A new elephant fossil from the late Pleistocene of Alghero: The puzzling question of Sardinian dwarf elephants

Endemic elephants, variously reduced in size, have been reported from a number of Mediterranean islands. Most of these originated from the mainland species Palaeoloxodon antiquus. A few dwarf mammoth remains are recorded from Crete and Sardinia. In Sardinia, a largely incomplete skeleton and a few mammoth teeth have been reported from localities believed to range in age from the late middle to the late Pleistocene. The chronology of colonisation by the ancestral species, the actual persistence through time of Mammuthus lamarmorai on the island, and the morphological and dimensional range of the species are, however, poorly known. This research aims to describe a distal portion of a left tibia of a dwarf elephant found in the Alghero area (NW Sardinia), showing some morphological traits and dimensions consistent with those of the endemic Sardinian mammoth (Mammuthus lamarmorai). The main unanswered questions about chronology, colonisation and population dynamics of endemic Sardinian elephants are highlighted and briefly discussed.     

Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications

Crocodylian remains are collected in 39 fossil-bearing localities but only in seven localities specimens with reliable taxonomic attributions, at least to genus level have been collected. Three species have been reported from the early Lutetian Purga di Bolca site: Pristichampsus cf. Pristichampsus rollinati, Asiatosuchus sp., Hassiacosuchus sp. (=Allognathosuchus sp.). The three crocodilians discovered at Purga di Bolca have been reported also from Geiseltal and Messel (Middle Eocene, Germany). Bolca at that time was part of a Tethysian archipelago and no mammals have been found there till now. Crocodilians and turtles clearly arrived from the European mainland across a marine water barrier. Among the other fossiliferous localities of Veneto, very interesting is the Monte Zuello site, of late Middle Eocene age, yielding a longirostrine crocodilian, Megadontosuchus arduini, a tomistomine species. Tomistomines are known in contemporaneous sediments of both Europe and Africa, but the European forms Dollosuchus and Kentisuchus seem the closest taxa. Remains of Oligocene age have been collected in Veneto and Liguria, but the fossils discovered in the second region are teeth or fragmented bones. The fossil crocodilians of Monteviale (Veneto), of Early Oligocene age, have been assigned to two species but they have been recently all identified as Diplocynodon ratelii, known from several European sites of Late Eocene, Oligocene and Miocene age. This species arrived in the Monteviale area from the European mainland across a narrow sea. Several crocodilian fossils of Miocene age are very fragmentary or represented by isolated teeth. In the Middle and Late Miocene of Sardinia, a well-established species, Tomistoma calaritanum is present. Remains of Tomistoma of the same age have been reported in some localities in Tuscany, Apulia, Sicily and Malta. In the Mediterranean area, the genus is known from European and African sites (of older age). The colonisation of Europe by this genus is the result of a dispersion from Africa (or less probably from Asia). During Late Miocene Sardinia and Tuscany belong to the same palaeobioprovince characterized by the Oreopithecus-Maremmia fauna. In Tuscany, a crocodilian identified as Crocodylus bambolii is present in the late Miocene site of Monte Bamboli. If the generic attribution of this form is correct, its ancestors must have arrived from Africa. Another fossil assemblage of Late Miocene age characterizes the Apulia-Abruzzi palaeobioprovince (Hoplitomeryx-Microtia fauna) and testifies complete isolation between the two palaeobioprovinces. In this last area, remains of Crocodylus sp. have been collected in coastal sandstones at Scontrone (Abruzzi) and in several fissure fillings of Gargano of slightly younger age. The ancestors of this species arrived from Africa while no African elements are present among the mammalian fauna. The dispersion of the genus Crocodylus in the Italian palaeoislands may have taken place once, with allopatric differentiation of the two populations (Tuscany-Sardinia and Apulia-Abruzzi) or twice with independent colonisation of each area.     

Establishing the eradication unit of Molara Island: a case of study from Sardinia, Italy

Molara is a small island belonging to the Marine protected Area Tavolara—Punta Coda Cavallo, in Sardinia. During 2006–2007, a bio-monitoring program reported a strong presence of the black rat, Rattus rattus, on Molara island. Rat predation has detrimentally affected the unique biodiversity of this island, thus, in 2008 an eradication campaign was conducted. Our eradication protocol included a pre-eradication genetic investigation, using 8 microsatellite loci, on a rat population of Molara as well as on neighbour islands within the Marine Protected Area (MPA). The main goal of this genetic investigation was to establish the correct borders of the eradication unit of Molara island. As several recent eradication campaigns have been unsuccessful, due to incomplete and unstable eradication, we also aimed to assess possible hidden sources of reinvasion. Specimens were also collected during post- eradication monitoring on Molara for genetic screening to establish their origin, and thus validate the effectiveness of our eradication campaign. According to our genetic analysis, within the MPA there are four different eradication units, corresponding to the islands of Molara, Tavolara, Piana and to the Sardinia mainland. Gene flow among these four units is more or less absent. The assignment and clustering tests performed on pre and post-eradication samples seem to indicate that the population of Sardinia mainland is a possible source of re-invasion for the Piana and Molara populations.     

The endemic bovids from Sardinia and the Balearic Islands: State of the art

Bovids are not so common in endemic insular faunas and are mainly recorded in Southeast Asia, Japan and some Mediterranean islands. In the Western Mediterranean, endemic bovids have been recorded during the late Miocene in the Tusco-Sardinian palaeobioprovince (Baccinello-Cinigiano basin, South Tuscany, and Fiume Santo, north-western Sardinia). In the latest Neogene and Quaternary, bovids showing highly endemic features were restricted to the Balearic Islands and Sardinia, while Bovini only slightly reduced in size were present on Pianosa, Malta and Sicily. On Sardinia, the richest bovid sample comes from Monte Tuttavista (Orosei), where at least three species have been identified: Asoletragus genthry, Nesogoral aff. N. melonii, and Nesogoral sp. 2. On Mallorca (Balearic Islands) six chronospecies belonging to the Myotragus endemic phylogenetic lineage have been described, spreading in age from the Early Pliocene to the Holocene. For decades, a close phylogenetic relationship between Nesogoral and Myotragus has been widely accepted by scholars. Morphological and biometrical differences shown by Balearic and Sardinian bovids have generally been regarded as the result of the evolution into two different island ecological systems, characterized by different inter and intra-guild selection pressures. Indeed, the more diversified environment of Sardinia, as well as the presence of other large mammals (similar-sized competitors belonging to the same guild and a running predator), increased the interspecific competition, forcing Sardinian bovids to exploit different resources and to occupy different niches, while Myotragus exploited under a monopoly regime the supply of resources available for large herbivores on the Eastern Balearic Islands. Nonetheless, new data suggest that Nesogoral and Myotragus possibly originated from different taxa.     

First European evidence for transcontinental dispersal of Crocodylus (late Neogene of southern Italy)

It is generally assumed that the Neogene crocodylian fauna of Europe has been represented only by brevirostrine alligatoroid Diplocynodon and longirostrine false gharials ( Gavialosuchus and/or Tomistoma ), which became extinct prior to 6 Mya. Although several lines of evidence suggest that Crocodylus originated in Africa during the Miocene and then promptly dispersed to other continents, the occurrence of this genus in Europe has never been rigorously proven and the traditional palaeontological approach failed to identify a monophyletic group of fossil Crocodylus (simply leading to a proliferation of extinct taxa). The new remains reported here, from an endemic insular fauna from southern Italy, Late Messinian to earliest Pliocene in age (5–6 million years old), represent the youngest European crocodylian, and allow, for the first time in a phylogenetic context, an unambiguous demonstration that Crocodylus dispersed into Europe, possibly during the Tortonian. If the peculiar morphology of the medial maxillary edge is interpreted as evidence for a medial dorsal boss, the southern Italian Crocodylus could be related to C. checchiai from the late Neogene of Libya. The presence of this African immigrant in Europe confirms the role of climate change for faunal dispersal and island colonization.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 293–307.     

Peopling of three Mediterranean islands (Corsica,Sardinia, and Sicily) inferred by Y-chromosome biallelic variability

An informative set of biallelic polymorphisms was used to study the structure of Y-chromosome variability in a sample from the Mediterranean islands of Corsica and Sicily, and compared with data on Sardinia to gain insights into the ethnogenesis of these island populations. The results were interpreted in a broader Mediterranean context by including in the analysis neighboring populations previously studied with the same methodology. All samples studied were enclosed in the comparable spectrum of European Y-chromosome variability. Pronounced differences were observed between the islands as well as in the percentages of haplotypes previously shown to have distinctive patterns of continental phylogeography. Approximately 60% of the Sicilian haplotypes are also prevalent in Southern Italy and Greece. Conversely, the Corsican sample had elevated levels of alternative haplotypes common in Northern Italy. Sardinia showed a haplotype ratio similar to that observed in Corsica, but with a remarkable difference in the presence of a lineage defined by marker M26, which approaches 35% in Sardinia but seems absent in Corsica. Although geographically adjacent, the data suggest different colonization histories and a minimal amount of recent gene flow between them. Our results identify possible ancestral continental sources of the various island populations and underscore the influence of founder effect and genetic drift. The Y-chromosome data are consistent with comparable mtDNA data at the RFLP haplogroup level of resolution, as well as linguistic and historic knowledge.     

Habitat requirements of Eurasian pine marten <Emphasis Type="Italic">Martes martes</Emphasis> in a Mediterranean environment

Evaluating presence and habitat requirements of small carnivores is essential for their conservation. The Eurasian pine marten Martes martes, often described as a habitat specialist associated primarily with forest habitats, has been recently found to live even in patchily wooded country and in shrublands. We evaluated the environmental factors that determine the distribution of the pine marten in a Mediterranean landscape on the island of Sardinia (central Italy). Camera trapping sessions and scat surveys were carried out to assess the presence of the species, then a potential distribution model was developed using ecological niche factor analysis (ENFA), which requires only presence data. The pine marten selected highest altitudes, shrublands, rocky areas, and woodlands, and avoided urban areas and arable lands. Our results indicate that pine marten distribution in our study area is constrained by these variables. The ENFA analysis provided important clues about the distribution range of M. martes and its preferential environmental conditions, updating knowledge of its ecological requirements in Italy.     

The origin of clonal diversity and structure of Populus alba in Sardinia: evidence from nuclear and plastid microsatellite markers

Background and Aims

Populus alba is a thermophilic forest tree present in the Mediterranean basin. Its habitat is highly fragmented and its distribution range has been subject to long-term human interference, resulting in debate surrounding whether certain populations are native or exotic in origin. In particular, populations from the islands of Corsica and Sardinia are of uncertain origin. While populations of P. alba mainly reproduce sexually, clonal reproduction is also common. The aims of this study were to locate and molecularly characterize the poorly studied island populations of P. alba and compare these with samples from various spatial scales, in order to provide information on the genetic structure and phylogeography of this species. This information will provide evidence on whether the species is native to Sardinia, which is important for the development of conservation strategies.

Methods

DNA extracts were obtained from the following P. alba trees: 159 from Sardinia, 47 from Ticino regional park (northern Italy), 15 acquired from an Italian Germoplasm Bank (IRC; Italian Reference Collection) and 28 from the Mediterranean basin (MB). Genetic polymorphisms were revealed at nuclear and chloroplast DNA (cpDNA) microsatellite loci, both at the island scale (Sardinia) and at broader scales, for comparative assessment of the genetic and genotypic diversity and phylogeography.

Key Results

Based on nuclear microsatellite loci, Sardinian white poplar consists of a small number of genets (26), each of which is represented by several ramets. Despite the uniqueness of the Sardinian haplotypes and the very low value of genetic diversity at the cpDNA level (v_K = 0·15), the H_T (0·60) and the A_R (3·61) values, estimated at the nuclear level for Sardinia, were comparable with those of the other populations and collections.

Conclusions

The uniqueness of the cpDNA haplotypes, the prevalence of clonality and the restricted number of genets recorded suggest that Sardinian white poplar could be a floristic relict of the native flora of the island, which has spread through available habitats on the island mainly by means of vegetative propagation and human activities.Key words: Populus alba, Sardinia, genets, ramets, phylogeography, native forest species     

Arundo micrantha,a new reed species for Italy,threatened in the freshwater habitat by the congeneric invasive A. donax

Arundo micrantha Lam., a reed species from southern Mediterranean area, has found in Sardinia. Its presence represents the first record for Italy and the third one for western Europe. Investigations on distribution and ecology of the Sardinian population have been carried out, with special focusing on synecology of the species. A. micrantha has found in 40 localities mainly distributed along temporary streams and permanent rivers in the central and southern part of the island. Most of Sardinian populations occurs on alluvial soils in Thermomediterranean bioclimatic context. The autonomy of Sardinian phytocoenoses was characterized with the definition of exclusive floristic taxa, biological and chorological elements, and syndinamic relationships with other assemblages. Comparing the community with other reed beds from the Mediterranean basin, a marked floristic differentiation arose between the phytocoenoses characterized by autochthonous Arundo species and those dominated by the invasive Arundo donax. Finally, management activities are proposed to ensure the conservation of this threatened autochthonous plant community in freshwater habitat.     

Genetics,geography, and culture: The population of S. Pietro Island (Sardinia,Italy)

An interesting aspect of the island of Sardinia (Italy) is the wide range of genetic variability within the island itself. The variability is widened by the presence of some populations of different ethnic origin who speak a language other than Sardinian. This work deals with the study of the genetic structure of the Carloforte population which inhabits the tiny island of S. Pietro 4 km off the southwest coast of Sardinia. S. Pietro was first populated in 1738 by emigrants coming from the island of Tabarka (Tunisia) who spoke an archaic form of the Ligurian dialect. Data on genetic polymorphisms in the Carloforte population are presented and discussed in relation to some Sardinian and Italian populations. Data on demographic and matrimonial structure are also presented. The genetic analyses show the Carloforte population as being clearly separated from both Sardinians and continental Italians. The isolation of Carloforte, highlighted by language diversity, endogamy, and consanguinity levels and marriage area, supports the idea of genetic diversity linked to cultural peculiarity. © 1996 Wiley-Liss, Inc.     

Reappearance of Aedes albopictus in Sardinia

Aedes (Stegomyia) albopictus is a vector of major human diseases. From 1979 established populations were found in the Mediterranean Region. The first record of Ae. albopictus in Italy goes back to 1990 and at present it is permanently established in several provinces of central and northern Italy. Ae. albopictus was detected for the first time in Sardinia in 1995, and a new record was reported in 1997. After these two isolated episodes, the "tiger-mosquito" was considered apparently eradicated. At the end of September 2006, two natural breeding populations including adults of Ae. albopictus were recorded in two different localities of South-Sardinia: in the botanical garden of downtown Cagliari and in a private garden of a small village, Teulada, on the south-western coast of Sardinia. The origin of this new introduction of Ae. albopictus in Sardinia is still unclear. In both cases the species has been found in artificial containers in densely populated areas.     

A new chinchilloid (Rodentia,Hystricognathi) from the early Miocene of the localities of Bryn Gwyn and Gran Barranca (Patagonia,Argentina)

A new genus and species of chinchilloid rodent is described here. It was found in Colhuehuapian levels (early Miocene) of the localities of Bryn Gwyn and Gran Barranca, Chubut Province, Argentina. The new taxon shows a unique combination of characters (e.g., protohypsodont teeth, upper molars trilophodont, m1 and m2 tri- or tetralophodont with the second crest in position of variable development, and m3 trilophodont, cement absent) that make it different from any other known chinchilloids. We performed a phylogenetic analysis to corroborate the relationships of the new taxon within the Chinchilloidea. Our results indicate that the new taxon is best classified as Chinchilloidea incertae sedis, together with Incamys, Garridomys, and Scotamys. These species form the basal stock that leads to the modern lineage Chinchillidae (chinchillas and viscachas). The presence of the new taxon in these localities increases the diversity of chinchilloids during the early Miocene and reduces the dissimilarity between the faunas found in Bryn Gwyn and Gran Barranca.     

Dental remains of Lower Miocene small rodents from Napak (Uganda): Afrocricetodontidae,Myophiomyidae, Kenyamyidae and Sciuridae

The Early Miocene sites of Napak (Uganda), which have been prospected by the Uganda Palaeontology Expedition since 1985, have yielded abundant fossil remains, including a rich and diverse rodent fauna. Ever since the work of Lavocat (1973) on the East African Miocene rodents, which focused mainly on the Kenyan remains (but included some material from Napak discovered by Bishop during the 1950s and 1960s), few studies have been made on the Ugandan specimens. This lack provides motivation for further study of Napak micro-mammals, especially the abundant rodents, which were collected at five sites, Napak IV, V, XV, XXX and XXXI. The fossils described here are attributed to six species representing four families: Afrocricetodontidae (Afrocricetodon songhori, Protarsomys macinnesi), Myophiomyidae (Myophiomys arambourgi), Kenyamyidae (Simonimys genovefae, Kenyamys mariae) and Sciuridae (Vulcanisciurus africanus). Comparison of the dentition of extant and extinct taxa allows us to propose dietary hypotheses (granivory, herbivory and omnivory) for these rodents. In addition, the fossil rodents associated with other fauna from the same localities suggest a more-or-less dense forest environment with clearings, and the probable presence of a humid climate at the time of deposition.     

The rodents from the Late Miocene Oreopithecus-bearing site of Fiume Santo (Sardinia,Italy)

The locality of Fiume Santo (Sardinia, Italy) represents the westernmost extension of endemic Oreopithecus-bearing faunas. Here we describe the rodent fauna recovered at this site, which only includes the murids Huerzelerimys oreopitheci and Anthracomys lorenzi, and the glirids Anthracoglis engesseri nov. sp. and Anthracoglis nov. sp. I. A. engesseri nov. sp. differs from Anthracoglis marinoi, the other species of this genus known so far, by its larger size, wider upper cheek teeth and more reduced accessory ridges, particularly in the upper cheek teeth. Anthracoglis nov. sp. I is only known by a single tooth and cannot be adequately characterized. The fauna seems to be a mixture of elements from the V-1, V-2 and V-3 local zones of the Baccinello-Cinigiano basin (Tuscany, Italy), with H. oreopitheci being characteristic from zone V-1 and Anthracoglis lorenzi from zone V-3. Nevertheless, a few remains of A. lorenzi have been also recovered from V-2 assemblages at Baccinello and A. engesseri nov. sp. is also recorded in Monte Bamboli, which is correlated to V-2 faunas. Accordingly, a correlation to V-2 zone of the Baccinello-Cinigiano basin is preferred. Finally, the structure and composition of the rodent assemblage is compared to those of the Tuscan sites. The Fiume Santo assemblage resembles the insular faunas of Tuscany by its low species richness, although it is more balanced. This may be related to slight chronological or environmental differences between the two areas or to the existence of a geographical barrier.     

Bryozoaires et Brachiopodes des “Calcaires bajociens a Bryozoaires” des Causses (France) et leur paléoécologie

Two Lower Bajocian new localities, with many silicified bryozoans and brachiopods have been discovered in Causses (France): Saint-Rome-de-Cernon (Aveyron) and Mende (Lozère). The study of the faunas allows to increase the anterior knowledges supplied by the study of localities in the South of Causses (neighbourhood of Alzon and Sumène = North of the Seuil sud-caussenard).The interest is, at first, the distance between the localities: Mende in the North, Saint-Rome-de-Cernon in the West. Moreover, in the second, well-preserved faunas were fossilized without transport. Bryozoans and brachiopods are very numerous. Large and complete bryozoan colonies have been get out by action of HCl. Fine internal structures like rhynchonellid Nannirhynchia or like thecidean transversarium and hemispondylium have been studied. At last, observations and paleoecological hypothesis are drawn, likewise comparison between three localities or locality-groups known in the Causses basin.The fauna from Saint-Rome-de-Cernon is biocoenosis, the substratum was soft, the hydrodynamism low and the depth, according to lamellibranchs, about 200 meters. The beds with bryozoans and small brachiopods contain a fauna with two predominant bryozoan species: Ceriocava straminea and Mesenteripora wrighti. These two species could built large colonies, in spite of fine granulometry of the sedimentary bottom. These colonies provided supports to less adapted bryozoans, to thecidids and to lamellibranch Lopha. The beds with terebratulids and bryozoans contain lesser bryozoans and thecidids. Terebratulids are the main group and are fixed with their peduncle to organic support above the irregular deposits. Beside these benthic animals, the rhynchonellid Nannirhynchia campestriensis is epibiontic on gorgonids or epiphytic on floating sea-weeds.On the contrary, all fossils from Mende have been more or less carried, excepted large sponges perhaps. The most numerous bryozoans (Ceriocava straminea, Mesenteripora wrighti) and numerous Lenticulines (Foraminifera) lived on hard bottom with moving water in community with crinoids. It is possible that a smaller part of bryozoans, requiring lower hydrodynamism, lived, either in sheltered parts of the crinoid community or closer to the sedimentation zone. Thecidids were fixed either on bryozoans in crinoid communities or on shelly remains and on bryozoans nearer to the sedimentation zone. Like in Saint-Rome-de-Cernon, Nannirhynchia campestriensis was epibiontic, or epiphytic, or both.In comparison with the whole localities from the Seuil sud-caussenard, we notice the simplicity of Saint-Rome-de-Cernon and Mende. The localities from the Seuil sud-caussenard were assemblages proceeding from 2 or 3 biotops. On the contrary, at Saint-Rome-de-Cernon we have a biocoenosis and at Mende a thanatocoenosis supplied by one biotop almost.     

Prominatherium dalmatinum from the late Eocene of Grancona (Vicenza,NE Italy). The oldest terrestrial mammal of the Italian peninsula

So far, the oldest terrestrial mammal associations in Italy dates to the beginning of the Oligocene, with Anthracotheriidae being the most represented taxon. Sites from northern Italy yielded remains of the genus Anthracotherium that spread from Asia to western Europe after the Grande Coupure. A finding at Grancona, which is Priabonian in age, implies that Anthracotheriidae family reached the Italian Peninsula before the Eocene–Oligocene boundary. Thus, the dispersal of this family in northern Italy is anterior than previously believed. The fossil consists of a poorly preserved right hemi-maxilla with well-preserved P4 and M3. The shape and the size of the teeth are not compatible with the genus Anthracotherium. On the contrary, the closer affinities with the Croatian species Prominatherium dalmatinum suggest a connection between the Balkan area and the Italian peninsula and a possible new way of dispersal for this family.