# Demographic structure and genetic diversity of Mauremys leprosa in its northern range reveal new populations and a mixed origin

Carmen Palacios, Cristina Urrutia, Nikolai Knapp, Marc Franch Quintana, Albert Bertolero, Gael Simon, Louis Du Preez, Olivier Verneau

<figure><img src="image_1.png" /></figure>

► To cite this version:

Carmen Palacios, Cristina Urrutia, Nikolai Knapp, Marc Franch Quintana, Albert Bertolero, et al. Demographic structure and genetic diversity of Mauremys leprosa in its northern range reveal new populations and a mixed origin. Salamandra, 2015, 51 (3), p. 221-230. [https://doi.org/10.3939/doi.1221019](https://doi.org/10.3939/doi.1221019)

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# Demographic structure and genetic diversity of *Mauremys leprosa* in its northern range reveal new populations and a mixed origin

CARMEN PALACIOS<fnref n="1" />,<fnref n="2" />,⁶, CRISTINA URRUTIA<fnref n="1" />,<fnref n="2" />, NIKOLAI KNAPP<fnref n="1" />,<fnref n="2" />, MARC FRANCH QUINTANA<fnref n="3" />, ALBERT BERTOLERO<fnref n="4" />, GAEL SIMON<fnref n="1" />,<fnref n="2" />, LOUIS DU PREEZ<fnref n="5" /> & OLIVIER VERNEAU<fnref n="1" />,<fnref n="2" />,<fnref n="5" />

Corresponding author: CARMEN PALACIOS, e-mail: carmen.palacios@univ-perp.fr

Manuscript received: 20 February 2014

Accepted: 2 February 2015 by PHILIPP WAGNER

**Abstract.** Freshwater turtle species are still poorly understood, and many species are in decline due to unsustainable trade as well as human alteration and degradation of freshwater ecosystems. *Mauremys leprosa* is a freshwater chelonian endemic to the Mediterranean Basin. Whereas the fossil record demonstrates that this species used to be distributed to well beyond the Spanish border in France, it is today restricted to the border region with Spain, at the Baillaury River in the Pyrenees, with some isolated observations from slightly farther into France. The species consequently holds an “Endangered” status according to the French IUCN Red list. Here we report for the first time the presence and demographic structure in its northern range and demonstrate that its distribution expands beyond the Pyrenees Mountains, throughout French Catalonia. Sequence analyses of the mitochondrial DNA (mtDNA) cytochrome b (cyt b) gene from 216 specimens mainly from France and Spanish Catalonia resulted in a patchwork pattern of haplotypes that supports a mixed origin of the species in France. We encountered two extreme haplotypes, with specimens with the endemic Spanish Catalanian haplotype A18 belonging to *M. leprosa leprosa* and others being clearly referable to *M. leprosa saharica* (cyt b haplotypes from clade B) that is otherwise typical from below the Atlas Mountain Range in Morocco. Short- and long-term directions for research as well as conservation management actions are suggested for this insufficiently studied species.

Key words. *Mauremys leprosa*, freshwater turtles, mtDNA cytochrome b, demography, species conservation, turtle trade.

## Introduction

Freshwater turtle species are still poorly understood and in decline due to human alteration and degradation of freshwater ecosystems (MOLL & MOLL 2004). Since the 1980s, freshwater chelonians have suffered from “the terrible turtle trade” (see, e.g., VAN DIJK et al. 2000), i.e., they are poached and sold as pets and sometimes released into the environment when they are no longer wanted, with the potential of becoming a threat to native species (CADI & JOLY 2004, POLO-CAVIA 2008, 2009, 2010a, 2010b, 2012). The Mediterranean pond turtle, *Mauremys leprosa* (SCHWEIGGER, 1812), is a freshwater species that mainly inhabits streams and ponds with riparian vegetation (DA SILVA 2002). The species is present in large parts of North Africa and on the Iberian Peninsula (IVERSON 1992, SEGURADO et al. 2005) and to a minimal extent in France. It is

classified as “Vulnerable” in both the European Red List of Reptiles (COX & TEMPLE 2009) and Spanish Red List (DA SILVA 2002) with its decline being due to the loss of suitable freshwater ecosystems and deteriorating water quality in the Mediterranean. Until recently, *M. leprosa* lacked a conservation status in France, but it is now listed as “Endangered” (UICN FRANCE & MNHN 2008) due to the fragmentation and scarcity of its populations (criterion B1a) and its continuous decline in its range as a result of habitat degradation and poor demographical structure (criterion B1b (i, ii, iii, iv, v)). Based on the fossil record, *M. leprosa* is believed to have originated in North Africa at least during the Pliocene (more than 2.5 Mya) and subsequently dispersed to the Iberian Peninsula in the early Pleistocene or late Pliocene (DE LAPPARENT DE BROIN 2001). Its genetic diversity has confirmed this demographic expansion (FRITZ et al. 2006). During the course of the species' expan-

<footnote n="1">¹) Univ. Perpignan Via Domitia, Centre de Formation et de Recherche sur les Environnements Méditèrranéens (CEFREM), UMR 5110, F-66860, Perpignan, France</footnote>

<footnote n="2">²) CNRS, Centre de Formation et de Recherche sur les Environnements Méditerranéens (CEFREM), UMR 5110, F-66860, Perpignan, France</footnote>

<footnote n="3">³) Univ. Barcelona, Dept. Biol. Animal, Av. Diagonal 645; 08028 Barcelona, Spain</footnote>

<footnote n="4">⁴) Ecosistemes Aquàtics IRTA, Ctra. Poble Nou km 5.5; 435840 Sant Carles de la Ràpita, Spain</footnote>

<footnote n="5">⁵) Unit for Environmental Sciences and Management, North-West University, Private Bag X6001, Potchefstroom, 2520, South Africa</footnote>
sion, only the Atlas Mountains in Morocco may have posed a barrier to its dispersion, impeding genetic exchange, as is shown by its genetic differentiation on both sides of the Atlas (FRITZ et al. 2005, 2006). In France, the oldest fossil remains date back to the Holocene, about 4,000 years ago (CHEYLAN 1982). Remains of the species dating back to the 1st century BC, the 2nd century AD (CHEYLAN & POITEVIN 2003) and the 11th century (MAUFRAS & MERCIER 2002) have also been found. These archaeological sites are all located in the Languedoc-Roussillon region, well beyond the Pyrénées-Orientales (PO) province to the northeast of the Pyrenees (Figs 1a and 1b). Given these data and the currently supposed scarcity of the species in France, CHEYLAN & VACHER (2010) considered *M. leprosa* the most endangered reptile in France.

Several *M. leprosa* subspecies have been proposed (SCHLEICH 1996, DOUCOTTERD & BOUR 2002) based on morphological characteristics, but according to genetic differences of the mitochondrial DNA cytochrome b gene (cyt b), only two subspecies are currently recognised (Fig. 2): *Mauremys leprosa leprosa* (SCHWEIGGER, 1812) (cyt b haploclade A) on the Iberian Peninsula and from north of Atlas Mountains in Morocco, and *M. l. sa-

harica* SCHLEICH, 1996 (haploclade B) in Algeria, Tunisia and mainly from south of the Atlas Mountains in Morocco (FRITZ et al. 2006, FRITZ & HAVAS 2007). The northern-most geographic limit of the species is the Baillaury River (France) in the Pyrenees Mountains on the border with Spain. This population is widely recognised as indigenous to France based on records dating from the previous century until today (KNOEPFFLER 1979b, GENIEZ & CHEYLAN 2005). Some individuals have also been observed or even captured in France at several localities beyond the Pyrenees Mountains, along the Mediterranean coast (COURMONT & RODRIGUEZ 2004, GENIEZ & CHEYLAN 2005, CHEYLAN & VACHER 2010, CHEYLAN & VERNEAU 2012). Nevertheless, no studies regarding the genetics of any French specimens, and/or the demographic structure of any French populations, have been published to date. In this paper, we report on the distribution and demographic structure of the species in France, identify the cyt b haplotype of specimens from France and Spanish Catalonia and compare them to other available *M. leprosa* sequences (FRITZ et al. 2006). The aim is to gain insights into the origin and genetic diversity of *M. leprosa* populations in its northern range in order to guide future research and conservation management.

<figure><img src="image_3.png" /><figcaption>Figure 1. Geographic locations and mtDNA cyt b haplotype abundances of the different populations of *M. leprosa* in the northern parts of its range. a) General overview of sample sites; b) sample sites from this study; c + d) sample sites of the two largest French metapopulations. Haplotype legend denotes “B” for B clade haplotypes and “New” for haplotypes that this study found to be new (see Table 1 for details). Raw data including GPS coordinates and individual specimen numbers are summarised in Table S1. Pie charts are sized proportionally to the number of mtDNA cyt b haplotypes found at each site except for Algeria, in a) magnified for improved visualisation. Maps were obtained from OpenStreetMap and vegetation layers from the Corine Land Cover dataset (European Environmental Agency).</figcaption></figure>
**Materials and methods**

Study area

In France, fieldwork was conducted in wetlands beyond the Pyrenees Mountains along the Mediterranean coast (Figs 1a and 1b), where *M. leprosa* was suspected to have expanded its range from Spain (CHEYLAN & VERNEAU 2012). The Baillaury River, from where the only French population is known, is a Mediterranean river with a typically irregular course that runs along the Albères Mountains in the Eastern Pyrenees, 11 km distant from the closest known Spanish population at the Orlina River (Fig. 1b) (KNOEPFFLER 1979a). Vegetation is composed here mainly of reeds, and a wild riparian forest develops between vineyards (Fig. 1c). Specimens from a turtle farm located in Sorède (PO province) and from Algeria (Figs 1a and b, Table 1) were also sequenced for comparison.

In Spanish Catalonia (Fig. 1a), where the species is widely distributed at altitudes below 200 m (LLORENTE et al. 1995), nine populations were sampled, covering a range of habitats typically occupied by this species. The Orlina is a river where an expansive riparian forest dominates the landscape (Fig. 1b). This locality has an abundant and well-preserved population of *M. leprosa* free of reintroductions. The Caldes de Malavella population inhabits a well conserved area that is probably free of reintroductions (FRANCH QUINTANA 2003). In the Delta de Llobregat plain, Bunyola, Ca l'Arana and Paretés de Murtra are localities composed of wetlands, ponds and canals. The Fauna Recovery Centres released specimens of different origins to reinforce these populations during the 1990s (FRANCH QUINTANA et al. 2007). The Riera de Canyelles is a typical Mediterranean seasonal stream that forms bodies of water only during autumn and spring. Local farmers have recently released some specimens of *M. leprosa* from the south of Spain into the area. Reeds dominate the Sèquia Major canal locality. According to local people, *M. leprosa* was not

known in this area until recently. At the Ebro River, three localities were sampled (Fig. 1b). The population inhabiting the Flix Dam (Reserva Natural de Fauna Salvatge de Sebes) and the small ponds and canals of Ulldecona are most likely unaffected by releases. In contrast, six individuals of unknown origin were sampled for blood before they were introduced to the Illà Audí (Reserva Natural de Fauna Salvatge de les Illes de l'Ebre) (Fig. 1b).

Fieldwork and demography structure

Field captures were performed in France and Spanish Catalonia between 2006 and 2010. Trapping of turtles was performed by using baited crayfish traps that were left overnight. Captured individuals were marked by cutting notches in the peripheral scutes of the carapace (PLUMMER, 1989) and sexed according to secondary sexual characters (PÉREZ et al. 1979). Accurate locality data of individuals captured in the field were obtained using a Global Position System (GPS) navigator eTrex Vista HCx (Garmin). Fifty to 200 µl of blood were obtained by means of the occipital sinus vein puncture technique (MARTÍNEZ-SILVESTRE et al. 2002) or otherwise by the coccygeal vein puncture technique using an insulin-type syringe. Blood was stored either pure at -80°C or fixed in 95% ethanol and then stored at -18°C until it was oven-dried at 40°C for 24-36 h. Straight carapace length was measured to facilitate the classification of specimens into age classes. Specimens were considered immature if the carapace length was less than 70 mm, because sex-indicative characters could not be discriminated below this size. We compared the size distributions of two populations according to carapace length by taking juveniles, males, and females together and performing a two-sample Kolmogorov-Smirnov test with R software (R Development Core Team 2011). Chi-square (χ²) tests to compare sex ratios between populations were also performed using R software.

<figure><img src="image_4.png" /><figcaption>Figure 2. The two *M. leprosa* subspecies. Left: *M. l. saharica* from Oued Massa at Toulou (29°57′15,87″ N, 9°39′15,70″ W) (photo by ANDREJ FUNK). Right: *M. l. leprosa* from Flix Dam (Reserva Natural de Fauna Salvatge de Sebes) (photo by AB).</figcaption></figure>
<table><caption>Table 1. Abundance matrix of mtDNA cyt b haplotypes of *M. leprosa* specimens in sampled populations. Their relative frequencies are indicated for French and Spanish Catalonia. Haplotypes A26–A29, A31, and A32 are new from this study.</caption><thead><tr><th rowspan="2">Populations</th><th colspan="12">Haplotypes</th></tr><tr><th>A16</th><th>A18</th><th>A24</th><th>B5</th><th>B6</th><th>A26</th><th>A27</th><th>A28</th><th>A29</th><th>A31</th><th>A32</th><th>Total</th></tr></thead><tbody><tr><td>Turtles farm</td><td>10</td><td>2</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>14</td></tr><tr><td>Algeria</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Ceyras</td><td>0</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>St. Gely du Fesc</td><td>0</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Narbonne</td><td>0</td><td>0</td><td>1</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td colspan="13">French Catalonia</td></tr><tr><td>St Hippolyte</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Agly</td><td>5</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Canet</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Basse (Thuir)</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Tech main watercourse</td><td>13</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>19</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>2</td><td>0</td><td>11</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>13</td></tr><tr><td>Baillaury (Banyuls)</td><td>32</td><td>5</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>41</td></tr><tr><td>SUBTOTAL</td><td>56</td><td>7</td><td>17</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>84</td></tr><tr><td>Haplotype relative frequency (%)</td><td>67</td><td>8</td><td>20</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td></tr><tr><td colspan="13">Spanish Catalonia</td></tr><tr><td>Orlina</td><td>2</td><td>19</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Caldes de Malavella</td><td>9</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Parets de Murtra</td><td>14</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>14</td></tr><tr><td>Ca L'Arana</td><td>17</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>17</td></tr><tr><td>La Bunyula</td><td>11</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>0</td><td>14</td></tr><tr><td>Riera de Canyelles</td><td>3</td><td>0</td><td>1</td><td>0</td><td>0</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Sequia Major</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Ebre (Flix)</td><td>15</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>15</td></tr><tr><td>Ebre (Ille Audi)</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Ulldecona</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>SUBTOTAL</td><td>81</td><td>19</td><td>3</td><td>0</td><td>0</td><td>5</td><td>1</td><td>1</td><td>1</td><td>1</td><td>0</td><td>112</td></tr><tr><td>Haplotype relative frequency (%)</td><td>72</td><td>17</td><td>3</td><td>0</td><td>0</td><td>4</td><td>1</td><td>1</td><td>1</td><td>1</td><td>0</td><td>0</td></tr><tr><td>TOTAL</td><td>147</td><td>28</td><td>22</td><td>7</td><td>2</td><td>5</td><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td>216</td></tr></tbody></table>

## DNA extraction, PCR, and sequencing

We extracted DNA from frozen blood following the Blood & Body Fluid DNA Protocol of the E.Z.N.A.® Tissue DNA or MicroElute Kits (Omega bio-tek), which is optimised for the use of fresh or frozen blood. Ethanol-dissolved blood samples were dried using a Speed Vac® Plus lyophiliser (Savant), and blood pellets were subsequently dissolved in 400 µl of PBS Buffer and stored at 4°C for 48 hours before DNA extraction. The concentration of extracted DNA was measured with a Nanodrop 2000 spectrophotometer (Thermo Scientific).

Before PCR amplification, all DNA samples were adjusted to a concentration of 30 ng/µl. The mtDNA cyt b gene was amplified with forward mt-a-new (5'-CTCCAGCCCCATCCAACATCTCAGCATGATGAAACT-

TCG-3') (LENK & WINK 1997) and reverse H-15909 (5'-AGGGTGGAGTCTTCAGTTTTTGGTTTACAAGACCAATG-3') primers (FRITZ et al. 2005). Amplification reactions consisted of a mix with a final volume of 30 µl containing 1 µl of each DNA sample, 200 µM of each dNTP (Promega), 0.4 µM of each primer, 1.5 mM MgCl₂, 6 µl Taq-buffer, and 1 unit of GoTaq FlexiDNA Polymerase (Promega). The PCR was conducted in a Mastercycler Ependorf® with the following settings: denaturation step of 2 min 30 sec at 95°C; 35 cycles of 30 sec each at 95°C, 40 sec at 49°C, 1 min 10 sec at 72°C; and one final extension step of 6 min at 72°C. The resultant PCR products were verified in 1% agarose gel, stained with ethidium bromide. The amplified DNA was then purified with the Wizard SV Gel and PCR Clean-up System (Promega) and sent for sequencing with both PCR primers to GATC (Biotech, France).
## Sequence analysis

Reverse and forward sequences were assembled by using the Sequencher™ software (Gene Codes Corporation, Ann Arbor, Michigan, USA). Sequence chromatograms were carefully inspected and sequences were corrected manually at ambiguous sites. The edited cyt b gene sequences were aligned with MEGA version 4 (TAMURA et al. 2007) to identify variable positions of different haplotypes. New haplotypes were named following the nomenclature outlined by FRITZ et al. (2006), which includes the clade plus a correlative number. New sequence haplotypes were submitted to GenBank (accession numbers are KF791559-KF791564). Intra-population diversity indices, such as mitochondrial haplotype (H) and nucleotide ($\pi_n$) diversities and mean number of pairwise differences ($\pi$), were calculated for each population by using ARLEQUIN 3.5.1.3 (EXCOFFIER & LISCHER 2010). French and Spanish intra-population diversity indices with and without A24 and/or B haplotypes were compared using a Mann-Whitney-Wilcoxon non-parametric test in R software. Given the slow evolutionary rate of the cyt b gene in this species, a network of haplotypes is sufficient to show their divergence. Haplotypes were connected in the most parsimonious way (minimum distance between haplotypes) by using NETWORK 4.6.0.0 (www.fluxus-engineering.com) and applying the Median Joining algorithm (BANDELT et al. 1999).

## Results

Demographic structure of *M. leprosa* in its northernmost distribution range

At the Baillaury River (Fig. 1c), the species was well represented with 108 individuals captured in 2010. The population presented an equilibrated age structure when compared to the structure of the little-impacted Orlina River population from Spanish Catalonia (two-sample Kolmogorov-Smirnov test D = 0.32, p = 0.30). With respect to gender composition, both populations show similar sex ratios biased in favour of males (Baillaury: 1:1.4; Orlina: 1:1.2; $\chi^2$ = 0.06, df = 1, p = 0.81). We also found forty-eight specimens at the Tech River, where only five specimens had been captured previously (COURMONT & RODRIGUEZ 2004). This is an irregularly routed but continuously flowing Mediterranean river in the Roussillon Plain adjoined by wetlands with vegetation dominated by dense reeds as well as a well-developed riparian forest (Fig. 1d). Along the Tech River's main bed, two turtles were captured each at two different sites, i.e., Riutec and Le Boulou, four at Ni dolères, and sixteen at La Falaise (Fig. 1d). At St. Jean Pla de Corts, which is an artificial pond annexed to the river that is used for recreational activities (Fig. 1d), 29 turtles were captured. Captures from 2009 to 2010 at the Tech River main course yielded only adults, with a sex ratio of 1:1.6 in favour of males (Fig. 3) that was not significantly differ-

<figure><img src="image_5.png" /><figcaption>Figure 3. Age class structure based on carapace lengths. Left: Age class structure for the major French *M. leprosa* populations from this study: the Baillaury River in 2010, the Tech River main course in 2009–10, and the artificial pond at St Jean Pla de Corts in 2010. Right: age class structure for two Spanish populations: the Orlina River and Delta de Llobregat (from FRANCH QUINTANA et al. 2007).</figcaption></figure>
ent from the Baillaury and Orlina populations (p = 0.92 and p = 0.70, respectively). The population at St. Jean Pla de Corts included juveniles and exhibited a heterogeneous age structure (Fig. 3). However, the sex ratio in favour of males (1:1.5) was not significantly different from the two previous populations (p = 0.84 and p = 0.95). Moreover, a two-sample comparison of carapace length distributions from the Baillaury and Tech Rivers (St. Jean Plat de Corts and main course taken together) populations produced an insignificant difference, too (D = 0.42, p = 0.07). In 2008, the presence of the species was recorded in an artificial channel in a swamp area at St. Hippolyte (Fig. 1b), where ten turtles were captured. In September 2010, no specimens were found at this site. Other singular catches of the species comprise eight individuals found at the Agly River in 2010, where stream water is calm forming anastomoses, six specimens at the Basse River in Thuir in 2008 and 2010, and one individual caught in a small artificial pond in Canet, which is annexed to the Têt River (Fig. 1b).

## Genetic analysis of specimens from France and Spanish Catalonia

Among the 216 specimens that were genetically investigated (Table 1), we identified 12 different haplotypes from 933 usable nucleotide sites of the cyt b gene (Tables 1 and S1). We assigned the majority of haplotypes to clade A (M. l. leprosa, Fig. 4), which is known to occur throughout the Iberian Peninsula as well as north of the Atlas Mountain range (FRITZ et al. 2006). However, some individuals had haplotypes referable to clade B and thus to M. l. saharica (Fig. 4, Table 1), occurring south of the Atlas Mountains (FRITZ et al. 2006). We found one specimen with haplotype B6 in Algeria (Fig. 1a). In France, haplotypes referable to clade B were found mainly in locations north of French Catalonia in Ceyras, Narbonne, and St. Gély du Fesc (Table 1, Fig. 1b), except for four specimens with haplotype B5 found at a single site on the Baillaury River (Table 1, Fig. 1c). The dominant haplotype from clade A was A16, which was detected in 81 specimens from Spanish Catalonia and 57 from French Catalonia, in addition to the ten from the turtle farm (Table 1, Fig. 1a). Seven new haplotypes from this study (Table 1) differed from A16 by only one mutation each (Fig. 4). Five of them, namely A27, A28, A29, A31, and A32, were encountered only once (Tables 1 and S1). A32 was found at the turtle farm (Table 1), while the remaining ones were detected in terrapins occurring at Spanish sites (Fig. 1b), including five specimens of the new A26-haplotype found at one locality, Canyelles (Table 1). We also found specimens with haplotypes A18 and A24. Haplotype A18, which was dominant in the Spanish border population of the Orlina River (Fig. 1b), was previously thought to be endemic there (FRITZ et al. 2006). We found A18-individuals at the sites Baillaury, Agly, and St. Hippolyte as well as at the turtle farm (Fig. 1b). Haplotype A24 had so far been encountered only in one individual in northern Morocco (FRITZ et al. 2006). It differs from A16 at

<table><caption>Table 2. Diversity indices (H – haplotype diversity; π_n – nucleotide diversity; π – mean number of pairwise differences) calculated for potentially natural populations with more than 5 individuals with and without exotic B5 and potentially exotic A24 haplotypes. Average diversity values are reported for French and Spanish Catalanian populations. Probability-values of the Mann-Whitney-Wilcoxon statistical test for differences between both groups of populations are also shown.</caption><thead><tr><th rowspan="3">Populations</th><th colspan="6">Diversity indexes</th></tr><tr><th colspan="3">With exotics</th><th colspan="3">Without exotics</th></tr><tr><th>H</th><th>π<sub>n</sub><br>(×10<sup>-3</sup>)</th><th>π</th><th>H</th><th>π<sub>n</sub><br>(×10<sup>-3</sup>)</th><th>π</th></tr></thead><tbody><tr><td colspan="7"><strong>Turtles farm</strong></td></tr><tr><td>Algeria</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Ceyras</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>St. Gely du Fesc</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Narbonne</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td colspan="7"><strong>French Catalonia</strong></td></tr><tr><td>St Hippolyte</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Agly</td><td>0.33</td><td>0.4</td><td>0.33</td><td>0.33</td><td>0.4</td><td>0.33</td></tr><tr><td>Canet</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Basse (Thuir)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Tech main water-course</td><td>0.47</td><td>1.00</td><td>0.94</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>0.36</td><td>0.70</td><td>0.73</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Baillaury (Banyuls)</td><td>0.38</td><td>2.00</td><td>1.84</td><td>0.24</td><td>0.26</td><td>0.24</td></tr><tr><td><strong>Mean values</strong></td><td>0.38</td><td>1.03</td><td>0.96</td><td>0.14</td><td>0.17</td><td>0.14</td></tr><tr><td colspan="7"><strong>Spanish Catalonia</strong></td></tr><tr><td>Orlina</td><td>0.18</td><td>0.19</td><td>0.18</td><td>0.18</td><td>0.19</td><td>0.18</td></tr><tr><td>Caldes de Malavella</td><td>0.35</td><td>0.60</td><td>0.55</td><td>0.35</td><td>0.60</td><td>0.55</td></tr><tr><td>Bunyula</td><td>0.39</td><td>0.70</td><td>0.67</td><td>0.17</td><td>0.18</td><td>0.17</td></tr><tr><td>Ca L'Arana</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Parets de Murtra</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Riera de Canyelles</td><td>0.64</td><td>1.00</td><td>1.00</td><td>0.54</td><td>0.57</td><td>1.00</td></tr><tr><td>Sequia Major</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Ebre (Flix)</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td></tr><tr><td>Ebre (Ille Audi)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Ulldecona</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td><strong>Mean values</strong></td><td>0.22</td><td>0.36</td><td>0.34</td><td>0.18</td><td>0.22</td><td>0.27</td></tr><tr><td>Mann-Whitney-Wilcoxon Test (p-values)</td><td>0.29</td><td>0.10</td><td>0.11</td><td>0.92</td><td>0.92</td><td>0.92</td></tr></tbody></table>

two mutational points (Fig. 4) and, together with A25, they are thought to represent an ancient branch within clade A that never crossed the Strait of Gibraltar (see Fig. 4). We found 17 specimens of haplotype A24 on the Tech River in France (Fig. 1d), three at the Spanish sample sites of Bunyola and Canyelles, one at the turtle farm, and another one in Narbonne (Fig. 1b, Table 1). Intra-population diversity indices (H, π_n, π) for all potentially natural populations with more than five individuals are shown in Table 2. The average gene diversity (H), or the probability of finding two different haplotypes in a sample when chosen ran-
domly, and the average molecular diversity indices ($\pi_n$, $\pi$), were much higher in French than in Spanish populations except when haplotypes A24 and B5 (Table 2) or A24 or B5 (not shown) were eliminated from the analysis. The Mann-Whitney-Wilcoxon test confirmed this difference (Table 2). P-values were close to 1 when A24 and B haplotypes were eliminated from the analysis, which indicates a small difference between both groups of populations.

## Discussion

Expansion of the Mediterranean pond turtle's known northern geographic distribution

Our results comprise the first survey of *M. leprosa* in France and demonstrate that the northern distribution expands beyond the Pyrenees Mountains all along the French Catalanian coast (Fig. 1b). We demonstrate that there are two

well-established populations. Firstly, the already known population on the Baillaury River (Fig. 1c) where more than 200 individuals have been captured since 1990 (VERNEAU 2007, VERNEAU 2009, HARDY 2010, VERNEAU 2010). Secondly, a population that was discovered in the course of this study at the Tech River. The species is spatially structured as metapopulations at both locations (Figs 1c, d), i.e., groups of spatially separated populations that are potentially connected by the main course of the river (HANSKI & SIMBERLOFF 1997). Both metapopulations showed an equilibrated demographical structure (Fig. 3) and no significant differences in sex ratio when compared to the well-preserved Orlina River population (Fig. 3). The small number of captures in spite of several survey campaigns and the apparent absence of juveniles on the Tech River main course (Fig. 3), lead us to hypothesise that the St. Jean Pla de Corts subpopulation might provide recruitments to the other Tech River subpopulations (Fig. 1d). *Mauremys leprosa* was

<figure><img src="image_6.png" /><figcaption>Figure 4. Minimum Spanning Network of *M. leprosa* mtDNA cyt b haplotypes. Coloured pies represent the distribution and frequency of identified haplotypes in Spanish and French Catalonia as well as in “Other” sites from this study (see Table 1). White circles represent known haplotypes of this species not observed in this study. A10, A11, A15, A17, A19, A20, A21, A22, and A23 are all distinct haplotypes that have been grouped for convenience; they all differ from A16 by one single mutation. Dashes represent missing haplotypes, each line between dashes or pies represents one mutational step.</figcaption></figure>
also captured farther north, at Canet on the Têt River, and at Agly and the Basse Rivers, and in a small pond at St. Hippolyte, in areas with habitats similar to the previously mentioned populations, i.e., those with relatively calm waters and dense riparian vegetation (Fig. 1b). Salty water, as indicated by the many marine crabs caught in the crayfish traps, may explain the absence of captures at St. Hippolyte in 2010, as the species does not tolerate salinity above a critical value (KELLER 1997, BERTOLERO & ORO 2009, MURAN 2010), indicating that the species might have vacated the site. Captures from this study in predictable habitats call for further demographic surveys to conclude the establishment of the species in those northern wetlands. Beyond the region of French Catalonia, captures are scarce and limited to isolated localities (Fig. 1b), which is most likely the result of random releases of turtles.

## A mixed origin in the northern range

Our genetic analyses aimed to shed light on the phylo-geography of the populations of *M. leprosa* living at its northern range limits. Its presence on the Baillaury River was already known prior to the previous century's turtle trade, and we have encountered several individuals with A18 haplotypes in this area (Fig. 1c). Given that A18 was previously only found in Spanish Catalonia (FRITZ et al. 2006), these results advocate the hypothesis of its presence on the Baillaury River being due to ancient dispersal from Spain, supporting the hypothesis that the Pyrenees Mountains have not always been an insurmountable geographical barrier for the species, as appears to be the case with the Atlas Mountains in Morocco (FRITZ et al. 2005, 2006). Furthermore, we found a similar relative abundance of the A16 haplotype between French and Spanish Catalanian populations (Table 1) and a widespread presence of A18 in French Catalonia (Figs 2, 4). These results favour the nativity hypothesis of the French populations, particularly on the Baillaury, Tech, Basse, and Agly Rivers (Fig. 1). We therefore promote two exclusive explanations for the origin of the species in France: i) an ancient natural origin in France expanding from Spain, as illustrated by the fossil record (CHEYLAN 1982, CHEYLAN & POITEVIN 2003), or ii) extinction and a more recent northern expansion. Nevertheless, a mixed origin in this country cannot be ruled out due to the presence of A24 and particularly B haplotypes in the Tech and Baillaury Rivers populations, respectively. With respect to A24, and given the limited data available from Spain and Morocco, we cannot discard the possibility that this haplotype may be more widespread in Europe than suspected. However, we found a higher relative abundance of the A24 haplotype in France (20%) than in Spain (3%, Table 2), and molecular diversity indices of French populations were exacerbated compared to those of Spanish populations except when A24 and/or B haplotypes were eliminated from the analysis (Table 2). Secondly, the B haploclade denotes another subspecies, *M. l. saharica* (Fig. 2). Translocation to reinforce natural populations or

the illegal trade in *M. leprosa* specimens and their unauthorised random release, as is known to happen with other turtle species (VAN DIJK et al. 2004, MOLL & MOLL 2004, VELO-ANTÓN et al. 2011), are therefore the most plausible explanations for the presence of these haplotypes in France. Furthermore, only singular individuals with A24 or B haplotypes have been encountered at isolated northern French locations (Fig. 1b), supporting the hypothesis that *M. leprosa* is present naturally only up to St. Hippolyte in the area known as French Catalonia.

## Conservation management and research perspectives in France

This is the first comprehensive study of *M. leprosa* in France, where it is classified as 'Endangered' (UICN FRANCE & MNHN 2008). Several allochthonous mtDNA cyt b haplotypes were detected. We found several individuals with haplotype B5 indicative of *M. l. saharica* (Figs 1c, 4) concentrated in one location at the Baillaury River (Fig. 1c). A high abundance of the most likely exotic haplotype A24 was also encountered on the Tech River (Table 1, Fig. 1b). This haplotype constitutes nowadays a paradox, as it has previously been found only in one individual in northern Morocco (FRITZ et al. 2006). A more thorough study of the genetic diversity of this species, particularly in Spain and Morocco, will likely help to explain this phenomenon. Even supposedly autochthonous haplotypes, such as A16 or A18, could potentially be the result of translocation, as corroborated in this study by the presence of these haplotypes at the turtle farm, where specimens are brought to by people who kept them as pets (Table 1). To identify the exact origin of French specimens, it will be necessary to study the genetic structure of the entire distribution range of the species using appropriate genetic markers. The dynamics of the Tech River metapopulation require further research to identify migration and gene flow patterns between subpopulations and understand the higher number of A24 individuals at St. Jean Pla de Corts. Subsequent modelling of population dynamics will better guide conservation management of the species in this area (OVASKAINEN et al. 2002). In order to guide decision-making regarding whether exotic haplotypes should be eliminated, it will be necessary to perform experimental research on the local adaptation of endemic haplotypes. If hybridisation between exotic and endemic haplotypes occurs, experimental research aiming to find signs of outbreeding depression or hybrid vigour will help with decisions concerning allochthonous haplotypes. If evidence of local adaptation or absent hybrid vigour is found, it would be advisable to eliminate exotic haplotypes (EDMANDS 2007, HUFF et al. 2011). Inversely, controlled hybridisation between endemic and exotic haplotypes would be advisable if signs of inbreeding depression are observed (FRANKHAM 1995, EDMANDS 2007). Nevertheless, due to the historical value of the Bailaury River population (KNOEPFFLER 1979b), it is advisable to extirpate specimens belonging to subspecies *M. l. saha-
*rica* from this area. Last but not least, efforts should be focused on informing the public of the risks associated with poaching or releasing turtles, as they might threaten native turtles for the reasons pointed out above.

Demographic structure and genetic diversity insights suggest a more widespread distribution of *M. leprosa* in France than recently reported (CHEYLAN & VACHER 2010), but one that is much smaller than suggested by the fossil record (CHEYLAN 1982). We propose to maintain the current 'Endangered' status of *M. leprosa* in France (UICN FRANCE & MNHN 2008) until research regarding the present distribution, possible population expansion and growth, and connectivity between populations will be completed.

## Acknowledgements

We are grateful for permits to capture turtles issued by the Servei de Protecció i Gestió de la Fauna (Environmental Department of the Catalan government) to AB, by the Port Aventura and Consorci of Llobregat Delta to MF, and by the Ministère de l'Ecologie, de l'Energie, du Développement Durable et de la Mer to OV. We are indebted to the herpetology group of Universitat de Barcelona, Ms. Malirach ("Vallée des Tortues", Sorède), Centre de Reproducció de Tortugues de l'Albera (Spain), The Reserva Natural de Fauna Salvatge de Sebes (especially P.J. JIMÉNEZ, M. VIÑAS), and to N. KAID (UPVD). This work has been supported by: projects entrusted to OV and LDP (CNRS PICS N°4837 2010-2012; Protea 2011-2012 from the Ministère des Affaires Étrangères et Européennes et de l'Enseignement Supérieur et de la Recherche), to OV (CEN L-R project 2010) and CP (CNRS PEPS N° FG/cb D156 2011); invited professorial grants to LDP (UPVD-L-R 2009-2010); and a project assigned to L. COURMONT (Groupe Ornitologique du Roussillon) and OV in 2010 for sampling in the Pyrenees Mountains. We also thank T. GENDRE and collaborators (CEN L-R) and V. RIGAUD (La Tartuga) for blood samples from Hérault and Aude.

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### Supplementary material

Additional information is available in the online version of this article at http://www.salamandra-journal.com

Supplementary table S1. Details of turtle specimens captured and sampled for genetic studies.
Online Supplementary data

PALACIOS, C., C. URRUTIA, N. KNAPP, M. F. QUINTANA, A. BERTOLERO, G. SIMON, L. DU PREEZ & O. VERNEAU: Demographic structure and genetic diversity of *Mauremys leprosa* in its northern range reveal new populations and a mixed origin. – Salamandra, **51**(3): 221–230.

<table><caption>**Supplementary table S1.** Details of turtle specimens captured and sampled for genetic studies: locality, turtle ID number, GPS coordinates, blood or tissue sampled for DNA extraction, DNA number, and mtDNA cyt b haplotype are indicated for all catches. Haplotypes in blue are new from this study. Note that for Spanish catches, GPS coordinates are indicated for the entire population.</caption><thead><tr><th>Population</th><th>Turtle Number</th><th>Site</th><th>GPS North</th><th>GPS East</th><th>Blood / Tissue</th><th>DNA-number</th><th>Haplotype</th></tr></thead><tbody><tr><td>Turtle Farm (Sorède)</td><td>1</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-625</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>2</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-457</td><td>A24</td></tr><tr><td>Turtle Farm (Sorède)</td><td>3</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-626</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>6</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-627</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>7</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-628</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>9</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-629</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>10</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-630</td><td>A18</td></tr><tr><td>Turtle Farm (Sorède)</td><td>11</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>MiAB75</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>17</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>MiAB76</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>25</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>MiAB79</td><td>A18</td></tr><tr><td>Turtle Farm (Sorède)</td><td>35</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>MiAB81</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>36</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>MiAB82</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>37</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>MiAB83</td><td>A16</td></tr><tr><td>Turtle Farm (Sorède)</td><td>119</td><td>Vallée heureuse</td><td>42°30′57.26″</td><td>2°57′28.54″</td><td>Fresh Blood</td><td>Mi-581</td><td>A32</td></tr><tr><td colspan="8"><strong>Outside Catalonia</strong></td></tr><tr><td>Algeria</td><td>No number</td><td>Oued Rhiou</td><td>35°58′25.35″</td><td>0°55′14.13″</td><td>Tissue</td><td>Mi-618</td><td>B6</td></tr><tr><td>Ceyras (France)</td><td>No number</td><td>a place called Le Pigné</td><td>43°38′56.69″</td><td>3°27′36.24″</td><td>Blood in alcohol</td><td>MiAB143</td><td>B5</td></tr><tr><td>Ceyras (France)</td><td>No number</td><td>a place called Le Pigné</td><td>43°38′56.69″</td><td>3°27′36.24″</td><td>Fresh Blood</td><td>MiAB147</td><td>B5</td></tr><tr><td>St. Gely du Fesc (France)</td><td>1</td><td>Mare 1735</td><td>43°42′22.80″</td><td>3°47′44.75″</td><td>Fresh Blood</td><td>MiAB145</td><td>B5</td></tr><tr><td>Narbonne (France)</td><td>1</td><td>Carrière</td><td>43°09′57.95″</td><td>2°56′50.93″</td><td>Fresh Blood</td><td>MiAB101</td><td>A24</td></tr><tr><td>Narbonne (France)</td><td>2</td><td>town centre</td><td>43°10′58.36″</td><td>3°00′03.73″</td><td>Fresh Blood</td><td>MiAB102</td><td>B6</td></tr><tr><td colspan="8"><strong>French Catalonia</strong></td></tr><tr><td>St. Hyppolite</td><td>10</td><td>............</td><td>42°48′17.21″</td><td>2°58′12.22″</td><td>Tissue</td><td>Mi-617</td><td>A18</td></tr><tr><td>Agly</td><td>71</td><td>GPS153</td><td>42°45′11.72″</td><td>2°56′42.32″</td><td>Fresh Blood</td><td>MiAB105</td><td>A16</td></tr><tr><td>Agly</td><td>72</td><td>GPS154</td><td>42°45′08.32″</td><td>2°57′00.99″</td><td>Fresh Blood</td><td>MiAB106</td><td>A16</td></tr><tr><td>Agly</td><td>73</td><td>GPS153</td><td>42°45′11.72″</td><td>2°56′42.32″</td><td>Fresh Blood</td><td>MiAB107</td><td>A16</td></tr><tr><td>Agly</td><td>74</td><td>GPS153</td><td>42°45′11.72″</td><td>2°56′42.32″</td><td>Fresh Blood</td><td>MiAB108</td><td>A16</td></tr><tr><td>Agly</td><td>75</td><td>GPS152</td><td>42°45′15.15″</td><td>2°56′29.71″</td><td>Fresh Blood</td><td>MiAB109</td><td>A18</td></tr><tr><td>Agly</td><td>76</td><td>GPS154</td><td>42°45′08.32″</td><td>2°57′00.99″</td><td>Fresh Blood</td><td>MiAB110</td><td>A16</td></tr><tr><td>Canet</td><td>1</td><td>............</td><td>42°42′28.55″</td><td>3°01′23.51″</td><td>Fresh Blood</td><td>Mi-612</td><td>A16</td></tr><tr><td>Basse (Thuir)</td><td>3</td><td>............</td><td>42°38′02.71″</td><td>2°46′28.43″</td><td>Fresh Blood</td><td>MiAB127</td><td>A16</td></tr><tr><td>Basse (Thuir)</td><td>4</td><td>............</td><td>42°38′02.71″</td><td>2°46′28.43″</td><td>Fresh Blood</td><td>MiAB128</td><td>A16</td></tr><tr><td>Basse (Thuir)</td><td>68</td><td>............</td><td>42°38′02.71″</td><td>2°46′28.43″</td><td>Fresh Blood</td><td>MiAB129</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>17</td><td>Nidolères</td><td>42°32′25.13″</td><td>2°51′39.57″</td><td>Fresh Blood</td><td>Mi-579</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>38</td><td>Nidolères</td><td>42°32′25.13″</td><td>2°51′39.57″</td><td>Fresh Blood</td><td>MiAB132</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>20</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-620</td><td>A24</td></tr><tr><td>Tech (main course)</td><td>22</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-824</td><td>A24</td></tr><tr><td>Tech (main course)</td><td>23</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-622</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>24</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB134</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>25</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB135</td><td>A16</td></tr></tbody></table>
<table><thead><tr><th>Population</th><th>Turtle Number</th><th>Site</th><th>GPS North</th><th>GPS East</th><th>Blood / Tissue</th><th>DNA-number</th><th>Haplotype</th></tr></thead><tbody><tr><td>Tech (main course)</td><td>26</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB136</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>27</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-624</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>28</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-825</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>29</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-826</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>30</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>Mi-455</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>32</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB138</td><td>A24</td></tr><tr><td>Tech (main course)</td><td>33</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB139</td><td>A24</td></tr><tr><td>Tech (main course)</td><td>34</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB140</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>35</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB141</td><td>A16</td></tr><tr><td>Tech (main course)</td><td>36</td><td>La Falaise</td><td>42°32′01.49″</td><td>2°50′55.24″</td><td>Fresh Blood</td><td>MiAB142</td><td>A24</td></tr><tr><td>Tech (main course)</td><td>37</td><td>Le Boulou</td><td>42°31′22.43″</td><td>2°50′10.28″</td><td>Fresh Blood</td><td>MiAB131</td><td>A24</td></tr><tr><td>Tech (main course)</td><td>39</td><td>Lac Riutec</td><td>42°30′14.44″</td><td>2°46′19.58″</td><td>Fresh Blood</td><td>MiAB111</td><td>A16</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>40</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB112</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>41</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB113</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>42</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB114</td><td>A16</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>43</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB115</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>44</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB116</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>45</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB117</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>46</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB118</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>47</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB119</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>48</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB120</td><td>A16</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>49</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB121</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>50</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB122</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>51</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB123</td><td>A24</td></tr><tr><td>Tech (St. Jean Pla de Corts)</td><td>66</td><td>St Jean Pla de Corts</td><td>42°31′0.76″</td><td>2°48′46.24″</td><td>Fresh Blood</td><td>MiAB124</td><td>A24</td></tr><tr><td>Baillaury (Banyuls)</td><td>219</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB90</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>220</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB96</td><td>B5</td></tr><tr><td>Baillaury (Banyuls)</td><td>221</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB98</td><td>B5</td></tr><tr><td>Baillaury (Banyuls)</td><td>222</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB100</td><td>B5</td></tr><tr><td>Baillaury (Banyuls)</td><td>223</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB95</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>224</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB92</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>226</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB99</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>227</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB87</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>230</td><td>GPS1</td><td>42°27′59.10″</td><td>3°06′10.14″</td><td>Fresh Blood</td><td>MiAB93</td><td>B5</td></tr><tr><td>Baillaury (Banyuls)</td><td>237</td><td>GPS2</td><td>42°28′8.52″</td><td>3°05′49.56″</td><td>Fresh Blood</td><td>MiAB94</td><td>A18</td></tr><tr><td>Baillaury (Banyuls)</td><td>239</td><td>GPS3</td><td>42°28′7.72″</td><td>3°05′47.33″</td><td>Fresh Blood</td><td>MiAB84</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>335</td><td>GPS4</td><td>42°28′6.54″</td><td>3°05′45.42″</td><td>Fresh Blood</td><td>MiAB88</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>106</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-827</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>138</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-596</td><td>A18</td></tr><tr><td>Baillaury (Banyuls)</td><td>155</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-597</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>179</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-598</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>184</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-582</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>186</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-608</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>198</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-599</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>199</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-600</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>203</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-454</td><td>A18</td></tr><tr><td>Baillaury (Banyuls)</td><td>204</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-828</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>205</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-601</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>206</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-602</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>207</td><td>Bridge</td><td>42°27′45.38″</td><td>3°05′26.70″</td><td>Fresh Blood</td><td>Mi-583</td><td>A16</td></tr></tbody></table>
<table><thead><tr><th>Population</th><th>Turtle Number</th><th>Site</th><th>GPS North</th><th>GPS East</th><th>Blood / Tissue</th><th>DNA-number</th><th>Haplotype</th></tr></thead><tbody><tr><td>Baillaury (Banyuls)</td><td>202</td><td>GPS5</td><td>42°27'44.58"</td><td>3°05'25.20"</td><td>Fresh Blood</td><td>Mi-609</td><td>A18</td></tr><tr><td>Baillaury (Banyuls)</td><td>208</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-603</td><td>A18</td></tr><tr><td>Baillaury (Banyuls)</td><td>209</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-604</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>210</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-605</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>211</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-584</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>212</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-606</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>213</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-829</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>214</td><td>Mas Abeilles</td><td>42°27'45.99"</td><td>3°05'08.77"</td><td>Fresh Blood</td><td>Mi-607</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>234</td><td>GPS6</td><td>42°27'39.72"</td><td>3°05'18.78"</td><td>Fresh Blood</td><td>MiAB97</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>215a</td><td>GPS7</td><td>42°27'39.84"</td><td>3°05'18.84"</td><td>Fresh Blood</td><td>Mi-610</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>231</td><td>GPS8</td><td>42°27'35.28"</td><td>3°05'17.64"</td><td>Fresh Blood</td><td>MiAB86</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>235</td><td>GPS8</td><td>42°27'35.28"</td><td>3°05'17.64"</td><td>Fresh Blood</td><td>MiAB126</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>232</td><td>GPS9</td><td>42°27'34.62"</td><td>3°05'17.70"</td><td>Fresh Blood</td><td>MiAB89</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>233</td><td>GPS9</td><td>42°27'34.62"</td><td>3°05'17.70"</td><td>Fresh Blood</td><td>MiAB85</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>216a</td><td>GPS10</td><td>42°27'29.22"</td><td>3°05'11.70"</td><td>Fresh Blood</td><td>Mi-611</td><td>A16</td></tr><tr><td>Baillaury (Banyuls)</td><td>236</td><td>GPS11</td><td>42°27'24.90"</td><td>3°04'56.22"</td><td>Fresh Blood</td><td>MiAB91</td><td>A16</td></tr><tr><td colspan="8"><strong>Spanish Catalonia</strong></td></tr><tr><td>Orlina</td><td>2157</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB296</td><td>A18</td></tr><tr><td>Orlina</td><td>2179</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB282</td><td>A18</td></tr><tr><td>Orlina</td><td>2216</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB440</td><td>A18</td></tr><tr><td>Orlina</td><td>2264</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB278</td><td>A18</td></tr><tr><td>Orlina</td><td>2303</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB281</td><td>A18</td></tr><tr><td>Orlina</td><td>2333</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB295</td><td>A18</td></tr><tr><td>Orlina</td><td>2365</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB290</td><td>A18</td></tr><tr><td>Orlina</td><td>2366</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB286</td><td>A18</td></tr><tr><td>Orlina</td><td>2372</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB436</td><td>A18</td></tr><tr><td>Orlina</td><td>2384</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB288</td><td>A18</td></tr><tr><td>Orlina</td><td>2415</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB273</td><td>A18</td></tr><tr><td>Orlina</td><td>2431</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB441</td><td>A16</td></tr><tr><td>Orlina</td><td>2433</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB276</td><td>A16</td></tr><tr><td>Orlina</td><td>2456</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB289</td><td>A18</td></tr><tr><td>Orlina</td><td>2479</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB280</td><td>A18</td></tr><tr><td>Orlina</td><td>2494</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB284</td><td>A18</td></tr><tr><td>Orlina</td><td>2496</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB292</td><td>A18</td></tr><tr><td>Orlina</td><td>2514</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB275</td><td>A18</td></tr><tr><td>Orlina</td><td>2522</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB277</td><td>A18</td></tr><tr><td>Orlina</td><td>2640</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB439</td><td>A18</td></tr><tr><td>Orlina</td><td>2649</td><td>Rabós d'Empordà</td><td>42°22'38</td><td>03°01'50"</td><td>Dried blood</td><td>MiAB437</td><td>A18</td></tr><tr><td>Caldes de Malavella</td><td>8223</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB194</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8242</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB195</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8243</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB196</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8244</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB197</td><td>A28</td></tr><tr><td>Caldes de Malavella</td><td>8245</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB198</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8246</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB189</td><td>A29</td></tr><tr><td>Caldes de Malavella</td><td>8248</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB187</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8248</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB188</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8267</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB191</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>8268</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB192</td><td>A16</td></tr><tr><td>Caldes de Malavella</td><td>9933</td><td>Riera de Santa Maria</td><td>41°49'30"</td><td>02°46'57"</td><td>Dried blood</td><td>MiAB193</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8304</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB240</td><td>A16</td></tr></tbody></table>
<table><thead><tr><th>Population</th><th>Turtle Number</th><th>Site</th><th>GPS North</th><th>GPS East</th><th>Blood / Tissue</th><th>DNA-number</th><th>Haplotype</th></tr></thead><tbody><tr><td>Parets de Murtra</td><td>8307</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB241</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8308</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB242</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8316</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB233</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8317</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB234</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8318</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB235</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8321</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB247</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8331</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB237</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8352</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB251</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8353</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB252</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8357</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB239</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8359</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB243</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8364</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB245</td><td>A16</td></tr><tr><td>Parets de Murtra</td><td>8365</td><td>Gavà</td><td>41°17'16"</td><td>02°00'54"</td><td>Dried blood</td><td>MiAB244</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>8113</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB213</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>8132</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB214</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4710</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB215</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>7123</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB216</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4719</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB217</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4733</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB218</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4740</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB219</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4811</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB220</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>8116</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB221</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>8112</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB222</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>8118</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB223</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4552</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB224</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4901</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB225</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4716</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB227</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4829</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB228</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4720</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB229</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4552</td><td>CA L'Arana</td><td>41°18'10"</td><td>02°07'44"</td><td>Dried blood</td><td>MiAB230</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>411</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB267</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4000</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB262</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4449</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB254</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4453</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB266</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4812</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB258</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4816</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB255</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4817</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB264</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4818</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB256</td><td>A24</td></tr><tr><td>El Prat de Llobregat</td><td>4819</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB261</td><td>A31</td></tr><tr><td>El Prat de Llobregat</td><td>4824</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB263</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4826</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB257</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>4828</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB260</td><td>A24</td></tr><tr><td>El Prat de Llobregat</td><td>4830</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB259</td><td>A16</td></tr><tr><td>El Prat de Llobregat</td><td>6117</td><td>Bunyula</td><td>41°18'31"</td><td>02°06'42"</td><td>Dried blood</td><td>MiAB253</td><td>A16</td></tr><tr><td>Riera de Canyelles</td><td>8134</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB205</td><td>A26</td></tr><tr><td>Riera de Canyelles</td><td>8137</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB206</td><td>A26</td></tr><tr><td>Riera de Canyelles</td><td>8146</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB202</td><td>A26</td></tr><tr><td>Riera de Canyelles</td><td>8147</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB203</td><td>A26</td></tr><tr><td>Riera de Canyelles</td><td>8149</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB210</td><td>A26</td></tr><tr><td>Riera de Canyelles</td><td>8155</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB212</td><td>A16</td></tr></tbody></table>
<table><thead><tr><th>Population</th><th>Turtle Number</th><th>Site</th><th>GPS North</th><th>GPS East</th><th>Blood / Tissue</th><th>DNA-number</th><th>Haplotype</th></tr></thead><tbody><tr><td>Riera de Canyelles</td><td>8163</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB201</td><td>A16</td></tr><tr><td>Riera de Canyelles</td><td>9133</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB200</td><td>A24</td></tr><tr><td>Riera de Canyelles</td><td>xx4x</td><td>Riera de Canyelles</td><td>41°17'19"</td><td>01°43'28"</td><td>Dried blood</td><td>MiAB208</td><td>A16</td></tr><tr><td>Sèquia Major</td><td>1001</td><td>La Pineda - Salou</td><td>41°04'37"</td><td>01°10'34"</td><td>Dried blood</td><td>MiAB183</td><td>A16</td></tr><tr><td>Sèquia Major</td><td>1002</td><td>La Pineda - Salou</td><td>41°04'37"</td><td>01°10'34"</td><td>Dried blood</td><td>MiAB184</td><td>A27</td></tr><tr><td>Sèquia Major</td><td>1003</td><td>La Pineda - Salou</td><td>41°04'37"</td><td>01°10'34"</td><td>Dried blood</td><td>MiAB185</td><td>A16</td></tr><tr><td>Sèquia Major</td><td>1007</td><td>La Pineda - Salou</td><td>41°04'37"</td><td>01°10'34"</td><td>Dried blood</td><td>MiAB186</td><td>A16</td></tr><tr><td>Ebre</td><td>1358</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB452</td><td>A16</td></tr><tr><td>Ebre</td><td>1360</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB464</td><td>A16</td></tr><tr><td>Ebre</td><td>1363</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB457</td><td>A16</td></tr><tr><td>Ebre</td><td>1364</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB450</td><td>A16</td></tr><tr><td>Ebre</td><td>1368</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB446</td><td>A16</td></tr><tr><td>Ebre</td><td>1369</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB455</td><td>A16</td></tr><tr><td>Ebre</td><td>1374</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB451</td><td>A16</td></tr><tr><td>Ebre</td><td>1376</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB461</td><td>A16</td></tr><tr><td>Ebre</td><td>1377</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB456</td><td>A16</td></tr><tr><td>Ebre</td><td>1379</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB466</td><td>A16</td></tr><tr><td>Ebre</td><td>1380</td><td>Flix</td><td>41°14'20"</td><td>0°31'28"</td><td>Dried blood</td><td>MiAB462</td><td>A16</td></tr><tr><td>Ebre</td><td>1381</td><td>Flix</td><td>41°14'10"</td><td>0°31'26"</td><td>Dried blood</td><td>MiAB465</td><td>A16</td></tr><tr><td>Ebre</td><td>1382</td><td>Flix</td><td>41°14'10"</td><td>0°31'26"</td><td>Dried blood</td><td>MiAB463</td><td>A16</td></tr><tr><td>Ebre</td><td>1391</td><td>Flix</td><td>41°14'10"</td><td>0°31'26"</td><td>Dried blood</td><td>MiAB454</td><td>A16</td></tr><tr><td>Ebre</td><td>1392</td><td>Flix</td><td>41°14'10"</td><td>0°31'26"</td><td>Dried blood</td><td>MiAB458</td><td>A16</td></tr><tr><td>Ebre</td><td>1213</td><td>Illa Audí</td><td>40°51'11"</td><td>0°31'21"</td><td>Dried blood</td><td>MiAB442</td><td>A16</td></tr><tr><td>Ebre</td><td>1306</td><td>Illa Audí</td><td>40°51'11"</td><td>0°31'21"</td><td>Dried blood</td><td>MiAB444</td><td>A16</td></tr><tr><td>Ebre</td><td>1396</td><td>Illa Audí</td><td>40°51'11"</td><td>0°31'21"</td><td>Dried blood</td><td>MiAB443</td><td>A16</td></tr><tr><td>Ebre</td><td>1397</td><td>Illa Audí</td><td>40°51'11"</td><td>0°31'21"</td><td>Dried blood</td><td>MiAB445</td><td>A16</td></tr><tr><td>Ebre</td><td>1398</td><td>Illa Audí</td><td>40°51'11"</td><td>0°31'21"</td><td>Dried blood</td><td>MiAB448</td><td>A16</td></tr><tr><td>Ebre</td><td>Unknown</td><td>Illa Audí</td><td>40°51'11"</td><td>0°31'21"</td><td>Dried blood</td><td>MiAB467</td><td>A16</td></tr><tr><td>Ulldecona</td><td>1395</td><td>............</td><td>40°38'38"</td><td>0°29'59"</td><td>Dried blood</td><td>MiAB447</td><td>A16</td></tr></tbody></table>