.net
The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae (Ephemeroptera, Baetidae). Martynov A. V. –
The life cycle types of mayflies of the subfamily Baetinae in Eastern Ukraine are determined. The scheme
for identification of larval age structure is given. The age structure changes of Baetis braaschi, B. nexus
and Alainites muticus larvae during the year are described in detail. The flight periods of imagoes through-
out the studied region are delimited for almost all species of the subfamily. B. braaschi is shown to have
the bivoltine life cycle, with its populations overwintering as the larval stage (MBws) independently on
temperature conditions in the water body. The peculiarities of oviposition are described for this species.
The following aspects of life cycles were revealed: nymphs, subimagoes and imagoes of different gener-
ations of B. braaschi differ considerably in size; the voltinity of B. vernus in the region can vary (Us and/or
MBss), mainly due to temperature conditions of the water bodies, where its development takes place;
geographically isolated populations of A. muticus in the Eastern Ukraine reproduce only by thelytoky.
Ke y wo r d s: mayflies, Ephemeroptera, life cycles, bionomy, parthenogenesis, Eastern Ukraine.
Æèçíåííûå öèêëû ïîä¸íîê Âîñòî÷íîé Óêðàèíû. Ïîäñåìåéñòâî Baetinae (Ephemeroptera, Baetidae).
Ìàðòûíîâ À. Â. – Óñòàíîâëåíû òèïû æèçíåííûõ öèêëîâ ïðåäñòàâèòåëåé ïîäñåìåéñòâà Baetinae,
êîòîðûå îíè äåìîíñòðèðóþò â ïðåäåëàõ Âîñòî÷íîé Óêðàèíû. Ïîäàíà ñõåìà îïðåäåëåíèÿ âîçðàñò-
íîé ñòðóêòóðû ëè÷èíîê. Äåòàëüíî îïèñàíî èçìåíåíèå âîçðàñòíîé ñòðóêòóðû ëè÷èíîê Baetis bra-
aschi, B. nexus è Alainites muticus íà ïðîòÿæåíèè ãîäà. Ïðèâåäåíû ïåðèîäû ë¸òà èìàãî â ðåãèîíå
ïðàêòè÷åñêè äëÿ âñåõ âèäîâ ïîäñåìåéñòâà. Óñòàíîâëåíî, ÷òî íåçàâèñèìî îò òåìïåðàòóðíîãî
ðåæèìà âîäîòîêà B. braaschi èìååò áèâîëüòèííûé æèçíåííûé öèêë ñ çèìîâêîé íà ñòàäèè ëè÷èí-
êè (MBws). Îïèñàíû îñîáåííîñòè îòêëàäêè ÿèö äàííûì âèäîì. Âûÿâëåíû ñëåäóþùèå àñïåêòû
æèçíåííûõ öèêëîâ: íèìôû, ñóáèìàãî è èìàãî ðàçíûõ ãåíåðàöèé B. braaschi ñóùåñòâåííî îòëè-
÷àþòñÿ ðàçìåðàìè; âîëüòèííîñòü B. vernus â ðåãèîíå ìîæåò âàðüèðîâàòü (Us è/èëè MBss) ãëàâ-
íûì îáðàçîì â çàâèñèìîñòè îò òåìïåðàòóðíîãî ðåæèìà âîäíîãî îáúåêòà, â êîòîðîì ïðîõîäèò åãî
ðàçâèòèå; ãåîãðàôè÷åñêè èçîëèðîâàííûå ïîïóëÿöèè A. muticus Âîñòî÷íîé Óêðàèíû ðàçìíîæàþò-
ñÿ èñêëþ÷èòåëüíî ïðè ïîìîùè òåëèòîêèè.
Êëþ÷ å âûå ñ ë î â à: ïîä¸íêè, Ephemeroptera, æèçíåííûå öèêëû, áèîíîìèÿ, ïàðòåíîãåíåç,
Âîñòî÷íàÿ Óêðàèíà.
Introduction
Despite the short period of detailed research on mayflies, much attention is devoted to their life cycles
and several classifications of life cycles are worked out by far (Landa, 1968; Sowa, 1975; Clifford, 1982). Clifford’s
(1982) classification is the most widely used at present: all known mayfly life cycles are divided into 14 groups,
belonging to 5 categories. The life cycle type of one species is shown to vary within its areal. For example,
Centroptilum luteolum in the range of its areal has two different life cycle types, Alainites muticus, whereas Baetis
vernus has four, and Cloeon dipterum has five life cycle types (Clifford, 1982). Considering that the variation of
life cycle is typical for most mayfly species, our research aimed to study life cycles of the Eastern Ukraine Baetinae.
The obtained data allow determining the most appropriate time for faunistic studies in the region. These data
could be used in the bioindicational exploration and to study variation of life cycles through the areal.
Material and methods
The life cycles classification of is used in this paper follows Clifford (1982). To study the age structure
changes of 3 species, the larvae were collected in model water bodies (table 1). Fourteen samples of each species
were taken with different periodicity from March 2010 to March 2011, but not less than once per month (except
Vestnik zoologii, 47(1): e-35—e-44, 2013
DOI 10.2478/vzoo-2013-0003
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of winter months in some cases). The coordinates of model water bodies are given according to Google Earth
(http://earth.google.com).
Changes of the age structure for mayfly larvae during the year were studied following Jop's (1981) sys-
tem with some alternations, which are displayed in fig. 1 and described below. The spreadsheet designed by
A. Buffangi on the base of Microsoft Office Excel (Tools, 2008) was used to generate histograms, which reflect
the larval age structure during the year.
Results and discussion
Age structure definition. To define the age structure of mayfly larvae, K. Jop (1981)
system was applied as the basis, except only 10 groups of larval ages are shown for the lar-
vae of Baetidae species (as for Leptophlebiidae): from L0 to L9 (fig. 1), as the larvae ready
to become winged with completely developed wings inside the pads, occurred in the L9—NIII
age groups of Jop’s system. All Baetidae larvae, whose wing pads by developmental degree
correspond to L9—NIII in Jop’s system were considered to be the variant of L9 in this study.
To show the larvae, which are ready to emerge (having completely developed wings inside
the pads), and/or winged stages the inclined black arrow was added above the dates of
sampling in the histograms of species age structure change (fig. 2, 7).
Life cycles. According to Martynov (2010), the subfamily Baetinae is represented in
the Eastern Ukraine by seven species: Baetis braaschi Zimmermann, 1980, B. vernus Curtis,
1834, B. fuscatus (Linnaeus, 1761), B. buceratus Eaton, 1870, B. nexus Navás, 1918,
Nigrobaetis digitatus (Bengtsson, 1912), and Alainites muticus (Linnaeus, 1758). The life
cycles of some of these species remain poorly studied throughout their whole areals, and
such data on all these species from the Eastern Ukraine were absent. Furthermore, some
of them appear to be situated at the margin of their areal and/or are presented by geo-
graphically separated populations in the studied region.
Baetis braaschi Zimmermann, 1980. Life cycle has not been previously studied in
details. Our preliminary studies carried out in the mountain regions of the Crimea, revealed
the bivoltine life cycle with wintering as the larval stage for the species (MBws) (Sroka
et al., 2012). The present paper contains the results of minute research on B. braaschi
life cycle throughout the Eastern Ukraine. Age structure changes of species during the
year were studied on different populations, which inhabit potamal and rhithral zones of
streams (table 1). The corresponding water temperature change in model water bodies
are presented in fig. 3, 4. As this study shows, such a big difference in temperature con-
ditions of model streams has no influence in species voltinity. B. braaschi has the bivol-
tine life cycle and overwinters as the larval stage (MBws) in both types of waterways (fig. 2).
These populations overwinter in elder, middle and junior larval ages. Larvae do not
grow actively during the wintering. As the spring comes, the growth rate increases and
e-36 A. V. Martynov
Ta b l e 1. Model water bodies and material used for analysis of age structure change of mayfly larvae during the
year
Òàáëèöà 1. Ìîäåëüíûå âîäîòîêè è îáú¸ì ìàòåðèàëà, èñïîëüçîâàííîãî äëÿ èçó÷åíèÿ èçìåíåíèé âîçðàñòíîé
ñòðóêòóðû ëè÷èíîê ïîä¸íîê íà ïðîòÿæåíèè ãîäà
Species Administrative location of water body Waterway type,
it’s characteristic Coordinates Number of
specimens
Baetis braaschi Donets’ka oblast, Yasynuvats’kyi
district, vicinity of Mineral’ne vil-
lage, the stream in gullied forest
rhithral stream 48°6'7'' N 37°49'44'' E 1145
Baetis braaschi Territory of Donets’k city,
Putilyvs’kyi park, the stream flowing
out from pond
potamal stream 48°3'53'' N 37°47'40'' E 745
Baetis nexus Donets’ka oblast, Yasynuvats’kyi
district, vicinity of Mineral’ne vil-
lage, the stream in the meadow
potamal stream 48°5'52'' N 37°49'52'' E 744
Alainites
muticus
Donets’ka oblast, Yasynuvats’kyi
district, vicinity of Debaltseve urban
village, Bulavina river
rhithral zone
of the river
48°18'55'' N 38°26'07'' E 622
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overwintered larvae of elder ages soon complete their germination. The larvae of first ge-
neration, been developing in the potamal zone, become winged from March to the end
of May. The larvae, whose germination took place in the rhithral zone, become winged
until the mid-June (fig. 2). The reason for such a long period, when the winged stages
and first generation larvae ready for emergence could be registered, is the wintering in
all larval ages from L0 to L8. The peak of winged stages emergence is in the end of April –
beginning of May. Emergence of the second generation subimago can insignificantly over-
lap with the end of the first generation or begins just after it, due to the long period, when
e-37The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae...
Fig. 1. The developmental degree of wing pads of larvae in different age groups: 1—3 – abdominal segments;
L0—L9 – age groups.
Ðèñ. 1. Ñòåïåíü ðàçâèòèÿ çà÷àòêîâ êðûëüåâ ó ëè÷èíîê ïîäåíîê ðàçëè÷íûõ âîçðàñòíûõ ãðóïï: 1—3 –
ñåãìåíòû áðþøêà; L0—L9 – âîçðàñòíûå ãðóïïû.
Fig. 2. The age structure change of Baetis braaschi during the year: 1 – rhithral stream (vicinity of Mineral’ne
village, 2010—2011); 2 – potamal stream (territory of Donets’k city, Putilyvs’kyi park, 2010—2011).
Ðèñ. 2. Èçìåíåíèå âîçðàñòíîé ñòðóêòóðû Baetis braaschi íà ïðîòÿæåíèè ãîäà: 1 – ðèòðàëüíûé ðó÷åé
(îêð. ñ. Ìèíåðàëüíîå, 2010—2011 ãã.); 2 – ïîòàìàëüíûé ðó÷åé (ã. Äîíåöê, Ïóòèëîâñêèé ïàðê,
2010—2011 ãã.).
1
2
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the first generation gets winged, the short period of egg stage existence and the rapid devel-
opment of the second generation larvae (owing to water temperature increase)
The winged stages of the second generation emerge until the end of October. The
most intensive emergence of winged stages of both generations is observed in the mid-
dle of day (12.00—14.00), the male swarming – from 12.00 to 17.00. The swarms are
usually seen not far from water surface or strait along the bank, and consist of no more
than 15 males. Small swarms (5—10 males) can be close to each other and remain sep-
arated. The undisturbed males do not fly up higher than 3—3.5 m in the swarm, and all
e-38 A. V. Martynov
Fig. 3. The water temperature changes in model water bodies during the year: 1 – the rhithral stream (vicin-
ity of Mineral’ne village, 2010—2011); 2 – the potamal stream (vicinity of Mineral’ne village, 2010—2011).
Ðèñ. 3. Èçìåíåíèå òåìïåðàòóðû âîäû â ìîäåëüíûõ âîäíûõ îáúåêòàõ íà ïðîòÿæåíèè ãîäà: 1 – ðèòðàëü-
íûé ðó÷åé (îêð. ñ. Ìèíåðàëüíîå, 2010—2011 ãã.); 2 – ïîòàìàëüíûé ðó÷åé (îêð. ñ. Ìèíåðàëüíîå,
2010—2011 ãã.).
Fig. 4. The water temperature change in potamal stream during a year (Donetsk City, Putilyvs’kyi park,
2010—2011).
Ðèñ. 4. Èçìåíåíèå òåìïåðàòóðû âîäû â ïîòàìàëüíîì ðó÷üå íà ïðîòÿæåíèè ãîäà (ã. Äîíåöê, Ïóòèëîâñêèé
ïàðê) (2010—2011 ãã.).
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the swarm stays at a height of 1.5—2.5 m. The number of swarming males is consider-
ably less closer to twilight.
For oviposition, B. braaschi female crawls (under the water) to the bottom side of
stones and other objects with strait surface (foam plastic blocks, flooded boards, etc.).
Then, female takes a position facing the upstream to increase the streamlining and oviposits
3000—3500 eggs. The eggs are aggregated in set, which resembles an elongated oval or
obtruncated pear (fig. 5). In general, the egg set is single-layered, double-layered only
in some parts. The female dies after the oviposition. The hatching of the first generation
larvae to overwinter, begins in September and lasts until the end of the autumn (fig. 2).
There is no considerable difference in duration of larval germination period and winged
stages flight between B. braaschi populations developed in rhithral and potamal zones
(fig. 2). The populations of species germinating in crenal zone (rheocrene) have not been
examined in this study. However, we can suggest, that thes species also have the bivol-
tine life cycle – MBws (overwintering as the larval stage) under these conditions.
The last generation larvae, as well as subimagoes and imagoes of different genera-
tions, considerably vary in size. Thus, the overwintered larvae of first generation are 2—2.5
times bigger than the second generation larvae (fig. 5) (Sroka et al., 2012). Such diffe-
rence in size of various generations is stipulated by temperature conditions, under which
the larval growth took place, and is recorded for most species with bivoltine life cycles
and larvae to overwinter (Cianciara, 1979; Humpesch, 1979; Hwang et al., 2005, etc.).
The size difference between both nymphs and winged stages of B. braaschi generations
developed in rhithral and potamal zones is absent.
Nigrobaetis digitatus (Bengtsson, 1912). The life cycle is poorly known. In Europe it
has monovoltine life cycle, with its populations overwinting as the larval stage (Uw) (Jop,
1981; Söderström, 1991; Bauernfiend, Soldán, 2012). It is difficult to determine the life
cycle type of N. digitatus in the Eastern Ukraine due to the lack of data and restricted field
period (spring). However, a lot of larvae ready for emergence and imagoes have been reg-
istered in the first decade of May. This fact can be an evidence of wintering as the larval
stage. There is no data on N. digitatus voltinity throughout the Eastern Ukraine. At pre-
sent we can only suppose that this species has monovoltine life cycle and overwinters as
the larval stage (Uw) in the studied region, as well as in other European counties.
Baetis vernus Curtis, 1834. Due to obtained data, the bivoltine life cycle of this species,
with its populations overwinting as the stage of egg (MBss) throughout the Eastern Ukraine,
can be confirmed. A similar life cycle B. vernus most often reveals in the majority of
European countries (Landa, 1968, 1969; Elliott et al., 1988; Soldán, Zahrádková, 2000;
Bauernfiend, Soldán, 2012), but in Germany this species can have two or three gener-
ations during the year (MB—MP) (Haybach, 1998). Similarly, B. vernus is shown to have
the monovoltine life cycle with wintering as the stage of egg or larva (Us, Uw) in Europe
(Bauernfiend, Soldán, 2012).
The hatching from overwintered eggs begins at the first days of April, and the winged
stages emergence gets started to the middle of May in the studied region. Emergence of
the second generation larvae is registered from the middle of July. There are no clear bounds
between the imagoes flight of first and second generations. Generally, the flight of B. ver-
nus imagoes was observed from the middle of May until the first decade of November
inclusive. As Bohle (1969) showed in the laboratory studies that duration of B. vernus
egg diapause varies from 1 to 5 months and depends on the water temperature. The lat-
ter circumstance can influence not only on the terms of generation development, but on
the species voltinity. Thus, this species has the monovoltine life cycle (Soldán, Zahrádková,
2000) in some cold-water bodies in the mountain regions of Czech Republic. In our opin-
ion, B. vernus can also have monovoltine life cycle, wintering as an egg (Us) in crenal
zone of the Eastern Ukrainian waterways, where the water temperature is stably low dur-
ing all the year. In addition, the quantitative sampling in rhithral zone of Bulavina River
during the year shows that second generation larvae of this species are much less nume-
e-39The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae...
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rous, than of the first one. Increasing of water temperature can cause of quantity reduc-
tion of second generation larvae, which is rather noxious for the species. Thus,
H. V. Bohle’s laboratory research revealed, that 15.6 ºÑ temperature causes the death of
75 % of diapausing eggs, and under 20 ºÑ about 85 % of them die. As the fig. 6 shows,
the temperature over 15 ºÑ has been registered in Bulavina river from middle of May
until the end of July. This is exactly the time, when the development of eggs (which orig-
inate the second generation of B. vernus) takes place in the river.
In addition, it is possible that the majority of eggs, laid by first generation females,
falls into a long diapause until the spring, and the small part of eggs gives rise for the sec-
ond generation larvae without any delay. If this assumption is true, some populations of
B. vernus and/or their parts have the monovoltine life cycles and overwinter as the egg
stage (Us) in the Eastern Ukraine.
Baetis fuscatus (Linnaeus, 1761). This species has a bivoltine life cycle, its po pula-
tions overwinter as the egg stage (MBss) throughout the Eastern Ukraine, as in the vast
part of Europe (Landa, 1968, 1969; Sowa, 1975; Elliott et al., 1988; Soldán, Zahrádková,
2000; Bauernfiend, Soldán, 2012). The represented phenological data are based on stud-
ies carried throughout Donetsk and Priazovskaya elevated areas. The larvae ready for emer-
gence and winged stages of the first generation were registered from the middle of May
until the end of June, and those of the second generation from the middle of August until
the first decade of October inclusively.
This species overwinters as a larva (MBws) (Clifford, 1982; Haybach, 1998) in some
European water bodies. Similarly, B. fuscatus has up to three generations in the year (MB-
e-40 A. V. Martynov
Fig. 5. The egg sets of Baetis braaschi and Alainites muticus, female nymphs of Baetis braaschi of the first and
second generations: 1 – general egg set shape of B. braaschi; 2 – group of egg sets of B. braaschi on the bot-
tom side of a stone; 3 – aggregation of Alainites muticus egg sets on the bottom surface of flooded log; 4 –
first generation nymph of B. braaschi; 5 – second generation nymph of B. braaschi. Scale bar 10 mm.
Ðèñ. 5. Êëàäêè Baetis braaschi è Alainites muticus, íèìôû ñàìîê Baetis braaschi ïåðâîãî è âòîðîãî ïîêî-
ëåíèé: 1 – îáùèé âèä êëàäêè B. braaschi; 2 – ãðóïïà êëàäîê B. braaschi íà íèæíåé ïîâåðõíîñòè êàìíÿ;
3 – ñêîïëåíèå êëàäîê Alainites muticus íà íèæíåé ïîâåðõíîñòè ïîãðóæåííîãî â âîäó áðåâíà; 4 – íèìôà
ïåðâîãî ïîêîëåíèÿ B. braaschi; 5 – íèìôà âòîðîãî ïîêîëåíèÿ B. braaschi. Ìàñøòàáíàÿ ëèíåéêà 10 ìì.
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MP) in Germany and in the big rivers of Czech Republic (Haybach, 1998; Soldán,
Zahrádková, 2000). The water temperature is a determinative factor in both cases. It is quite
possible that the species has three generations per year and/or overwinters as the larval stage
in such big rivers of the Eastern Ukraine as Siverskyi Donets, Oskol, Aydar and others.
Baetis buceratus Eaton, 1870 has two generations per year, its wintering takes place
on the stage of larva (MBws) throughout the main part of Europe, as in the Ukrainian
Carpathians. Thus, it can reveal three generations per year (MP) in some European coun-
tries under favorable conditions (Godunko, 2001; Landa, 1968, 1969; Elliott et al., 1988;
Haybach, 1998; Soldán, Zahrádková, 2000; Bauernfiend, Soldán, 2012). B. buceratus has
the bivoltine life cycle, with its population wintering as the larval stage (MBws) in the
studied region. The larvae ready for emergence and winged stages of the first generation
were recorded from the end of April until the middle of July, of the second generation –
during whole September. The first generation first instar larvae (overwintered) were observed
at the second decade of October.
Baetis nexus Navás, 1918. The life cycle has not been studied in detail due to species
rarity through the main part of Europe. Casual data were recounted in the paper on the
species population from the vicinity of Lion (France) (Camousseight, Fontaine, 1990);
the bivoltine life cycle (population overwinter in the larval stage (MBws)) was record-
ed for B. nexus . It is considered to be one of the most commonly occurring rheobionts
(Martynov, 2012) in the Eastern Ukraine, where lies current eastern boundary of it dis-
tribution. In the Eastern Ukraine, B. nexus has the life cycle similar to that in France –
MBws. Study of the age structure changes of B. nexus larvae in the potamal stream (fig. 7)
shows that it overwinters as the larvae of first and middle age (the water temperature change
in model water body is shown in fig. 3). The active growth of larvae was not observed
during the winter. When spring comes, the growth rate is restored, and the active flight
of winged stages of the first generation is registered until the middle of May. Mass hatch-
ing of the first generation larvae, characterised by rather rapid growth, begins from the
second half of May. There is no clear border between the imago flight period of the first
and second generations. The general flight period lasts from the middle of May until the
middle of September. The first generation larvae, which are going to overwinter, emerge
from October to November.
Alainites muticus (Linnaeus, 1758). Generally, the species has the bivoltine life cycle,
with its populations overwintering as the larval stage (MBws) in Europe (Landa, 1968,
e-41The Life Cycles of Mayflies of the Eastern Ukraine. Subfamily Baetinae...
Fig. 6. The water temperature change in model section of Bulavina river during the year (vicinity of Debaltseve)
(2010—2011).
Ðèñ. 6. Èçìåíåíèå òåìïåðàòóðû âîäû â ìîäåëüíîì ó÷àñòêå ðåêè Áóëàâèíà íà ïðîòÿæåíèè ãîäà
(îêð. Äåáàëüöåâî) (2010—2011).
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1969; Sowa, 1975; Soldán, Zahrádková, 2000; Bauernfiend, Soldán, 2012), but it has one
generation per year (Uw) in Scandinavia, high-mountain regions of Czech Republic and
in Alps, and tree in Atlantic Pyrenees (MP) (Ulfstrand, 1968; Thibault, 1971; Soldán,
Zahrádková, 2000).
Alainites muticus reveals the bivoltine life cycle, with its populations overwintering
as the larval stage (MBws) in the Eastern Ukraine. The age structure changes of its lar-
vae during the year has been studied on the most numerous A. muticus population, which
inhabits the rhithral zone of Bulavina River (fig. 7). The species overwinters as larvae of
junior and middle ages. One can find the junior age larvae (L0—L1) in March and April
due to hatching of small part of population in spring or due to very slow developmental
and growth rates of larvae in winter months (fig. 7). The winged stages of the first gen-
eration are registered from the middle of May until the middle of June.
The females oviposit on the bottom surface of stones and on the sunken logs.
Meanwhile, the density of egg sets can be very high and sets can be arranged one over
another (fig. 5) in the areas with a few stones on the bottom (with the shortage of places
suitable for oviposition). The second generation larvae begin to emerge in first days of
June. They develop very quickly, probably, due to the high water temperature in this peri-
od (fig. 6). The flight of winged stages of the second generation lasts from the second
half of July until the end of August. The hatching period of the first generation larvae is
rather long: it begins in August and lasts during all the autumn months.
It should be pointed out, that all known A. muticus populations in the Eastern Ukraine
reproduce by thelytoky (a form of parthenogenesis, with the unfertilized ovi develop into
females). Degrange (1960) has reported this species to propagate from time to time by
facultative parthenogenesis. In Funk's opinion, most of (maybe even all) bisexual species
of mayflies are able to reproduce by facultative parthenogenesis (Funk et al., 2010).
Though, virginal reproduction in bisexual species is generally registered occasional-
ly, being alternated with reproduction with fertilization (Harker, 1997; Funk et al., 2010).
At present, the obligate parthenogenesis is declared only for a few mayfly species from
North America, South America and Africa (Froehlich, 1969; Gibbs, 1977; Bergman,
Hilsenhoff, 1978; Gillies, Knowles, 1990, etc.). Five-year long studies on A. muticus po-
pulation in the rhithral zone of Bulavina River revealed that all its generations were repro-
ducing only by parthenogenesis.
e-42 A. V. Martynov
Fig. 7. The age structure change of Baetis nexus è Alainites muticus during the year: 1 – Baetis nexus, the pota-
mal stream (vicinity of Mineral’ne village, 2010—2011); 2 – Alainites muticus, the rhithral zone of Bulavina
river (vicinity of Debaltseve urban village, 2010—2011).
Ðèñ. 7. Èçìåíåíèå âîçðàñòíîé ñòðóêòóðû Baetis nexus è Alainites muticus íà ïðîòÿæåíèè ãîäà: 1 – Baetis
nexus, ïîòàìàëüíûé ðó÷åé (îêðåñòíîñòè ñ. Ìèíåðàëüíîå, 2010—2011 ãã.); 2 – Alainites muticus, ðèòðàëü-
íàÿ çîíà ðåêè Áóëàâèíà (îêð. ïãò Äåáàëüöåâî, 2010—2011 ãã.).
1
2
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In our opinion, A. muticus population in Bulavina River, as other populations of this
species in the Eastern Ukraine, has completely switched to obligate parthenogenetic repro-
duction.
Other populations of A. muticus in the Eastern Ukraine have not been subjected to
long-term investigation yet, but periodical sampling have not revealed males in the lar-
val, subimaginal and imaginal stages of the species.
By far, the mechanisms causing such a switch remain unknown. The studies in col-
laboration with I. A. Kozeretska and S. V. Serga (Taras Shevchenko National University
of Kyiv) to investigate the infectiousness of A. muticus populations from Bulavina River
by Wolbachia sp. and Spiroplasma sp. causing parthenogenesis and male killing (Werren
et al., 2008) gave negative results.
In addition, populations of species through the studied region are geographically iso-
lated from the main part of their areals. It is possible that in this case the display of geo-
graphical parthenogenesis is observed, which is typical for insular and isolated from the
main parts of areal populations (Victorov, 2006).
There is one more fact confirming this: we revealed the parthenogenetical popula-
tion of Nigrobaetis niger (Linnaeus, 1761) in the mountain region of the Crimea, which
is geographically isolated from the main areal. It is also possible that parthenogenesis is
caused by specific conditions observed during larval development (temperature in the first
place), which, in some authors’ opinions, mostly influence on appearance of partheno-
genesis in mayflies (Humpesch, 1980, 1981).
Conclusions
1. The life cycle types of mayflies in the subfamily Baetinae are determined. Most
species in the region of studies have bivoltine life cycles (MBws or MBss) and only some
of them have other cycles (Us or Uw). We failed to determine the life cycle types for
several species due to their rarity in the studied region.
2. B. braaschi was shown to have the bivoltine life cycle, with its populations over-
wintering as a larval stage (MBws) regardless of temperature conditions in water body.
Females of this species oviposit under the water and their eggs appear to be aggregated
into sets. Nymphs, subimagoes and imagoes of different generations of this species con-
siderably differ in size.
3. The voltinity of B. vernus in the region can vary (Us and/or MBss), mainly due
to temperature conditions of the water body, where its development takes place.
4. Geographically isolated populations of A. muticus in the Eastern Ukraine repro-
duce only by thelytoky.
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Received 10 September 2012
Accepted 21 November 2012
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