Glacial refugia of mammals in Europe: evidence from fossil records

• 1 Glacial refugia were core areas for the survival of temperate species during unfavourable environmental conditions and were the sources of postglacial recolonizations. Unfortunately, the locations of glacial refugia of animals and plants are usually described by models, without reference to facts about real geographical ranges at that time.
• 2 Careful consideration of the faunal assemblages of archaeological sites from the Younger Palaeolithic, which are precisely dated to the Last Glacial Maximum (LGM), gives indications about the distribution of species during the LGM (23 000–16 000 bp ) and provides evidence for the locations of glacial refugia for mammalian species in Europe.
• 3 In Europe, 47 LGM sites, dating from 23 000 to 16 000 bp and containing typical temperate mammal species, have been described. The geographical range of these archaeological sites clearly shows a distribution which differs from the hypothesized traditional refuge areas of the temperate fauna. A considerable number of sites situated in the Dordogne in south‐western France and the Carpathian region contain records of red deer Cervus elaphus, roe deer Capreolus capreolus, wild boar Sus scrofa and red fox Vulpes vulpes.
• 4 The faunal composition of the majority of the evaluated Palaeolithic sites in the southern European peninsulas (with the exception of Greece), as well as France and the Carpathian region, indicates the co‐occurrence of these temperate species with cold‐adapted faunal elements such as mammoth Mammuthus primigenius and/or reindeer Rangifer tarandus.
• 5 The survival of species in Central European refugia would have significant consequences for phylogeography and would be revealed by the dominant distribution of haplotypes, originating from this region. A Carpathian refuge could also be the reason for the very early records of small mammals or mustelids from the Late‐Glacial or Interstadials before the LGM in regions like southern Germany.

     

The fossil record of North American mammals: evidence for a Paleocene evolutionary radiation

Paleontologists long have argued that the most important evolutionary radiation of mammals occurred during the early Cenozoic, if not that all eutherians originated from a single common post-Cretaceous ancestor. Nonetheless, several recent molecular analyses claim to show that because several interordinal splits occurred during the Cretaceous, a major therian radiation was then underway. This claim conflicts with statistical evidence from the well-sampled latest Cretaceous and Cenozoic North American fossil record. Paleofaunal data confirm that there were fewer mammalian species during the latest Cretaceous than during any interval of the Cenozoic, and that a massive diversification took place during the early Paleocene, immediately after a mass extinction. Measurement data show that Cretaceous mammals were on average small and occupied a narrow range of body sizes; after the Cretaceous-Tertiary mass extinction, there was a rapid and permanent shift in the mean. The fact that there was an early Cenozoic mammalian radiation is entirely compatible with the existence of a few Cretaceous splits among modern mammal lineages.     

The tri-segmented limbs of therian mammals: kinematics, dynamics, and self-stabilization--a review

The evolution of therian mammals is to a large degree marked by changes in their motion systems. One of the decisive transitions has been from the sprawled, bi-segmented to the parasagittal, tri-segmented limb. Here, we review aspects of the tri-segmented limb in locomotion which have been elucidated in our research groups in the last 10 years. First, we report the kinematics of the tri-segmented therian limb from mouse to elephant in order to explore general principles of the therian limb configuration and locomotion. Torques will be reported from a previous paper (Witte et al., 2002. J Exp Biol 205:1339-1353) for a better understanding of the anti-gravity work of all limb joints. The stability of a limb in z-configuration will be explained and its advantage with respect to other potential solutions from modeling will be discussed. Finally, we describe how the emerging concept of self-stability can be explained for a tri-segmented leg template and how it affects the design of the musculoskeletal system and the operation of legs during locomotion. While locomotion has been considered as mainly a control problem in various disciplines, we stress the necessity to reduce control as much as possible. Central control can be cheap if the limbs are "intelligent" by means of their design. Gravity-induced movements and self-stability seem to be energy-saving mechanisms.     

Body size of insular carnivores: evidence from the fossil record

Aim Our goals here are to: (1) assess the generality of one aspect of the island rule – the progressive trend towards decrease in size in larger species – for fossil carnivores on islands; (2) offer causal explanations for this pattern and deviations from it – as far as fossil carnivores are concerned; and (3) estimate the speed of this trend. Location Oceanic and oceanic‐like islands world‐wide. Methods Body size estimates of fossil insular carnivores and of their phylogenetically closest mainland relative were obtained from our own data and the published literature. Our dataset consisted of 18 species from nine islands world‐wide. These data were used to test whether the body size of fossil insular carnivores varies as a function of body size of the mainland species in combination with characteristics of the island ecosystem. Results Dwarfism was observed in two canid species. Moderate decrease in body mass was observed in one hyena species. Gigantism was observed in one otter species. Moderate body mass increase was observed in two otter species, one galictine mustelid and perhaps one canid. Negligible or no change in body mass at all was observed in five otter species, three galictine mustelids and one genet. Size changes in teeth do not lag behind in comparison to skeletal elements in the dwarfed canids. The evolutionary speed of dwarfism in a canid lineage is low. Main conclusions Size change in fossil terrestrial insular carnivores was constrained by certain ecological conditions, especially the availability of prey of appropriate body size. When such alternative prey was not available, the carnivores retained their mainland size. The impact of competitive carnivores seems negligible. The case of (semi‐)aquatic carnivores is much less clear. The species that maintained their ancestral body mass may have changed their diet, as is evidenced by their dentition. Among the otters, one case of significant size increase was observed, perhaps best explained as being due to it entering the niche of an obligate aquatic otter. Dwarfism was not observed in otters. The island rule seems to apply to fossil carnivores, but with exceptions. The dependency of the island rule on resource availability is emphasized by the present study.     

The shape of pterosaur evolution: evidence from the fossil record

Abstract Although pterosaurs are a well‐known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon‐specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased – this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.     

Comparative patterns of postcranial ontogeny in therian mammals: an analysis of relative timing of ossification events

Data on the relative sequence of ossification of postcranial elements for eight therian mammals (Myotis lucifugus, Homo sapiens, Rattus norvegicus, Mus musculus, Mesocricetus auratus, Cavia porcellus, Didelphis albiventris, and Sminthopsis macroura) and three outgroups (Chelydra serpentina, Alligator mississippiensis, and Lacerta vivipara) were taken from the literature. For each species, a matrix was constructed in which the relative timing of the onset of ossification in 24 elements was summarized. This resulted in 276 event pairs (characters) for each species. Thirty-three (33.3)% of the characters examined are uniform across all taxa, 16.3% are variable but uninformative in the phylogeny, and 50.4% potentially deliver diagnostic features for clades of two or more taxa. In all species examined, the clavicle is the first bone to appear. Placentalia is not unequivocally diagnosed by the state of any event pair, while Marsupialia has the largest amount of autapomorphies with 18. The acceleration in the timing of ossification of the scapula in relation to the hindlimb in marsupials is most probably causally correlated to movements after birth and during early phases of pre-weaning life. Marsupials are almost unique among amniotes in that the earliest onset of ossification of at least one element among carpals and among tarsals is simultaneous. Three parsimony analyses, each one with a different reptilian taxon as outgroup, were performed using event pairs as characters. In all cases, results were incongruent with the phylogeny of the studied taxa.     

Metabolic profile of the perivertebral muscles in small therian mammals: implications for the evolution of the mammalian trunk musculature

In order to gain a better understanding of the ancestral properties of the perivertebral muscles of mammals, this study investigated the fiber type composition of these muscles in six small, extant therians (two metatherians and four eutherians) similar in body shape to early mammals. Despite a few species-specific differences, the investigated species were very similar in their overall distribution of fiber types indicating similar functional demands on the back muscles in mammals of this body size and shape. Deep and short, mono- or multisegmental muscles (i.e., mm. interspinales, intermammillares, rotatores et intertransversarii) consistently showed the highest percentage of slow, oxidative fibers implying a function as local stabilizers of the vertebral column. Superficial and large, polysegmental muscles (i.e., mm. multifidus, sacrospinalis, iliopsoas et psoas minor) were predominantly composed of fast, glycolytic fibers suggesting they function to both globally stabilize and mobilize the spine during rapid non-locomotor and locomotor activities. Some muscles contained striking accumulations of oxidative fibers in specific regions (mm. longissimus et quadratus lumborum). These regions are hypothesized to function independently from the rest of the muscle belly and may be comparable in their functionality to regionalized limb muscles. The deep, central oxidative region in the m. longissimus lumborum appears to be a general feature of mammals and likely serves a proprioceptive function to control the postural equilibrium of the pelvic girdle and lumbar spine. The potential functions of the m. quadratus lumborum during ventilation and ventral stabilization of the vertebral column are discussed. Because representatives of the stem lineage of mammals were comparable in their body proportions and probably also locomotor parameters to the species investigated here, I suggest that the described fiber type distribution is representative of the ancestral condition in mammals. The origin of mammals was associated with a substantial enlargement of the epaxial muscles and the addition of subvertebral muscle mass. Because this novel muscle mass is mainly composed of fast, glycolytic fibers in extant species, it is plausible that these changes were associated with the evolution of increased sagittal mobility in the posterior trunk region in the therapsid ancestors of mammals. The caudally increasing role of sagittal bending in body propulsion is consistent with the overall increase in the percentage of glycolytic fibers in the cranio-caudal direction. The evolution of mammals was also associated with a loss of ribs in the posterior region of the trunk. This loss of ribs is thought to have decreased the stability of the posterior trunk, which may explain the observed greater oxidative capacity of the caudal local stabilizers. The increased need for postural feedback in the more mobile lumbar region may also explain the evolution of the proprioceptive system in the m. longissimus lumborum. Furthermore, the anatomical subdivision of the transversospinal muscle into several smaller muscle entities is suggested to facilitate their functional specialization.     

Paleoequatorial rain forest of western India during the EECO: evidence from Uvaria L. fossil and its geological distribution pattern

During the early Eocene, Rajasthan was positioned near the equator and had a warm and humid tropical climate dominated by tropical rainforests like the present-day equatorial forests of South India. Many of the plants retrieved as fossils from Rajasthan are growing there as refugee. This study further strengthens this view as it reports a new species of Uvaria L. from the early Eocene sediments of Bikaner (Rajasthan) showing its best resemblance with the extant U. zeylanica Deless. ex DC., which is presently growing in the evergreen forests of South India and Sri Lanka. The genus is thought to have originated in Africa, and the present finding gives an idea about its geologic distribution in Asia and Australasia via India relying on ‘stepping stone’ hypothesis during the Early Eocene Climatic Optimum (EECO) when climatic conditions were favourable for the luxuriant growth of tropical vegetation. A general cooling trend after EECO and change in the configuration of land and sea affected the climate on the regional scale causing total devastation of tropical evergreen forests that existed in western India during the depositional time; this change is ultimately responsible for creating dry and desertic conditions prevailing in the area at present.     

Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber

We describe a new species of extinct riodinid butterfly, Voltinia dramba, from Oligo-Miocene Dominican amber (15-25 Myr ago). This appears to be the first butterfly to be taxonomically described from amber, and the first adult riodinid fossil. The series of five specimens represents probably the best-preserved fossil record for any lepidopteran. The phenomenon of extant Voltinia females ovipositing on arboreal epiphytes probably explains the discovery of multiple female V. dramba specimens in amber. Voltinia dramba appears to be one of many extinct butterfly species on Hispaniola. The northwestern Mexican distribution of the explicitly hypothesized sister species, the extant V. danforthi, supports the hypothesis that V. dramba reached Hispaniola by the 'proto-Greater Antillean arc', dating the divergence of V. dramba and V. danforthi to 40-50 Myr ago. This date is contemporaneous with the oldest known butterfly fossils, and implies a more ancient date of origin for many of the higher-level butterfly taxa than is often conceded.     

The first 50Myr of dinosaur evolution: macroevolutionary pattern and morphological disparity

The evolutionary radiation of dinosaurs in the Late Triassic and Early Jurassic was a pivotal event in the Earth's history but is poorly understood, as previous studies have focused on vague driving mechanisms and have not untangled different macroevolutionary components (origination, diversity, abundance and disparity). We calculate the morphological disparity (morphospace occupation) of dinosaurs throughout the Late Triassic and Early Jurassic and present new measures of taxonomic diversity. Crurotarsan archosaurs, the primary dinosaur 'competitors', were significantly more disparate than dinosaurs throughout the Triassic, but underwent a devastating extinction at the Triassic-Jurassic boundary. However, dinosaur disparity showed only a slight non-significant increase after this event, arguing against the hypothesis of ecological release-driven morphospace expansion in the Early Jurassic. Instead, the main jump in dinosaur disparity occurred between the Carnian and Norian stages of the Triassic. Conversely, dinosaur diversity shows a steady increase over this time, and measures of diversification and faunal abundance indicate that the Early Jurassic was a key episode in dinosaur evolution. Thus, different aspects of the dinosaur radiation (diversity, disparity and abundance) were decoupled, and the overall macroevolutionary pattern of the first 50Myr of dinosaur evolution is more complex than often considered.     

Molecular cloning and characterization of the major allergen Myr p II from the venom of the jumper ant Myrmecia pilosula: Myr p I and Myr p II share a common protein leader sequence

A major allergen Myr p II of the Australian jumper ant Myrmecia pilosula has been cloned, immunocharacterized and nucleotide sequenced. An open reading frame of 225 bases was identified and found to encode a deduced amino acid sequence of 75 residues which contained a typical hydrophobic peptide leader sequence. Expressed fusion proteins of Myr p II in both phage and plasmid vectors bind high levels of ant venom-specific IgE and the expressed clones are recognised by 35% of ant venom-allergic individuals. IgE antibodies that recognise the expressed clone have been shown to recognise IgE-binding bands in blots of native venom after separation by SDS-PAGE. The amino acid sequence of Myr p II shares close structural homology with the other major jumper ant allergen Myr p I, differing by only three amino acids in the first 47 residues of both sequences. However, N-terminal analysis of IgE-binding bands derived from Tricine-SDS-PAGE gel blots indicates that both Myr p I and Myr p II undergo extensive post-translational proteolytic processing to unique peptides of 45 and 27 residues, respectively.     

Mediterranean landscape changes: evidence from old postcards

Recent, rapid and often underestimated landscape changes have occurred over large areas in Mediterranean Europe. They are the result of major rural depopulation. Old photographs of landscapes taken at the beginning of the twentieth century (i.e. old postcards) and present-day photographs taken at the same places were compared in a 2500-km² area of southern France. Vegetation changes were analysed using transition matrices. During the 80-year study period, land uses and vegetation changed dramatically. Woodland cover and tree height increased; but in contrast, the extent of cropped lands and rangelands decreased. Forest spread was heterogeneous, depending on initial composition of the vegetation, and locally dominant ecological and socio-economic conditions. Our data show that a Mediterranean forest can re-establish under humid climatic conditions and spread within a century, despite severe prior exploitation over several decades. These dramatic changes are liable to have biological and ecological consequences (e.g. spread of woodland species, threat against open habitat species, fire regime modification, deterioration in water resources), some of them being already perceptible.     

Fossil Ericaceae from New Zealand: Deconstructing the use of fossil evidence in historical biogeography

The Australasian Ericaceae epitomize many problems in understanding the biogeography of the southern hemisphere, especially the relative contributions of Gondwanan vicariance and dispersal. Late Cretaceous fossil pollen of the family suggests extreme antiquity of the group in Australasia, but recent phylogenetic evidence suggests much younger histories for most of the groups in that region. This paper documents two new species of latest Oligocene-Early Miocene macrofossils of Ericaceae from New Zealand. Cyathodophyllum novae-zelandiae G.J.Jord. & Bannister gen. and sp. nov. is the oldest record of the tribe Styphelieae, but is of a clade now extinct in New Zealand, possibly related to the Tasmanian genus Cyathodes. Richeaphyllum waimumuensis G.J.Jord. & Bannister sp. nov. is a member of Richeeae, but it is ambiguous as to whether it is a member of the impressive modern New Zealand radiation in Dracophyllum. These fossils emphasize the fact that at least some of the fossil pollen of Ericaceae may have been derived from extinct lineages and therefore should not be used as evidence for the antiquity of any modern New Zealand clade of Ericaceae. New fossils and/or detailed analysis of fossil and extant pollen may help resolve such uncertainty.