L J Borkin, A V Korshunov, G A Lada - Mass occurrence of polyploid green frogs (rana esculenta complex) in eastern ukraine - страница 1

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Russian Journal of Herpetology

Vol. 11, No. 3,2004, pp. 194-213

MASS OCCURRENCE OF POLYPLOID GREEN FROGS (Rana esculenta COMPLEX) IN EASTERN UKRAINE

Leo J. Borkin,[1] Alexey V. Korshunov,[2] Georgiy A. Lada,1,3 Spartak N. Litvinchuk,[3] Jury M. Rosanov,[4] Dmitry A. Shabanov,[5] and Alexander I. Zinenko[6]

Submitted August 20, 2004.

In eastern Ukraine, the Rana esculenta complex consists of three species: R. lessonae, R. ridibunda, and hybrid R. esculenta. The first one was rare, whereas two latter frog taxa were very common. Based on DNA flow cyto-metry, mass occurrence of the triploidy in Rana esculenta has been revealed in 14 localities of Kharkov, Do­netsk, and Lugansk Provinces. One hybrid specimen from Kharkov Province was tetraploid. All polyploids were recorded along the middle part of Seversky Donets River (above 450 km). Triploids comprised two groups with different genome composition (LLR and LRR), and were found in three types of population systems (E, R-E, and L-E-R). Geographic distribution of polyploidy in European green frogs is briefly outlined. Different meth­ods of ploidy level identification are discussed. The chromosome count and nuclear DNA cytometry provide the most reliable data.

Keywords: hybridogenetic frogs, triploidy, tetraploidy, Rana esculenta complex, DNA flow cytometry, Ukraine.

INTRODUCTION

Three taxa of European green frogs (Rana esculenta complex) are distributed in eastern Europe. These are two "normal" species - the lake frog, Rana ridibunda Pallas, 1771, and the pool frog, R. lessonae Camerano, 1882, as well as the edible frog, R. esculenta Linnaeus, 1758, which arose as a result of hybridization between the former two parental species. The complex exhibits an unusual mode of speciation, characterized by compli­cated genetic mechanisms, involving hybridization, so-called hemiclonal (or meroclonal) inheritance, polyploi-dy, unisexual and bisexual population systems of vari-

3 Department of Zoology, Tambov State University, ul. Internatsio-nal'naya, Tambov 392622, Russia. E-mail: esculenta@mail.ru.

ous kinds (e.g., Uzzell et al., 1977, 1980; Borkin et al., 1987; Vinogradov et al., 1988, 1990, 1991; Graf and Polls Pelaz, 1989a; Berger, 1990; Bucci et al., 1990; GUnther, 1990; Tunner and Heppich-Tunner, 1991; Hotz et al., 1992; Caune and Borkin, 1993; Lada, 1995; Rag-ghianti et al., 1995). Rana esculenta is considered to be a klepton (in Greek: thief), a special category of taxa of the species group, the character of which does not coin­cide with that of the biological species (Dubois and Gunther, 1982; Dubois, 1990; Gunther, 1991).

Although hybridogenetic frogs (R. esculenta) are widely known across temperate Europe from France in the west to Volga River in the east (Gunther, 1997; Bor-kin et al., 2003a, 2003b), natural triploidy has been re­corded in many populations distributed in western and central parts of Europe only (e.g., Gunther, 1970, 1975a; Knudsen and Scheel, 1975; Ebendal and Uzzell, 1982; Regnier and Neveu, 1986; Blommers-Schlosser, 1990; Plotner and Klinkhardt, 1992; Tunner and Heppich-Tunner, 1992; Berger and Berger, 1994; Mikulicek and Kotlik, 2001; Ogielska et al., 2001; Rybacki and Berger, 2001).

During two last decades, we examined by means of DNA flow cytometry numerous samples of green frogs from the territory of the former Soviet Union (Fig. 1). However, no record of triploidy in wild populations has

1026-2296/2004/1103-0194 © 2004 Folium Publishing Company

Fig. 1. The distribution of hybrid Rana esculenta in European part of the former Soviet Union (based on genome size data).

not yet been published (Borkin et al., 1987; Vinogradov et al., 1988, 1990, 1991; Caune and Borkin, 1993; Lada et al., 1995; Borkin et al., 2003a, 2003b). An enigmatic tetraploid frog only was evidenced by karyotyping from Latvia (Borkin et al., 1979). An occasional triploid, taken by S. N. Litvinchuk from Kaliningrad Province, the westernmost enclave of Russia situated between Po­land and Lithuania, has been mentioned by Borkin

(2001).

In this contribution, we present the results of our study on green frogs of eastern Ukraine, obtained main­ly on the basis of DNA flow cytometry. The aim of the paper is to give the evidence of mass occurrence ofvari-ous kinds of polyploid frogs (R. esculenta) in local populations.

MATERIAL AND METHODS

Since 1989, a total of 551 green frogs were collected in eastern Ukraine and examined by DNA flow cyto-metry. In 1989 we studied 11 specimens from Lopan River, Kharkov City, presented by A. M. Rudik. In June 1995 we sampled 24 frogs from three other localities of Kharkov Province (Zmiyov and Kharkov Districts). In June 1996 we examined 43 specimens from two locali­ties of Zmiyov and Zachepilovka Districts, Kharkov Province, Ukraine.

In 2002 - 2004 our sampling has been more exten­sive, and 473 frogs were taken from 38 localities of Kharkov, Donetsk, and Lugansk Provinces (Fig. 2, Table 1). The majority of specimens were collected at the "juvenile" stage, which covered both froglets before

Fig. 2. The distribution of members of the Rana esculenta complex in eastern Ukraine (based on genome size data). The triangle marks the Khar­kov City sample with R. ridibunda studied in 1989. The asterisk denominates the Russkii Orchik sample with three frog species studied in 1996.

TABLE 1. Genome Size Variation (in picograms) in Green Frogs collected from Eastern Ukraine in 2002 - 2004

No.      Locality Latitude (N) Longitude (E)     Type N Mean±SD Range CV, %

Kharkov Province

1

Kozievka

50°08'

35°13'

RR

1j

16.10

 

 

 

2

Gorodnoe

50°06'

35°15'

RR

16j

16.11 ±0.04

16.03 -

16.18

0.3

3

Gubarevka

50°10'

35°21'

RR

1f+10j

16.15 ±0.06

16.05 -

16.27

0.4

4

Mikhailovskoe

50°01'

35°05'

LL

3j

14.13 ±0.06

14.09-

14.20

0.4

 

 

 

 

LL?

1j

13.68

 

 

 

 

 

 

 

RR

1j

16.19

 

 

 

5

Sharovka

50°02'

35°28'

LR

3j

15.01 ±0.05

14.96-

15.04

0.3

6

Liptsy

50°13'

36°23'

LR

6m

14.89 ±0.03

14.86-

14.92

0.2

 

 

 

 

RR

3m

16.06 ±0.07

15.99-

16.14

0.5

7

Murom

49°24'

37°11'

LR

1j

14.94

 

 

 

 

 

 

 

RR

9j

16.06 ±0.09

15.88 -

16.15

0.6

8

Russkaya Lozovaya

50°10'

36°19'

LR

7j

14.97 ±0.05

14.88 -

15.04

0.3

9

Lesnoe

50°06'

36°16'

RR

1j

16.18

 

 

 

10

Tsirkuny

50°06'

36°27'

RR

2j

16.22

16.20-

16.25

 

11

Pyatikhatka

50°05'

36°17'

RR

2j

15.98

15.98 -

15.98

 

12

Ol'khovka

49°58'

36°31'

RR

1f

16.12

 

 

 

13

Bezlyudovka

49°52'

36°17'

LR

11j

15.02 ±0.07

14.93 -

15.12

0.5

14

Izbitskoe

50°12'

36°44'

LR

2f+4j

15.01 ±0.04

14.95 -

15.06

0.3

 

 

 

 

LLR

1m+ 1f+ 7j

21.88 ±0.08

21.73 -

22.01

0.4

15

Verkhnyaya Pisarevka

50°11'

36°51'

RR

1m+5j

16.15 ±0.04

16.10-

16.20

0.2

16

Pershotravnevoe

50°00'

36°55'

RR

1m+ 9j

16.18 ±0.05

16.12-

16.26

0.3

17

Martovoe

49°56'

36°58'

RR

1f+6j

16.08 ±0.06

15.97-

16.18

0.4

18

Artemovka

49°56'

37°00'

RR

8j

16.28 ±0.05

16.21 -

16.35

0.3

19

Pechenegi

49°53'

37°00'

LR

1j

14.93

 

 

 

 

 

 

 

RR

2m+ 2f+ 12j

16.20 ±0.07

16.05 -

16.31

0.4

TABLE 1 (continued)

No.

Locality

Latitude (N)

Longitude (E)

Type

N

Mean ± SD

Range

CV, %

20

Pyatnitskoe

49°56'

36048'

RR

3j

16.17 ±0.05

16.11 -16.21

0.3

21

Kitsevka

49°52'

36049'

LR

1j

15.10

 

 

22

Merefa

49048'

36005'

LR

7j

14.96 ±0.10

14.85 -15.14

0.6

23

Levkovka

49°44'

36019'

RR

4j

16.23 ±0.03

16.20-16.26

0.2

24

Zhadanovka

50°13'

36023'

LR

2j

14.98

14.95 -15.01

 

 

 

 

 

LLR

1f

21.90

 

 

 

 

 

 

RR

15j

16.06 ± 0.09

15.86-16.23

0.6

25

Biostantsiya

49037'

36020'

LL

1j

14.01

 

 

 

 

 

 

LR

1m + 9j

14.99 ± 0.09

14.83 -15.09

0.6

 

 

 

 

LR?

2j

15.40

15.37-15.44

 

 

 

 

 

LRx

1j

22.36

 

 

 

 

 

 

LRR

2j

22.81

22.71 -22.91

 

 

 

 

 

RR

11j

16.10 ±0.04

16.04-16.17

0.3

26

Gaidary

49039'

36016'

LR

8m + 1f+40j

14.98 ±0.07

14.72-15.09

0.5

 

 

 

 

LR?

1j

15.27

 

 

 

 

 

 

LLR

3j

21.76 ± 0.13

21.61 -21.87

0.6

 

 

 

 

LRR

1f+14j

23.04 ± 0.17

22.65 -23.25

0.7

 

 

 

 

RR

3f+54j

16.13 ±0.07

15.80-16.26

0.5

27

Gomolshanskii

49034'

36016'

LR

10j

14.96 ± 0.04

14.90-15.03

0.3

 

 

 

 

LLR

4j

21.76 ± 0.09

21.64 -21.84

0.4

 

 

 

 

LRR

9j

22.91 ± 0.12

22.64 -23.02

0.5

 

 

 

 

LLRR?

1j

29.58

 

 

 

 

 

 

RR

3j

16.17 ±0.04

16.12-16.20

0.3

28

Velikaya Gomolsha

49033'

36014'

LL

1j

13.97

 

 

 

 

 

 

LR

3j

14.96 ±0.08

14.90-15.04

0.5

 

 

 

 

LLR

1f

21.67

 

 

 

 

 

 

RR

1j

15.99

 

 

29

Sukhaya Gomolsha

49032'

36020'

LR

2m + 3f

14.97 ± 0.09

14.88 -15.09

0.6

 

 

 

 

LRR

1j

22.96

 

 

 

 

 

 

RR

2j

16.07

16.02-16.12

 

30

Kreidyanka

49026'

36048'

LR

5m+8j

14.91 ±0.10

14.79-15.15

0.7

 

 

 

 

LR?

1f

15.44

 

 

 

 

 

 

LLR

2f

21.61

21.61 -21.62

 

 

 

 

 

LRR

2m + 1j

22.80 ±0.12

22.73 -22.93

0.5

 

 

 

 

RR

1f+1j

15.95

15.90-16.00

 

31

Chervonaya Gusarovka

49025'

36052'

LR

4j

14.91 ±0.07

14.80-14.97

0.5

 

 

 

 

LLR

1j

21.54

 

 

 

 

 

 

LRR

2j

22.75

22.72 -22.79

 

 

 

 

 

RR

9j

15.96 ±0.10

15.74-16.07

0.6

32

Ol'khovatka

49025'

36053'

LR

5m+ 1f+3j

14.84 ±0.05

14.76-14.91

0.3

 

 

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L J Borkin, A V Korshunov, G A Lada - Mass occurrence of polyploid green frogs (rana esculenta complex) in eastern ukraine