Josep Marmi

Phylogeny, evolutionary history and taxonomy of the Mustelidae based on sequences of the cytochrome b gene and a complex repetitive flanking region

The Mustelidae is a diverse family of carnivores which includes weasels, polecats, mink, tayra, martens, otters, badgers and, according to some authors, skunks. Evolutionary relationships within the family are under debate at a number of different taxonomic levels, and incongruencies between molecular and morphological results are important. We analysed a total of 241 cytochrome b (cyt b) gene sequences and 33 sequences of a complex repetitive flanking region from 33 different species to compile an extensive molecular phylogeny for the Mustelidae. We analysed these sequences and constructed phylogenetic trees using Bayesian and neighbor‐joining methods that are evaluated to propose changes to the taxonomy of the family. The peripheral position of skunks in phylogenetic trees based on both loci suggests that they should be considered a separate family, Mephitidae. The subfamily Melinae is the basal group within the Mustelidae and trees based on the cyt b gene suggest that the American badger, Taxidea taxus, should be considered a separate monotypic subfamily, Taxidiinae. Otters classified within the genera Lutra, Amblonyx and Aonyx are grouped within the same clade in cyt b and combined partial cyt b and flanking region trees and show reduced levels of inter specific divergence, suggesting that they could be classified together under a single genus, Lutra. The Bayesian tree based on combined data from both loci supports the idea that subfamily Mustelinae is paraphyletic, as otters (subfamily Lutrinae) are included in this subfamily. Low levels of genetic divergence among European polecat, Mustela putorius, steppe polecat, Mustela eversmannii, and European mink, Mustela lutreola, suggest that these species could be considered subspecies within a single species, Mustela putorius. Our results are consistent with a rapid diversification of mustelid lineages in six different radiation episodes identified since the Early Eocene, the oldest events being the separation of subfamilies and the split of marten (Martes, Gulo) and weasel (Mustela) lineages in the Early Middle Miocene. The separation of New World from Old World lineages and the split of the remaining genera are estimated to have occurred in Late Miocene. The most recent events have been the differentiation of species within genera and this probably occurred in four radiation episodes at the end of Late Miocene, Early Pliocene, Late Pliocene and Pleistocene epochs.

Nuclear and mitochondrial phylogenies provide evidence for four species of Eurasian badgers (Carnivora)

Del Cerro, I., Marmi, J., Ferrando, A., Chashchin, P., Taberlet, P. & Bosch, M. (2010). Nuclear and mitochondrial phylogenies provide evidence for four species of Eurasian badgers (Carnivora). —Zoologica Scripta, 39, 415–425.

Mitochondrial DNA diversity and origins of South and Central American goats.

We have analysed the genetic diversity of South and Central American (SCA) goats by partially sequencing the mitochondrial control region of 93 individuals with a wide geographical distribution. Nucleotide and haplotype diversities reached values of 0.020 +/- 0.00081 and 0.963 +/- 0.0012 respectively. We have also observed a rather weak phylogeographic structure, with almost 69% of genetic variation included in the within-breed variance component. The topology of a median-joining network analysis including 286 European, Iberian, Atlantic and SCA mitochondrial sequences was very complex, with most of the haplotypes forming part of independent small clusters. SCA sequences showed a scattered distribution throughout the network, and clustering with Spanish and Portuguese sequences occurred only occasionally, not allowing the distinguishing of a clear Iberian signature. Conversely, we found a prominent cluster including Canarian, Chilean, Argentinian and Bolivian mitochondrial haplotypes. This result was independently confirmed by constructing a Bayesian phylogenetic tree (posterior probability of 0.97). Sharing of mitochondrial haplotypes by SCA and Canarian goats suggests that goat populations from the Atlantic archipelagos, where Spanish and Portuguese ships en route to the New World used to stow food and supplies, participated in the foundation of SCA caprine breeds.

Allodaposuchus palustris sp. nov. from the Upper Cretaceous of Fumanya (South-Eastern Pyrenees, Iberian Peninsula): Systematics, Palaeoecology and Palaeobiogeography of the Enigmatic Allodaposuchian Crocodylians

The controversial European genus Allodaposuchus is currently composed of two species (A. precedens, A. subjuniperus) and it has been traditionally considered a basal eusuchian clade of crocodylomorphs. In the present work, the new species A. palustris is erected on the base of cranial and postcranial remains from the lower Maastrichtian of the southern Pyrenees. Phylogenetic analyses here including both cranial and postcranial data support the hypothesis that Allodaposuchus is included within Crocodylia. The studied specimen suggests little change in postcranial skeleton along the evolutionary history of crocodylians, except for some bone elements such as the axis, the first caudal vertebra and the ilium. The specimen was found in an organic mudstone corresponding to a coastal wetland environment. Thus, A. palustris from Fumanya is the first Allodaposuchus reported in lacustrine-palustrine settings that expand the ecological range for this genus. The S-DIVA palaeobiogeographic reconstruction of ancestral area suggests that early members of Crocodylia rapidly widespread for the Northern Hemisphere landmasses no later than the Campanian, leading the apparition of endemic groups. In that way “Allodaposuchia” represents an endemic European clade probably originated in the Ibero-Armorican domain in the late Campanian and dispersed by the Southern European archipelago prior to the early Maastrichtian.

50,000 years of genetic uniformity in the critically endangered Iberian lynx

Low genetic diversity in the endangered Iberian lynx, including lack of mitochondrial control region variation, is thought to result from historical or Pleistocene/Holocene population bottlenecks, and to indicate poor long‐term viability. We find no variability in control region sequences from 19 Iberian lynx remains from across the Iberian Peninsula and spanning the last 50 000 years. This is best explained by continuously small female effective population size through time. We conclude that low genetic variability in the Iberian lynx is not in itself a threat to long‐term viability, and so should not preclude conservation efforts.

Oldest Records of Stingray Spines (Chondrichthyes, Myliobatiformes)

JOSEP MARMI,* 1# BERNAT VILA,1 # ORIOL OMS,2 ANGEL GALOBART,1 and HENRI CAPPETTA3; institut Catala de Paleontologia, Edifici ICP, Campus de la Universitat Autonoma de Barcelona, 08193, Bellaterra, Spain, josep.marmi@icp.cat; bernat.vila@icp.cat; angel.galobart@icp.cat; 2Universitat Autonoma de Barcelona, Facultat de Ciencies (Geologia), 08193, Bellaterra, Spain, joseporiol.oms@uab.cat; 3ISE-M, Universite de Montpellier II-Sciences et Techniques du Languedoc, Ce 064, Place Eugene Bataillon, 34095, Montpellier Cedex 5, France, henri.cappetta@univ-montp2.fr

Phylogenetic inference and comparative evolution of a complex microsatellite and its flanking regions in carnivores.

We sequenced locus Mel 08, with complex short repetitive motifs, in 24 carnivore species belonging to five different families in order to explore mutational changes in the region in the context of locus and species evolution. This non-coding locus includes up to four different parts or repetitive motifs showing size variability. The variability consists of repeat additions and deletions; substitutions, insertions and/or deletions creating interruptions in the repeat; and substitutions, insertions and deletions in the flanking regions. The locus has different repeat expansions in different carnivore subfamilies. We hypothesize that the complexity of this locus is due to a high mutation rate at an ancestral DNA sequence and, thus, prompts the emergence of repeats at mutational hotspots. High levels of homoplasy were evident, with nine electromorphs representing 28 haplotypes never shared across species. The variability in flanking regions was informative for phylogenetic inference and their evolutionary content. Tree topologies were congruent with relevant hypotheses on current conflicts in carnivore phylogenies, such as: (i) the monophyly of Lutrinae, (ii) the paraphyly of Mustelinae, (iii) the basal position of the Eurasian badger, Meles meles , in the Mustelidae, (iv) the classification of skunks as a separate family, Mephitidae, and (v) the placement of the red panda, Ailurus fulgens , as a monotypic family, Ailuridae, at a basal position in the Musteloidea.

The youngest sauropod evidence in Europe

Titanosaur sauropods constituted one of the major dinosaur faunal components in Europe along the latest Cretaceous. Although they were abundant in the late Campanian–early Maastrichtian, titanosaurs apparently demised in terms of diversity and abundance in the late Maastrichtian Ibero-Armorican domain. Thus, any finding in this crucial period provides new clues to understand the biodiversity trend of the group prior to the K–Pg boundary. Here, we report the occurrence of a vertebral element attributed to a titanosaur in fluvial deposits dated as uppermost Maastrichtian (C29r chrone). The specimen corresponds to the right posterior part of a posterior cervical vertebra. CT-scan and histological analyses revealed high pneumatisation with laminar structures composed of compact bone, strongly remodelled with at least three generations of secondary osteons. These evidences suggest a relatively advanced age for the individual at death. Given that the partial vertebra was recovered from the uppermost portion of the late Maastrichtian, it might represent the youngest evidence for titanosaur sauropods in Europe discovered so far. Further, alongside to other remains (i.e. eggshells and footprints), the studied partial vertebra provides evidence for the persistence of this taxonomic group at the very end of the Cretaceous in southwestern Europe.

Oldest European occurence of Meles (Mustelidae, Carnivora) from the Middle Pliocene (Mn16) of Almenara-Casablanca-4 Karstic Site (Castellón, Spain)

The origin of badgers (Subfamily Melinae) and the early evolutionary history of the genus Meles Brisson, 1762 remain still unresolved. Fossil, morphologic and genetic evidence suggest that only Meles and Arctonyx Cuvier, 1825 are closely related among extant badger genera (Petter, 1971; Bryant et al., 1993; Koepfli et al., 2008). Petter (1971) suggested that Meles, Arctonyx and Arctomeles Stach, 1951 evolved from an ancient “Meles” stock during the beginning of Late Pliocene. The origin of this lineage probably occurred in Asia during the Early Pliocene; afterwards, the Meles lineage spread to the west into Europe between the Late Pliocene and Early Pleistocene (Neal and Cheeseman, 1996). Recently, Koepfli et al. (2008) pointed that Arctonyx and Meles diverged in Eurasia between 4.4 and 3.6 MYA. At present, the Eurasian badger (Meles meles Linnaeus, 1758) has a Palearctic distribution, ranging from the Iberian Peninsula to Japan, including Palestine, southern China, Western Siberia and Great Britain, showing a wide morphologic variability across its range, which complicates its taxonomy at the intraespecific level. On the other hand, the hog badger (Arctonyx collaris Cuvier, 1825) is distributed across South-eastern Asia (China, Bhutan, Assam, Myanmar, Thailand, Vietnam and Sumatra) (Neal and Cheeseman, 1996). Historically, the oldest record of the genus Meles in Europe corresponded to Meles gennevauxi Viret, 1939 from the Early Pliocene of Montpellier (Viret, 1939). Recently, some authors have transferred this form to the genus Arctomeles (Tedford and Harrington, 2003 and references therein). Arctomeles pliocaenicus Stach, 1951, from the Early Pliocene of Weze-1 (Poland) (Stach, 1951), is the type species of this genus, where Parameles ferus Roshchin, 1949 must be also included (Roshchin, 1949; Tedford and Harrington, 2003 and references therein). Therefore according to this revision, the genus Arctomeles includes these five taxa: A. pliocaenicus, A. gennevauxi, A. suillus Teilhard and LeRoy, 1945, A. ferus and A. sotnikovae Tedford and Harrington, 2003. Up to now, six species and one subspecies of the genus Meles have been reported from the Late Pliocene to the Middle Pleistocene: M. thorali Viret, 1951 from Saint-Vallier (France) and Vatera (Greece); M. iberica Arribas and Garrido, 2007 from Fonelas P-1 (Spain); M. dimitrius Koufos, 1992 from Gerakarou and Apollonia-1 (Greece); M. hollitzeri Raebeder, 1976 from Untermassfeld (Germany) and Deutsch-Altenburg 2 (Austria); M. thorali spelaeus Bonifay, 1971 from Lunel-Viel (France); M. atavus Kormos, 1914 from Beftia 5 (Hungary) and the extant badger M. meles (Kormos, 1914; Viret, 1951; Bonifay, 1971; Raebeder, 1976; Koufos, 1992; Wolsan, 2001; Arribas and Garrido, 2007). The latter species is the most frequent mustelid in the European late Palaeolithic subfossil record suggesting a widespread distribution of this species during the Last Glacial (Sommer and Benecke, 2004). However, the distribution of the badger lineage throughout the European Plio-Pleistocene is poorly known. Forms similar to Meles thorali from the mid Villafranchian of China suggest that this species was widespread throughout the Paleartic region since its origins (Neal and Cheeseman, 1996). The discovery of badger remains close to the Plio-Pleistocene boundary is highly significant to elucidate the ancient history of Meles, especially in Western Europe, far away from the most likely area of origin of this genus. In the Early Pleistocene from the Iberian Peninsula, Meles has been reported from Venta Micena, Barranco León 5, Fuente Nueva 3 and Trinchera Dolina (TDE5) in Atapuerca (Garcı́a, 2003; Martı́nez-Navarro et al., 2003 and references therein). However, the scarcity of remains from these localities has thus far prevented their determination at the species level. In this paper, we report new material of Meles cf. thorali from the Middle Pliocene (MN16) of Almenara-Casablanca-4 (ACB-4) (Castellón, Spain). Abbreviations—ACB, Almenara-Casablanca, Almenara, Spain; IPS, Institut de Paleontologia M. Crusafont, Sabadell, Spain; ICP, Institut Català de Paleontologia, Cerdanyola del Vallès, Spain.

The taxonomic status of European Plio-Pleistocene badgers

ABSTRACT The taxonomic status of European Plio-Pleistocene badgers is currently uncertain, due to the relative scarcity of their fossil remains. Here we describe craniodental remains from the Iberian locality of Vallparadís, indicating that the extant European badger (Meles meles) was distributed throughout Europe during the late Villafranchian. On the basis of morphological and morphometrical comparisons, we attribute the studied specimens, together with other late Villafranchian and Epivillafranchian badgers from Europe, to M. meles atavus, thereby considering that M. hollitzeri and M. dimitrius (in part) are subjective junior synonyms of the former. Only the early to middle Villafranchian M. thorali can be considered a distinct species, with M. iberica and M. dimitrius (in part) being subjective junior synonyms of this other species. It is concluded that extant European badger species must have diverged from Asian badgers before the top of the Olduvai subchron. Available molecular and paleontological data are consistent with an Asian origin of the Meles lineage, and with M. thorali being the ancestral species that dispersed into Europe attaining a wholly Palearctic distribution by the early Villafranchian. Later, before the top of the Olduvai subchron, M. thorali might have given rise to both the European and Asian badger lineages through a vicariance process prompted by paleoclimatic changes.