Tertiary Taxa Research Articles

The Grotta Grande of Scario (Salerno, Italy): Archaeology and environment during the last interglacial (MIS 5) of the Mediterranean region

Archeological and paleo-environmental researches carried on the Grotta Grande site illustrate the importance of a multidisciplinary approach among archeologists, palynologists and paleontologists. The archaeology, fauna, pollen and micro-charcoal recovered in two short sedimentary successions (trenches A, F) located close to the entrance of the cave are discussed. The cave opens directly on the Tyrrhenian Sea, 2 km from Scario (Salerno, Campania, Southern Italy). The morphology of the cave and sedimentary processes were controlled by eustatic fluctuations during the late Middle Pleistocene and the early Late Pleistocene. The sea repeatedly occupied the cave. The cave was frequented by humans of Middle Palaeolithic culture. Archaeological and faunal record of the two trenches can be positioned within the climatic fluctuation posterior to the warm interglacial MIS 5e peak. Pollen have been retrieved in the older series (b–c) of trench A, referable to MIS 7–6. The most important archaeological finds are the occurrence of structures that indicate the living space (trench F) together with hearths and lithic industry. The latter is characterised by the presence of the Levallois system and by the prevalence of sidescrapers among tools. The systematic use of limestone is observed only in trench F. Large and small mammal remains recovered within the two trenches show that in the neighbourhood of the site, a diversified assemblage occurred, including eleven large mammals, among which the hippo, the straight tusked elephant and the narrow-nosed rhino are noteworthy, and fourteen small mammal taxa, mainly rodents, with a significant amount of glirids. The fauna is indicative of a temperate, forested Mediterranean environment. Variation of faunal composition suggests that environments underwent some minor fluctuations towards cooler and/or drier landscapes. Pollen indicates vegetation organized in different belts, mainly Mediterranean evergreen forest/maquis and mixed forest with conifers and deciduous broadleaved trees. Fresh water plant communities are mainly represented in trench A. Some records of “Tertiary taxa” suggest stands of survival of these plants in the surroundings of the cave. Over the time span, the landscape become more open, due to some episodes of steppe vegetation spread caused by variation in temperature and humidity. Deposit of this age are rather uncommon even in the Italian peninsula, and therefore the integration of sedimentological, archaeological, faunal and palynological data provides an important piece of information to the puzzling reconstruction of the Late Pleistocene Mediterranean environments before the onset of the glaciation.

The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis

The present study was undertaken in order to effect a comprehensive phylogenetic analysis of the order Pholidota, examining seven of the eight currently recognized extant species (absent is Manis culionensis, formerly recognized as a subspecies of Manis javanica) and nearly all the well-known fossil taxa, and employing a wide range of osteological characters from the entire skeleton. In addition, the relationship of pangolins to several putative early Tertiary relatives, including palaeanodonts and the enigmatic “edentate” Eurotamandua joresi, were investigated. The goal of the study was to improve understanding of the systematics and the biogeographic and evolutionary history of the pangolins. A computer-based cladistic analysis of phylogenetic relationships among seven extant species of pangolins, five extinct pangolin species (including all but one of the well-preserved taxa), as well as Eurotamandua and two genera of metacheiromyid palaeanodonts, Palaeanodon and Metacheiromys, was performed based upon 395 osteological characteristics of the skull and postcranial skeleton. Characters were polarized via comparison to the following successive outgroups: the basal feliform carnivoran Nandinia binotata and the hedgehog Erinaceus sp., a eulipotyphlan laursiatherian placental. A revised classification is presented based on the results of the analysis. The results support the monophyly of Pholidota and Palaeanodonta by providing new anatomical characters that can serve to diagnose a pangolin/palaeanodont clade, termed here Pholidotamorpha. Pholidota is defined so as to include all living and fossil pangolins, including all three taxa of middle Eocene “edentates” from the Messel fauna of Germany, among them Eurotamandua joresi. The results do not support the monophyly of the remaining two Messel “edentates” originally placed in the same genus Eomanis, which is restricted to the type species Eomanis waldi. Euromanis, new genus, is named with Eomanis krebsi Storch and Martin, 1994, as the type species, to form a new combination Euromanis krebsi (Storch and Martin, 1994). The analysis strongly supports the monophyly of a crown clade of pangolins diagnosed by many anatomical synapomorphies, the family Manidae. This crown clade is sister to the family Patriomanidae, which includes two Tertiary taxa, Patriomanis americana and Cryptomanis gobiensis, within the superfamily Manoidea. The relationship of the Tertiary European pangolin Necromanis to these two families is unresolved. Within Manidae, the extant species are divided into three well-supported, monophyletic genera, Manis for the Asian pangolins, Smutsia for the African ground pangolins, and Phataginus for the African tree pangolins. The latter two form a monophyletic African assemblage, the subfamily Smutsiinae. The biogeographic implications of this phylogeny are examined. A European origin for Pholidota is strongly indicated. The fossil record of pangolins would seem to support a European origin for the modern forms, with subsequent dispersal into sub-Saharan African and then to southern Asia, and the phylogeny produced in this analysis is consistent with such a scenario.

An ecological tardis: the implications of facilitation through evolutionary time

It is often challenging to link ecological processes to evolution because different temporal scales are studied and necessarily inferred. In a recent paper by Valiente-Banuet et al., Quaternary plant taxa are shown to 'pull' more ancient Tertiary taxa through evolutionary time by facilitation. This finding has profound implications for population and community ecology. Positive species interactions are of wider importance than was assumed previously in determining community composition by enhancing long-term biodiversity, mediating climate change and providing an interdependent set of selection processes in addition to the environment. Future experiments should consider the evolutionary history of species, manipulate species interactions explicitly to test for environmental effects and re-evaluate the adaptive significance of traits in the context of other species.

Chapter 7: Phylogeny and Divergence of Basal Glires

Phylogenetic analyses based on morphological data support monophyly of Glires, but not a link between Glires and zalambdalestids. Glires are more closely related to several Tertiary taxa, including primates, leptictids, pseudictopids, anagalids, and macroscelideans. Phylogenetically constrained distributions of Glires support the conventional view for a post K-T boundary radiation of modern orders of placental mammals and disagree with conclusions of some molecular studies that divergence of Rodentia and Lagomorpha at infraordinal, ordinal, and certain supraordinal levels occurred in the Cretaceous. Current hypotheses employed to explain the discrepancy between the fossil record and the molecular clock hypothesis are not supported by phylogenetic and distributional evidence of Glires. There is no compelling evidence that close relatives of Glires were present in the Cretaceous.

New remains of Sphagesaurus huenei (Crocodylomorpha: Mesoeucrocodylia) from the Late Cretaceous of Brazil

ABSTRACT Recent discovery of an almost complete skull of Sphagesaurus huenei, a previously poorly known form from the Late Cretaceous of southern Brazil, shows unexpected morphology for a crocodyliform, including an extreme heterodonty with small lower incisors, large upper caniniforms, and reversed triangle-like postcanines; edentulous anterior region of premaxilla; and maxilla with posterior wall over the anterior margin of the suborbital fenestra. The postcanine teeth have extensive wear facets, implying an alternate unilateral occlusion and suggesting the presence of both lateral and fore-aft movement of the jaw. These characteristics appear unique within the crocodyliform lineage, showing that feeding diversity was wider than thought in this clade. The phylogenetic relationships of Sphagesaurus huenei are evaluated through a parsimony analysis. Sphagesaurus lies within a monophyletic group formed by Cretaceous and Tertiary taxa traditionally referred as notosuchians and sebecosuchians. Interestingly, several mammal-like tooth morphologies were present in this clade of terrestrial crocodyliforms.

Phenotypic experiments into new pelagic niches in early Danian planktonic foraminifera: aftermath of the K/T boundary event

Abstract In the aftermath of the Cretaceous-Tertiary (K/T) boundary event it appears that unusual phytoplankton blooms were boosted by the abundant nutrient levels left over from a short-lived ‘Strangelove Ocean’ period, which possibly gave rise to the negative-to-positive δ 13 C surface-to-bottom gradient in the Danian. The highly plastic morphology of the planktonic foraminifera at the base of the Palaeocene appeared, most likely, as an adaptive response of functional morphology to the pelagic habitat of the species and its mode of life. The diversification episodes of Danian planktonic foraminifera appear to correspond to times of reduced oceanic mixing, stratified water masses, recovery of the surface water productivity and a progressive expansion of the trophic-resource continuum, with periods of maximum diversity corresponding to widespread oceanic surface-water oligotrophic conditions. These suggest a parallel adaptive trend of feeding strategies caused by the changing availability of nutrients in the water column, with probable increasing reliance in symbiosis under low nutrient conditions later in the Danian. Guembelitria cretacea , one of the few Cretaceous survivors, has been recorded in northeastern Brazil up to the lower part of the P2 Zone. A triserially coiled planktonic morphotype, this species started to diversify immediately after the K/T transition. After only a few thousand years, in the earliest Danian, the first Tertiary taxa (the Guembelitria- Woodringina lineage) appeared; these are microperforate nonspinose forms that were probably restricted to shallow epipelagic waters. There is a trend towards uncoiling of the triserial pattern by progressively changing the positioning of the aperture, without any detectable ecophenotypic preference at first. The first specimens of the Eoglobigerina-Pseudosubbotina, Eoglobigerina-Subbotina and Praemurica lineages, cancellate spinose and non-spinose forms, also evolved in the Danian in a nearly coeval and parallel evolutionary trend from a probably ancestral Hedbergella stock. A revised phylogeny is suggested for the early Palaeocene planktonic foraminifera based on recorded stratigraphic ranges and morphological affinities (chamber arrangement and surface texture ornamentation).