Acta Herpetologica 13(2): 101-108, 2018 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-22886 Yellow patterns polymorphism of the fire salamander Salamandra salamandra in Poland Anna Najbar1,3,*, Agnieszka Konowalik1,3, Bartłomiej Najbar2, Maria Ogielska1 1 Department of Evolutionary Biology and Conservation of Vertebrates, University of Wrocław, Sienkiewicza 21, 50-335 Wrocław, Poland. *Corresponding author. E-mail: anna.najbar@uwr.edu.pl 2 Faculty of Biological Sciences, University of Zielona Góra, Prof. Z. Szafrana 1, 65-516 Zielona Góra, Poland 3 A. Najbar and A. Konowalik contributed equally to this work Submitted on: 2018, 13th March; revised on: 2018, 15th May; accepted on: 2018, 9th July Editor: Marco Sannolo Abstract. We analysed variation of dorsal yellow patterns in 2077 individuals that represented 23 populations from the northern parts of the Sudetes and the Carpathian Mountains. We distinguished four types of yellow patterns: spotted (50.1%), spotted-and-striped (42.8%), zig-zag (5.8%), and striped (1.3%). Spotted form dominated in the east (69.2%, the Carpathians) and its frequency decreased to the west (46.4%, the Sudetes), where spotted-and-striped forms became more common. The rarest in both mountain ranges (0.9% in the Carpathians, 1.3% in the Sudetes) was striped type. The exception was the westernmost population represented by 15.8% of the striped individuals. We did not find evidence of differences between yellow colouration variants and genotypes of 557 individuals defined by 10 microsatellite loci. No differences were found between females and males. We concluded that patterning does not con- stitute a unique feature at population and mountain ranges levels. Keywords. Amphibians, colouration, dorsal yellow patterns, Poland, Salamandra salamandra. INTRODUCTION The fire salamander Salamandra salamandra (Lin- naeus, 1758) is highly polytypic and widely distributed urodelan in Europe (Kuzmin et al., 2009). The species consists of over a dozen divergent geographical subspe- cies traditionally distinguished by various patterns of yellow dots and/or stripes on the black background of the dorsal side, with a great variety observed at region- al, local, and individual levels (e.g., Freytag, 1955; Eiselt, 1958; Veith, 1992; Thiesmeier, 2004; Velo-Antón and Buckley, 2015). For many years, yellow-black colouration had been explained as a cryptic colouration closely linked to climate and quality of the substrate or background of habitats (e.g., Frish, 1920). Nowadays, it is known that phenotype depends on heritable genetic variation, and functions as an aposematic colouration (Thiesmeier, 2004; Thiesmeier and Grossenbacher, 2004). The differ- ences among populations are explained by evolutionary processes, such as natural selection. Among presumed factors, e.g., van Alphen and Arntzen (2016) mentioned predation intensity or thermal conditions. Current classification of Salamandra species and sub- species was supported by molecular studies, and some mor- phologically recognizable subspecies were also genetically differentiated (Steinfartz et al., 2000; Steinfartz and Tautz, 2003). On the other hand, phenotypical similarities exist between closely related species e.g., S. salamandra and S. infraimmaculata, or subspecies e.g., S. s. salamandra and S. s. terrestris (Degani, 1986; Thiesmeier, 2004). Colour poly- morphism is mostly evident among the fire salamanders inhabiting the Iberian Peninsula (Alcobendas et al., 1996; Thiesmeier and Grossenbacher, 2004; Reis et al., 2011; Velo- Antón and Buckley, 2015), but no genetic or geographical 102 Anna Najbar et alii divergence was recently revealed between phenotypically differentiated subspecies S. s. bernardezi or S. s. alfred- schmidti (Beukema et al., 2016). Therefore the authors sug- gested to revoke subspecies status of the latter taxon. Poland is inhabited by the nominative subspecies S. s. salamandra, characterized by yellow, or rarely orange, irregular spots and stripes on the dorsal part of the body (Juszczyk, 1987; Zakrzewski, 2007). The classification of dorsal yellow patterns described by Eiselt (1958) was fur- ther applied by Juszczyk (1987) and Zakrzewski (2007) for individuals collected in the Carpathians. Originally it contained four forms: spotted, striped, striped-spotted, and spotted-striped. Furthermore Paluch and Profus (2004) proposed to distinguish the fifth, i.e., zig-zag form. According to available data, spotted and striped-spotted forms dominate, and striped one is the rarest among the fire salamanders in Poland (Juszczyk, 1987; Paluch and Profus 2004; Zakrzewski, 2007). In addition, zig-zag form was described as extremely rare (Zakrzewski, 2007). Here, we present diversity and distribution of dorsal yellow patterns of the fire salamanders from the species entire range in Poland. The aim of this study was to char- acterize differences at population and mountain range levels, and between females and males. We assume that a gradient of dorsal patterns exists with domination of spotted form in the east (the Carpathians), and striped form in the west (the Sudetes). We also tested the pos- sible relationship between pattern types and genotypes based on 10 microsatellite loci. MATERIAL AND METHODS Sampling and study area 2077 individuals were collected during active seasons (April-October) in the years 2004-2016 from 23 populations in Poland. The study area represents the Western, Central and Eastern Sudetes Mountains (16 populations), and the Outer Western and Outer Eastern Carpathian Mountains (seven pop- ulations). They constitute two main mountain ranges in the country, isolated by the Moravian Gate (depression between the Eastern Sudetes and the Western Carpathians). Polish populations are located at the northern margin of the species range and inhabit mountainous areas including their foothills Table 1. Characteristic of sampling sites, years and the number of the collected individuals of the fire salamander Salamandra salamandra from Poland. Population number Mountain range Locality Altitude (m a.s.l.) Years of samples collection (number of collected individuals) 1 the Sudetes Eastern Upper Lusatia 283 2015 (22), 2016 (16) 2 the Sudetes Izera Mts. 470 2008 (51), 2009 (59) 3 the Sudetes Katzbach Mts. 494 2008 (24), 2009 (71) 4 the Sudetes Kaczawskie Piedmont 392 2008 (46), 2009 (57) 5 the Sudetes Kaczawskie Piedmont 394 2005 (116), 2006 (20), 2008 (7), 2009 (53) 6 the Sudetes Wałbrzyskie Mts. 509 2008 (26), 2009 (58) 7 the Sudetes Stołowe Mts. 589 2008 (40), 2009 (64) 8 the Sudetes Ślęża Massif 263 2004 (139), 2005 (28) 9 the Sudetes Bardzkie Mts. 407 2008 (42), 2009 (60) 10 the Sudetes Bardzkie Mts. 356 2006 (59), 2007 (38) 11 the Sudetes Bardzkie Mts. 279 2008 (41), 2009 (52) 12 the Sudetes Bardzkie Mts. 418 2008 (45), 2009 (67) 13 the Sudetes Bardzkie Mts. 294 2008 (40), 2009 (53) 14 the Sudetes Śnieżnik Massif 372 2007 (5), 2008 (42), 2009 (54) 15 the Sudetes Golden Mts. 496 2007 (122), 2009 (57), 16 the Sudetes Opawskie Mts. 401 2014 (44), 2015 (18) 17 the Carpathians Bielsko-Biała city, Silesian Piedmont 394 2015 (35), 2016 (43) 18 the Carpathians Rożnów Piedmont 351 2014 (4), 2016 (38) 19 the Carpathians the Ciężkowice Piedmont 378 2014 (8), 2015 (9), 2016 (41) 20 the Carpathians Strzyżów Piedmont 326 2014 (5), 2015 (2) 21 the Carpathians Lower Beskids 417 2014 (2), 2016 (42) 22 the Carpathians Lower Beskids 440 2014 (12), 2015 (40) 23 the Carpathians Bieszczady Mts. 615 2016 (60) 103Polymorphism of S. salamandra from Poland (Głowaciński and Zakrzewski, 2003). Dominant vegetation component in the Sudetes sampling sites are mixed or decidu- ous forests (including Dentario enneaphylli-Fagetum) with pre- dominance of the European beech Fagus sylvatica and admix- ture of e.g. spruce Picea sp., fir Abies sp. or sycamore Acer sp. (see Ogrodowczyk et al., 2010). In the Carpathians, salamanders inhabit large beech Dentario glandulosae-Fagetum or mixed forests with F. sylvatica as dominant species and highly shaded bottom with thick layer of litter. The exception is population no. 17 that inhabits small, highly isolated and contaminated hetero- geneous forest, located within urbanized area of Bielsko-Biała city (for details see Najbar et al., 2017). Although the Carpathians reach higher altitudes than the Sudetes, populations from both ranges are located at an aver- age altitude of 400 m above the sea level. Altitudes of each study site were determined in the center of a sampled site using Google Earth software (Google Inc.). Detailed description of the study area is presented in Table 1 and Fig. 1. Straight line minimum distance was approx. 1.1 km from population no. 11 to population no. 13 in the Sudetes, and maximum dis- tance reached approx. 580.6 km between population no. 1 and 23 (Fig. 1). Distances between populations were designated in Quantum GIS 2.18.2 software (QGIS Development Team 2013). In each population, transects were established by a breeding stream and salamanders were collected in the buffer of approx. 50 m up to 100 m on its both sides. Statistical and molecular analysis Statistical analyses were performed in STATISTICA 12 software (Statsoft, USA). Pearson’s χ2 and post-hoc χ2 were used to find differences in compositions of dorsal yellow patterns of individuals: a) from the Sudetes and the Carpathians, b) from westernmost population no. 1, the Sudetes and the Carpathians, and c) between males and females. Pearson’s χ2 was not used to test differences between populations due to small number or absence of striped and zig-zag forms, but frequencies of dor- sal yellow patterns are presented in Table 2 and Supplementary Table S1. To determine possible relationship between genetic varia- tion and type of dorsal colour pattern we used polymorphism of 10 microsatellite loci: Sal3, Sal23, Sal29, SalE5, SalE6, SalE7, SalE8, Sal E11, SalE12, SalE14 (Steinfartz et al., 2004), that were obtained in our previous study concerning population genetics of S. salamandra in Poland (for protocol see Najbar et al., 2015; Konowalik et al., 2016). Genotypes of 557 individuals represent- ing 19 populations from the entire Polish range were grouped according to five forms (see below preliminary results) and four forms in applied classification (Table 3). Values of FST between pairs of colour variants were calculated in ARLEQUIN 3.5 (Excoffier and Lisher, 2010). Their significance was tested with 10 000 permutations. Dorsal yellow patterns classification According to the criteria of Eiselt (1958), then applied by Juszczyk (1987), Paluch and Profus (2004), and Zakrzewski (2007), the classification of S. salamandra dorsal yellow pat- terning includes five distinct forms: spotted, striped, zig-zag, striped-spotted and spotted-striped (Fig. 2, Supplementary Table S1). However, the last two (striped-spotted and spotted- Fig. 1. Salamandra salamandra distribution range (Kuzmin et al., 2009) and the study sites in the Sudetes (left, populations 1 – 16) and the Carpathians (right, populations 17 – 23) in Poland. MG – the Moravian Gate. Map was created in CorelDRAW X3 software (Corel Corpora- tion, Canada). 104 Anna Najbar et alii striped) reveal difficulties in their unquestionable recogni- tion because intermediate forms may occur. To avoid signifi- cant mistakes while comparing various studies, we considered whether we were allowed to pool striped-spotted and spotted- striped forms into one spotted-and-striped form. Preliminary results supported simplification of the classification because 1) their frequencies were comparable in the entire studied area (Supplementary Table 2), and 2) no relationship between geno- types and patterning existed (Table 3A). Although we found significant differences in frequencies of striped-spotted and spotted-striped forms between the Sudetes and the Carpathians (post hoc χ2 = 13.13, P = 0.001), the above arguments support- ed this procedure. Therefore, in this study we applied the simplified classifi- cation of yellow patterns including four forms: A) spotted, B) striped, C) zig-zag, and D) spotted-and-striped (Fig. 2). All the collected salamanders were documented in the pic- tures and checked in order to avoid analysis of the same individ- ual twice or more times. Moreover, selected salamanders from the entire range were marked by toe clipping for further research what helped to recognize previously sampled individuals. RESULTS Spotted form dominated in the entire S. salamandra range and constituted 50.1% of all the collected individu- als. However, its frequency was higher in the Carpathi- ans (69.2%) than in the Sudetes (46.4%). The frequency of the remaining forms in the range was: 42.8% of spot- ted-and-striped form, 5.8% of zig-zag, and only 1.3% of striped form. The exception was population no. 1 where 15.8% of individuals represented striped form and the Table 2. Frequency of dorsal yellow patterns of the fire salamander Salamandra salamandra within populations, mountain ranges and sexes. The Carpathian populations are presented in bold. Location n Frequency [%] spotted striped zig-zag spotted-and-striped the entire species range in Poland 2077 50.1 1.3 5.8 42.8 the Sudetes 1736 46.4 1.3 6.5 45.8 the Carpathians 341 69.2 0.9 2.3 27.6 in each population: 1 38 13.2 15.8 13.2 57.9 2 110 60.0 0.0 4.5 35.5 3 95 46.3 1.1 4.2 48.4 4 103 65.0 0.0 2.9 32.0 5 196 27.0 0.0 1.0 71.9 6 84 57.1 1.2 10.7 31.0 7 104 47.1 1.9 1.0 50.0 8 167 47.3 0.6 4.8 47.3 9 102 43.1 2.0 3.9 51.0 10 97 50.5 0.0 14.4 35.1 11 93 52.7 1.1 8.6 37.6 12 112 51.8 1.8 13.4 33.0 13 93 50.5 1.1 9.7 38.7 14 101 43.6 1.0 12.9 42.6 15 179 46.4 0.0 6.7 46.9 16 62 32.3 8.1 1.6 58.1 17 78 55.1 2.6 1.3 41.0 18 42 73.8 0.0 0.0 26.2 19 58 77.6 0.0 8.6 13.8 20 7 42.9 0.0 0.0 57.1 21 44 63.6 0.0 0.0 36.4 22 52 76.9 1.9 1.9 19.2 23 60 76.7 0.0 1.7 21.7 females 936 47.9 1.0 5.9 45.3 males 884 46.5 1.9 5.4 46.2 juveniles 252 70.2 0.0 7.1 22.6 105Polymorphism of S. salamandra from Poland lowest frequency (13.2%) of spotted salamanders was found. In addition, population no. 16 also revealed higher number (8.2%) of striped forms. Detailed predominance and frequency of dorsal colour patterns within popula- tions are presented in Table 2 and Fig. 3A, and Supple- mentary Table S1. Statistical analyses showed significant differences in the frequencies of dorsal yellow patterns between sala- manders from the Sudetes and the Carpathians (χ2 = 60.72, P < 0.001), and while comparing population no. 1 with western (the Sudetes) and eastern (the Carpathians) groups of populations (χ2 = 138.48, P < 0.001). Post-hoc Table 3. Dorsal yellow patterns pairwise FST (below diagonal) for: A) classification of Eiselt (1958), applied by Juszczyk (1987), Paluch and Profus (2004) and Zakrzewski (2007) for Polish populations, and B) simplified classification used in this study. Above diagonal FST P values are presented. A n spotted striped zig-zag striped-spotted spotted-striped 297 spotted - 0.758 0.887 0.412 0.273 8 striped -0.004 - 0.849 0.703 0.915 22 zig-zag -0.003 -0.008 - 0.368 0.934 84 striped-spotted 0.001 -0.005 -0.002 - 0.719 146 spotted-striped 0.001 -0.009 -0.004 -0.001 - B n spotted striped zig-zag spotted-and-striped 297 spotted - 0.763 0.886 0.158 8 striped -0.004 - 0.841 0.842 22 zig-zag -0.003 -0.008 - 0.870 230 spotted-and-striped 0.001 -0.007 -0.003 - Fig. 2. Classification of colour patterns of the fire salamander Salamandra s. salamandra. A – spotted form: irregular spots also on vertebral line; B – striped form: two rows of full or discontinuous stripes, vertebral line remains black without any spots; C – zig-zag form: continuous, rarely interrupted in one or two places, linear pattern that covers vertebral line; D – spotted-and-striped form: spots and striped, vertebral line remains black or may be disturbed by yellow spots; this intermediate form consists of striped-spotted and spotted-striped forms described by Eiselt (1958), and applied for Polish populations of the species by Juszczyk (1987), Paluch and Profus (2004) and Zakrzewski (2007). 106 Anna Najbar et alii χ2 results revealed differences between mountain rang- es in the proportion of spotted (χ2 = 59.46, P < 0.001), zig-zag (χ2 = 9.00, P = 0.003), and spotted-and-striped individuals (χ2 = 38.69, P < 0.001). Population no. 1 dif- fered in the proportion of spotted (χ2 = 17.23, P < 0.001) and striped (χ2 = 62.18, P < 0.001) forms with salaman- ders from the Sudetes, and in the proportion of all forms with individuals collected in the Carpathians (for all P < 0.001). We did not find differences between males and females (χ2 = 3.36, P = 0.340). Values of pairwise FST between forms of dorsal pat- terning varied from -0.008 to 0.001 (for all P > 0.05; Table 3B). The results suggested no evidence of genetic differences between individuals that represent defined patterns at least according to variation of selected molec- ular markers used in this study. DISCUSSION It has been recognized that spotted forms of S. sala- mandra are characteristic of Eastern Europe, while striped individuals with two yellow bands are domi- nant in the West (e.g., Eisely, 1958; Thiesmeier, 2004). In Poland, S. s. salamandra is mainly represented by indi- viduals with irregular spots and/or stripes on the dorsal side of the body. In our study, spotted form was the most common, whereas striped form was the rarest (Table 2). Predomination of spotted and striped-spotted forms (a part of spotted-and-striped form according to clas- sification applied for this study, Fig. 2D) was previously reported by Juszczyk (1987), Paluch and Profus (2004), and Zakrzewski (2007). This result complies with dis- tribution of S. salamandra subspecies recognized on the basis of variety in regionally observed colouration. In relation to other populations, substantial pre- dominance of striped form, characteristic of S. s. terrestris (Lacépede, 1788), was observed in site no. 1. This popu- lation is located at the Polish-German border along the Lusatian Neisse (Najbar and Najbar, 2015), where ranges of these two subspecies contact (Thiesmeier, 2004). Fre- quency of striped form in this site was higher than in the remaining sites (Table 2), while spotted and zig-zag rep- resented the minority of observed individuals. Significant dominance of striped patterns in the extremely western population no. 1 may suggest natural admixture with S. s. terrestris, as was also suggested by Thiesmeier (2004). Such results indicated an increase in the frequency of striped form towards the West. On the other hand, rela- tively large number of striped salamanders was observed also in population no. 16 located in the Opawskie Moun- tains (Fig. 1). Likewise, Paluch and Profus (2004) report- ed 4.8% of salamanders with the striped dorsal pattern in population from Morzyszów, which is distanced of about 130 km to the east of population no. 1. Although we observed striped and zig-zag salamanders in both the Sudetes and the Carpathians (Table 2), these patterns were confirmed to be rare similarly to other studies from Poland (Juszczyk, 1987; Paluch and Profus, 2004; Zakrze- wski, 2007). Phenotypes of several Salamandra species and their subspecies tend to be misleading in their recognition (e.g., Bickford et al., 2007), and the fire salamanders col- lected in this study resemble at least two subspecies, e.g. S. s. terrestris and S. s. beschkovi (Obst, 1981). Kowalski and Młynarski (1965) reported similarities in yellow pat- terning between some of the collected individuals from Poland and those representing western European subspe- cies S. s. terrestris, whereas Juszczyk (1987) found simi- larities to S. s. algira, S. s. infraimmaculata or S. s. bejarae Fig. 3. Frequency of dorsal yellow patterns: A) in the Carpathians and the Sudetes, and B) the Carpathians, the Sudetes (excluding population no. 1), and the westernmost population no. 1. 107Polymorphism of S. salamandra from Poland according to subspecies classification of Eiselt (1958). The first two do not exist in the current systematics anymore (Joger and Steinfartz, 1995; Steinfartz et al., 2000) and were recognized as distinct species: S. algira (Bedriaga, 1883) and S. infraimmaculata (Mertens, 1948). On the other hand, the existence of several distinct phenotypes may be observed in Spanish populations of S. s. bernarde- zi (Beukema et al., 2016). Sexual polymorphism has been frequently report- ed for amphibian species, including body size, colour dichromatism, pigmentation, quality or quantity of pat- terns (e.g., Kupfer, 2007; Davis and Grayson, 2008; Hoff- man and Blouin, 2008). In S. salamandra, sex-related morphological features, such as size and shape of the body, were usually observed (e.g., Zakrzewski, 2007; Labus et al., 2013; Balogová and Uhrin, 2015). Our study did not reveal relationship between the occurrence of exact dorsal patterns and sexes, and correspond to pre- vious conclusions of Zakrzewski and Wójcik (1982) that females and males did not differ in the topography of spots. However, a recent study of Balogová and Uhrin (2015) from the Slovakian Carpathians revealed the exist- ence of sexual dichromatism in S. s. salamandra, and concluded that males were covered with larger yellow spots on dorsal side than females. Genetic diversity in 10 microsatellite loci did not showed correlation with a variety of dorsal yellow pat- terns. However, the westernmost population no. 1 was not included to genetic analyses because the site was dis- covered later than when we conducted population genet- ics study (Najbar et al., 2015; Konowalik et al., 2016). Veith (1992) excluded the possibility to recognize of S. s. salamandra and S. s. terrestris by color patterns in wide zone of hybrid populations from Germany. Therefore, further genetic analyses should be performed to confirm our hypothesis about possible occurrence of the second subspecies, S. s. terrestris, in Poland. ACKNOWLEDGEMENTS We thank Editor and two anonymous Reviewers for useful comments that helped to improved our manu- script. We also thank Wiesław Babik for useful sugges- tions, and Tomasz Januszkiewicz, Ewa Kunicka-Najbar, Monika Malinowska, Joanna Górajewska, Katarzyna Serwa, and Anna Szwedor for help with samples col- lection. The project was approved by Polish Ministry of Environment (permit No. DOPog.-4201-02-74/05kl), General Directorate for Environmental Protection (per- mit No. DZP-WG.6401.02.7.2014.JRO), Regional Direc- torate for Environmental Protection in Wrocław (permit No. WPN.6401.211.2015.MR.2), and the II Local Ethics Commission for Animal Experiments in Wrocław (Reso- lution No. 63/2008, 78/2014 and 68/2015). The study was financed by grants KBN NN 303 4147737, DS 1076/S/ IBŚ/2014, and 0420/1409/16. SUPPLEMENTARY MATERIAL Supplementary material associated with this article can be found at manuscript number 22886. REFERENCES Alcobendas, M., Dopazo, H., Alberch, P. (1996): Geo- graphic variation in allozymes of populations of Sala- mandra salamandra (Amphibia: Urodela) exhibiting distinct reproductive modes. J. Evol. Biol. 9: 83-102. van Alphen, J.J.M., Arntzen J.W. (2016): Paul Krammerer and the inheritance of acquired characteristics. Contr. Zool. 85: 457-470. Balogová, M., Uhrin, M. (2015): Sex-biased dorsal spot- ted patterns in the fire salamander (Salamandra sala- mandra). Salamandra 51: 12-18. Beukema, W., Nicieza, A.G., Lourenço, A., Velo-Antón, G. (2016): Colour polymorphism in Salamandra sala- mandra (Amphibia: Urodela), revealed by a lack of genetic and environmental differentiation between distinct phenotypes. J. Zool. Syst. Evol. Res. doi: 10.1111/jzs.12119. Bickford, D., Lohman, D.J., Sohdi, N.S., Ng, P.K.L., Meier, R., Winker, K., Ingram, K.K., Das, I. (2007): Cryptic species as a window on diversity and conservation. Trends. Ecol. Evol. 22: 148-155. Degani, G. (1986): Plasma proteins and morphology of Salamandra salamandra in Israel. Amphibia-Reptilia 7: 105-114. Eiselt, J. (1958): Der Feuersalamander (Salamandra sala- mandra L.) Beiträge zu einer taxonomischen Syn- these, Abh. Ber. Mus. Magdeburg B.X.6: 77-154. Excoffier, L., Lisher H.L. (2010): Arlequin suite ver 3.5: a new series of programs to perform population genet- ics analyses under Linux and Windows. Mol. Ecol. Res. 10: 564-567. Frisch, K. (1920): Über den Einfluss der Bodenfarbe auf die Fleckenzeichnung des Feuersalamanders. Biologis- ches Zentralblatt 40: 390-414. Głowaciński, Z., Zakrzewski, M. (2003): Salamandra plamista Salamandra salamandra (Linnaeus, 1758). In: Głowaciński, Z., Rafiński, J. (eds), Atlas rozmiesz- 108 Anna Najbar et alii czenia płazów i gadów Polski. Status – Rozmieszc- zenie – Ochrona 27-29. Wydawnictwo IOŚ i IOP PAN, Kraków. Hoffman, E.A., Blouin, M.S. (2000): A review of colour and pattern polymorphisms in anurans. Biol. J. Linn. Soc. 70: 633-665. Joger, U., Steinfartz, S. (1995): Protein electrophoretic data on taxonomic problems in East Mediterranean Salamandra (Urodela: Salamandridae). In: Llorente, G.A. (eds), Scientia Herpetologica. Barcelona: Aso- ciación Herpetológica Española: 33-36. Juszczyk, W. (1987): Płazy i gady krajowe. PWN, War- szawa. Kupfer, A. (2007): Sexual size dimorphism in amphibians: an overview. In: Fairbairn, D.J., Blanckenhorn, W.U., Szekely, T. (eds), Sex, size and gender roles: evolution- ary studies of sexual size dimorphism. Oxford Univer- sity Press, Oxford. Kuzmin, S., Papenfuss, T., Sparreboom, M., Ugurtas, I.H., Anderson, S., Beebee, T., Denoël, M., Andreone, F., Anthony, B., Schmidt, B., Ogrodowczyk, A., Ogielska, M., Bosch, J., Tarkhnishvili, D., Ishchenko, V. (2009): Salamandra salamandra. The IUCN Red List of Threatened Species. Version 2012.1. www.iucnredlist. org [accessed on 17 May 2017]. Konowalik, A., Najbar, A., Babik, W., Steinfartz, S., Ogiel- ska, M. (2016): Genetic structure of the fire salaman- der Salamandra salamandra in the Polish Sudetes. Amphibia-Reptilia 37: 405-415. Kowalski, W., Młynarski, M. (1965): Uwagi o płazach i gadach Pienińskiego Parku Narodowego. Ochrona Przyrody 31: 87-115. Kraków. Labus, N., Cvijanovič, M., Vukov, T. (2013): Sexual size and shape dimorphism in Salamandra salamandra (Amphibia, Caudata, Salamandridae) from the Cen- tral Balkans. Arch. Biol. Sci. Belgrade. 65: 969-975. Najbar, A., Najbar, B. (2015): Stanowisko salamandry plamistej Salamandra salamandra na Pogórzu Zacho- dniosudeckim. Chrońmy Przyrodę Ojczystą 71: 300- 303. Najbar, A., Babik, W., Najbar, B., Ogielska, M. (2015): Genetic structure and differentiation of the fire sala- mander Salamandra salamandra at the northern mar- gin of its range in the Carpathians. Amphibia-Reptilia 36: 301-311. Najbar, A., Rusek, A., Najbar, B. (2017): Zagrożenia i propozycje ochronne salamandry plamistej Sala- mandra salamandra w zurbanizowanym siedlisku w Bielsku-Białej. Chrońmy Przyrodę Ojczystą 71: 300- 303. Ogrodowczyk, A., Ogielska, M., Kierzkowski, P., Maślak, R. (2010): Występowanie salamandry plamistej Sala- mandra s. salamandra Linnaeus, 1758 na Dolnym Śląsku. Przyroda Sudetów 13: 179-192. Paluch, A., Profus, P. (2004): Status i rozmieszczenie salamandry plamistej Salamandra salamandra (Lin- neus, 1758) w Polsce ze szczególnym uwzględnieniem populacji gatunku w Górach Bardzkich (Sudety Środkowe). Chrońmy Przyrodę Ojczystą 60: 49-77. Reis, D.M., Cunha, R.L., Patrão, C., Rebelo, R., Castilho, R. (2011): Salamandra salamandra (Amphibia: Cau- data: Salamandridae) in Portugal: not all black and yellow. Genetica 139: 1095-1105. Steinfartz, S., Veith, M., Tautz, D. (2000): Mitochondrial sequence analysis of Salamandra taxa suggest old splits of major lineages and postglacial recolonizations of Central Europe from distinct source populations of Salamandra salamandra. Mol. Ecol. 9: 397-410. Steinfartz, S., Tautz, D. (2003): The Fire Salamander: Source for New Species. German Research. Life Sci- ences 25: 14-16. Steinfartz, S., Küesters, D., Tautz, D. (2004): Isolation and characterization of polymorphic tetranucleotide microsatellite loci in the fire salamandra Salamandra salamandra (Amphibia: Caudata). Mol. Ecol. Notes 4: 626-628. Thiesmeier, B. (2004): Der Feuersalamander. Laurenti- Verlag. Bielefeld. Thiesmeier, B., Grossenbacher, K. (2004): Salamandra salamandra (Linnaeus, 1758) – Feuersalamander. Pp. 1059-1132 in: Thiesmeier B., Grossenbacher K. (Eds.) Handbuch der Reptilien und Amphibien Europas. Schwanzlurche IIB. Aula Verlag, Wiebelsheim. Veith, M. (1992): The fire salamander, Salamandra sala- mandra L., in central Europe: subspecies distribution and intergradation. Amphibia-Reptilia 13: 297-313. Velo-Antón, G., Buckley, D. (2015): Salamandra común – Salamandra salamandra (Linnaeus, 1758). In: Salva- dor, A., Martínez-Solano, I. (Eds.): Enciclopedia Vir- tual de los Vertebrados Españoles. Museo Nacional de Ciencias Naturales, Madrid. Zakrzewski, M., Wójcik, S. (1982): Charakterystyka ubarwienia rasy nominalnej salamandry plamistej Salamandra salamandra, (L.) pochodzącej z trzech różnych populacji. Rocznik Naukowo-Dydydaktyczny. Prace Zoologiczne IV (82): 131-139. WSP Kraków im. Komisji Edukacji Narodowej w Krakowie. Zakrzewski, M. (2007): Salamandra plamista. Rozmiesz- czenie, biologia i zagrożenia. Wydawnictwo Naukowe AP, Kraków. Acta Herpetologica Vol. 13, n. 1 - June 2018 Firenze University Press