Pelagic neonatal fossils support viviparity and precocial life history of Cretaceous mosasaurs

Mosasaurs were large marine squamates that inhabited all of the world's oceans during the Late Cretaceous. Their success as apex predators has been attributed to their rapid acquisition of aquatic adaptations, which allowed them to become fully pelagic. However, little is known about the breeding biology of derived, flipper‐bearing mosasaurs, as the record of neonatal mosasaur fossils is extremely sparse. Here, we report on the fragmentary cranial remains of two neonatal mosasaurs from the Niobrara Formation, referred to Clidastes sp. Comparison with other preliminary reports of neonatal mosasaurs reveals that these specimens are among the smallest individuals ever found and certainly represent the smallest known Clidastes specimens. The recovery of these extremely young specimens from a pelagic setting indicates that even neonatal mosasaurs occupied open oceanic habitats and were likely born in this setting. These data shed new light on the ecology of neonatal mosasaurs and illustrate the degree to which size‐related taphonomic and collection biases have influenced our understanding of the early life history of these iconic marine reptiles.     

Unusual histology and morphology of the ribs of mosasaurs (Squamata)

We report the presence of two previously unrecognized features in the dorsal ribs of mosasaurs: first, the presence of extremely dense, pervasive extrinsic fibres (anchoring soft tissue to bone, sometimes called Sharpey's fibres); and second, high intraspecific variation in costal bone compactness. Extensive extrinsic fibres are developed in the dorsal ribs of the mosasaurs Tylosaurus proriger and Eonatator sternbergi. The dorsal ribs of these mosasaurs are also characterized by a longitudinally ridged texture that almost completely covers the bone. Pervasive extrinsic fibres and ridged textures are absent in the mosasaur Selmasaurus russelli as well as the dorsal ribs of extant semi‐aquatic reptiles (e.g. crocodyliforms) and mosasaurs' close extant relatives and analogues (e.g. snakes and varanids). Similar ridged textures characterize the dorsal ribs of several other mosasaur taxa but are developed to a lesser extent (e.g. Mosasaurus, Clidastes, Platecarpus and Ectenosaurus), but in no other taxa have pervasive extrinsic fibres been reported. We interpret these osteohistological features in T. proriger and E. sternbergi as evidence of tendinous attachment of extensive and highly differentiated axial musculature capable of producing great stresses, most likely related to stabilization of the trunk relative to contralateral movements of the tail during carangiform locomotion. We also report the compactness indices (percentage of space occupied by bone rather than cavities) for these large mosasaur ribs, which are much higher than previously reported. This suggests high intraspecific variation in bone compactness that complicates its use in reconstructing mosasaur palaeoecology.     

The effects of food and density on the nocturnal behaviour ofArion ater andAriolimax columbianus (Pulmonata: Stylommatophora)

The effects of food supply and population density on the nocturnal behaviour of Arion ater and Ariolimax columbianus were investigated. Density did not significantly affect A. ater's level of activity or short-term movement, resting, or feeding. A. columbianus was more active and moved, rested, and fed more frequently when slug density was high. A. ater foraged and rested more often, but fed less when good food was unavailable. Ariolimax's only response to the food regime was to feed more when good food was available. Seasonal changes in the level of activity and behaviour of Arion were evident, whereas Ariolimax's activity and behavioural repertoire were not similarly affected. Arion ater's nightly activity appeared to be mainly food oriented, while Ariolimax columbianus seemed most responsive to slug density during its nocturnal activity periods.     

The Skull of Stephanomys and a Review of Malpaisomys Relationships (Rodentia: Muridae): Taxonomic Incongruence in Murids

The cranial morphology of the extinct murid genus Stephanomys, previously known only by dental remains, is described here on the basis of partial skulls of three species of Pliocene age. Important cranial characters of the genus are a robust rostrum, a high zygoma, a wide zygomatic arch, a narrow interorbit, a large orbit, and an optic foramen in the backward position. In addition to some dental characters, Stephanomys shares most of these cranial traits with the extinct Malpaisomys from the Canary Islands. Some of these traits may be linked to the development of large eyes and life in a rocky environment. The peculiar dental pattern of Stephanomys (stephanodonty) is also present in some recent murids (Oenomys and Thamnomys) having a different skull morphology. A comparison with nine other extant genera of murids verified the relationship among Malpaisomys, Stephanomys, and Acomys, supporting our previous conclusion. Phenetic and cladistic analyses of 17 cranial and 23 dental characters show that skull morphology is phylogenetically informative but highly convergent and incongruent with other partial evidence based on dental and biochemical characters. The combined analyses of skull and teeth illustrate a case of mosaic evolution in murids.     

水位波动对洞庭湖越冬小天鹅家域的影响

洞庭湖是小天鹅在我国的重要越冬地之一,为探讨洞庭湖水位变动对越冬小天鹅活动的影响,本研究于2014-2015年利用卫星跟踪技术对洞庭湖自然保护区18只越冬小天鹅活动轨迹进行跟踪,采用最小凸多边形（MCP）和核心密度估计（KDE）方法估算了越冬中、后期不同水位条件下小天鹅的家域。同时,基于MODIS遥感数据提取洞庭湖水体面积、计算水体淹没时间指数（STI）,进而开展了水位与家域关系、栖息地选择模式等方面的分析和探讨。结果表明：水位对小天鹅家域起制约作用,随着水位上升小天鹅家域范围会明显减小;越冬期间小天鹅喜好在空间相邻的浅水和草地综合性区域活动,食物资源分布和可获得性是小天鹅活动区发生改变的主要原因。     

Postnatal ontogenetic scaling of Nesodontine (Notoungulata,Toxodontidae) cranial morphology

Cassini G.H., Flores D.A. and Vizcaíno S.F. 2012. Postnatal ontogenetic scaling of Nesodontine (Notoungulata, Toxodontidae) cranial morphology. —Acta Zoologica (Stockholm) 93 : 249–259. Toxodontidae is a clade of endemic South American ungulates that comprises medium to very large animals, including strict megammamals, i.e., 1000 kg or more. Adinotherium at approximately 120 kg and Nesodon at 550 kg are, respectively, the smallest and the largest Nesodontinae of Santacrucian age (early Miocene). The large number of specimens recorded and the quality of preservation (including very young animals) permit a morphometric study of cranial ontogeny. We measured 17 cranial variables on an ontogenetic series of 23 specimens of Adinotherium ovinum and 11 of Nesodon imbricatus. Bivariate analysis (standardized major axis) was performed to obtain the coefficients of allometry using skull length as the independent variable. Results indicate that eight of 16 variables show an isometric trend, seven exhibit positive allometry, and only the height of the orbit in N. imbricatus exhibits negative allometry. Contrary to expectation, neurocranial variables are positively allometric or isometric. With respect to the splanchnocranium, most variables related to the rostrum, palate, and masticatory muscles show positive allometry, suggesting a strengthening of masticatory system in adults of both taxa. The splanchnocranial allometric trends fail to support previous inferences of specialized herbivory, suggesting generalized herbivory in nesodontines.     

The effects of food and density of the movement ofArion ater andAriolimax columbianus (Pulmonata: Stylommatophora) between habitats

Experiments were conducted to determine if slug density, or food quantity and quality could cause dispersal by slugs. Arion ater dispersed quickly from overcrowded areas, where high mortality was incurred by this species. Poor food conditions, caused more dispersal than good food conditions, but food had no effect on the mortality of A. ater. Neither food nor density factors had significant effects on the dispersal of Ariolimax columbianus. However, more A. columbianus died in the high- than low-density treatments. A. columbianus mortality was unaffected by the food factor. Dispersing Arion were heavier than non-dispersers, but there was no difference between the weights of dispersing and non-dispersing Ariolimax. There were significant seasonal changes in the dispersal rate, mortality and weight of dispersing Arion, whereas only the mortality of Ariolimax changed significantly during the experiments. Since Arion ater did not markedly reduce its activity during unsuitable weather it was able to leave areas of poor food and high slug density fairly rapidly. Ariolimax columbianus' large size allowed it to remain inactive during unfavourable weather and permitted this species to be relatively unresponsive to the experimental treatments.     

Phenotypic diversification and island evolution of pipistrelle bats (Pipistrellus pipistrellus group) in the Mediterranean region inferred from geometric morphometrics and molecular phylogenetics

Aim The Mediterranean Basin is a centre of radiation for numerous species groups. To increase our understanding of the mechanisms underlying speciation and radiation events in this region, we assessed the phenotypic variability within the Pipistrellus pipistrellus–pygmaeus–hanaki species complex. Although bats form the second largest mammalian order, studies of insular evolution in this group are scarce. We approached this problem from a microevolutionary perspective and tested for the recurrence of the insular syndrome. Location The Mediterranean Basin, with a special focus on isolated populations from Corsica, the Maghreb, Cyprus, Cyrenaica and Crete. Methods Phenotypic variability was assessed by cranial morphometrics using the coordinates of 41 3D landmarks and associated geometric‐morphometric methods. We analysed 125 specimens representing all of the lineages in the species complex. Differences between taxa and between insular and continental populations in cranial size, shape, form and allometries were tested using analyses of variance and visualized using boxplots and canonical variate analysis. Relationships between molecular data from a previous study (cytochrome b sequences) and morphometric data were tested with co‐inertia analyses (RV test) and multivariate regressions. Results The three species were relatively well differentiated in cranial size and shape, and each species showed a significant amount of inter‐population variability. Comparisons of pairs of insular versus continental populations revealed heterogeneities in cranial patterns among island phenotypes, suggesting no recurrent insular syndrome. Molecular and phenotypic traits were correlated, except for molecular and lateral cranium shape. Main conclusions The Pipistrellus pipistrellus – pygmaeus – hanaki species complex exhibits phenotypic variability as a result of the fragmentation of its distribution (especially on islands), its phylogenetic and phylogeographic history and, most probably, other evolutionary factors that were not investigated in this study. We found no recurrent pattern of evolution on islands, indicating that site‐specific factors play a prevailing role on Mediterranean islands. The correlation between molecular and phenotypic data is incomplete, suggesting that factors other than phylogenetic relationships, potentially connected with feeding ecology, have played a role in shaping cranial morphology in this species complex.     

Endothermic mosasaurs? Possible thermoregulation of Late Cretaceous mosasaurs (Reptilia,Squamata) indicated by stable oxygen isotopes in fossil bioapatite in comparison with coeval marine fish and pelagic seabirds

The thermoregulatory style of Late Cretaceous mosasaurs has become a highly controversial subject in vertebrate palaeontology. These extinct marine reptiles have previously been described as poikilothermic, endothermic or gigantothermic. Here we analyse three genera of mosasaurs from the Mooreville Chalk in Alabama (USA) of differing body mass, and compare their δ¹⁸O_PO4 derived body temperatures (T_b) with those of coeval poikilothermic fish (Enchodus) and endothermic pelagic seabirds (Ichthyornis). Results show that all mosasaurs, Clidastes (T_b = 33.1°C), Platecarpus (T_b = 36.3°C), and Tylosaurus (T_b = 34.3°C), had elevated average body temperatures in relation to those of the fish (T_b = 28.3°C) and were closer to those of Ichthyornis (T_b = 38.6°C). The temperatures calculated for Enchodus compare well with previously reported temperature estimates for the Mooreville Chalk and the T_b of Ichthyornis compares well with temperatures that have been reported for modern seabirds, suggesting that this method provides accurate results. Finally, although there are small differences of body temperature among mosasaur genera, these are independent of size, and thus inferred body mass, suggesting that mosasaurs were not gigantotherms, but rather endotherms.     

Lung morphology of cursorial and non-cursorial mammals: Lagomorphs as a case study for a pneumatic stabilization hypothesis

Gross lung morphology is examined in representative species from four genera within the order Lagomorpha (Lepus californicus, Sylvilagus nuttali, Oryctolagus cuniculus, Ochotona princeps), and compared with a representative rodent out-group (Spermophilus richardonsii). Examination of pulmonary morphology reveals several correlations between the thoracic morphology and locomotor behavior. Lepus, the most cursorial species, exhibits a distinct suite of characteristics: 1) tissue of the right cranial lobe interposed between the heart and sternum; 2) well-defined grooves in the lung tissue for both the aorta and ribs; 3) a fibrous pericardial attachment to the sternum; 4) relatively large heart and lung mass. Sylvilagus, a sprinter, exhibits these features to a lesser degree, whereas Oryctolagus and Ochotona, non-cursorial species, lack most of these features. This same suite of pulmonary features is also observed in a wide range of unrelated cursorial taxa (including selected Artiodactlya, Perissodactyla, Carnivora). Corrosion casts of the internal airways demonstrate that the cursorial and non-cursorial taxa examined here have similar branching patterns despite their variable external morphologies. The juxtaposition of pulmonary lobes, heart, and ribs leads to the hypothesis that the lungs themselves provide mechanical support of the heart and visceral mass during locomotion. Analyses of cineradiographic and pneumotachographic data obtained from Oryctolagus tend to support a pneumatic stabilization hypothesis: the lungs themselves, intimately associated with the chest walls and positively pressurized during landing, may provide some mechanical support to the viscera. This mechanism may be important in stabilizing the relatively large hearts of the most cursorial species during running. © 1996 Wiley-Liss, Inc.     

Craniometric sexual dimorphism inLeontopithecus Lesson, 1840 (Callitrichidae, Primates)

Lion tamarins are among the World's most critically endangered primates. Many studies have been produced under guidance of the International Management Committees for the preservation and management of these tamarins. Primates present morphological sexual differences in a wide range of characteristics, including cranial morphology. Studies of sexual dimorphism in the cranial morphology of theLeontopithecus are few in number and contradictory in their results. In order to check for the existence of sexual dimorphism in lion tamarins the present study analyzed 17 craniometric distances on 56 crania of three species of lion tamarins (Leontopithecus): 20L. rosalia (14 females and 6 males); 13L. chrysomelas (6 females and 7 males); and 23L. chrysopygus (8 females and 15 males). All crania are housed in the CPRJ-FEEMA collection (Primatological Center of Rio de Janeiro) and came from animals born in captivity.L. chrysopygus was more sexually dimorphic (10/17 measurements, 59%) thanL. chrysomelas (9/17 measurements, 53%) orL. rosalia (7/17 measurements, 41%). In all three species, male values are greater than the female ones, except for orbital breadth (m7) inL. rosalia. However, this distance is not sexually dimorphic in this species. This study reveals that some cranial distances, especially in the facial region, are sexually dimorphic in lion tamarins.     

A new species of Acynodon (Crocodylia) from the upper cretaceous (Santonian–Campanian) of Villaggio del Pescatore,Italy

Abstract: The new species Acynodon  adriaticus is described on the basis of remains from the Santonian–Campanian of Villaggio del Pescatore (Trieste, NE Italy). This species differs in several cranial features from Acynodon  iberoccitanus, the only other Acynodon species whose cranial osteology is known in detail. The absence of maxillary and dentary caniniform teeth coupled with the presence of enlarged molariform teeth suggests that Acynodon probably fed on slowly moving hard‐shelled prey. Moreover, the new materials reveal for the first time the morphology of some postcranial elements of Acynodon: in particular, medial‐most paravertebral osteoderms that are characterized by two keels. A new cladistic phylogenetic analysis resolves the previously reported polytomy among the basal Globidonta: Acynodon is recognized as the most primitive globidontan. This genus may represent the geologically oldest known globidontan. The fact that Acynodon has been found only in Europe and that the outgroup of Globidonta, the Diplocynodontinae, is mainly known from Europe, suggests that globidontans may have originated in Europe and not in North America as previously supposed.     

Towards a phylogeny of Cyclops (Copepoda): (in)congruences among morphology,molecules and zoogeography

The phylogeny of Cyclops (~30 spp.), a predominantly Palearctic cold‐adapted genus, was reconstructed based on morphological and molecular characters. The morphological analysis used extensive taxon sampling from the entire Holarctic range of the genus and included 53 morphological characters. Polymorphic traits were coded by the “unordered,” “unscaled” and “scaled” methods; maximum parsimony criterion was applied in tree building. Molecular phylogenetic reconstructions utilized partial nuclear 18S and 28S ribosomal genes, mitochondrial cytochrome oxidase I and complete internal transcribed spacer regions I and II, albeit with limited taxon sampling. Bayesian inference and maximum likelihood were used in these tree reconstructions. The molecular characters were used both in combination with morphology and as an independent test of the basal relationships inferred from morphology. Monophyly of the genus received strong support in both the morphological and molecular phylogenies; the basal relationships remain unresolved. The morphology‐based phylogenies, along with the geographic distribution patterns and ecological traits, supported monophyly of the ankyrae?ladakanus clade, scutifer‐clade (C. scutifer, C. jashnovi, C. columbianus), kolensis‐clade (C. kolensis, C. kikuchii, C. vicinus, C. furcifer, C. insignis, C. alaskaensis), abyssorum‐clade (C. abyssorum s. str., C. abyssorum larianus, C. ricae, C. sevani) and divergens‐clade (South Carpathian “Cyclops sp. Y,” C. mauritaniae, C. divergens, C. bohater, C. lacustris). Relationships among European and North American populations of C. scutifer and C. columbianus based on partial sequences of the 12S mitochondrial gene show C. scutifer to be paraphyletic, suggesting two independent invasions into North America via the Bering Land Bridge from Siberia to Alaska.     

Reconstruction and re-evaluation of the skull ofHomo antiquus (hominoidea: Homininae) from Hadar

The skull ofHomo antiquus Ferguson, 1984, represented by cranial remains of a fossilized skeleton, A.L. 288-1, from the Plio/Pleistocene of Hadar in Ethiopia, is reconstructed and the procedure described. Re-evaluation of the skull shows that it is apparently the smallest, normal, unequivocal hominid skull known; and that its cranial morphology is not Australopithecine, but Hominine.     

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best‐known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well‐developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 145?169.     

Primate cranial diversity

Many studies in primate and human evolution focus on aspects of cranial morphology to address issues of systematics, phylogeny, and functional anatomy. However, broad analyses of cranial diversity within Primates as an Order are notably absent. In this study, we present a 3D geometric morphometric analysis of primate cranial morphology, providing a multivariate comparison of the major patterns of cranial shape change during primate evolution and quantitative assessments of cranial diversity among different clades. We digitized a set of 18 landmarks designed to capture overall cranial shape on male and female crania representing 66 genera of living primates. The landmark data were aligned using a Generalized Procrustes Analysis and then subjected to a principal components analysis to identify the major axes of cranial variation. Cranial diversity among clades was compared using multivariate measurements of variance. The first principal component axis reflects differences in cranial flexion, orbit size and orientation, and relative neurocranial volume. In general, it separates strepsirrhines from anthropoids. The second axis reflects differences in relative cranial height and snout length and primarily describes differences among anthropoids. Eulemur, Mandrillus, Pongo, and Homo are among the extremes in cranial shape. Anthropoids, catarrhines, and haplorhines show a higher variance than prosimians or strepsirrhines. Hominoids show the highest variance in cranial shape among extant primate clades, and much of this diversity is driven by the unique cranium of Homo sapiens. Am J Phys Anthropol 142:565–578, 2010. © 2010 Wiley‐Liss, Inc.     

Genetic differences among subspecies of the saddle-back tamarin (Saguinus fuscicollis):evidence from hybrids

The subspecies of saddle-back tamarins (Saguinus fuscicollis) are known to be chromatically and morphologically diverse but little is known of the genetic basis for the observed morphological variation. The morphology of first generation subspecific hybrids can be compared to that of the parental subspecies to provide information on the extent and nature of genetic differences in morphology between subspecies. We compare two groups of saddle-back tamarin hybrids (S. f. illigeri × S. f. lagonotus and S. f. illigeri × S. f. leucogenys) to pure-bred members of their parental subspecies. These crosses were examined for heterosis, caused by allele frequency differences between the subspecies in combination with directional dominance. Thirty-nine craniofacial measurements were derived from three-dimensional coordinates of landmarks on 355 adult tamarin skulls. These measurements were corrected for sex differences and differences due to environment (wild-derived vs. laboratory-born) prior to analysis of hybridity. Sex differences were minimal for these traits. Environment had a more significant effect on craniofacial morphology. Laboratory environments produce larger faces but smaller orbits, anterior cranial vaults, and cranial bases. Significant heterosis was found for many individual traits and for the first principal component representing size and size-related shape measurements in the S. f. illigeri × S. f. lagonotus cross. The smaller samples involved in the S. f. illigeri× S. f. leucogenys cross led to a much lower number of statistically significant results, although most traits did display heterosis. Heterosis for craniofacial size was nearly statistically significant. These results suggest that there are large differences in allele frequencies among these subspecies of saddle-back tamarin for genes affecting craniofacial morphology. Based on these data we suggest that these subspecies are likely to be independent, largely isolated, evolutionary units. © 1993 Wiley-Liss, Inc.     

The cranial anatomy,ontogeny, and relationships of Karpinskiosaurus secundus (Amalitzky) (Seymouriamorpha,Karpinskiosauridae) from the Upper Permian of European Russia

The Upper Permian seymouriamorph tetrapod Karpinskiosaurus from European Russia includes two species: Karpinskiosaurus secundus and Karpinskiosaurus ultimus. Karpinskiosaurus secundus is represented by two specimens with skull lengths of about 75 mm. All specimens of K. ultimus are smaller than those of K. secundus. Revision of the cranial anatomy of all previously known and several new specimens of Karpinskiosaurus shows that the specimens of K. secundus and most of the specimens of K. ultimus represent the ontogenetic series of one species: K. secundus. The holotype specimen of K. ultimus requires revision, with the aim to find out whether it represents a second species of Karpinskiosaurus or not. The available material permits new reconstructions of the largest, holotype skull, and one smaller skull with a length of about 36 mm. Karpinskiosaurus secundus is included in a cladistic analysis for the first time here. The analysis shows it to form a sister taxon to Discosauriscidae. The clade comprising Karpinskiosaurus secundus plus Discosauriscidae forms a sister group to Seymouriidae. Karpinskiosaurus secundus has a large postorbital and a short preorbital region, and the orbits are placed in the posterior portion of the anterior half of the skull length. Among all seymouriamorphs, such cranial proportions are exhibited only by the largest known specimens of Discosauriscus austriacus. None of the specimens of K. secundus described here exhibits the presence of sensory grooves; thus, all specimens composing the ontogenetic sequence of K. secundus are considered to be terrestrial. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Description of the Type Specimen of the Extinct Tenerife Giant Rat (Canariomys bravoi)

The holotype of the Tenerife (Canary Islands) giant rat, Canariomys bravoi, an almost complete cranium from the Late Pleistocene site of Cueva de las Palomas, is described for the first time. This species is characterized by its large size, robust skull with a short rostrum, dorsal inflation at the level of the infraorbital foramen, and moderately high-crowned upper molars which develop partial stephanodonty at advanced wear stages. Canariomys tamarani from Gran Canaria Island, the only other species of this genus known so far, is slightly smaller and further differs in its higher crowned molars with cusps arranged in a lamellar pattern. However, their crania are remarkedly similar and present a distinctive anatomy of the zygomatic plate, which is very high on the rostrum, so that its dorsal border is nearly on level with the zygomatic process of the maxilla instead of considerably below it as usually seen in murines. This is also observed in other insular, often large-sized, murines but certainly evolved in parallel as an adaptation to herbivory. Molar morphology is congruent with recent analyses of ancient molecular data of C. bravoi which place it in the tribe Arvicanthini (mostly African murines), more concretely within the Arvicanthis niloticus species complex. Even though genetic data indicate that both species diverged very recently, just 650,000 years ago, cranial and dental anatomy of C. bravoi are very derived. Conversely, C. tamarani presents a molar morphology reminiscent of that of A. niloticus albeit associated with a similarly highly derived cranial anatomy.