Basic and Applied Herpetology 33 (2019) 81-91

A catalogue of of Monfragüe National Park (Spain), with molecular characterization of populations of Wagler, 1830 in this protected area

Daniel Fernández-Ortín1, Gregorio Sánchez-Montes2, Íñigo Martínez-Solano2,*

1 Betania, 1, bajo G, 10003 Cáceres, Spain. 2 Departamento de Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales, José Gutiér- rez Abascal, 2, 28006 Madrid, Spain.

*Correspondence: E-mail: [email protected]

Received: 17 July 2019; returned for review: 29 October 2019; accepted 06 November 2017.

Monfragüe National Park (Cáceres, Extremadura, Spain) is a protected area in central‐western Iberia,including some of the best preserved primary Mediterranean vegetation. Legal protection dates back to 1979 (first as a Natural Park and then as a National Park), but knowledge about its communities is so far limited to sparse records. In this paper we present an updated list based on 521 records covering 163 1x1 km UTM grids in the study area, compiled in the period 2000‐2019. We detected 20 native species, representing 71.4% of the reptile fauna in Extremadura and 35% of the ibero‐balearic reptile fauna. Additionally, based on molecular analyses populations of the amphisbaenid Blanus in the study area are assigned to the oriental Iberian taxon, B. cinereus. The new records extend the known distribution of the different reptile species in the study area in 56 10x10 km UTM grids. Species presenting more restricted distributions in Monfragüe are Lacerta schreiberi (one 1x1 grid), Emys orbicularis, and Acanthodactylus erythrurus (four 10x10 grids each).

Key words: Blanus cinereus; Extremadura; Iberian Peninsula; Monfragüe National Park; reptiles.

The Iberian Peninsula hosts a great ripheral Protected Area (“Zona Periférica diversity of habitats and ecosystems, a de Protección”, ZPP) conform a study area legacy of its deep geological history and of 116 000 ha (Fig. 1), which includes other topographic complexity. National parks protected areas like ZEPA “ES0000014 provide legal protection for representative Monfragüe y las Dehesas del Entorno”, natural areas including singular geological ZEC “ES4320077 Monfragüe” and a Bio‐ characteristics, flora, and fauna. Monfra‐ sphere Reserve. güe National Park is located in the prov‐ Monfragüe National Park has been ince of Cáceres, in the region of Extrema‐ legally protected since 1979 (first as a Nat‐ dura, in central‐western Iberia, occupying ural Park and since 2007 as a National an extension of 18 396 ha of rugged terrain Park), but there is little information about with steep slopes, and includes some of its reptile communities. Apart from the the best preserved primary Mediterranean study of Álvarez‐Vasserot (1998), the vegetation. The National Park and its Pe‐ information about the reptile fauna of

DOI: http://dx.doi.org/10.11160/bah.173 Supplementary material available online FERNÁNDEZ-ORTÍN ET AL.

Figure 1: Location of the study area.

Monfragüe consists of sparse records, with mostly focusing on amphibians. Other in‐ the exception of a survey conducted by formation about reptiles in national parks biology students from the University of comprises specific reports on conservation Extremadura (MuÑoz del Viejo, 2005). The plans for threatened species in the Canary poor knowledge about reptiles in Monfra‐ islands or isolated records, like the first güe is in line with the situation in most of report of the presence of Lacerta schreiberi the 15 national parks in Spain (16 in Iberia Bedriaga, 1878 in Monfragüe (Fernández‐ considering Peneda Gerês National Park Ortín Toboso‐Borrella, 2014). in Portugal). Of all these protected areas, Of all reptile taxa present in Monfragüe there are only detailed studies and species National Park, genus Blanus Wagler, 1830 catalogues in the national parks of Islas is the only case presenting a conflict re‐ Atlánticas (Galán Regalado, 2003), garding specific characterization of its Peneda Gerês (Soares et al., 2005), Picos populations. At present, two lineages are de Europa (AyllÓn et al., 2010), and Do‐ known to occur in Iberia: an occidental ñana (Andreu, 2014). In other national lineage, in the Iberian southwest, and an parks like Cabañeros, Sierra de Guadarra‐ oriental one distributed across the rest of ma and Sierra Nevada there are monitor‐ Iberia. In 2009, Albert Fernández de‐ ing programs of herptile populations, but scribed a new blanid species in Iberia,

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Blanus mariae Albert Fernández, 2009. güe National Park, and new molecular Its distribution encompasses the south‐ evidence regarding the specific status of western quadrant of Iberia, thus represent‐ Blanus populations in this protected area. We ing the occidental lineage, with its north‐ provide >500 new records documenting ern range border in the province of Cáce‐ the presence of 20 native reptile species in res (Albert Fernández, 2009). Albert et Monfragüe, and show the presence of B. al. (2007) identified the closest Blanus rec‐ cinereus in two localities in the study area ords to Monfragüe as Blanus cinereus based on the analysis of sequences of the Vandelli, 1797, at the southern Tajo river mitochondrial DNA gene ND4. margin, in the locality of Garciaz in Cáce‐ Materials and Methods res. Nevertheless, Sampaio et al. (2014) recorded the presence of B. mariae north This study is based on a compilation of of the Tajo river, in the portuguese locality observations of reptiles from surveys con‐ of Carvalhão, suggesting that the two spe‐ ducted between 2000 and June 2019, docu‐ cies of Iberian Blanus could coexist in the mented in a database including photo‐ vicinities of Monfragüe National Park, graphic records. These records were ob‐ since both lineages occupy both the north‐ tained through direct observation, photo‐ ern and southern margins of the Tajo river. graphs, and, occasionally, identification in Based on alleged nomenclatural prob‐ hand. No individual was sacrificed for lems in Albert Fernández’s (2009) de‐ identification or tissue sampling. scription of B. mariae, Ceríaco Bauer The study area includes Monfragüe (2018) concluded that B. mariae is a junior National Park and its ZPP; as geographic synonym of B. cinereus and described reference we used the system UTM_MGRS Blanus vandellii Ceríaco Bauer, 2018 as a ED50, zones 29N and 30N, codes EPSG: new species. Thus, according to these au‐ 23029 and 23030, and referenced observa‐ thors, all B. cinereus records should be tions in 1x1 km UTM grids. For georefe‐ ascribed to the new species, B. vandellii, rencing, we used cartographic viewers and B. mariae records should be assigned (“Sistema de Información Geográfica de to B. cinereus. However, the existence of Parcelas Agrícolas del Ministerio de Agri‐ another problematic taxon, B. rufus cultura y Pesca, Alimentación y Medio (Hemprich, 1820), with type locality in Ambiente, SIGPAC” (Mapa.gob.es, s.f.), “southern Spain”, an area where the two and “Instituto Geográfico Nacional, IBER‐ Iberian lineages co‐occur, suggests that the PIX” (Instituto Geográfico Nacional de new species, B. vandellii, could be likewise España, s.f.). Toponyms were obtained considered a junior synonym of B. rufus. from 1:25 000 maps in both cartographic This circumstance will not be resolved un‐ servers or from local denominations. til the type specimen of B. rufus can be The orography of Monfragüe is defined assigned to either the occidental or the by two major mountain chains running in oriental Iberian Blanus lineage. parallel NW‐SE, with the highest peaks at In the current study, we present an up‐ Pico de Miravete (839 m a.s.l.) and Sierra dated reptile species catalogue for Monfra‐ de Enmedio (870 m a.s.l.), in Casas de

83 FERNÁNDEZ-ORTÍN ET AL.

Figure 2: Sampling effort (grey squares) and biogeographic regions in the study area, showing 10 x 10 km UTM grids.

Miravete and Deleitosa, respectively. Nev‐ part of the Western Iberian Mediterranean ertheless, most of the study area comprises biogeographic province, “Luso‐ elevations ranging from 220 m to 600 m. Extremadurense” subprovince, “Toledano‐ The two most prominent hydrographic Tagano” sector, and include five districts: features are rivers Tajo and Tiétar, embed‐ “Talaverano”, “Vereño” and “Coriano”, in ded in steep margins and dammed along the “Talaverano‐Placentino” subsector; their full course in the study area. The hy‐ and “Cacereño” and “Villuerquino”, in the drological network in Monfragüe is com‐ “Oretano” subsector (Fig. 2). Average an‐ plemented with second order streams as‐ nual rainfall ranges from 868 mm in Mal‐ sociated with these two major river basins; partida de Plasencia to 551 mm in Torrejón most of them are temporary, with higher el Rubio. Most of the study area presents water flow in the spring, and low flow climatic values associated with the Ocean‐ during the drier months. The water table ic Pluvistational Mediterranean biocli‐ only surfaces at fountains and springs, mate, except the southeastern part, where with no relevant endorreic ponds except climatic data from the station at Campillo Laguna de Cantalgallo, in Jaraicejo, and de Deleitosa (Fig. 1) provides values with‐ Lagunas de Mesas Caveras, in Deleitosa. in the Continental Pluvistational Mediter‐ According to Belmonte LÓpez (2008) ranean type. Regarding thermal and om‐ Monfragüe National Park and its ZPP are brotypes, almost all the study area is with‐

84 REPTILES OF MONFRAGÜE AND MOLECULAR ANALYSIS OF BLANUS in the Mesomediterranean (“Inferior‐Seco‐ mendations of Asociación Herpetológica Superior”) realm, except some spots at the Española (Carretero et al., 2018). Thus, southeastern border, which present char‐ regarding genus Blanus, we follow their acteristics within the Mesomediterranean decision to accept B. mariae as a valid tax‐ (“Superior‐Subhúmedo‐Inferior”) realm. on, with B. cinereus representing the ori‐ With respect to temperatures, the station ental lineage in Iberia. For estimates of of Campillo de Deleitosa is singular, with ibero‐balearic species richness we exclud‐ periods of freezing every year, but average ed allochthonous taxa and marine species. annual temperatures in the study area Observations recorded during field sur‐ range from 18ºC in Torrejón el Rubio to veys were contrasted with literature rec‐ 15ºC in Campillo de Deleitosa. ords (Pleguezuelos et al., 2002), and the We used specific methodologies to lo‐ database of amphibian and reptile records cate different species all year round, like maintained by Asociación Herpetológica nocturnal surveys for mauritani- Española (S.I.A.R.E., s.f.). Biogeographic ca Linnaeus, 1758 and Hemidactylus turci- classification of species follows Salvador cus (Linnaeus, 1758), or inspecting rocks Milla (2002). and other refugia, which was useful to For the molecular characterization of locate fossorial species like B. cinereus. We samples of Blanus in Monfragüe we ana‐ also recorded road‐killed individuals dur‐ lysed sequences of a fragment of mito‐ ing road surveys. When planning these chondrial gene ND4 and adjacent tRNAs prospections, we took into consideration in two individuals collected in Torrejón el the environmental requirements of target Rubio (accession code, Tissue and DNA species, and the bioclimatic characteristics collection, Museo Nacional de Ciencias of different parts of the study area. For Naturales: MNCN:ADN: 94916), and instance, Lacerta schreiberi requires espe‐ Casas de Miravete (accession code: cially humid areas, and thus sampling MNCN:ADN: 94917) (see location in the sites were selected using other taxa as bio‐ study area in Supplementary Material). indicators, like the Iberian frog Rana We extracted DNA from tissue samples iberica Boulenger, 1879 or the trees Alnus stored in ethanol with a commercial kit glutinosa (L.) Gaertn. and Prunus lusitani- (QIAquick, QIAGEN, Germany). Then, we ca L., which are typical from humid biocli‐ amplified the target fragment via polymer‐ mates. Prospected 1 x 1 km grids were se‐ ase chain reaction (PCR) with primers lected based on their accessibility and the ND4 and LEU (Arévalo et al., 1994), fol‐ presence of favourable habitats for rep‐ lowing the conditions described in Albert tiles. We did not perform systematic sur‐ et al. (2007), with minor modifications. veys, but many sites have been visited of‐ Amplified products were sequenced in ten. Thus, we acknowledge that many spe‐ both directions with PCR primers and se‐ cies records correspond to chance observa‐ quences were assembled and edited with tions. software Sequencher (Gene Codes, USA). Systematic treatment of the species fol‐ New sequences were compared with those lows the most recent taxonomic recom‐ available in GenBank (https://

85 FERNÁNDEZ-ORTÍN ET AL. www.ncbi.nlm.nih.gov/genbank/) using GenBank showed similarity scores ≥ the BLAST algorithm (https:// 94.75% with sequences of Blanus cinereus, blast.ncbi.nlm.nih.gov/Blast.cgi), which and ≤ 89.1% with Blanus mariae, confirm‐ identifies the most similar sequences in the ing the presence of the former species in database. Monfragüe National Park in the two stud‐ ied localities. Results Discussion Based on our fieldwork we compiled a database with 521 records in 163 1x1 km We found all the species previously UTM grids, 12.4% of the study area (Fig. cited in Monfragüe, representing nine out 2), and detected 20 native species (Table 1, of the 13 reptile families present in the Ibe‐ see maps in Supplementary Material), rep‐ rian Peninsula and the Balearic Islands. Of resenting 71.4% of the reptile species in the 11 reptile families present in Extrema‐ Extremadura and 35% of the ibero‐balearic dura, only and are reptile fauna. These records expand our missing. Families Testudinidae and Cha‐ knowledge of the distribution of reptiles in maeleonidae, absent in Extremadura, are Monfragüe in 55 new 10x10 km UTM grids not present in Monfragüe either. (see maps in Supplementary Material). In Of all reptile families present in Mon‐ addition, in 2004 we found an adult indi‐ fragüe, has the lowest represen‐ vidual of the alien, invasive species Tra- tation of its ibero‐balearic diversity, with chemys scripta elegans (Wied, 1838), but it just 20% of species present in the study was not seen again in subsequent years. area. This is probably related with the low There are eight reptile species in Extre‐ altitudinal range in Monfragüe, with alti‐ madura that have not been reported to tudes mostly between 220 and 600 m a.s.l. occur in Monfragüe, and did not appear in This can explain the absence of species our surveys (Table 2). Two of them have with strict orophylous requirements, like been reported in localities close to the genus Iberolacerta Arribas, 1997, which is study area: Hemidactylus turcicus and characterized by high taxonomic diversity. Podarcis guadarramae (Boscá, 1916), which Likewise, another taxonomically diverse could therefore be present in Monfragüe. but poorly represented genus in Monfra‐ Hemidactylus turcicus has been reported in güe is Podarcis Wagler, 1830, with a sin‐ Cañaveral (S.I.A.R.E., s.f.), near the west‐ gle representative, Podarcis virescens Ge‐ ern limit of the study area, whereas there niez, Sá‐Sousa, Guillaume, Cluchier is a small, isolated population of P. gua- Crochet, 2014. The most likely reason is darramae in Trujillo (Geniez et al., 2014), also the lack of ecological heterogeneity, in south of the study area. addition to the phylogenetic characteristics We obtained two sequences of 848 base of the group, formed by cryptic species pairs for the two individuals of Blanus an‐ with parapatric distributions (Geniez et alysed (GenBank accession numbers: al., 2007; Kaliontzopoulou et al., 2012). MK978773‐MK978774). Comparison of The faunistic composition of the reptile these sequences with those available in communities of Monfragüe is strictly Med‐

86 REPTILES OF MONFRAGÜE AND MOLECULAR ANALYSIS OF BLANUS

Table 1: Taxonomic diversity of reptiles in Monfragüe National Park. Sp = number of species in Monfragüe, P1 = proportion of species with respect to Extremadura and P2 = proportion of species with respect to the Iberian Peninsula and the Balearic Islands, 1x1 = number of 1x1 km UTM grids where the species was found in this work, 10x10 = number of 10x10 km UTM grids with records for the species, Distr. = distribution, Pal = Palearctic, Med = Mediterranean, End = Iberian endemic.

Order Family Species 1x1 10x10 Distr. CHELONII Emydidae Emys orbicularis 4 4 Pal Sp: 2 Sp: 1; P1:100%; P2:100% P1: 100% Geoemydidae Mauremys leprosa 23 23 Med P2: 50% Sp: 1; P1:100%; P2:100% Phyllodactylidae 23 17 Med Sp: 1; P1:100%; P2:100% Scincidae Chalcides bedriagai 4 5 End Sp: 2; P1:100%; P2:100% Chalcides striatus 20 10 Med Acanthodactylus 1 4 Med erythrurus Lacerta schreiberi 1 1 End Podarcis virescens 27 12 End Lacertidae Psammodromus Sp: 6; P1:60%; P2:20% 59 23 Med algirus Psammodromus 12 9 End occidentalis Timon lepidus 56 24 Med Sp: 18 Blanidae 1 Blanus cinereus 18 16 End P : 69,2% Sp: 1; P1:50%; P2:50% P2: 34% Hemorrhois 18 13 Med hippocrepis Zamenis scalaris 18 15 Med Sp: 6; P1:85.7%; Coronella girondica 12 11 Med P2:54.5% Macroprotodon brevis 12 9 Med Natrix maura 27 23 Med Natrix astreptophora 8 8 Med Lamprophiidae Malpolon 24 18 Med Sp: 1; P1:100%; P2:100% monspessulanus Sp: 1; P1:100%; Vipera latastei 7 6 Med P2:33.3%

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Table 2: Taxonomic diversity of species present in Extremadura but not found in the study area. Distr. = distribution, EurSib = Eurosiberian, Med = Mediterranean, End = Iberian endemic.

Order Family Species Distr.

Blanidae Blanus mariae Albert & Fernández, 2009 End

Anguidae Anguis fragilis Linnaeus, 1758 EurSib

Gekkonidae Hemidactylus turcicus (Linnaeus, 1758) Med

SQUAMATA Iberolacerta cyreni (Müller & Hellmich, 1937) End

Iberolacerta martinezricai (Arribas, 1996) End Lacertidae Podarcis guadarramae (Boscá, 1916) End Podarcis carbonelli Pérez‐Mellado, 1981 End

Colubridae Coronella austriaca Laurenti, 1768 EurSib

iterranean, with a single taxon exhibiting Chalcides bedriagai (Boscá, 1880), and Ch. a wider chorology: Emys orbicularis striatus (Cuvier, 1829); and Lamprophi‐ (Linnaeus, 1758), with a Palearctic distri‐ idae, with one representative, Malpolon bution. Families Colubridae and Viperi‐ monspessulanus (Hermann, 1804), all of dae clearly illustrate this affinity of rep‐ which have 100% representation in Mon‐ tiles in Monfragüe for thermal environ‐ fragüe. ments, since only species with a Mediter‐ With just five species, the endemic rep‐ ranean distribution in these families are tile fauna of Monfragüe is below levels of present in Monfragüe: Hemorrhois hip- endemism in Extremadura and the ibero‐ pocrepis (Linnaeus, 1758), Zamenis scalaris balearic fauna: 25% endemisms in Mon‐ (Schinz, 1822), Coronella girondica fragüe compared to 35.7% in Extremadu‐ (Daudin, 1803), Macroprotodon brevis ra, and 33.33% in Iberia and the Balearic (GÜnther, 1862), Natrix maura (Linnaeus, Islands. This is probably a result of the 1758), and Natrix astreptophora (LÓpez‐ absence of genus Iberolacerta and the low Seoane, 1884) among Colubridae, and representation of genus Podarcis in Mon‐ Vipera latastei Boscá, 1878 in Viperidae. fragüe, since both genera include a high Families most represented in Monfra‐ number of endemic taxa. güe are typically Mediterranean and with Taxonomically complex species in the few elements in the ibero‐balearic context. study area include: B. cinereus, P. virescens This is the case of Emydidae, with a single and Psammodromus occidentalis Fitze, representative, E. orbicularis; Geoemydidae, González‐Jimena, San‐José, San Mauro with one representative, Mauremys lepro- Zardoya, 2012. The presence of B. cinere- sa (Schweigger, 1812); Phyllodactylidae, us in Monfragüe has been confirmed with with one representative, Tarentola mauri- mtDNA sequences in this study. Regard‐ tanica; Scincidae, with two representatives, ing Podarcis, Geniez et al. (2014) cite a

88 REPTILES OF MONFRAGÜE AND MOLECULAR ANALYSIS OF BLANUS subadult individual of P. virescens in ar‐ tics of the single locality for L. schreiberi royo de La Vid in Torrejón el Rubio, with‐ in Monfragüe were described in Fernán‐ in the ZPP of the National Park. Finally, dez‐Ortín Toboso‐Borrella, 2014. We Fitze et al. (2012) record P. occidentalis visited the area often, with new observa‐ in Aldea del Obispo, just a few km from tions in dates: 25/06/14, 16/04/17, 23/06/17 the southern limit of the ZPP. Species of and 15/07/17, all in the same site. these three genera (Blanus, Podarcis and The new information presented in this Psammodromus) generally have parapatric study can inform conservation actions for distributions, and therefore, all records in reptiles in Monfragüe, especially for spe‐ the study area could in principle be as‐ cies presenting relict populations in the signed to the referred taxa. However, area. Future studies should include sys‐ some published records of B. mariae and tematic surveys in order to detect the po‐ P. guadarramae are close to Monfragüe, so it’s tential presence of additional species, and not unlikely that these species will be re‐ set the basis for a long‐term monitoring of ported within the limits of the National some of the best examples of reptile com‐ Park in future studies. munities in Mediterranean areas. According to the maps presented in Acknowledgement the Supplementary Material, the most widespread species in the study area are We thank X. Santos and an anony‐ Timon lepidus (Daudin, 1802), recorded in mous reviewer for constructive com‐ 24 of the 25 10x10 UTM grids in Monfra‐ ments. José María Benítez, Ángel Bláz‐ güe, M. leprosa, P. algirus (Linnaeus, 1758), quez, Óscar Díaz, Adrián Fernández, Inés and N. maura (23 grids each). On the con‐ García, José María Jiménez and Francisco trary, the species with a more restricted Pérez provided data or helped in field distribution in Monfragüe are L. surveys. Special thanks to Pedro Holgado schreiberi, found in a single 1x1 km UTM for sharing some records of Vipera lat- grid, E. orbicularis, and Acanthodactylus astei. Isabel Rey and Beatriz Álvarez of the erythrurus (Schinz, 1833), in four 10x10 Tissue and DNA collection at Museo km UTM grids each. Nacional de Ciencias Naturales facilitated Of all reptile species in Monfragüe, L. access to samples and molecular analyses. schreiberi is the most threatened (Fernández‐ Finally, we thank managers of Monfragüe Ortín Toboso‐Borrella, 2014). In Ibe‐ National Park and park rangers the facili‐ ria, Marco Pollo (1993) associate the ties provided to conduct field work. presence of the species with the Eurosibe‐ References rian and Mediterranean biogeographic Albert, E., Zardoya, R. García‐París, M. regions, with 95,6% of the species records (2007). Phylogeographical and speciation in the Supra‐ and Oromediterranean bio‐ patterns in subterranean worm of climatic regions. The Mesomediterranean the genus Blanus (: Blani‐ region, characteristic in most of Monfra‐ dae). Molecular Ecology 16(7): 1519‐1531. güe, only accounts for 4.42% of the spe‐ Albert, E. Fernández, A. (2009). Evidence of cies records. Environmental characteris‐ cryptic speciation in a fossorial reptile:

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