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Archaeological evidence for the introduction of Emys orbicularis (Testudines: Emydidae) in the Balearic Islands, Western Mediterranean

In: Amphibia-Reptilia
Authors:
Alejandro Valenzuela 1Departament de Biodiversitat i Conservació, Institut Mediterrani d’Estudis Avançats (IMEDEA, CSIC-UIB), c/Miquel Marqués 21, 07190 Esporles, Mallorca, Spain
2Equip de Recerca Arqueològica i Arqueomètrica, Universitat de Barcelona (ERAAUB), Departament de Prehistòria, Història Antiga i Arqueologia, Facultat de Geografia i Història, c/Montalegre 6-8, 08001 Barcelona, Spain

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Miguel Ángel Cau 2Equip de Recerca Arqueològica i Arqueomètrica, Universitat de Barcelona (ERAAUB), Departament de Prehistòria, Història Antiga i Arqueologia, Facultat de Geografia i Història, c/Montalegre 6-8, 08001 Barcelona, Spain
3Institució Catalana de Recerca i Estudis Avançats (ICREA), Barcelona, Spain

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Josep Antoni Alcover 1Departament de Biodiversitat i Conservació, Institut Mediterrani d’Estudis Avançats (IMEDEA, CSIC-UIB), c/Miquel Marqués 21, 07190 Esporles, Mallorca, Spain

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Among the different anthropogenic processes that affected the current distribution of the European pond turtle (Emys orbicularis), the timing of the human-mediated translocations is rarely known. Here we present data derived from an archaeological specimen of E. orbicularis obtained at the Roman site of Pollentia (Mallorca, Balearic Islands) using direct radiocarbon dating. These remains correspond to the early Roman period and represent the first reliable evidence for the ancient introduction of this turtle species in a Western Mediterranean Island.

Introduction

After the last glacial, the European pond turtle (Emys orbicularis) naturally recolonized the northerly parts of the Western Palaearctic from a few southern refugia (Sommer et al., 2007, 2009). The Holocene records attest a rapid post-glacial range expansion that was favored by significantly increased mean temperatures (e.g., Sommer et al., 2011). However, the natural Holocene range has been significantly altered by humans from prehistoric times onwards (e.g., Fritz, 1996; Sommer et al., 2007). Holocene climate deterioration, human-induced habitat loss and overharvesting have been the predominant threats leading to range losses of this reptile (Fritz, 1996; Lever, 2003; Kraus, 2008; Sommer et al., 2007, 2009). In wide areas of western, central and northern Europe, native populations of E. orbicularis are now extinct (e.g., Stuart, 1979; Fritz, 1996; Cheylan, 1998; Sommer et al., 2007, 2009). Alongside this, other anthropogenic processes have been operating, such as human-mediated translocation, which has led to the introduction of this species into new areas (e.g. Fritz et al., 2004). However, the chronology and extent of these events are still insufficiently known.

Currently, E. orbicularis is widely distributed in the Mediterranean region, occurring there throughout the European coasts, in the western part of the basin also including the Maghreb, and in the east also along the coasts of Anatolia (e.g., Schleich, Kastle and Kabisch, 1996; Lenk et al., 1999; Lever, 2003; Fritz et al., 2009). Among the main islands, it inhabits Corsica, Sardinia, Sicily, Mallorca and Menorca. On Corsica and Sardinia, the fossil record argues for its continuous presence since the Late-Middle Pleistocene (Caloi et al., 1988; Vigne, Bailon and Cuisin, 1997; Hervet and Salotti, 2000). However, population genetic investigations show that the extant populations on Corsica and Sardinia are introduced, suggesting the extinction and later replacement of the original populations (Pedall et al., 2011; Vamberger et al., 2015). Pond turtles inhabiting Sicily represent another species, E. trinacris. This endemic species is recorded at least from the Pleistocene onwards (Bonfiglio et al., 2002). It has recently been described on the basis of its genetic distinctiveness (Fritz et al., 2005) and the lack of gene flow with the widespread mainland species E. orbicularis (Pedall et al., 2011; Vamberger et al., 2015).

For the Balearic Islands, no fossil records of E. orbicularis have ever been reported, despite the presence of several sites that yield rich late Pleistocene and Holocene fossil deposits that include small vertebrates as Alytes and Podarcis (Alcover, Moyà-Solà and Pons-Moyà, 1981; Mayol, 1985; Fritz et al., 1998; Bover, 2011). In return, fossil remains of Chelonia are present in early Pliocene sites of Mallorca (Bover et al., 2014), even when this paleontological record is scarce in the island. The absence of E. orbicularis in the Pleistocene-Holocene deposits, along with the present lack of genetic differences compared to mainland populations (Fritz et al., 1998; Velo-Antón, García-París and Cordero-Rivera, 2008), suggests that the Balearic pond turtles are introduced (Fritz, 2001; Lever, 2003). In fact, out of all the extant herpetofauna on the Balearic Islands, just two reptiles and one amphibian are native (Mayol, 1985; Pinya and Carretero, 2011). The vast majority of species (19 reptiles and 4 amphibians) has been introduced since the human colonization (2350-2150 cal BC; Alcover, 2008). However, the exact chronology of most of these introductions remains unresolved. Hence, these Mediterranean islands provide an excellent framework for the study of the human impact on the evolution of the vertebrate fauna (i.e., processes of extinction, translocation, dispersal disruption, etc.).

The chronology of the introduction and source region of Emys orbicularis in Mallorca and Menorca is an unresolved topic. Genetic data show that two distinct lineages are present on Mallorca (lineages II and V) and one on Menorca (lineage V; Lenk et al., 1999); both of these two lineages occur along the northwestern Mediterranean coast (II, V) and western Italy (V; Pedall et al., 2011; Vamberger et al., 2015).

There are some claims about the occurrence of E. orbicularis on Menorca in the 9th-11th centuries AD (Keller and Andreu, 2002; Mateo, 2015), but they are based on a misreading of an earlier reference (Dürigen, 1897). Ramis (1814) constitutes the earliest mention of this turtle on Menorca, while in Mallorca it was firstly cited by Barceló i Combis (1876). Although some authors suggest that the species could have been introduced in Roman or prehistoric times (e.g., Mayol, 1985; Pleguezuelos, 2002), until now archaeological evidence was lacking.

Recent research has allowed to obtain the first archaeological remains of E. orbicularis in two sites in Mallorca (Pollentia and Mestre Ramon; unpublished) and in one in Menorca (Cornia Nou; Ferrer, Plantalamor and Anglada, 2014). Prehistoric sites in the Balearic Islands frequently include highly disturbed contexts (e.g., Chapman and Grant, 1995; Morales and Rofes, 2008; Valenzuela and Alcover, 2013), making direct 14C dating the most suitable method to guarantee the establishment of chronologies of particular bones. In this study, we report the first direct radiocarbon dating of a bone of E. orbicularis from an archaeological site of the Balearic Islands. This new evidence sheds light on the time of introduction and sets a historical framework around the possible agents and on source of origin.

Figure 1.
Figure 1.

Location and context of Emys orbicularis remains from Pollentia. (1) Map showing the location of the Roman city of Pollentia on the island of Mallorca. (2) Location of the cesspit E-107 in the forum of Pollentia. (3) Section of the cesspit E-107 and stratigraphical context of the dated E. orbicularis specimen.

Citation: Amphibia-Reptilia 37, 2 (2016) ; 10.1163/15685381-00003049

Figure 2.
Figure 2.

Archaeological remains of Emys orbicularis recovered from Pollentia and its anatomical position. (A) Dorsal view of the left pleural plate (IMEDEA/P-2325). (B) Ventral view of the left fifth peripheral plate (IMEDEA/P-907). This figure is published in colour in the online version.

Citation: Amphibia-Reptilia 37, 2 (2016) ; 10.1163/15685381-00003049

Material and methods

Two bones of European pond turtle were retrieved from the archaeological layers of the Roman city of Pollentia (Alcúdia, Mallorca; see fig. 1). They correspond to fragments of disjointed plates from the carapace and have been identified at the Institut Mediterrani d’Estudis Avançats (IMEDEA).

Table 1.

Radiocarbon measurement results of the dated Emys orbicularis specimen (IMEDEA/P-2325) from Pollentia.

Table 1.

Both elements could be attributed to a single turtle (fig. 2), but the stratigraphical context suggests two different individuals that could be separated by at least two hundred years. IMEDEA/P-2325 is a left pleural plate of the carapace and was retrieved from a refill layer (SU-5538) of a cesspit (E-107). This structure is located in front of a shop (taberna) in the forum of the ancient city and has been dated around the first century BC (Rivas, 2004; Doenges, 2005). IMEDEA/P-907 is a left fifth peripheral plate of the carapace that retains the bridge suture. The bone was retrieved from a pavement layer (SU-5387) dated by ceramic association to the 3rd century AD (Orfila, 2000; Orfila, Cau and Chávez, 2005). This stratigraphical unit was located in the same sector of the cesspit E-107, but c. 2 m above its entrance (Rivas, 2004; Doenges, 2005). Evidence of anthropogenic alteration has not been documented for the specimens. A taphonomical alteration has only been documented for IMEDEA/P-907 presenting a greenish staining, probably due to contact with bronze oxide.

Radiocarbon dating

The sample from the cesspit E-107 (IMEDEA/P-2325) was selected for radiocarbon dating. It was measured at the AMS 14C facility of the Royal Institute for Cultural Heritage (RICH) in Brussels, Belgium (Boudin et al., 2015). The first step in the pretreatment of the sample was extracting collagen following Longin (1971), but adapted for accelerator mass spectrometry (AMS) samples. A supplementary NaOH step was added. Graphite was prepared using a Fe/H2 reaction and measured at Kiel (Nadeau et al., 1998). The dating results are reported as conventional 14C dates in 14C yr BP (Stuvier and Polach, 1977). Calibrated dates have been obtained using OxCal version 4.2 (Bronk Ramsey, 2009) with IntCal13 (Reimer et al., 2013) and are quoted as cal BC/AD.

Results and discussion

The European pond turtle remains from Pollentia are one of the scarce evidences for an introduced chelonian in the archaeological record of the Balearic Islands. Previous records come from the archaeological sites of S’Illot (Sant Llorenç, Mallorca), where a possible Testudo was retrieved, without identification of its chronological context (Uerpmann, 1971) and Cornia Nou (Menorca), where E. orbicularis has been mentioned (Ferrer, Plantalamor and Anglada, 2014).

The obtained 14C date of IMEDEA/P-2325 (RICH-22210) is presented in table 1. The C:N ratio of the sample (3.3) was in the accepted range of well-preserved bones (DeNiro, 1985), and therefore can be considered as reliable to determine when this species was introduced. The 2σ-calibrated 14C age of the bone, 342-328 cal BC (2.0%) and 205-47 cal BC (93.4%), attests the occurrence of E. orbicularis in Mallorca between the 4th-1st century BC, most probably between 205 and 47 cal BC. This radiocarbon dating agrees with the 1st century BC chronology of the archaeological context established by the pottery study.

Two different scenarios could explain the occurrence. Firstly, this species could have arrived by natural oversea dispersal. This dispersal mode has been directly observed for islands close to the mainland (Jelić, Budinski and Lauš, 2012) and inferred by genetic means across the Adriatic Sea (Vamberger et al., 2015) and narrow sea straits (Stuckas et al., 2014; Vamberger et al., 2015). However, the Gymnesic Islands (i.e., Mallorca and Menorca) are the most isolated islands of the Mediterranean (e.g., Alcover, 2008; Vogiatzakis, Pungetti and Mannion, 2008). In fact, since the end of the Messinian salinity crisis (c. 5.33 Myr ago), no terrestrial or limnic animal species is known to have arrived across the sea on these islands. Therefore, this possibility seems highly unlikely, and the other scenario, human-mediated translocation, has to be favored to explain the present occurrence of E. orbicularis.

Taking the archaeological context and dating results into account, the species has most probably been introduced by the Romans. A previous introduction would theoretically been possible by the original inhabitants of the Gymnesic islands (i.e., the Talaiotic culture) or their Carthaginian neighbors from Eivissa (Pytiusic Islands). However, the Talaiotics were mainly inland people (Hernández-Gasch et al., 2002; Rihuete, 2003; Van Strydonck, Boudin and Ervynck, 2002; Van Strydonck et al., 2005) and had no seafaring technology. Moreover, the absence of E. orbicularis on the nearby island Eivissa, with a Carthaginian colony since 654 BC, does not support that Carthaginians introduced the Gymnesic turtle populations. This is also supported by the fact that in the North African core region of the Carthaginians another genetic lineage occurs (Stuckas et al., 2014) than on Mallorca and Menorca.

This suggests that the Romans have introduced E. orbicularis. The Romans conquered the Balearic Islands in 123 BC, having a significant social and natural impact. The introduction of several alien Balearic species by the Romans has already been recognized (e.g., Alcover, 1979, 2010; Reumer and Sanders, 1984; Sanders and Reumer, 1984; Mayol, 1985; Valenzuela and Alcover, 2015), and we have now the first direct evidence for an introduced reptile species. The Roman Empire comprised along the northwestern Mediterranean coasts and in western Italy many potential source regions where pond turtles occur with the same genetic lineages as on the Balearic Islands (Lenk et al., 1999; Pedall et al., 2011; Vamberger et al., 2015).

In ancient times, chelonian shells were appreciated and used to make objects such as musical instruments (Lawergren, 1998). The meat of turtles and tortoises was also consumed, and chelonians played in prehistoric and historic times an important role as ‘living cans’ for provisioning seafarers. This has led worldwide to many cases of human-mediated dispersal of chelonians to islands (Vamberger et al., 2011; Fritz et al., 2013). Yet, Toynbee (1973) states that in Roman times turtle meat was only consumed exceptionally, but this may be not true for lower social classes and galley slaves. From later centuries, it is known that during the time of the French Triangular Trade, slaves seemingly have been fed on ships with turtles, leading to the introduction of an African species (Pelusios castaneus) to the Lesser Antilles (Fritz et al., 2013), and perhaps a similar situation could be true for Roman times, with surplus animals being abandoned locally after the arrival of galleys at Pollentia. In any case, the ultimate reasons behind this translocation remain open.

The combination of our archaeological data with the genetic and ecological knowledge about the extant Balearic pond turtles thus provides new insights. From a biogeographical point of view, the presence of the turtle records in Pollentia is in concordance with its current distribution. This Roman city is located next to S’Albufera d’Alcúdia, the largest wetland of the Balearic Islands (Martínez and Mayol, 1995). It is the single site in Mallorca where E. orbicularis occurs today (Mayol, 1985). During the early Roman period (2nd-1st century BC), a maritime port was established there to connect the island to the Roman trade network (Zucca, 1998; Orfila, 2008). At that time, most of the commercial products came from the Italic peninsula, as documented by the imported pottery (Orfila, 2000; Rivas, 2004; Orfila, Cau and Chávez, 2008). Therefore, the harbor of Pollentia is likely to have facilitated the introduction of E. orbicularis.

Conclusions

The record and radiocarbon dating of E. orbicularis from the archaeological levels of Pollentia suggests that the Romans introduced this species to Mallorca. Our results emphasize the value of the archaeological record for non-native reptiles to unravel their introduction dates. When integrated with the molecular and ecological data, this data has the potential to provide essential insights into complex human-animal interactions.

Acknowledgements

The authors would like to thank Dr Uwe Fritz and an anonymous reviewer for their helpful and constructive comments that greatly contributed to improving the final version of the paper. This research was conducted as part of Research Project “Cambios holocénicos en la biodiversidad animal de las islas de la Macaronesia y de Baleares”/(CGL2012-8087) of the Spanish Ministerio de Ciencia e Innovación. This is also part of the research activities of the Equip de Recerca Arqueològica i Arqueomètrica de la Universitat de Barcelona (ERAAUB), Consolidated Group (2014 SGR 845), thanks to the support of the Comissionat per a Universitats i Recerca del DIUE de la Generalitat de Catalunya, as well as of the project of archaeological research at the Roman city of Pollentia, funded by the Consorci de la ciutat romana de Pollèntia and the Ministerio de Educación, Cultura y Deportes.

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