ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah Acta Herpetologica 8(1): 53-57, 2013 First evidence of a paedomorphic population of the smooth newt (Lissotriton vulgaris) in the Czech Republic Václav Gvoždík1,2, Veronika Javůrková3,4, Oldřich Kopecký5,* 1 Department of Zoology, National Museum, Cirkusová 1740, 193 00 Prague, Czech Republic 2 Laboratory of Molecular Ecology, Institute of Animal Physiology and Genetics, Academy of Sciences of the Czech Republic, 277 21 Liběchov, Czech Republic 3 Department of Zoology, Faculty of Science, Charles University in Prague, Viničná 7, 128 44, Prague, Czech Republic 4 Institute of Vertebrate Biology v. v. i., Academy of Sciences of the Czech Republic, Květná 8, 603 65 Brno, Czech Republic 5 Department of Zoology and Fisheries, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences, Kamýcká 957, 165 21 Prague, Czech Republic. * Corresponding author. E-mail: kopeckyo@af.czu.cz Submitted on 2012, 20th December; revised on 2013, 13th March; accepted on 2013, 27th March. Abstract. Facultative paedomorphosis is an environmentally induced polymorphism that is well known for many caudate species including newts. Although facultative paedomorphosis has been documented in some smooth-newt populations, records of entirely paedomorphic populations outside the Balkans are limited. Here we present the first evidence of a paedomorphic population of the smooth newt in the Czech Republic with discussion of potential causes that need to be further tested. Keywords. Facultative paedomorphosis, urodeles, “Triturus”, Central Europe. Facultative paedomorphosis, the case of phenotypic heterochrony when some sexually mature individuals from a population (or some populations of a species) retain some states of larval morphology, has been widely documented for many caudates (review in Denoël et al., 2005; Laudet, 2011). In Europe, facultative paedomor- phosis regularly occurs in newts (former genus “Tritu- rus”) with high frequency in the Alpine newt (Ichthyosau- ra alpestris), especially in some areas of the Balkans and Alps (Denoël et al., 2001). Beside the Alpine newt pae- domorphosis has also been recorded in other newt spe- cies (summarized by Wells, 2007): Lissotriton helveticus, L. italicus, L. vulgaris, Triturus cristatus, T. macedonicus, and Ommatotriton ophryticus (Başkale et al., 2011). In the smooth newt (L. vulgaris), this phenomenon is also rela- tively common, although usually only a small part of a population or single individuals are paedomorphic. Cases of peadomorphosis in the smooth newt have been report- ed from various places within almost the whole distri- bution range (review in Schmidtler and Franzen, 2004), but entirely paedomorphic populations of L. vulgaris are rarely reported (Dely, 1967; Dolmen, 1981; Denoël et al., 2009). One such population from northern Hungary was described like a different subspecies, Triturus vulgaris tataiensis (Dely, 1967), but validity of this subspecies was rejected (Raxworthy, 1990). Here, we report a presumably almost completely paedomorphic population of L. vulgaris from the Czech Republic. This is the first formally reported recent record of urodelan paedomorphosis for the country. Paedomor- phic smooth newts (Fig. 1) were discovered (by VG and VJ) on 28 April 2012 in a water reservoir in Stará Lysá (50°13’31.6”N, 14°47’57.1”E, ~190 m a.s.l.). The finding place is situated in “Polabí”, which is a lowland area along the Labe (Elbe) River in the Central Bohemian Region. This part of the Czech Republic is one of the warmest, with dry and mild winters and average annual tempera- ture between 8-9 °C (Quitt, 1971). The local landscape 54 Václav Gvoždík et al. Fig. 1. Smooth newts observed in the Stará Lysá water reservoir, Czech Republic: A) Paedomorphic male in nuptial colouration; B) paedo- morphic female in nuptial colouration; C) close-up view on another paedomorphic male showing the structure of gills; D) aquatic imma- ture specimen with open gill slits (A-D photographed on 7 May 2012); E) paedomorphic male, and F) female later in the season with already reduced gills; G) different pair (adult male and female) demonstrating variation in the level of persistence of larval traits; H) imma- ture paedomorphic specimen with large gills; I) metamorph-like aquatic immature specimen with open gill slits (E-I photographed on 23 August 2012); J) Stará Lysá water reservoir. 55Paedomorphic smooth newts in the Czech Republic has been historically used for agriculture and so the locality is from three quarters surrounded by crop fields and from one quarter by a relatively dry cultivated pine forest (cf. Covaciu-Marcov et al., 2011). Newts inhabit a water reservoir (originally used for fire protection) sized 25 × 15 m with maximal water depth of 1.5 m made from concrete blocks (Fig. 1J). Shorter sides of the reservoir are formed by vertical walls while longer sides slope in approximate 45°. The reservoir is supplied from a deep borehole (~60 m), and water is changed approximately once per three years and occasionally refilled (last change in 2010; V. Vaněk, pers. comm.). No fish were observed. The reservoir is with- out vegetation, only filamentous algae grow later in the season over submerged objects and newts use them for wrapping eggs. Apart from newts, several adult individu- als of marsh frogs (Pelophylax ridibundus) were observed and their numerous offspring later in the season. The water is rich on planktonic micro-crustaceans. Generally, the reservoir meets conditions for occurrence of paedo- morphic newts: does not freeze to the bottom in winter, do not dry in summer, few predators (no fish), abun- dant food resources (Litvinchuk et al., 1996; Denoël and Poncin, 2001). The locality has been visited in several occasions (April, May, July, August, September, October 2012), and opportunistic visual surveys and dip-netting have been done. Paedomorphic individuals were observed during all visits, with a visually assessed abundance in the spring. During the assumed peak of reproductive season (7 May 2012), we observed 70 adult smooth newt individuals when counting specimens visible along the reservoir walls during one walk around the tank. Except of two seem- ingly metamorphic females (but without success to catch them and inspect for morphology) and one almost meta- morphic juvenile (but with open gill slits; Fig. 1D), all individuals were clearly paedomorphic (Fig. 1A-C) and only three of them were males. Sex ratio bias toward the females is common in paedomorphic individuals of this species (e.g., Kalezić and Džukić, 1986; Litvinchuk et al., 1996; Çiçek and Ayaz, 2011), however the observed ratio is unusual. Also situations when paedomorphic smooth newts outnumber metamorphic individuals have been only rarely reported (e.g., Kalezić and Džukić, 1986; Rot- Nikcević et al., 2000) compared to the more common metamorph-biased population ratio in this species (e.g., Banks, 1985; Kalezić and Džukić, 1985; Çiçek and Ayaz, 2011). However, it is important to mention that we can- not make clear conclusion about the ratio of peadomorphs vs. metamorphs in our smooth-newt population as we did not make a quantitative survey. Potential bias toward pae- domorphs might be also caused by differences in behav- iour of the morphs with respect to the relatively high water depth. During the spring visits, many paedomorphic newts were resting in the water column along the walls to 50 cm below water surface and were easily counted. To observe reproductive behaviour, three paedomor- phic individuals (one male, two females) were taken into captivity (7 May 2012). Water temperature in the aquari- um ranged 16.7-19. 4 °C and newts were kept under nat- ural light conditions. The newts were housed together in an aquarium (50 × 25 × 25 cm) filled by settled tap water and with several waterweed plants (Egeria densa). Dur- ing the following days we had observed the male’s court- ship behaviour (fan display), which is in newts similar for both morphs (Denoël, 2002), and deposition of eggs by both females. As we expected (cf. Banks, 1985; Litvin- chuk et al., 1996; Litvinchuk, 2001), the male metamor- phosed after 19 days, followed by the females which met- amorphosed after 42 and 45 days in captivity, respectively (including closed gill slits). Later in the season (23 August 2012), all captured adults (Table 1) were already almost without crests and fin appendices, and with small, sometimes rather rudi- mental gills, nonetheless, still in aquatic phase and with open gill slits (Fig. 1E-G). Immature individuals, accord- ing to the body size and shape and level of development of the cloaca, were of two types: some had large gills and fins (Fig. 1H), while some others resembled the adults, but being substantially smaller, by having rudimentary gills or only open gill slits (Fig. 1I). Beside these paedo- morphic specimens (open gill slits as a common charac- ter), some typical larvae were also observed. It seems that adult individuals reduce their larval traits (gills) toward the winter (cf. Denoël, 2003). However, it is currently unknown if the same individuals grow the larval traits again in the next season. A study of the demographic structure is currently ongoing using the capture-mark- recapture method and shall shed more light onto this and other questions in near future. Despite relatively strong evidence for the benefi- cial role of facultative paedomorphosis as an alternative Table 1. Morphometric characteristics in cm (SVL - snout-vent length, Lt - length of tail, Dbl - distance between front and hind limbs) of adult peadomorphic smooth newts (females n = 7; males n = 3) from Stará Lysá, Czech Republic. mean min. max. SD females SVL 3.45 3.13 3.82 0.27 Lt 3.10 2.80 3.47 0.22 Dbl 1.69 1.41 1.93 0.18 males SVL 3.68 3.34 4.01 0.34 Lt 3.50 3.19 3.82 0.32 Dbl 1.67 1.57 1.77 0.10 56 Václav Gvoždík et al. ontogenetic pathway in many tailed amphibians, impor- tance and influence of particular factors lying behind its evolution is not fully resolved. Whilst paedomorphosis can be a highly heritable trait (e.g., Harris et al., 1990; Voss et al., 2003), the occurrence of facultative paedo- morphosis in individuals and its maintenance in a popu- lation is most probably a result of interplay between gen- otype and heterogeneity of the environment (reviewed by Denoël et al., 2005). Among many factors, streambed or pond-bank structures (Bennett and Chippindale, 2006), slow desiccation rate (Semlitsch et al., 1990), and high food availability in water (Ryan and Semlitsch, 2003) have been observed to induce occurrence of pae- domorphs and the maintenance of resulted polyphenism in a population. All the above-listed factors, particularly high food availability and resource partitioning between different phenotypes seem to be feasible as possible fac- tors responsible for the maintenance of paedomorphosis in our smooth-newt population. Retention of larval feed- ing apparatus in paedomorphs may allow more efficient foraging on plankton and aquatic prey (Whiteman et al., 1996; Denoël, 2004). As a result, both microhabitat segregation and utilization of different food has been observed in particular morphs, with paedomorphs main- ly foraging on plankton and distributed in both shallow and deep parts of water column compared to foremost shallower distributed metamorphs (Denoël and Joly, 2001; Denoël and Schabetsberger, 2003). As the water reservoir is rich in planktonic micro-crustaceans and areas of deep water column, we suppose the predomi- nance of paedomorphs in our population to be inherent in these factors. It is however worth noting that predom- inance of paedomorphs in our smooth newt population could also be a result of an environmental-hormonal interplay, as both stress and steroid hormones were observed to significantly affect metamorphosis of larvae and paedomorphs to the adult stage (reviewed by Lau- det, 2011). Detailed endocrinological studies are needed to shed light on these proximate mechanisms. ACKNOWLEDGEMENTS The study was carried out in accordance to the permit SZ-092744/2012KUSK/3 issued by the Regional Office of the Central Bohemian Region of the Czech Republic and with a support by Vladimír Vaněk (the mayor of Stará Lysá). We also would like to thank Jiří Moravec (National Museum in Prague) and an anonymous reviewer for valuable discussion and com- ments, respectively. The research was supported by the Ministry of Culture of the Czech Republic (DKRVO 2013/13, National Museum, 00023272). VJ is also grateful to the Institutional Research Support of the Charles University in Prague and pro- ject SVV-2012-265-206. REFERENCES Banks, B. (1985): Observations on neoteny in the smooth newt. Br. Herpetol. Soc. Bull. 12: 37-38. Başkale, E., Sayim, F., Kaya, U. (2011): Body size and reproductive characteristics of paedomorphic and metamorphic individuals of the northern banded newt (Ommatotriton ophryticus). Acta Herpetol. 6: 19-25. Bennett, R.M., Chippindale, P. T. (2006): Streambed microstructure predicts evolution of development and life history mode in the plethodontid salamander Eurycea tynerensis. BMC Biol. 4: 1-12. Çiçek, K., Ayaz, D. (2011): New data on facultative pae- domorphism of the smooth newt, Lissotriton vulgaris, in Western Anatolia, Turkey. J. Freshwater Ecol. 26: 99-103. Covaciu-Marcov, S.D., Sas, I., Cicort-Lucaciu, A.Ş., Bog- dan, H.V. (2011): Lissotrion vulgaris paedomorphs in south-western Romania: a consequence of a human modified habitat? Acta Herpetol. 6: 15-18. Dely, O.G. (1967): Neuere Angaben zur Kenntnis des neotenischen Teichmolches (Triturus vulgaris L.). Acta Zool. Acad. Sci. Hungar. 13: 253-270. Denoël, M. (2002): Paedomorphosis in the Alpine newt (Triturus alpestris): decoupling behavioural and mor- phological change. Behav. Ecol. Sociobiol., 52: 394- 399. Denoël, M. (2003): How do paedomorphic newts cope with lake drying? Ecography, 26: 405-410. Denoël, M. (2004): Feeding performance in heterochro- nic Alpine newts is consistent with trophic habits and maintenance of polymorphism. Ethology 110: 127- 136. Denoël, M., Joly, P. (2001): Adaptive significance of facul- tative paedomorphosis in Triturus alpestris (Amphib- ia, Caudata): resource partitioning in an alpine lake. Freshw. Biol. 46: 1387-1396. Denoël, M., Poncin, P. (2001): The effect of food on growth and metamorphosis of paedomorphs Triturus alpestris apuanus. Archiv. Hydrobiol. 152: 661-670. Denoël, M., Schabetsberger, R. (2003): Resource parti- tioning in two heterochronic populations of Greek Alpine newts, Triturus alpestris veluchiensis. Acta Oecol. 24: 55-64. Denoël, M., Joly, P., Whiteman, H.H. (2005): Evolution- ary ecology of facultative paedomorphosis in newts and salamanders. Biol. Rev. 80: 663-671. Denoël, M., Duguet R., Džukić, G., Kalezić, M.L., Maz- zotti S. (2001): Biogeography and ecology of paedo- morphosis in Triturus alpestris (Amphibia, Caudata). J. Biogeogr. 28: 1271-1280. Denoël, M., Ficetola, G.F., Ćirović, R., Radović, D., Džukić, G., Kalezić, M.L., Vukov T.D. (2009): A mul- 57Paedomorphic smooth newts in the Czech Republic ti-scale approach to facultative paedomorphosis of European newts in the Montenegrin karst: distribu- tion pattern, environmental variables and conserva- tion. Biol. Conserv. 142: 509-517. Dolmen, D. (1981): Distribution and habitat of the Smooth Newt, Triturus vulgaris (L.), and the warty newt, T. cristatus (Laurenti), in Norway. Proc. Eur. Herp. Symp. Oxford 1980: 127-139. Harris, R.N., Semlitsch, R.D., Wilbur, H.M., Fauth, J.E., (1990): Local variation in the genetic-basis of pedo- morphosis in the salamander Ambystoma talpoideum. Evolution 44: 1588-1603. Kalezić, M.L., Džukić, G. (1985): Ecological aspects of the smooth newt (Triturus vulgaris) paedomorphosis from Montenegro. Arhiv. Bioloških. Nauka, Beograd 37: 43-50. Kalezić, M.L., Džukić, G. (1986): The frequent occurence of paedomorphosis in the smooth newt (Triturus vul- garis) population from the Submediterranean area of Yugoslavia. Amphibia-Reptilia 7: 86-89. Laudet, V. (2011): The origins and evolution of vertebrate metamorphosis. Curr. Biol. 21: 726-737. Litvinchuk, S.N. (2001): First record of paedomorphosis for the smooth newt (Triturus vulgaris) from Ukraine. Russ. J. Herpetol. 8: 77-78. Litvinchuk, S.N., Rudyk, A.M., Borkin, L.J. (1996): Obser- vation of paedomorphic newts (Triturus vulgaris) from the former Soviet union. Russ. J. Herpetol. 3: 39-48. Quitt, E. (1971): Klimatické oblasti Československa. Aca- demia. Studia Geographica, GÚ ČSAV., Brno. Raxworthy, C.J. (1990): A review of the smooth newt (Triturus vulgaris) subspecies, including an identifica- tion key. Herpetol. J. 1: 481-492. Ryan, T.J., Semlitsch, R.D. (2003): Growth and life cycle polymorphism in the salamander Ambystoma talpoi- deum. Biol. J. Linn. Soc. (London) 80: 639-646. Rot-Nikcević, I., Kalezić, M.L., Džukić, G. (2000): Paedo- genesis, life history traits and sexual dimorphism: a study case of the smooth newt, Triturus vulgaris, from Pannonia. Folia Zool. 49: 41-52. Schmidtler, J.F., Franzen M. (2004): Triturus vulgaris Lin- naeus, 1758 – Teichmolch. In: Handbuch der Reptili- en und Amphibien Europas, Bd. 4/2b, Schwanzlurche (Urodela) IIb, pp. 847-967. Grossenbacher, K., Thies- meier B., Eds, Aula Verlag, Wiesbaden. Semlitsch, R.D., Harris, R.N., Wilbur, H.M. (1990): Pae- domorphosis in Ambystoma talpoideum: maintenance of population variation and alternative life-history pathways. Evolution 44: 1604-1613. Voss, S.R., Prudic, K.L., Oliver, J.C., Shaffer, H.B. (2003): Candidate gene analysis of metamorphic timing in ambystomatid salamanders. Mol. Ecol. 12: 1217–1223. Wells, K.D. (2007): The Ecology and Behavior of Amphibians. The University of Chicago Press, Chica- go and London. Whiteman, H.H., Wissinger, S.A., Brown, W.S. (1996): Growth and foraging consequence of facultative pae- domorphosis in the tiger salamander Ambystoma tigrinum nebulosum. Evol. Ecol. 10: 433-446. 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