C. R. Palevol 18 (2019) 317–324 Contents lists available at ScienceDirect Comptes Rendus Palevol www.sciencedirect.com General Palaeontology, Systematics, and Evolution (Vertebrate Palaeontology) First evidence of faunal succession in terrestrial vertebrates of the Plio-Pleistocene of the Balearic Islands, western Mediterranean Première évidence d’une succession faunique chez les vertébrés terrestres du Plio-Pléistocène des îles Baléares, Méditerranée occidentale Josep Quintana Cardona a,b,∗ , Jordi Agusti c a Institut Català de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain b Carrer Gustau Mas, 79-1er, 07760 Ciutadella de Menorca, Illes Balears, Spain c ICREA, Institut català de Paleoecologia Humana I Evolució Social (IPHES), Universitat Rovira I Virgili, Carrer Marcel·lí Domingo s/n (Edifici W3), Campus Sescelades, 43007 Tarragona, Spain a r t i c l e i n f o a b s t r a c t Article history: Terrestrial vertebrates from the karst filling in Cap d’Artrutx, situated in the extreme Received 19 October 2018 Southwest of Menorca (Balearic Islands, western Mediterranean) have yielded interesting Accepted after revision 14 February 2019 information on the migration and extinction dynamics in the terrestrial island vertebrate Available online 23 March 2019 faunas in Mallorca and Menorca between the late Pliocene and early Pleistocene. In this site, remains of the Menorcan giant tortoise (“Testudo” gymnesica Bate, 1914) and two mammals Handled by Lars van den Hoek Ostende (Nesiotites sp. and Hypnomys sp.) have been found. The remains of the Menorcan giant tor- toise represent the youngest record of this species and of the so-called second faunistic Keywords: group of the island. The morphology and size of Nesiotites sp. are comparable to those of Palaeokarst filling Nesiotites aff. ponsi Reumer, 1979 from the Plio-Pleistocene of Pedrera de S’Ònix (Mallorca) Second and third faunistical groups and Nesiotites meloussae Pons and Moyà, 1980 (= Nesiotites aff. ponsi sensu Pons-Monjo Minorcan giant tortoise Migration and extinction et al., 2012) from the lower Pleistocene of Binigaus (Menorca). Hypnomys sp. can belong to Climatic changes an archaic species not yet described. They represent the first evidence of the entrance of the third faunistic group, which originally evolved on the neighbouring island of Mallorca. This association is interpreted, therefore, as the first evidence of a faunal succession in the Plio-Pleistocene in the Balearic Islands. © 2019 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved. r é s u m é Mots clés : L’étude d’ossements de vertébrés terrestres du gisement d’origine karstique du cap Remplissage de palaeokarst d’Artrutx, situé dans l’extrême Sud-Ouest de Minorque (îles Baléares, en Méditerranée occi- Deuxième et troisième groupes fauniques dentale), ont fourni des données très intéressantes relatives à la dynamique de migration Tortue géante de Minorque et d’extinction des faunes de vertébrés terrestres insulaires de Majorque et Minorque pen- Migration et extinction Changements climatiques dant le Pliocène supérieur et le Pléistocène inférieur. Dans le gisement ont été découverts des restes de la tortue géante de Minorque (« T. » gymnesica) et de deux mammifères ∗ Corresponding author. Institut Català de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain. E-mail address: picoguevo@gmail.com (J.Q. Cardona). https://doi.org/10.1016/j.crpv.2019.02.001 1631-0683/© 2019 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved. 318 J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 (Nesiotites sp. et Hypnomys sp.). Dans ce contexte, les restes de tortue géante de Minorque représentent l’enregistrement le plus récent de cette espèce et du soi-disant deuxième groupe faunique de l’île, alors que les deux espèces de mammifères correspondent à la première évidence de l’arrivée d’un troisième groupe faunique originaire de Majorque, île voisine où il a évolué. Cette association est donc interprétée comme une première évidence de succession faunique dans le Plio-Pléistocène des îles Baléares. © 2019 Académie des sciences. Publié par Elsevier Masson SAS. Tous droits réservés. 1. Introduction Myotragus sp. include re-sedimented red limestones stones with remains of “Testudo” gymnesica (Quintana, 1998). Of the three islands that form the Balearic archipelago, This paper arises from the study of two known Menor- Menorca is the second largest in size (702 km2 ) and the can sites (Cala en Bastó and Bol de Ses Sírvies; Quintana, island which is located furthest from the European and 1995, 1998) and an unpublished one (Cap d’Artrutx). At African continents, as it is situated in the centre of the west- first, the main purpose of this study was to find deposits ern Mediterranean. Its reduced size and isolation have had with remains of the ancestor of the Menorcan giant rab- a notable influence during the periods of colonization of bit. In Cala en Bastó and Bol de Ses Sirvies, no remains of the different groups of terrestrial vertebrates between the mammals have been found. However, the study of the Cap middle Miocene and the Pleistocene (Quintana and Agustí, d’Artrutx deposit has provided very few but very interest- 2007; Quintana and Moncunill-Solé, 2014). ing remains, the study of which is presented now. In the case of the Balearic Islands, the reciprocal influ- ences among the faunas of each island is evident from, 2. Methods principally, the palaeontological record preserved in the red limestones deposited in the interior part of caves. The The remains studied were recovered by chemical influence between islands is not surprising, given their means, treating the matrix with acetic acid diluted to 10%. proximity (Mallorca, which is located in the centre of Once separated from the matrix, the remains were sub- the archipelago is separated from Eivissa and Menorca merged in water to neutralize the residual acid and calcium by 81.5 and 36.5 km, respectively). Because of this, Mal- acetate. Subsequently the fossils were dried and consol- lorca presents some common faunistic elements with idated by applicating Paraloid B-72. The acid treatment the so-called early faunas of Eivissa (Bover et al., 2014; affected, unexpectedly, the most fragile parts of one of the Quintana and Moncunill-Solé, 2014). Until now, Mallorca examples (the mandibular branch of Nesiotites sp.), in such was considered to be the origin of the Pleistocene fauna in a way that the chemical treatment was stopped, in order Menorca (Moyà-Solà and Pons-Moyà, 1980), which, seem- to avoid breakage of the mandibular body. Once consoli- ingly abruptly, appears in the fossil record of the island. dated, the mandible was separated using manual methods, In contrast to what occurred in Mallorca and Eivissa, up reducing the matrix with a rounded file of a small caliber to now, no sites have been found in Menorca preserving (1 mm in diameter). evidence of the arrival of the early faunas of the Messinian The measurements of the specimens were done based or early Pliocene, which should include, hypothetically, on photographs, using a digital camera incorporated in a remains from the ancestor of the Menorcan giant rabbit stereoscopic microscope with a micrometric eyepiece. The (Nuralagus rex, Quintana, Köhler and Moyà-Solà, 2011). measurements made on the photographs were taken with Until now, there was no “early evidence” of the first arrival a digital calibrator with a ± 0.01 mm margin of error. The in Menorca of the Pleistocene fauna from Mallorca, that is, specimens are stored in the Museu Municipal des Bastio de the concurrence, at the same site, of remains belonging to sa Font (acronym: MMBF) (Ciutadella de Menorca, Balearic the two faunistic groups. Islands, Spain). In this way, the fauna described at Binigaus (Es Migjorn Gran, Menorca) (Agustí, 1980; Moyà-Solà and Pons-Moyà, 2.1. Geographical and geological context 1980; Pons-Moyà and Moyà-Solà, 1980) cannot be consid- ered as an early evidence, given that, only remains from one faunistic group have been found. The fauna present in the The studied site is located on the eastern margin of promontory of Bajolí described by Bate (1914) (equivalent Punta des Bregarol, to the east of Cap d’Artrutx (Ciutadella to sites 15 and 16 in the Punta Esquitxador area, located in de Menorca) (Fig. 1). The red limestones with the remains the Northeast of Bajolí) (Quintana, 1998) cannot be consid- of the vertebrates are found inside a small cave (Fig. 2) very ered an early evidence site either, given that the remains of close to the sea. The red limestones cover the base of the the Menorca giant tortoise and Myotragus sp. were found cave and the west wall, where they are better exposed. in two clearly different outcrops, both from a geological On the floor, the thickness is less, as it has suffered more and temporal point of view. A confirmation of the latter wear due to erosion, as can be deduced from the irregular is that, on occasions, the clear coloured limestones with morphology in the area. J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 319 Fig. 1. Geographical situation of the Punta des Bregarol cave (Menorca, Balearic Islands). Fig. 1. Situation géographique de la grotte de Punta des Bregarol (Minorque, îles Baléares). The Miocene calcarenites in which the cave is exca- tentatively assigned the type specimens stored at the vated are interpreted as belonging to a reef formation of National History Museum (London, United Kingdom) to aff. Messinian age by Rosell et al. (1989). Moreover, Obrador Titanochelon gymnesica (Bate, 1914). The genus Cheirogaster and Pomar (2004: fig. 4.1) consider that the margin of the Bergounioux, 1935, which was used by various authors southwestern end of Menorca (where the studied site is (Jiménez Fuentes and Martín de Jesús, 1991; Luján et al., located) is included in the Reef Unit, which Pomar (2001 2014; Quintana, 1995, 1998), is only valid for one species and Pomar et al., 1996) situate in the upper Tortonian- from the Eocene in France (Pérez-García et al., 2016). lower Messinian. Taking into consideration the geological Despite their fragmentary state, the attribution of the evidence and the chronology that this provides, it is only remains to the giant turtle of Menorca is safe, because possible to say that the reddish silts and the fauna that they of their relatively large size and their characteristic bone contain are, at least, after the lithification, emersion and structure. carstification of the limestones of the Reef Unit, which is to say, after the early Messinian. However, the chronolog- Nesiotites sp. (Fig. 3) ical context of the site is established more precisely from Material. A right mandible (MMBF/CA-1), which is miss- its faunal peculiarities and the relation with the different ing the anterior end of the incisor and the condylar process. groups of terrestrial vertebrates present in the Neogene in Description. The premolars and molars show no sig- the Balearic Islands, as will be discussed in more detail in nificant morphological differences when compared with the section on biogeographical implications. Nesiotites aff. ponsi Reumer, 1979, Nesiotites meloussae Pons and Moyà, 1980 (= Nesiotites aff. ponsi sensu Pons-Monjo 2.2. Systematic Palaeontology et al., 2012) and Nesiotites hidalgo Bate, 1944. Notably, there are cusps on the oblique ridge of the m1 and the m2 “Testudo” gymnesica Bate, 1914 of Nesiotites sp., although it is true that this characteristic The giant tortoise from the Pliocene in Menorca is has also been observed in the aforementioned species undergoing taxonomic revision. Therefore, in this paper (Pons-Monjo et al., 2012). The size of the molars of Nesi- the original name is preserved. Luján et al. (2017) otites sp. coincide with the range of variation of Nesiotites 320 J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 Fig. 2. (A) Panoramic view of the coastal area in which the cave is located with the remains of vertebrates, marked with a white arrow. In the background, towards the west, the lighthouse of Cap d’Artrutx. (B) Cross section of the cave (east–west direction), excavated in the Miocene calcarenites. The grey areas inside the cave represent the red silts in which the remains of vertebrates are conserved. Fig. 2. (A) Vue panoramique de la zone côtière où est située la grotte avec les restes de vertébrés, marquée par une flèche blanche. À l’arrière-plan, vers l’ouest, le phare du cap d’Artruts. (B) Coupe est–ouest de la grotte creusée dans les calcarénites miocènes. La zone grise dans la grotte représente les silts rouges dans lesquels les restes de vertébrés sont conservés. Table 1 Dental measurements of Nesiotites sp. (MMBF/CA-1) from Cap d’Artrutx compared with Nesiotites aff. ponsi (Pedrera de s’Onix, Mallorca), Nesiotites hidalgo (Cova de Canet, Mallorca) and Nesiotites meloussae (= Nesiotites aff. ponsi sensu Pons-Monjo et al., 2012) (Barranc de Binigaus, Menorca). Data source: Pons-Monjo et al., 2012. L: length; taw: talonid width; triw: trigonid width; W: width. Tableau 1 Mesures dentaires de Nesiotites sp. (MMBF/Ca-1) du cap d’Artrutx, comparées à celles de Nesiotites aff. ponsi (Perdrera de s’Onix, Majorque), Nesiotites hidalgo (cova de Canet, Majorque) et Nesiotites meloussae (= Nesiotites aff. ponsi sensu Pons-Monjo et al., 2012) (Barranc de Binigaus, Minorque). Source des données : Pons-Monjo et al., 2012. L : longueur ; taw : largeur du talonide ; triw : largeur du trigonide ; W : largeur. Artrutx Ònix Binigaus Canet n n max–min mean SD n max-min mean SD n max–min Mean SD m1 L 1 2 25 2.20–1.85 2 0.08 25 2.19–1.93 2.04 0.07 25 2.31–2.09 2.17 0.06 taw 1 1.36 25 1.40–1.04 1.27 0.08 25 1.38–1.16 1.3 0.05 25 1.55–1.37 1.43 0.04 triw 1 1.19 25 1.29–1.02 1.15 0.08 25 1.22–1.03 1.14 0.06 25 1.35–1.13 1.23 0.05 m2 L 1 1.71 25 1.96–1.72 1.84 0.07 25 1.95–1.75 1.87 0.06 25 2.10–1.88 1.99 0.07 taw 1 1.12 25 1.18–1.03 1.12 0.04 25 1.25–1.02 1.12 0.06 25 1.26–1.09 1.21 0.04 triw 1 1.04 25 1.15–1.01 1.09 0.05 25 1.23–1.00 1.09 0.06 25 1.29–1.12 1.21 0.04 m3 L 1 1.35 25 1.45–1.28 1.36 0.06 21 1.55–1.35 1.44 0.05 25 1.73–1.38 1.5 0.078 W 1 0.87 25 0.94–0.73 0.83 0.05 21 0.90–0.72 0.81 0.05 25 0.98–0.77 0.86 0.05 aff. ponsi (Pedrera de s’Onix, Mallorca) and Nesiotites Hypnomys sp. (Fig. 4; Table 2) meloussae (Barranc de Binigaus, Menorca) In their smaller Material. An incomplete left mandible with p4-m3 size, they are clearly different from those of Nesiotites (MMBF/CA-2) and a right mandible, also incomplete, with hidalgo (Table 1). m1-m3 (MMBF/CA-3). J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 321 Fig. 3. Right mandible (MMBF/CA-1) of Nesiotites sp. in oclusal (A), lingual (B), and lateral (C) views. Scale bar: 3 mm. Fig. 3. Mandibule droite (MMBF/CA-1) de Nesiotites sp. En vues occlusale (A), linguale (B) et latérale (C). Barre d’échelle : 3 mm. Fig. 4. Left (MMBF/CA-2, A) and a right mandible (MMBF/CA-3, B) of Hypnomys sp. Scale bar: 3 mm. Fig. 4. Mandibules gauche (MMBF/CA-2, A) et droite (MMBF/CA-3, B) d’Hynomys sp. Barre d’échelle : 3 mm. 322 J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 Table 2 Dental measurements of Hypnomys sp. from Cap d’Artrutx. L: length; W: width. Tableau 2 Mesures dentaires d’Hypnomys sp. du cap d’Artrutx. L : longueur ; W : largeur. p4 m1 m2 m3 L W L W L W L W MMBF/CA-2 1.43 1.52 2.25 2.21 – 2.58 2.17 2.1 MMBF/CA-3 – – 2.37 2.1 2.36 2.33 2 1.8 Description. p4: The only available p4 is lingually dam- 3. Biogeographical implications aged. Even so, four transverse ridges can be distinguished: anterolophid, metalophid, mesolophid, and posterolophid. The finding of three species belonging to two faunal The anterolophid and metalophid are almost fused episodes in the same site (Cap d’Artrutx) establishes the and can hardly be distinguished. The mesolophid and first evidence of faunal succession in the terrestrial verte- posterolophid, which are labially connected, are well brates in the Plio-Pleistocene in Menorca. In this site, an developed. endemic vertebrate from the island can be found, belong- The m1 has six transverse ridges: anterolophid, met- ing to the so-called “second faunal episode of Menorca” alophid, centrolophid, mesolophid, posterotropid, and (the giant tortoise “Testudo” gymnesica) together with posterolophid. The anterolophid is medially interrupted. some species belonging to the “second faunal episode of The metalophid is short, its lingual end connecting the lin- Mallorca” (Nesiotites sp. and Hypnomys sp.). These two gual segment of the anterolophid. The centrolophid is long, mammals form, along with the genus Myotragus Bate, 1909 its labial end connecting with the mesolophid and forming (not present in Cap d’Artrutx), the so-called “classic” fauna a “Y”-like pattern. The long mesolophid reaches the labial of the Plio-Pleistocene in Mallorca and the Pleistocene in wall of the tooth. The isolated posterotropid is short in Menorca. The presence of Nesiotites sp. and Hypnomys sp. one case and long in the other. The hypoconid is anteriorly together with the Menorcan giant tortoise is very relevant directed, although it does not connect to the mesoconid, from a biogeographical point of view, given that it is early even at a high level of wear. evidence of this faunal succession. As in the m1, the m2 has six transverse ridges: Cap d’Artrutx is comparable, to a certain extent, with the anterolophid, metalophid, centrolophid, mesolophid, pos- Holocene site located at Pas den Revull (Barranc d’Algendar, terotropid and posterolophid. Other than in the m1, there is Ferreries, Menorca) where remains of Myotragus baleari- in the two cases a continuous anterolophid along the ante- cus Bate, 1909 occur together with Oryctolagus cuniculus rior wall of the tooth. Long metalophid, that in one case (Linnaeus, 1758), clearly of anthropogenic origin (Quintana reaches the metaconid. et al., 2003, 2016). This represent, therefore, early evidence The m3 has four transverse ridges: anterolophid, of the entrance into the island of the first domestic fau- metalophid, mesolophid and posterolophid. A short pos- nas. Radiocarbon dating indicates a difference in age of terotropid can be distinguished in one of the specimens. 1170 years in comparing two samples of Myotragus baleari- Anterolophid and metalophid are both fused labially and cus, and between 3655 and 2485 years in comparing the lingually, as it is also the case of the mesolophid and samples of Myotragus balearicus with Oryctolagus cunicu- posterolophid. In one case, the mesoconid is anteriorly lus. Thus, the presence of different species in the same site projected, not reaching the protoconid. Metaconid and does not imply, necessarily, the temporal coexistence of the entoconid are connected, forming a continuous endolophid species. Radiocarbon datings reflect, clearly, a short interval along the lingual wall of the tooth. of time during which the endemic species became extinct, Remarks. Although provisionally included in the genus supposedly, and domestic species were introduced. Hypnomys, the two mandibles from Cap d’Artrutx display In the specific case of Cap d’Artrutx, it is unknown some features which are different from those observed in whether Nesiotites sp., Hypnomys sp. and “Testudo” gym- that taxon, particularly the medially fused centrolophid nesica are contemporaries and to what extent the age and mesolophid, which is present in all of the molars differences are significant. In spite of the lack of absolute of both mandibles. This characteristic is present in the dating, several hypotheses have been presented which try lower molars of Margaritamys llulli Mein and Adrover, to explain the reasons why most of the representatives of 1982, an endemic glirid from the middle Miocene of Mal- the second faunal episode of Menorca became extinct. lorca, described from the sites of Santa Margarita (Mein In this sense, Alcover et al. (1999) considers that Myotra- and Adrover, 1982) and San Lorenzo (Adrover et al., gus and the accompanying fauna (Nesiotites and Hypnomys) 1985). However, as a difference with the species from Cap would have colonized Menorca during the first Pliocene d’Artrutx, the lower molars of Margaritamys never present glaciations, being responsible for the extinction of Nurala- a posterotropid, a characteristic which is always present in gus rex and the majority of the vertebrates from the second Hypnomys. Therefore, we assume that the specimens from faunal episode in Menorca (among which “Testudo” gym- Cap d’Artrutx can belong to an archaic, not yet described, nesica), due to a displacement of competitive nature. In species of Hypnomys. More material will be needed in order contrast to this opinion, Quintana and Moncunill-Solé to clarify the relationships of the Cap d’Artrutx species with (2014) consider that it was not a displacement of com- respect other species included in that genus. petitive nature: the extinction of “Testudo” gymnesica was J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 323 motivated by a climate change and was prior to the arrival chronological point of view, the site at Cap d’Artrutx is situ- of Myotragus, Nesiotites and Hypnomys from Mallorca. ated, therefore, among the Pliocene sites of Menorca (Punta The scarcity of remains of micromammals in Cap Nati, Binibèquer, among others), where the remains of “Tes- d’Artrutx also is not only interesting from a taphonomic tudo” gymnesica are abundant and Binigaus, where the point of view, but also because of its biogeographical impli- Menorcan giant tortoise appears to be definitively extinct. cations. The rarity of these might reflect the existence of populations with few individuals at the moment of for- Acknowledgements mation of the site or it may be due to the lack of vectors responsible for the concentration of these remains, despite The development of this paper is a part of a project the existence of high densities of individuals (e.g., absence, titled “Estudi de les faunes vertebrades de tres jaciments in the vicinity of the site of nocturnal birds of prey). càrstics del Plio-Plistocè de Menorca” financed by research Notably, the site considered to have the oldest represen- assistance provided by the “Institut Menorquí d’Estudis” tatives of the genus Nesiotites, Hypnomys and Myotragus in (IME, Maó, Menorca) in the year 2014 and CGL2016-80000- Menorca up to now, Barranc de Binigaus (Es Migjorn Gran; P (Ministerio de Economía y Competitividad, España) 2017 Agustí, 1980; Moyà-Solà and Pons-Moyà, 1980; Pons-Moyà SGR 859 (AGAUR, Generalitat de Catalunya). In the same and Moyà-Solà, 1980) is characterized for being extraor- way, the authors are especially grateful to Lars van den dinarily rich in remains of small vertebrates and by the Hoek Ostende and an anonymous reviewer, whose com- absence of “Testudo” gymnesica. Moyà-Solà and Pons-Moyà ments and suggestions have greatly improved the original (1980) place it, tentatively, in the Early Pleistocene. The manuscript. characteristics of the site at Binigaus seem to indicate that, once the Menorcan giant tortoise became extinct, there was a significant increase in the density of micromammals References originating from Mallorca, whose populations were already Adrover, R., Agustí, J., Moyà, S., Pons, J., 1985. Nueva localidad de micro- established, definitively, in the early Pleistocene. mamíferos insulares del Mioceno medio en las proximidades de San Lorenzo en la isla de Mallorca. Paleontologia i Evolució 18, 121–129. 4. Conclusions Agustí, J., 1980. Hypnomys eliomyoides nov. sp. Nuevo glírido (Roden- tia, Mammalia) del Pleistoceno de Menorca (Islas Baleares). Endins 7, 49–52. The fauna from the site at Cap d’Artrutx, located in the Alcover, J.A., Seguí, B., Bover, P., 1999. Extintions and local disappearances extreme Southeast of the island of Menorca, consists of two of vertebrates in the western Mediterranean islands. In: MacPhee, mammals (Nesiotites sp. and Hypnomys sp.) and the Menor- R.D.E. (Ed.), Extinctions in near time. Plenum Press, New Cork, pp. 165–188. can giant tortoise (“Testudo” gymnesica). The Menorcan Bate, D.M.A., 1914. The Pleistocene ossiferous deposits from the Balearic giant tortoise is, along with Nuralagus rex, one of the most Islands. Geol. Mag., N.S., Dec. 6 (1), 347–354. representative taxa of the Pliocene and the so-called second Bate, D.M.A., 1944. Pleistocene shrews from the larger western Mediter- ranean islands. Ann. Mag. Nat. Hist. 11 (11), 738–769. faunal episode in Menorca, while Nesiotites sp. and Hypno- Bover, P., Rofes, J., Bailon, S., Agustí, J., Cuenca-Bescós, G., Torres, E., Alcover, mys sp. belong to the third faunal episode. This association J.A., 2014. Late Miocene/Early Pliocene vertebrate fauna from Mallorca is interpreted as the first evidence of a faunal succession (Balearic Islands, Western Mediterranean): an update. Integrative Zool. 9, 183–196. in the Plio-Pleistocene in Menorca. It establishes, there- Jiménez Fuentes, E., Martín de Jesús, S., 1991. Ejemplares-tipo de quelo- fore, early evidence of the presence of the genus Nesiotites nios fósiles españoles. Rev. Española de Paleontología 6, 98–106. and Hypnomys in Menorca and, apparently, the last record Luján, À.H., Alba, D.M., Fortuna, J., Carmona, R., Delfino, M., 2014. First craneal remains of Cheirogaster richardi (Testudines: Testudinidae) of “Testudo” gymnesica on the island. The two mammals from the Late Miocene of Ecoparc de Can Mata (Vallès-Penedès Basin, form part of the faunal “stock” which evolved in Mallorca NE Iberian Peninsula): taxonomic and phylogenetic implications. J. from the Messinian until the Holocene. Their arrival in Syst. Palaeontol. 12, 833–864. Menorca from Mallorca would have occurred in the inter- Luján, À.H., Alcover, J.A., Ivanov, M., Torres, E., Alba, D.M., 2017. Revisión taxonómica de “Testudo” gymnesica Bate, 1914 (Testudines, Testu- val of time situated between the late Pliocene and the early dinidae) a partir de la descripción del material tipo de Menorca (Islas Pleistocene. Baleares). Spanish J. Palaeontol. 32 (2), 261–278. The presence of “Testudo” gymnesica at Cap d’Artrutx Mein, P., Adrover, R., 1982. Une faunule de mammifères insulaires dans le Miocene moyen de Majorque (îles Baléares). Geobios Mem. spec. 6, indicates that this site is older than the site at Binigaus, 451–463. which was until now considered to be the site with the old- Moyà-Solà, S., Pons-Moyà, J., 1980. Una nueva especie del género Myotra- est representatives of the third faunal episode in Menorca gus Bate, 1909 (Mammalia, Bovidae) en la isla de Menorca: Myotragus binigausensis nov. sp. Implicaciones paleozoogeográficas. Endins 7, (Nesiotites meloussae, Hypnomys eliomyoides Agustí, 1980 7–47. and Myotragus binigausensis Moyà and Pons, 1980) and in Obrador, A., Pomar, L., 2004. El Miocè del Migjorn. In: Fornós, J.J., Obrador, which, moreover, remains of the Menorcan giant tortoise A., Rosselló, V.M. (Eds.), Història Natural del Migjorn de Menorca: el medi físic i l’influx humà. Soc. d’Hist. Nat. Balears, Palma de Mallorca, had not be found. Cap d’Artrutx is also different from Bini- pp. 73–92. gaus in the shortage of remains recovered from Nesiotites Pérez-García, A., Ortega, F., Jiménez Fuentes, E., 2016. Taxonomy, system- sp. and Hypnomys sp. and by the absence of Myotragus. Such atics and diversity of the European oldest testudinids. Zool. J. Linn. Soc. 177, 648–675. characteristics could indicate that Cap d’Artrutx would rep- Pons-Moyà, J., Moyà-Solà, S., 1980. Nuevo representante del género Nesi- resent a moment in which the populations of Nesiotites otites Bate, 1944, Nesiotites meloussae nov. sp. (Insectivora, Soricidae) and Hypnomys sp. were composed of a low number of de los rellenos cársticos del “Barranc de Binigaus” (Es mercadal, individuals, at a time in which it is not possible to know Menorca). Endins 7, 53–56. Pomar, L., 2001. Ecological enhancement of sedimentary accomodation: precisely whether the Menorcan giant tortoise was extinct evolution from a carbonate ramp to rimmed shelf, Upper Miocene, and whether Myotragus was present on the island. From a Balearic Islands. In: Camoin, G. (Ed.), Paleoceanology of reefs and 324 J.Q. Cardona, J. Agusti / C. R. Palevol 18 (2019) 317–324 carbonate platforms: Miocene to Modern. Palaeogeogr., Palaeoclima- Quintana, J., Bover, P., Ramis, D., Alcover, J.A., 2003. Cronologia de la tol., Palaeoecol., special issue 175. , pp. 259–272. desaparició de Myotragus balearicus Bate 1909 a Menorca. Endins 25, Pomar, L., Ward, W.C., Green, D.G., 1996. Upper Miocene Reef Complex of 155–158. the Llucmajor area, Mallorca, Spain. In: Franseen, E., Esteban, M., Ward, Quintana, J., Moncunill-Solé, B., 2014. Hypolagus balearicus Quintana, W.C., Rouchy, J.M. (Eds.), Models for Carbonate Stratigraphy from Bover, Alcover, Agustí & Bailon, 2010 (Mammalia: Leporidae): new Miocene Reef Complexes of the Mediterranean regions, 5. S.E.P.M. data from the Neogene of Eivissa (Balearic Islands, Western Mediter- Concepts in Sedimentology and Paleontology Series, pp. 191–225. ranean). Geodiversitas 36 (2), 283–310. Pons-Monjo, G., Moyà-Solà, S., Furió, M., 2012. New data on the origin of Quintana, J., Ramis, D., Bover, P., 2016. Primera datació d’un mamífer Nesiotites (Soricidae, Mammalia) in Menorca (Balearic Islands, Spain). no autòcton (Oryctolagus cuniculus [Linnaeus, 1758)]) (Mammalia: C. R. Palevol 11 (5), 393–401. Lagomorpha) del jaciment holocènic del Pas d’en Revull (barranc Quintana, J., 1995. Fauna malacológica asociada a Cheirogaster gymnesica d’Algendar, Ferreries). Rev. Menorca 95, 185–200. (Bate, 1914). Implicaciones biogeográficas. Boll. Soc. Hist. Nat. Balears Reumer, J., 1979. On the new micromammals from the Pleistocene of Mal- 38, 95–119. lorca. Proc. Konink. Ned. Akad. Wetenschappen, ser. B 82 (4), 473–482. Quintana, J., 1998. Aproximación a los yacimientos de vertebrados del Rosell, J., Gómez-Gras, D., Elizaga, E., 1989. Mapa Geológico de España, Mio-Pleistoceno de la isla de Menorca. Boll. Soc. d’Hist. Nat. Balears escala 1:25.000. Santandria: 645 bis-I, 646-IV. Instituto Tecnológico 41, 101–117. Geominero de España, Madrid. Quintana, J., Agusti, J., 2007. Los mamíferos insulares del Mioceno medio y superior de Menorca (Islas Baleares, Mediterráneo occidental). Geo- bios 40, 677–687.