NORTH-WESTERN JOURNAL OF ZOOLOGY 14 (1): 50-59 ©NWJZ, Oradea, Romania, 2018 Article No.: e171504 http://biozoojournals.ro/nwjz/index.html
New data and distribution of common spadefoot toad Pelobates fuscus (Laurenti, 1768) (Anura: Pelobatidae) in Western Balkans
Ana ĆURIĆ1*, Adnan ZIMIĆ2, Tomislav BOGDANOVIĆ3 and Dušan JELIĆ4
1. Society for Research and Protection of Biodiversity, Brace Potkonjaka 16, 78000 Banja Luka, Bosnia and Herzegovina. 2. Herpetological Association in Bosnia and Herzegovina ATRA, Alipašina 207, 71000 Sarajevo, Bosnia and Herzegovina. 3. Croatian Odonatological Socieety “Croatocordulia”, Vijenac Hrvatske Republike 9, 31550 Valpovo, Croatia. 4. Croatian Institute for Biodiversity, Croatian Herpetological Society Hyla, Lipovac I. 7, 10000 Zagreb, Croatia. *Corresponding author A. ĆURIĆ, E-mail: [email protected] / [email protected]
Received: 15. June 2016 / Accepted: 26. March 2017 / Available online: 28. July 2017 / Printed: June 2018
Abstract. Based on known literature data and data collected during field research in 2014 and 2015, we present an updated distribution map of the common spadefoot toad (Pelobates fuscus) in Western Balkans (Bosnia and Herzegovina and Croatia). Pelobates fuscus is listed as least concern on the global IUCN Red List, data deficient in Croatian Red Book and the populations are in constant decline. Until year 2014 this species was only suspected to inhabit Bosnia and Herzegovina. Today we have confirmed localities in Posavina region (western, central and eastern part) presented with precise coordinates and elevation. In this paper we also present first findings of tadpoles, more precisely, the reproductive sites of the common spadefoot toad in Bosnia and Herzegovina. In Croatia, this species is found along the rivers Mura, Drava and Sava, including most lowland areas up to 300 m above sea level. Historical records of Pelobates fuscus in Adriatic region are discussed and compared with the distribution area of the Italian isolated population.
Key words: conservation, decline, Posavina, Pannonian Plain, common spadefoot toad.
Introduction ations happened from 115,000 - 10,000 years ago with the last glacial maximum (LGM) about 26,000 years ago and it ended The common spadefoot toad, Pelobates fuscus (Laurenti 1768), around 19,000 years ago (Sibrava et al. 1986, Clark et al. is a wide-ranging European anuran species, which can be 2009). As the ice sheet spread south, populations migrated found in lowlands and hilly areas of central, eastern and from mentioned refuges and back. This was followed with southeastern Europe, up to western Siberia in the west and the rapid increase of temperature 13,000 years ago, when the northwestern Kazakhstan in the east (Hutchins et al. 2003). increase was 8-20 °C in both summer and winter tempera- By the last studies of genome variation there can be identi- tures. Range of Italian population spread to the Eastern fied two main groups: P. f. fuscus as East European and P. f. Europe and Balkan Peninsula, but in the 1970’s the species vespertinus as West European group (Litvinchuk et al. 2013). vanished from this area (Džukić et al. 2005, Crottini et al. Considering the Balkans, the species range is quite frag- 2007). mented (Džukić et al 2005, Sillero et al. 2014). Balkan low- Main aims of this study were: 1) map the distribution of land areas are preferred by the common spadefoot toad and P. fuscus in Posavina region in Bosnia and Herzegovina and it can be found in northern and eastern parts of this region. Croatia, 2) discuss the historical records of P. fuscus in Medi- Also, there are isolated areas in Dalmatia and Istria which terranean biogeographical region (Rijeka town and Dalma- are considered to be a part of the species historical range tia), 3) to compare species ecology based on temperature, (Džukić et al. 2005 and references therein), but no precise precipitation and altitudinal preference. data are given and these records have not been confirmed yet. Until the year 2014 there was only one unreliable record of P. fuscus from Bosnia and Herzegovina found in analyzed Materials and methods sample of digestive tract of Black-crowned night heron Posavina region was divided into three areas for easier organization (Nycticorax nycticorax (Linnaeus 1758)) (Obratil 1981). In 2014 of the search – western, central and eastern part (Fig. 1). Research three more records were published (Crnobrnja-Isailović et al. was aimed towards lowland areas with suitable water bodies (shal- 2014, Šukalo et al. 2014) reconfirming its presence within low ponds, canals, meadows, temporary and permanent ponds, Bosnia and Herzegovina. In Croatia, P. fuscus was found in lakes) and with nearby arable lands or natural sandy or loose soil the eastern part of the country (Slavonija, Baranja) and along (Boulenger 1897, Nöllert et al. 2012, Rannap et al. 2015, Stevens et al. the Sava and Drava river floodplains, with the central gap 2015). ranging from the town of Zagreb to Papuk Mountain In years 2014 and 2015, from March to October, listed regions were visited on multiple occasions. Field research timing depended (Džukić et al. 2008, Jelić et al. 2012). on the common spadefoot toad activity period. In the early spring Four glacial refuges of species P. fuscus are suggested: a) period (from February to April) we were field searching and map- area between Black and Caspian Seas as origin of the „east- ping suitable habitats. In the breeding period (April and May) we ern lineage“ of P. f. fuscus, b) Danube system as a center of heard distinctive male calling sounds in/under the water by. During diversity for the „western lineage” of P. f. fuscus, c) the Po night (10:00 pm to 02:00 am) we revisited all favorable sites and Valley with P. f. fuscus (northeastern Italian subspecies P. f. looked for adults around possible breeding areas. Adults and insubricus has been revoked based on genetic data) (Crottini subadults were searched for at night with hand/head lamps in men- tioned sites. In the summer period (June, July, August) we detected et al. 2007, Litvinchuk et al. 2013) and d) area between the the presence of P. fuscus by tadpole water survey (using hand nets) Caspian and Azov Seas as the origin of expansion of subspe- and determination of caught tadpoles. Waterbodies were selected cies P. f. vespertinus (Litvinchuk et al. 2013). Würm glaci- due to mentioned species habitat suitability and preferences such as
New data and distribution of Pelobates fuscus in Western Balkans 51
Figure 1. Research areas in Posavina region in Bosnia and Herzegovina with impor- tant localities presented.
low predatory impact, shallow and stagnant water with aquatic tal of 9999 times, following the procedure in PAST software. Climatic vegetation in the entire Posavina region. At the same period we also data and Digital elevation model were downloaded from WorldClim surveyed terrestrial habitats for adults and metamorphosed indi- - Global Climate Data version 1.4 (Free climate data for ecological viduals. The final mapping was done in late summer and fall (Sep- modeling and GIS; http://www.worldclim.org/) web service which tember, October) when we were only conducting night survey of is based on weather conditions recorded from 1950 to 2000. Data adults and subadults. We also collected new data by determination were analyzed with PAST 3.06. paleontological statistical software, of photographs provided by local people and known experts from where we used PCA analysis to compare monthly mean tempera- both countries. All literature and personal data were collected and tures (tmean_1 = January, tmean_2 = February, etc.), annual mean tem- presented with precise name of locality, country, reference, coordi- perature (tmean_Annual), monthly mean precipitation (precmean_1 = nates, elevation, year, UTM fields (10 x 10 km) and sample size (Ta- January, precmean_2 = February, etc.), annual mean precipitation ble 1). Most literature data was provided only with locality or area (prec_Annual) and elevation (Altitude) of the localities in two gen- name and elevation. T-test was done comparing climate from locali- eral regions (Continental and Mediterranean). All precipitation data ties in Continental (this study) and Adriatic (historical range) re- are expressed in mm/cm, temperatures as °C and elevation in m gions. Personal data with exact coordinates and altitudes were used a.s.l. Elevation data were corrected for localities where we had con- for comparing localities in terms of annual mean temperature, pre- fident field measurements with a GPS device. cipitation and elevations to get the overview of suitable habitats due to those parameters. For Rijeka and Dalmatia regions only three his- torical records and localities were available and we used them for Results ecological niche comparison with the mainland localities concerning the above three environmental parameters. We also used two his- With literature data and data collected during the field re- torical localities from Italy, both from Trieste area, for more compre- search (two year period) of P. fuscus in Bosnia and Herzego- hensive statistical results regarding historical ranges within the vina, as well as literature and personal data from Croatia, we Adriatic group (Džukić et al. 2005). In order to determine the poten- tial lower and upper variable limit (for annual mean temperature, present an updated distribution map of the common spade- annual mean precipitation and elevation) in the small Adriatic (his- foot toad (P. fuscus) in the Western Balkans (Bosnia and Her- torical) group (n = 5), we bootstrapped the observed sample for a to- zegovina and Croatia) (Fig. 2). The total number of sites is
Figure 2. Distributional range and locali- ties of Pelobates fuscus in Bosnia and Her- zegovina (BA) and Croatia (HR); sug- gested historical records are presented for reference
Table 1. Data on distribution of European common spadefoot toad (Pelobates fuscus) in Bosnia and Herzegovina and Croatia (X and Y coordinates are in WGS84, in Decimal Degrees). For the better overview of lo- calities approximated coordinates and elevation are given and this is indicated with parentheses. (AD – Adults, SUBAD – Subadults, TAD – tadpoles, N/A – not applicable, ♂ - male, ♀ - female, > - more than certain number.
Locality Country Reference X Y Elev (m.a.s.l.) Year UTM Sample size Brod BiH This study, leg. Ruškić, S. 45.143047 18.012459 83 2014 BR60 1 ♀ AD Brvnik BiH This study, leg. Ćurić, A. 45.047444 18.579635 80 2015 CQ09 6 SUBAD Čardak BiH This study, leg. Ćurić, A. 44.958280 18.405363 93 2014 BQ98 20 SUBAD Čardak BiH This study, leg. Ćurić, A. 44.958614 18.404472 94 2014 BQ98 39 SUBAD Čardak BiH This study, leg. Ćurić, A., Zimić, A. 44.958589 18.405828 94 2015 BQ98 2 ♀ AD; 24 ♂ AD; 3 SUBAD Čardak BiH This study, leg. Ćurić, A. 44.958244 18.401279 95 2015 BQ98 6 ♀ AD; 13 ♂ AD; 9 SUBAD Donje Ledenice, Okanovac BiH This study, leg. Ćurić, A. 44.929334 18.408811 97 2015 BQ97 2 SUBAD; 1 ♀ AD Donje Ledenice, Okanovac BiH This study, leg. Ćurić, A. 44.930456 18.409064 97 2015 BQ97 3 SUBAD Gromiželj BiH This study, leg. Ćurić, A. 44.872448 19.310834 79 2015 CQ67 2 SUBAD Jošik BiH This study, leg. Ćurić, A. 45.213929 16.862567 92 2015 XL40 1 ♀ AD Liješće BiH This study, leg. Ćurić, A. 45.071105 18.078641 85 2015 BQ69 1 AD Liješće BiH This study, leg. Ćurić, A., Jelić, D. 45.074751 18.077833 85 2014 BQ79 1 ♀ AD; 1 ♂ AD Liješće BiH This study, leg. Ćurić, A., Zimić, A. 45.086671 18.078906 86 2014 BQ79 1 AD Srbac, Kaoci BiH This study, leg. Jelić, D. 45.105485 17.577171 103 2012 YK09 1 AD Modriča BiH This study, leg. Ćurić, A. 44.963247 18.319962 105 2014 BQ88 2 AD Ostojićevo, Brodac Gornji BiH This study, leg. Ćurić, A. 44.858524 19.239464 83 2015 CQ66 2 SUBAD Ostojićevo BiH This study, leg. Ćurić, A. 44.915156 19.170806 78 2015 CQ57 1 ♂ AD Šamac BiH This study, leg. Ćurić, A. 45.055754 18.467266 85 2015 CQ09 1 ♀ AD;1 ♂ AD; 1 SUBAD Liješće, Bosanski Brod BiH Šukalo, G.et al (2014) 45.084665 18.079077 107 2014 BQ79 3 Sitneši, Srbac BiH Šukalo, G.et al (2014) 45.051826 17.537221 118 2014 XK99 1 Posavina BiH Crnobrnja-Isailović, J., (2014) N/A N/A N/A 2014 N/A N/A Bardača, Brajinci BiH Obratil, S. (1981) N/A N/A N/A 1981 XK99 N/A Batina HR This study, leg. Jelić, D. 45.828595 18.840799 138 2009 CR37 50 AD; 5 TAD Beravci - Berava HR This study, leg. Bogdanović, T. 45.149218 18.452962 86 2010 BR90 > 2 Bobota HR This study, leg. Bogdanović, T. 45.429951 18.850094 88 2010 CR33 > 1 Budimci HR This study, leg. Bogdanović, T. 45.460185 18.338166 100 2010 BR93 > 1 Budrovci fishpond HR This study, leg. Bogdanović, T. 45.284488 18.440592 87 2010 BR91 > 1 Road Koprivnica - Ðelekovec HR This study, leg. Samardžić, M., Jovanović, O., Šafarek, G. 46.228702 16.852929 125 2010 XM42 1 Divoševci - Kupina HR This study, leg. Bogdanović, T. 45.137052 18.339057 87 2010 BR90 > 2 Donji Miholjac, pond Bajer HR This study, leg. Jelić, M. 45.747715 18.183453 92 2008 BR87 TAD Dopsin - Lipovac HR This study, leg. Bogdanović, T. N/A N/A 88 2010 CR03 > 1 Durgutovica - woods HR This study, leg. Bogdanović, T. 45.323965 18.667667 107 2010 CR12 > 1 Čajkovci - Biđ HR This study, leg. Bogdanović, T. 45.176177 18.364619 86 2010 BR90 > 1 Jaruge HR This study, leg. Bogdanović, T. 45.121999 18.441282 86 2010 BR90 > 2 Kapela Podravska HR This study, leg. Šalamon, D. N/A N/A N/A 2007 XM32 1 AD Kešinačka woods HR This study, leg. Bogdanović, T. 45.361853 18.630104 85 2010 CR12 > 1 Kešinci - Zgorelica HR This study, leg. Bogdanović, T. 45.329951 18.550929 107 2010 CR02 > 1 Klisa HR This study, leg. Bogdanović, T. 45.461887 18.859269 88 2010 CR33 > 1
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Table 1 (continued)
Locality Country Reference X Y Elev (m.a.s.l.) Year UTM Sample size Koprivna HR This study, leg. Bogdanović, T. 45.408969 18.611058 90 2010 CR13 > 1 Koprivnica HR This study, leg. Jelić, D. 46.186202 16.867794 128 2010 XM41 1 AD Koritna HR This study, leg. Bogdanović, T. 45.387335 18.589526 94 2010 CR02 > 1 Koritna (dam) HR This study, leg. Bogdanović, T. 45.370017 18.562181 93 2010 CR02 > 1 Mužilovčica, Lonjsko polje HR This study, leg. Mihoković, N. N/A N/A N/A 2002 XL32 > 2; 1 ♂ AD Laslovo (ada) HR This study, leg. Bogdanović, T. 45.404724 18.685147 88 2010 CR13 > 1 Lonjsko polje, Drenov Bok HR This study, leg. Janev Hutinec, B. 45.266022 16.830069 90 1991 XL41 > 4 Marjančaci, Vučica HR This study, leg. Bogdanović, T. 45.636859 18.364547 89,6 2010 BR95 > 1 Metlinci, Vučica HR This study, leg. Bogdanović, T. 45.631450 18.422568 90,5 2010 BR95 > 1 Mrzović HR This study, leg. Bogdanović, T. 45.322118 18.586246 109 2010 CR12 > 1 Murščak, Dekanovec HR This study, leg. Jovanović, O., Šaferek, G. 46.456603 16.616136 148 2010 XM24 TAD Bicycle path, primary school - Bilje HR This study, leg. Šalomon, D. 45.571470 18.720484 78 2007 CR25 1 SUBAD Novi Perkovci HR This study, leg. Bogdanović, T. 45.240390 18.349212 100 2010 BR91 > 1 Paulin Dvor HR This study, leg. Bogdanović, T. 45.436525 18.641364 87 2010 CR13 > 1 Piškorevci HR This study, leg. Bogdanović, T. 45.246675 18.394991 90 2010 BR91 > 1 Požarike HR This study, leg. Bogdanović, T. 45.389291 18.502177 93 2010 CR02 > 1 Šelevrtak HR This study, leg. Bogdanović, T. 45.373586 18.486693 94 2010 CR02 > 1 Široko Polje HR This study, leg. Bogdanović, T. 45.392221 18.496852 93 2010 CR03 > 1 Slokovec HR This study, leg. Šalomon, D. N/A N/A N/A 2007 XM22, XM 32 1 ♂ AD Strizivojna HR This study, leg. Bogdanović, T. 45.204833 18.446480 96 2010 BR91 > 1 Tikveš HR This study, leg. Mikuša, T. N/A N/A N/A 2006 CR36 > 1 Vajda HR This study, leg. Bogdanović, T. 45.458674 18.937133 86 2010 CR33 > 1 Vrbica - Cerik HR This study, leg. Bogdanović, T. 45.295432 18.530847 116 2010 CR01 > 1 Vrbica - Gaj HR This study, leg. Bogdanović, T. 45.302415 18.632912 116 2010 CR12 > 1 Vrpolje - Biđ HR This study, leg. Bogdanović, T. 45.180454 18.414024 84 2010 BR90 > 2 Vuka HR This study, leg. Bogdanović, T. 45.440881 18.504961 93 2010 CR03 > 1 Zmajevac, Baranja HR This study, leg. Jelić, D. 45.804007 18.820079 89 2009 CR37 4 TAD; 3 ♂ AD; 3 ♀ AD Zmajevac, Baranja HR This study, leg. Jelić, D. 45.789990 18.829177 97 2009 CR37 38 SUBAD Zmajevac, Baranja HR This study, leg. Jelić, D. 45.804335 18.819690 86 2009 CR37 1 Zokovica - fishpond HR This study, leg. Bogdanović, T. 45.315036 18.461016 113 2010 CR02 > 1 Ivanovo Selo, Grubišno Polje HR This study, leg. Jelić, D. 45.668614 17.264180 141 2014 XL75 1 Nova Gradiška HR This study, leg. Barišić, F. N/A N/A N/A 2015 XL81 1 Rajići, Bjelovar HR This study, leg. Veljković, M. 45.900727 16.700470 113 2015 XL38 1 ♀ AD Osijek, tvrđava HR This study, leg. Bogdanović, T. 45.561864 18.697342 88 2014 CR24 2 AD Valpovo, Toplice HR This study, leg. Bogdanović, T. 45.648000 18.442579 92 2014 CR05 1 AD Županja, old brickyard HR This study, leg. Bogdanović, T. 45.062300 18.674658 81 2014 CQ19 2 AD Village Nard, old basement HR This study, leg. Bogdanović, T. 45.660968 18.484154 88 2015 CR05 3 AD VIllage Nard, old orchard HR This study, leg. Bogdanović, T. 45.663241 18.480231 87 2015 CR05 2 AD Slokovec - Kapela road HR CHS- Hyla, (2007), project N/A N/A N/A 2007 XM32 N/A
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Table 1 (continued)
Locality Country Reference X Y Elev (m.a.s.l.) Year UTM Sample size Baranja HR CHS-Hyla, (2006/07), project N/A N/A N/A 2006 CR36 N/A Baranja, Suza HR Džukić, G. et al (2008) N/A N/A 97 2008 CR27 N/A Donji Miholjac HR Džukić, G. et al (2008) N/A N/A 97 2008 BR77 N/A Koprivnica, Peteranec HR Džukić, G. et al (2008) N/A N/A 133 2008 XM41 N/A Nova Gradiška, Vrbova HR Džukić, G. et al (2008) N/A N/A 125 2008 YL00 N/A Rijeka, Grobničko polje HR Džukić, G. et al (2008) (Depoli, 1898) N/A N/A 281 1898 VL52 N/A Rijeka, Grobnik HR Džukić, G. et al (2008) (Depoli, 1898) N/A N/A 466 1898 VL52 N/A Baranja, Kazuk, Siga HR Džukić, G. et al (2008) (Džukić & Pasuljević 1983) N/A N/A 84 1893 CR37 N/A Čazma HR Džukić, G. et al (2008) (Džukić & Pasuljević 1983) N/A N/A 144 1983 XL26 N/A Turopolje, Orle HR Džukić, G. et al (2008) (Džukić,1981) N/A N/A 99 1981 WL95 N/A Našice HR Džukić, G. et al (2008) (Karaman, 1921) N/A N/A 157 1921 BR64 N/A Rijeka, small pond near Šurinj HR Džukić, G. et al (2008) (Matisz, 1896) N/A N/A 345 1896 N/A N/A Baranja, Petlovac HR Džukić, G. et al (2008) (Mikuška & Vuković, 1980) N/A N/A 93 1980 CR07 N/A Belje HR Džukić, G. et al (2008) (Mojsisovics, 1897) N/A N/A 95 1897 CR15 N/A East Baranja, Batina HR Džukić, G. et al (2008) (Mojsisovics, 1897) N/A N/A 85, 89, 105 1897 CR37 N/A Zagreb HR Džukić, G. et al (2008) (Pavletić, 1964) N/A N/A 135 1964 N/A N/A Zagreb HR Džukić, G. et al (2008) (Pavletić, 1964) N/A N/A 135 1964 N/A N/A Slavonija, Bošnjaci HR Džukić, G. et al (2008) (Pozzi, 1966) N/A N/A 85 1966 CQ29 N/A Našice HR Karaman, St. (1921) N/A N/A 110 1921 BR74 TAD Našice HR Karaman, St. (1921) N/A N/A 110 1921 BR74 TAD Jasik, stream Županijac, Virovitica HR Kletečki, E. (2009) N/A N/A N/A 2008 YL08 TAD Mali Sakadaš, Kopački Rit HR Kletečki, E. (2009) N/A N/A 78 2008 CR25 TAD Novo Selo HR Kletečki, E. (2009) N/A N/A N/A 2008 BR76 TAD Osijek, Donji Miholjac HR Kletečki, E. (2009) N/A N/A 95 2008 BR77 N/A Vladimirovac, Nova Gradiška HR Kletečki, E. (2009) N/A N/A N/A 2008 YL08 TAD Donji Miholjac HR Kletečki, E. (2009) N/A N/A 95 2008 BR77 N/A Lonjsko polje, Krapje Đol, Jasenovac HR Kobašlić, A. (2002) N/A N/A 91 1985 XL41 N/A Peščenica, Turopolje, Vratovo HR Kobašlić, A. (2002) N/A N/A 111 1978 WL95 N/A Peščenica, Turopolje, Vratovo HR Kobašlić, A. (2002) N/A N/A 111 1986 WL95 N/A Našice HR Pavletić, 1964 N/A N/A 110 1964 BR74 N/A Zagreb (surrounding) HR Pavletić, 1964 N/A N/A N/A 1954 WL77 N/A Zagreb (surrounding) HR Pavletić, 1964 N/A N/A N/A 1943 WL78 N/A Zagreb (surrounding) HR Pavletić, 1964 N/A N/A N/A 1935 WL79 N/A Zagreb (surrounding) HR Pavletić, 1964 N/A N/A N/A 1934 WL80 N/A Zagreb (surrounding) HR Pavletić, 1964 N/A N/A N/A 1931 WL81 N/A Štrogač HR Reeder, D. (2004) N/A N/A N/A 1905 XM61 1 ♂ AD Šuma Repaš, Drava HR Reeder, D. (2004) N/A N/A N/A 1905 XM61 TAD Sava, Sava River valley near Prelošćica HR Strijbosch et al. (1986) N/A N/A 97 1986 XL13 N/A
New data and distribution of Pelobates fuscus in Western Balkans 55
Figure 3. Combination of terrestrial and water habitats of found Pelobates fuscus individuals a) Čardak, Bosnia and Herzegovina, b) Osto- jićevo, Bosnia and Herzegovina.
112 (suggested historical ranges are excluded). In two years (old river stream) is accessible in regards to canals where of active study (2014, 2015) we recorded 12 new sites in Bos- most metamorphosed individuals were found. Tadpoles nia and Herzegovina and three in Croatia. All together more were caught in 2014 and 2015 at the same site, in two sepa- than 30 unpublished new localities are listed for Croatia. rated pools. Due to heavy rain in 2014 this population had a Data presented as localities under UTM fields have four seasonal breeding explosion and the number of tadpoles, in UTM fields where literature and new data localities overlap different fazes and sizes, was several times greater than in (UTM fields: BQ79, CR37, XL41, XM41) (Table 1). Rijeka 2015 when we noticed absence of spawning and eggs. It is town data are presented with precise localities on the map, very likely that spawning season started earlier in 2015 and unlike data for Dalmatia region where precise locality is not that the number of amplexus was lower while only ad- provided (Fig. 2). vanced stages of tadpoles (possible last season tadpoles) During mapping in Posavina region we found surpris- were found in early summer in waterbodies in the Čardak ingly many individuals and confirmed that the species is lo- locality. cally common in Bosnia and Herzegovina. We managed to Data for Croatia were collected from 2002 until the pre- confirm P. fuscus in all three investigated regions: West, Cen- sent, with several data from year 1991. New significant lo- tral and East Posavina region (Fig. 1). calities in Croatia are confirmed in Grubišno Polje, Ivanovo In the western part of Posavina we found the common Selo (45.668614 °N 17.26418 °E) and Bjelovar, Rajići spadefoot toad in Jošik village (45.213929 °N 16.862567 °E), (45.900727 °N 16.70047 °E), connecting the Drava and Sava municipality of Kozarska Dubica (Fig. 1) (one female found River populations in the central part of Croatia (Fig. 1). on macadam road close to canal and arable land in 2015 – There it was a large number of tadpoles found in a small 01:05 am, temperature 19.5 °C, humidity 68.7 %). The most pond (Bajer Pond) in Donji Miholjac in June 2008. Many lo- suitable localities were found in the central part of Posavina calities were confirmed during field mapping in July, 2010, and new sites were found in municipalities of: Brod (Brod under UTM cells BR90, BR91, BR93, BR95. and Liješće), Modriča (Modriča, Čardak), Gradačac (Donje According to analyzed data from 81 localities, the low- Ledenice), Šamac (Šamac) and Domaljevac (Brvnik). In this land range of P. fuscus in Bosnia and Herzegovina and Croa- part, the common spadefoot toads were found in the areas tia varies from 75 m a.s.l. (Ostojićevo, Bosnia and Herzego- with arable land as well as several different types of water vina) to 148 m a.s.l. (Murščak, Dekanovec, Croatia) where bodies (old river stream, different types of canals). All indi- the larger proportion of localities is between 82 m a.s.l. and viduals were found at night between 21:30 pm and 04:00 am 97 m a.s.l. Most localities had an annual mean temperature and 134 individuals were observed. Most of the adult indi- of 11.5 °C. In the researched areas this species mostly occu- viduals were collected in Čardak (Fig. 3a). During 2014 and pies areas with annual mean precipitation between 58 mm 2015 we found only two DOR (Dead on road) individuals in and 68 mm (Fig. 4). Also, with available data we analyzed Čardak (44.958589 °N 18.405828 °E) (Fig. 1) and three DOR the same parameters for historical localities of the Adriatic in Liješće (45.086671 °N 18.078906 °E) (Fig. 1). There is a group in Mediterranean Biogeographical Region (Rijeka and much higher percentage of other DOR species (e.g. Pelophy- Dalmatia) and compared them with data collected in this lax sp., Bufo bufo, Bufotes viridis), although in 2014, during the study (Table 2). Due to climate differences between localities big explosive breeding period for P. fuscus, there was a huge in Mediterranean Biogeographical Region (historical data) number of DOR subadults. In the eastern part of Posavina and Continental Biogeographical Region (new data), the region we found three new sites in municipality of Bijeljina, data vary in all parameters (Fig. 5). In general, the Mediter- in villages Gromiželj (44.872448 °N 19.310834 °E) and Osto- ranean Biogeographical Region (Rijeka town surroundings jićevo (44.915156 °N 19.170806 °E) (Fig. 1, 3b). Subadult indi- and Dalmatia) has higher values of annual mean tempera- viduals were found at night (00:00 – 01:30 am) in the vicinity ture, annual mean precipitation and mean altitude (Table 2). of arable land, canals and permanent ponds. Presence of Bootstrap analyses (with 9999 resamplings) revealed that the tadpoles was not confirmed in this area due to short period potential annual mean temperature limits for the Mediterra- of research. In total, we found, examined and photographed nean Biogeographical Region are 12-15 °C, potential annual 134 individuals: 40 males, 17 females and 77 subadults. mean precipitation limits for the Mediterranean area is 67- Tadpoles were found only in municipality of Modriča, 140 mm/m2 and potential upper elevation limit is 692 m a.s.l. Čardak locality, Bosnia and Herzegovina. This water body Analysis of WorldClim climatic data and altitude (Fig. 5) 56 A. Ćurić et al.
sociated with topo-climatic variables explained 93 % of the total variation (Fig. 5). First principal component represents 74 % of total variation and it was mainly positively influ- enced by altitude and precipitation during winter months (September until December). Second principal component represents 19 % of total variation and it was negatively in- fluenced by altitude and positively by monthly mean tem- peratures during summer months (June – August). PCA analysis revealed that the two groups do not overlap in the ecological niche space and that the Adriatic group is proba- bly more connected to Po River populations in Italy. But this could also be the result of a smaller input sample size for the Adriatic group.
Discussion
The results of this study show that there is a large distribu- tional range of P. fuscus in the Pannonian lowlands of Croa- tia and Bosnia and Herzegovina. It seems that the range in North Bosnia is quite narrow and restricted, mainly corre- sponding to the Sava River plains. The reason for this is mountainous areas, which form a natural barrier (Dinaric Arc barrier; Fig. 2) for species dispersal. Probably the species is also locally present further to the south, in lowland valleys of large tributaries of Sava River (Una, Vrbas, Bosna, Drina, etc.). But upstream these rivers very quickly lose their low- land characteristics and form steep cliffs and canyons. The central part of the Posavina region in Bosnia and Herzego- vina had only three known localities for P. fuscus (Crnobrnja- Isailović et al. 2014, Šukalo et al. 2014) and all three were re- confirmed. Only the record for village of Brajinci (possibly Bajinci today), published by Obratil (1981), was not recon- firmed in this study. With new data collected in the area around Ivanovo Selo village and Bjelovar town, the Drava and Sava river populations are confirmed to be connected in the central part of Croatia, leaving the only gap in the area of the Papuk Mountain due to its high altitude. There are several historical locations mentioned for Adriatic region but none has been reconfirmed recently. There are at least three historical localities around and within the city of Rijeka, and habitat alternation is mainly mentioned as the reason for possible disappearance of this species. On the other hand, there are no exact locality data for Dalmatia, ex- cept that the species was found in northern parts (Ravni Ko- tari; Depoli 1898, Džukić et al. 2005 and references therein). If there ever was or possibly is a population of P. fuscus in those areas, it is more likely that they were/are a part of pa- leo Po population dispersed from northeastern Italy. Accord- ing to paleo-coastline reconstruction of the Adriatic Sea dur-
ing the LGM the water level was 115 m lower (Sikora et al. Figure 4. Pelobates fuscus histograms of this study data (n = 81) for a) 2014), and that suggests that the Po River Valley area was altitude (m a.s.l.), b) annual mean precipitation (mm) and c) annual dry land connected all the way to Zrmanja and possibly the mean temperature (°C) in Continental biogeographical region (data Krka River estuary. The northern part, from today’s Po River interpolated from WorldClim ≈ 1 km2 grid). delta to the north side of the Mid-Adriatic Deep (MAD) formed an alluvial plain, suitable for P. fuscus (Maselli et al. revealed that Adriatic group localities have higher altitudes 2011, Moscon et al., 2015). Geographically, the entire area of and higher precipitation (especially during winter months). northern Italy and Croatian coastline were connected. That Continental group localities are more uniform, represented gives us a clear picture of possible presence not just of P. fus- by lower altitude values and smaller amount of precipita- cus but many other species along the Croatia coastline that tion. The first two principal components (PC1 and PC2) as- today are characteristic for the Apennine Peninsula. Due to a
New data and distribution of Pelobates fuscus in Western Balkans 57
Table 2. Comparison of this study data for Continental biogeographical region and historical range data for Mediterranean biogeographical region for three main topo-climatic parameters: annual mean temperature, annual mean precipitation and altitude in Pelobates fuscus records.
This study data (Continental) Historical range data (Adriatic) t-test (df = 80) Parameter N Mean (SD) Min - Max N Mean (SD) Min - Max P Annual mean temperature (°C) 81 11.0 (0.5) 8 - 12 5 13.2 (0.84) 12 - 14 <0.0001 Annual mean precipitation (mm/m²) 81 64.2 (6) 53 - 78 5 99.6 (15.6) 82 - 116 <0.0001 Altitude (m a.s.l.) 81 94.8 (14.6) 75 - 148 5 171.4 (197) 0 - 466 <0.0001
Figure 5. PCA diagram for com- pared Pelobates fuscus Continen- tal and Mediterranean Adriatic (historical) population localities according to monthly mean tem- peratures (tmean_1-12), annual mean temperature (tmean_AN- NUAL), monthly mean precipita- tion (prec_1-12), annual mean precipitation (prec_ANNUAL) and elevation (Altitude).
lower level of the Adriatic Sea, P. fuscus would have dis- ing mean results of three selected topo-climatic parameters persed along the historical Po Valley all the way to Istria, for suggested historical Adriatic range with Continental Kvarner and North Dalmatia. During the LGM, the perma- range, all three differ significantly. There is a statistically nent ice border was located somewhere north of Drava River significant difference between annual mean temperature (Willis 1996), so continental populations of P. fuscus could which varies around 2 °C, which in general means that the survive in the Balkan refugia. climate in the Mediterranean area is characterized by mild This connection with the Po River Valley area is still winters and hot summers. It is indicative that the Italian iso- visible in the biogeography of many other species of ani- lated population inhabits areas with very similar climate mals, especially fish. There are several species found in (Scali & Gentilli 2003). The temperature in which main activ- Adriatic area whose distribution today belongs to western ity occurs was observed. According to Nöllert et al. (2012), and south-western Europe and can be defined as former pa- significant temperature values varied from 4 °C to 24 °C leo Po distribution during glacial conditions. Some of those which means that mean temperature for species activity species are Rana latastei Boulenger, 1879 (Gasc et al. 2004, span is 14.0 (SD=1.35) °C. Considering this study data for the Kuljerić 2011), three fish species, Barbus plebejus Bonaparte, Mediterranean Biogeographical Region, annual mean tem- 1839, Leucos aula (Bonaparte, 1841), Romanogobio benacensis perature of 13.2 (SD=0.84) °C is appropriate for species dis- (Pollini, 1816) (Kottelat & Freyhof 2007, Bianco & Ketmaier tribution. If there are still surviving populations in the men- 2014, Jelić et al. 2016) and two historical records of Vipera tioned area (Adriatic range), according to temperature pa- aspis francisciredi Laurenti, 1768, and Chalcides chalcides (Lin- rameters, they would probably be active earlier in the spring naeus, 1758), deposited in the Natural History Museum of and have low activity period during the summer. Annual Milano and Natural History Museum in Vienna (Dolce 1977, mean precipitation is also higher in Mediterranean region, Tome 2003, Torkar 2003, Miras et al. 2009, Jelić 2014). Due to which is a known preference of the common spadefoot toads paleo Po reconstruction, today’s rivers of the north Adriatic (Scali & Gentilli 2003, Nöllert et al. 2012). According to Scali basin in Croatia were tributary rivers of Po River (Maselli et & Gentilli (2003) The isolated Italian population mainly fa- al. 2011) and the populations maintained in Croatia are pos- vors high precipitation during its reproduction period, while sible remnants of those historical populations now isolated afterwards it is not so important for their activity. Maximum by the Adriatic Sea. altitude recorded during our study was 466 m a.s.l. (locality T-test was done comparing climate from localities in belongs to historical range in Adriatic region), while species Continental (this study) and Adriatic (historical range) re- can even be found up to 810 m a.s.l. (Czech Republic) (Za- gions, but with large deviation of sample sizes we failed to vadil et al. 1995, Nöllert et al. 2012). get a confident result. Bootstrapped data showed higher po- PCA analyses of all 27 topo-climatic variables confirmed tential variable span in all three parameters, thus further in- that there is no overlap between the characteristics of locali- creasing the general difference between the groups. Compar- ties from the Mediterranean and Continental region. This 58 A. Ćurić et al. further supports the theory that Adriatic populations are the Acknowledgements. We would like to thank our colleges who remnant of a formerly widely distributed population of P. provided many data for Pelobates fuscus species and contributed to fuscus in the Po drainage. Similar unexplained distributional this research. gaps were recently published, as mentioned, for Vipera aspis francisciredi and Chalcides chalcides (Jelić 2014) and Romanogo- bio benacensis (Jelić et al. 2016). 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Springer, Heidelberg. nental region, presented in this paper, in the near future we Crottini, A., Andreone, F., Kosuch, J., Borkin, L.J., Litvinchuk, S.N., Eggert, C., can expect visible range and habitat quality decline. Field Veith, M. (2007): Fossorial but widespread: the phylogeography of the observations have confirmed that huge lowland areas are common spadefoot toad (Pelobates fuscus), and the role of the Po Valley as a major source of genetic variability. Molecular Ecology 16: 2734-2754. currently being converted to extensive agricultural lands. Dávid, A., Schäffer, D.A., Pirohoffer, E., Horváth, G., Purger, J.J. (2006): Water bodies (stagnant and flowing) are being dried out or Distribution of common Spadefoot toad (Pelobates fuscus) and soil types in changed permanently to reduce flooding. Habitats are being Hungary. Book of Abstracts, 1st European Congress of Conservation Biology, Hungary. rapidly destroyed due to degradation, developing infrastruc- Depoli, G. (1898): I Rettili e gli Anfibi del territorio di Trieste. Rivista Italiana di ture, intensive monocultures, use of pesticides and fertiliz- Scienze Naturali 18: 47-50. ers, etc. Pannonian lowlands in Croatia and Bosnia and Her- Džukić, G., Beškov, V., Sidrovska, V., Codălinceanu, D., Kalezić, M.L. (2008): Contemporary chorology of the spadefoot toads (Pelobates spp.) in the zegovina are the source of the majority of both countries’ ag- Balkan Peninsula. Zeitschrift für Feldherpetologie 15: 61-78. ricultural yield due to fertile soil. Extensive agricultural land Džukić, G., Beškov, V., Sidrovska, V., Cogãlniceanu, D., Kalezić, M.L. (2005): has proved to be a good replacement for the natural habitats Historical and contemporary ranges of the spadefoot toads Pelobates spp. of this species in the region (e.g. Nyström et al. 2002, Dávid (Amphibia: Anura) in the Balkan Peninsula. Acta Zoologica Cracoviensia 48A(1-2): 1-9. et al. 2006, Rannap et al. 2011) and we can also certainly con- Dolce, S. (1977): L' erpetofauna del Friuli, della Venezia Giulia, dell'Istria e della firm this from our data. Many natural habitats are lost, but Dalmazia nella collezione del Museo civico di Storia Naturale di Trieste. the species has managed to adapt to similar anthropogenic Catalogo ragionato. Parte I: Amphibia. Atti del Museo Civico di Storia Naturale di Trieste 30(2): 209-240. sites. So the key action for maintenance and conservation of Gasc, J.P., Cabela, A., Crnobrnja-Isailovic, J., Dolmen, D., Grossenbacher, K., these P. fuscus populations is to keep the water bodies in Haffner, P., Lescure, J., Martens, H., Martínez Rica, J.P., Maurin, H., Oliveira, natural or semi-natural conditions. For example, the original M.E., Sofianidou, T.S., Veith, M., Zuiderwijk, A. (eds.) (2004): Atlas of amphibians and reptiles in Europe. Reedition. Museum national d'Histoire course of Tolisa River in Bosnia and Herzegovina has a big naturelle, Paris. potential as a natural habitat, but in two years of our study Gislen, T., Kauri, H. (1959): Zoogeography of the Swedish amphibians and we noticed significant water depletion. This change has also reptiles with notes on their growth and ecology. Acta Vertebratica 1: 195- 398. been confirmed by the locals. We suspect that, in the near fu- Hutchins, M., Duellman, W.E., Schlager, N. (2003): Grzimek’s Animal Life ture, this site would be completely lost due to water overex- Encyclopedia. 2nd edition. Volume 6, Amphibians. Gale Group, Farmington ploitation and eutrophication. In many other countries of Hills, Michigan. Europe, decline of the common spadefoot toad is already Jehle, R., Hödl, W., Thonke, A. (1995): Structure and dynamics of central European amphibian populations: a comparison between Triturus dobrogicus confirmed (France: Nöllert et al. 2012 and references therein; (Amphibia, Urodela) and Pelobates fuscus (Amphibia, Anura). Australian Belgium: Rappè 1982; Netherlands: Zekhuis & Ottburg 2008; Journal of Ecology 20: 362-366. Denmark: Gislen & Kauri 1959; Sweden: Berglund 1998, Jelić, D. (2014): Checklist of Croatian amphibians and reptiles with bibliography of 250 years of research. Natura Sloveniae 16(2): 17-72. Nyström et al. 2002; Italy: Andreone et al. 2004). Bosnia and Jelić, D., Kuljerić, M., Koren, T., Treer, D., Šalamon, D., Lončar, M., Podnar- Herzegovina represents the southern border in distribution Lešić, M., Janev Hutinec, B., Bogdanović, T., Mekinić, S., Jelić, K. (2012): Red range of this species and there is a presumption that the book of amphibians and reptiles of Croatia. Ministry of Environment and Nature Protection, State Institute for Nature Protection, Zagreb. border could move further to the north, reducing the species Jelić, D., Jelić, M., Žutinić, P., Šimunović, I., Zupančič, P., Naseka, A. (2016): range (Jehle et al. 1995, Hutchins et al. 2003, Džukić et al. Distribution of endangered Italian gudgeon Romanogobio benacensis 2005). Based on our study we may say that populations in (Cypriniformes, Cyprinidae, Gobioninae) with remarks on distinguishing morphological characters. ZooKeys 729: 103-127. this region are still stabile and possible reasons for the future Litvinchuk, S.N., Crottini, A., Federici, S., de Pous, P., Donaire, D., Andreone, decline are already discussed above. F., Kalezić, M.L., Džukić, G., Veith, M., Lada, G.A., Borkin, L.J., Rosanov, J.M. (2013): Phylogeographic patterns of genetic diversity in the common spadefoot toad, Pelobates fuscus, reveals evolutionary history, postglacial range expansion and secondary contact. 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