Fossil brains provide evidence of underwater feeding in early seals.

Pinnipeds (seals and related species) use their whiskers to explore their environment and locate their prey. Today they live mostly in marine habitats and are adapted for a highly specialised amphibious lifestyle with their flippers for locomotion and a hydrodynamically streamlined body. The earliest pinnipeds, however, lived on land and in freshwater habitats, much like mustelids today. Here we reconstruct the underwater foraging behaviour of one of these earliest pinnipeds (Potamotherium), focusing in particular on how it used its whiskers (vibrissae). For this purpose, we analyse the coronal gyrus of the brain of 7 fossil and 31 extant carnivorans. This region receives somatosensory input from the head. Our results show that the reliance on whiskers in modern pinnipeds is an ancestral feature that favoured survival of stem pinnipeds in marine habitats. This study provides insights into an impressive ecological transition in carnivoran evolution: from terrestrial to amphibious marine species. Adaptations for underwater foraging were crucial for this transition.

Dwarfism and gigantism drive human-mediated extinctions on islands.

Islands have long been recognized as distinctive evolutionary arenas leading to morphologically divergent species, such as dwarfs and giants. We assessed how body size evolution in island mammals may have exacerbated their vulnerability, as well as how human arrival has contributed to their past and ongoing extinctions, by integrating data on 1231 extant and 350 extinct species from islands and paleo islands worldwide spanning the past 23 million years. We found that the likelihood of extinction and of endangerment are highest in the most extreme island dwarfs and giants. Extinction risk of insular mammals was compounded by the arrival of modern humans, which accelerated extinction rates more than 10-fold, resulting in an almost complete demise of these iconic marvels of island evolution.

The macroevolutionary impact of recent and imminent mammal extinctions on Madagascar.

Many of Madagascar's unique species are threatened with extinction. However, the severity of recent and potential extinctions in a global evolutionary context is unquantified. Here, we compile a phylogenetic dataset for the complete non-marine mammalian biota of Madagascar and estimate natural rates of extinction, colonization, and speciation. We measure how long it would take to restore Madagascar's mammalian biodiversity under these rates, the "evolutionary return time" (ERT). At the time of human arrival there were approximately 250 species of mammals on Madagascar, resulting from 33 colonisation events (28 by bats), but at least 30 of these species have gone extinct since then. We show that the loss of currently threatened species would have a much deeper long-term impact than all the extinctions since human arrival. A return from current to pre-human diversity would take 1.6 million years (Myr) for bats, and 2.9 Myr for non-volant mammals. However, if species currently classified as threatened go extinct, the ERT rises to 2.9 Myr for bats and 23 Myr for non-volant mammals. Our results suggest that an extinction wave with deep evolutionary impact is imminent on Madagascar unless immediate conservation actions are taken.

Breaking the constraint on the number of cervical vertebrae in mammals: On homeotic transformations in lorises and pottos.

Mammals almost always have seven cervical vertebrae. The strong evolutionary constraint on changes in this number has been broken in sloths and manatees. We have proposed that the extremely low activity and metabolic rates of these species relax the stabilizing selection against changes in the cervical count. Our hypothesis is that strong stabilizing selection in other mammals is largely indirect and due to associated pleiotropic effects, including juvenile cancers. Additional direct selection can occur due to biomechanical problems (thoracic outlet syndrome). Low metabolic and activity rates are thought to diminish these direct and indirect effects. To test this hypothesis within the primates, we have compared the number of cervical vertebrae of three lorisid species with particularly low activity and metabolic rates with those of more active primate species, including with their phylogenetically closest active relatives, the galagids (bushbabies). In support of our hypothesis, we found that 37.6% of the lorisid specimens had an abnormal cervical count, which is a higher percentage than in the other nine primate families, in which the incidence varied from zero to 2.2%. We conclude that our data support the importance of internal selection in constraining evolvability and of a relaxed stabilizing selection for increasing evolvability. Additionally, we discuss that there is no support for a role of the muscularized diaphragm in the evolutionary constraint.

Global raster dataset on historical coastline positions and shelf sea extents since the Last Glacial Maximum.

Motivation: Historical changes in sea level caused shifting coastlines that affected the distribution and evolution of marine and terrestrial biota. At the onset of the Last Glacial Maximum (LGM) 26 ka, sea levels were >130 m lower than at present, resulting in seaward-shifted coastlines and shallow shelf seas, with emerging land bridges leading to the isolation of marine biota and the connection of land-bridge islands to the continents. At the end of the last ice age, sea levels started to rise at unprecedented rates, leading to coastal retreat, drowning of land bridges and contraction of island areas. Although a growing number of studies take historical coastline dynamics into consideration, they are mostly based on past global sea-level stands and present-day water depths and neglect the influence of global geophysical changes on historical coastline positions. Here, we present a novel geophysically corrected global historical coastline position raster for the period from 26 ka to the present. This coastline raster allows, for the first time, calculation of global and regional coastline retreat rates and land loss rates. Additionally, we produced, per time step, 53 shelf sea rasters to present shelf sea positions and to calculate the shelf sea expansion rates. These metrics are essential to assess the role of isolation and connectivity in shaping marine and insular biodiversity patterns and evolutionary signatures within species and species assemblages. Main types of variables contained: The coastline age raster contains cells with ages in thousands of years before present (bp), representing the time since the coastline was positioned in the raster cells, for the period between 26 ka and the present. A total of 53 shelf sea rasters (sea levels <140 m) are presented, showing the extent of land (1), shelf sea (0) and deep sea (NULL) per time step of 0.5 kyr from 26 ka to the present. Spatial location and grain: The coastline age raster and shelf sea rasters have a global representation. The spatial resolution is scaled to 120 arcsec (0.333° × 0.333°), implying cells of c. 3,704 m around the equator, 3,207 m around the tropics (±30°) and 1,853 m in the temperate zone (±60°). Time period and temporal resolution: The coastline age raster shows the age of coastline positions since the onset of the LGM 26 ka, with time steps of 0.5 kyr. The 53 shelf sea rasters show, for each time step of 0.5 kyr, the position of the shelf seas (seas shallower than 140 m) and the extent of land. Level of measurement: Both the coastline age raster and the 53 shelf sea rasters are provided as TIFF files with spatial reference system WGS84 (SRID 4326). The values of the coastline age raster per grid cell correspond to the most recent coastline position (in steps of 0.5 kyr). Values range from 0 (0 ka, i.e., present day) to 260 (26 ka) in bins of 5 (0.5 kyr). A value of "no data" is ascribed to pixels that have remained below sea level since 26 ka. Software format: All data processing was done using the R programming language.

No evidence for widespread island extinctions after Pleistocene hominin arrival.

The arrival of modern humans into previously unoccupied island ecosystems is closely linked to widespread extinction, and a key reason cited for Pleistocene megafauna extinction is anthropogenic overhunting. A common assumption based on late Holocene records is that humans always negatively impact insular biotas, which requires an extrapolation of recent human behavior and technology into the archaeological past. Hominins have been on islands since at least the early Pleistocene and Homo sapiens for at least 50 thousand y (ka). Over such lengthy intervals it is scarcely surprising that significant evolutionary, behavioral, and cultural changes occurred. However, the deep-time link between human arrival and island extinctions has never been explored globally. Here, we examine archaeological and paleontological records of all Pleistocene islands with a documented hominin presence to examine whether humans have always been destructive agents. We show that extinctions at a global level cannot be associated with Pleistocene hominin arrival based on current data and are difficult to disentangle from records of environmental change. It is not until the Holocene that large-scale changes in technology, dispersal, demography, and human behavior visibly affect island ecosystems. The extinction acceleration we are currently experiencing is thus not inherent but rather part of a more recent cultural complex.

Size matters: micro-evolution in Polynesian rats highlights body size changes as initial stage in evolution.

Microevolutionary patterns in populations of introduced rodent species have often been the focus of analytic studies for their potential relevance to understanding vertebrate evolution. The Polynesian rat (Rattus exulans) is an excellent proxy species because of its wide geographic and temporal distribution: its native and introduced combined range spans half the globe and it has been living for at least seven centuries wherever it was introduced. The objective of this study was to assess the effects of long-term isolation (insularity; up to 4,000 years) and geographic variables on skull shape variation using geometric morphometrics. A sample of 513 specimens from 103 islands and four mainland areas was analysed. This study, to my knowledge the first to extensively sample introduced rats, analysed 59 two-dimensional landmarks on the skull. Landmarks were obtained in three separate aspects (dorsal, lateral, ventral skull view). The coordinate data were then subjected to a multivariate ordination analysis (principal components analysis, or PCA), multivariate regressions, and a canonical variates analysis (CVA). Three measures of disparity were evaluated for each view. The results show that introduced Polynesian rats evolve skull shapes that conform to the general mammalian interspecific pattern of cranial evolutionary allometry (CREA), with proportionally longer snouts in larger specimens. In addition, larger skulls are more tubular in shape than the smaller skulls, which are more balloon-shaped with a rounder and wider braincase relative to those of large skulls. This difference is also observed between the sexes (sexual dimorphism), due to the slightly larger average male size. Large, tubular skulls with long snouts are typical for Polynesia and Remote Oceania, where no native mammals occur. The greater disparity of Polynesian rats on mammal species-poor islands ('exulans-only' region) provides further insight into how diversity may affect diversification through ecological release from predators and competitors.

Effect of isolation on coat colour polymorphism of Polynesian rats in Island Southeast Asia and the Pacific.

Populations of vertebrate species introduced onto islands regularly develop similar phenotypic changes, e.g., larger or smaller body size, shortened limbs, duller coats, as well as behavioural changes such as increased tameness and reduced flight-initiation distance. These changes overlap in part with those associated with the 'domestication syndrome', especially tameness and changes in coat patterns, and might indicate a similar neural crest involvement in the concurrent development of multiple phenotypic traits. Here I examine long-term data on free-living populations of wild Polynesian rats from seven mainland countries and 117 islands (n = 3,034), covering the species' native and introduced range. Mainland populations showed no aberrant coat patterns, with the exception of one albino, whereas aberrant coat patterns were found in 12 island populations. Observed coat colour polymorphisms consisted of leucistic (including singular white patches), melanistic (darkly pigmented) and piebald (mixed) coat patterns. After isolation for at least seven centuries, wild Polynesian rat populations on islands seem to exhibit a trend towards a higher incidence of aberrant coat patterns. These phenotypic changes are here explained as a neutral, non-adaptive process, likely part of the 'domestication syndrome' (via the commensal pathway of domestication), in combination with genetic drift, little or no gene flow between the islands and/or the mainland and a relaxed selection (as a result of the weakening or removal of competitor/predator pressure) under commensality.

Paradise lost: Evidence for a devastating metabolic bone disease in an insular Pleistocene deer.

PURPOSE: This communication reports skeletal pathology in a Pleistocene endemic deer from the Mavromouri caves of Crete. MATERIALS: 287 bones and bone fragments from Mavromouri caves are compared to 2986 bones from Liko Cave. METHODS: Bones were evaluated macroscopically, and measurements were made of morphometric characteristics of limb long bones. Representative bone specimens were examined radiographically and histologically. RESULTS: Macroscopic hallmarks were loss of bone mass and increased porosity. The long bones were brittle, some of them having thin cortices, and others reduction of medullary cavities that contain dense Haversian tissue. The flat bones were spongy and fragile. Erosions of the metaphyses and articular surfaces were noted. Histological findings included: sub-periosteal resorption; loss of lamellar bone; enlargement of vascular canals; and remodeling of cortical bone. Two types of fibrous osteodystrophy were recognized in skeletal remains, subostotic and hyperostotic. CONCLUSIONS: The deer of Mavromouri caves were affected by severe metabolic bone disease, likely nutritional secondary hyperparathyroidism. We hypothesize a multifactorial cause, including overgrazing, flora senescence, soil mineral deficiencies, and a prolonged period of climate extremes, degrading the Cretan deer habitat. VALUE: This is the first evidence of a metabolic bone disease causing this level of destructive pathology in an insular fossil deer. LIMITATIONS: The lack of absolute chronometric dates for the site limits potential linking with the prevailing environmental conditions. SUGGESTIONS FOR FURTHER RESEARCH: Investigation of similar skeletal pathologies at other islands or isolated habitats is advised.

Changing Invaders: trends of gigantism in insular introduced rats.

The degree and direction of morphological change in invasive species with a long history of introduction is insufficiently known for a larger scale than the archipelago or island group. Here, I analyse data for 105 island populations of Polynesian rats, Rattus exulans, covering the entirety of Oceania and Wallacea to test whether body size differs in insular populations and if so what biotic and abiotic features are correlated with it. All insular populations of this rat, except one, exhibit body sizes up to twice the size of their mainland conspecifics. Body size of insular populations is positively correlated with latitude, consistent with thermoregulatory predictions based on Bergmann's rule. Body size is negatively correlated with number of co-occurring mammalian species, confirming an ecological hypothesis of the island rule. The largest rats are found in the temperate zone of New Zealand as well as on mammalian species-poor islands of Polynesia and the Solomon Islands. Carnivory in the form of predation on nesting seabird colonies seems to promote 1.4- to 1.9-fold body size increases.

High incidence of cervical ribs indicates vulnerable condition in Late Pleistocene woolly rhinoceroses.

Mammals as a rule have seven cervical vertebrae, a number that remains remarkably constant. Changes of this number are associated with major congenital abnormalities (pleiotropic effects) that are, at least in humans, strongly selected against. Recently, it was found that Late Pleistocene mammoths (Mammuthus primigenius) from the North Sea have an unusually high incidence of abnormal cervical vertebral numbers, approximately ten times higher than that of extant elephants. Abnormal numbers were due to the presence of large cervical ribs on the seventh vertebra, indicating a homeotic change from a cervical rib-less vertebra into a thoracic rib-bearing vertebra. The high incidence of cervical ribs indicates a vulnerable condition and is thought to be due to inbreeding and adverse conditions that may have impacted early pregnancies in declining populations. In this study we investigated the incidence of cervical ribs in another extinct Late Pleistocene megaherbivore from the North Sea and the Netherlands, the woolly rhinoceros (Coelodonta antiquitatis). We show that the incidence of abnormal cervical vertebral numbers in the woolly rhinoceros is unusually high for mammals (15,6%, n = 32) and much higher than in extant Rhinoceratidae (0%, n = 56). This indicates that woolly rhinoceros lived under vulnerable conditions, just like woolly mammoths. The vulnerable condition may well have contributed to their eventual extinction.

Mammalian bone palaeohistology: a survey and new data with emphasis on island forms.

The interest in mammalian palaeohistology has increased dramatically in the last two decades. Starting in 1849 via descriptive approaches, it has been demonstrated that bone tissue and vascularisation types correlate with several biological variables such as ontogenetic stage, growth rate, and ecology. Mammalian bone displays a large variety of bone tissues and vascularisation patterns reaching from lamellar or parallel-fibred to fibrolamellar or woven-fibred bone, depending on taxon and individual age. Here we systematically review the knowledge and methods on cynodont and mammalian bone microstructure as well as palaeohistology and discuss potential future research fields and techniques. We present new data on the bone microstructure of two extant marsupial species and of several extinct continental and island placental mammals. Extant marsupials display mainly parallel-fibred primary bone with radial and oblique but mainly longitudinal vascular canals. Three juvenile specimens of the dwarf island hippopotamid Hippopotamus minor from the Late Pleistocene of Cyprus show reticular to plexiform fibrolamellar bone. The island murid Mikrotia magna from the Late Miocene of Gargano, Italy displays parallel-fibred primary bone with reticular vascularisation and strong remodelling in the middle part of the cortex. Leithia sp., the dormouse from the Pleistocene of Sicily, is characterised by a primary bone cortex consisting of lamellar bone and a high amount of compact coarse cancellous bone. The bone cortex of the fossil continental lagomorph Prolagus oeningensis and three fossil species of insular Prolagus displays mainly parallel-fibred primary bone and reticular, radial as well as longitudinal vascularisation. Typical for large mammals, secondary bone in the giant rhinocerotoid Paraceratherium sp. from the Late Oligocene of Turkey is represented by dense Haversian bone. The skeletochronological features of Sinomegaceros yabei, a large-sized deer from the Pleistocene of Japan closely related to Megaloceros, indicate a high growth rate. These examples and the synthesis of existing data show the potential of bone microstructure to reveal essential information on life history evolution. The bone tissue and the skeletochronological data of the sampled island species suggest the presence of various modes of bone histological modification and mammalian life history evolution on islands to depend on factors of island evolution such as island size, distance from mainland, climate, phylogeny, and time of evolution.

Systematic revision of the family Hoplitomerycidae Leinders, 1984 (Artiodactyla: Cervoidea), with the description of a new genus and four new species.

Six species of the cervoid genus Hoplitomeryx are currently recognized from the Late Miocene sites of Gargano and Scontrone, in Italy: H. matthei Leinders, 1984, H. apruthiensis Mazza & Rustioni, 2011, H. apulicus Mazza & Rustioni, 2011, H. falcidens Mazza & Rustioni, 2011, H. magnus Mazza & Rustioni, 2011, and H. minutus Mazza & Rustioni, 2011. These species are interpreted as members of an anagenetic series in these two localities, which are considered as part of the same bioprovince but with different geological ages. Comparative analysis of postcranial, dental, and cranial material from Hoplitomerycidae resulted in the reinterpretation of this current taxonomic arrangement. Two distinct genera can be distinguished. The new genus Scontromeryx is restricted to Scontrone (Early Tortonian) and is characterised by the presence of second upper and lower premolars and the absence of a nasal (median) horn. Hoplitomeryx is restricted to Gargano (Middle and/or Late Tortonian), and is characterized by the loss of the second premolar and presence of a nasal horn. Both genera are characterized by orbital appendages in some species, but the morphology of these appendages differs between the genera. Six species can be recognized for Scontromeryx gen. n.: S. minutus (type species), S. falcidens, S. apulicus, S. apruthiensis, S. magnus (new combinations) and the newly described S. mazzai sp. n.. Hoplitomeryx is represented by the H. matthei (type species) and 3 newly described species H. devosi sp. n., H. macpheei sp. n. and H. kriegsmani sp. n.. These two multispecies assemblages are best explained as independent adaptive radiations with the two genera as sister taxa. There is no evidence that the two localities were connected during the Late Miocene.

Parallel patterns and trends in functional structures in extinct island mammals.

Endemic mammalian species on islands are generally known to have followed a different evolutionary pathway than their mainland relatives. General patterns, such as body size trends, have been described regularly. However, most island mammal species are unique and each of them is adapted to a specific local niche as part of an equally specific ecological assemblage. Therefore, comparing island species across taxa, islands and time is inherently dangerous without understanding the adaptational value of the studied feature in the compared taxa and without taking the ecological setting of the taxa into account. In this contribution, general and recurring patterns are described per taxon. Some features, like body mass change and sturdy limbs, are relatively general, whereas most features, like bone fusions and change of orbital axis, occur only in a very few taxa. Some features are even contradictory, such as brain size and degree of hypsodonty, with each taxon having its own particular design. In conclusion, general patterns are more often than not just trends and need to be applied with caution.