The external gills and ornamentation of skull roof bones of the Lower Permian tetrapodDiscosauriscus (Kuhn 1933) with remarks to its ontogeny

Until now, the largest specimens ofDiscosauriscus (and alsoUtegenia andAriekanerpeton- the Lower Permian members of the family Discosauriscidae) have been considered to be adult individuals. The largest specimens ofDiscosauriscus, in which the external gills are still present, had skull lengths of about 22-25 mm. Metamorphosis was believed to have commenced at this size in discosauriscids and specimens with skull lengths of about 50-54 mm were considered to be already adult or senile. New finds ofDiscosauriscus demonstrate the presence of external gills in specimens with skull lengths of up to 32 mm. On the base of this, together with the ornamentation or the skull roof bones and osteological features of new large specimens, it is here concluded that 1)Discosauriscus represents the larval, metamorphic and early juvenile stages of a reptiliomorph tetrapod, the adults of which are yet unknown and 2) two other discosauriscids(Ariekanerpeton andUtegenia) also represent early ontogenetic stages.     

The subdivisions and fusions of the exoskeletal skull bones ofDiscosauriscus austriacus (Makowsky 1876) and their possible homologues in rhipidistians

Two localities in the Boskovice Furrow region of Moravia (Czecho-Slovakia) have produced new, well preserved material of the Lower Permian tetrapodDiscosauriscus austriacus (Makowsky 1876). A relatively large number of specimens have been found with some dermal skull roof bones partly or fully subdivided, and/or fused. These are the first records of such subdivisions and fusions in this tetrapod and the following bones are discussed: frontal, postfrontal, parietal, intertemporal, supratemporal, tabular and postparietal. The subdivided bones within the skull roof ofDiscosauriscus (and some dissorophoids) are situated at the same places as those found in Devonian rhipidistians, and may relate to the homology of the bones of the cranial exoskeletal roof inDiscosauriscus and in osteolepiforms. It shows that the “orthodox” terminology of the skull roof bones used in osteolepiforms is correct.     

Spore morphology of the genus Dryopteris Adans. (Dryopteridaceae) in Korea

Spore morphology of the Korean members of the genus Dryopteris was examined by scanning electron microscopy. In particular, spores of D. hangchowensis, D. woodsiisora, D. saxifragivaria, and D. subexaltata were examined here for the first time. Three perispore types were recognized on the basis of the shape of macro-ornamentations on perispore: these include rugate, echinate, and spinose. The rugate perispore type appeared to be the most common in the genus, and probably represents plesiomorphic condition in the genus Dryopteris. The echinate perispore type is found in D. expansa, and spinose perispore type in D. formosana. The results of this study demonstrate that the perispore characteristics, particularly the shape of macro-ornamentation and surface texture, have diagnostic value at the species level. Indeed, most of the Dryopteris taxa in Korea could be distinguished by their perispore characters. In addition, spores of sexual diploid individuals of D. saxifraga were much smaller than those of agamosporous triploid individuals, supporting the contention that the size of spores in ferns is generally related to ploidy levels.     

First record of Sorex (Drepanosorex) margaritodon (Mammalia, Soricidae) in Western Europe: biostratigraphy, biogeography and evolution of the species

The taxonomy of the extinct European subgenus Drepanosorex (Mammalia, Soricidae) has traditionally been problematic, but it seems to comprise five valid species from the Pleistocene: Sorex (Drepanosorex) praearaneus, S. (D.) savini, S. (D.) margaritodon, S. (D.) austriacus and S. (D.) rupestris. The first record of Sorex (Drepanosorex) margaritodon in Western Europe comes from the Lower Red Unit (levels TE7–14) of Sima del Elefante (Sierra de Atapuerca, Burgos, Spain), which is dated to ca. 1.1–1.5 Ma. Detailed morphological comparisons and morphometric analyses have allowed us to assign the items from Sima del Elefante indisputably to this taxon. Sorex (D.) margaritodon is now firmly dated to the Early Pleistocene of Central Europe and the north of the Iberian Peninsula, which makes it a very useful biochronological tool in the continental context. Sorex (D.) margaritodon and S. (D.) savini could have evolved in Central Europe from the more primitive S. (D.) praearaneus, which could also be the potential ancestor of the Caucasian Sorex (D.) rupestris. Sorex (D.) margaritodon, the Drepanosorex species with the westernmost distribution in Europe, may be regarded as the most probable ancestor of S. (D.) austriacus.     

Phylogeny and bionomics of Lasius austriacus (Hymenoptera,Formicidae)

Summary Phylogenetic analysis based on sequence data of the mitochondrial COI gene confirms the species status of the recently described Lasius austriacus. The five haplotypes of L. austriacus do not cluster according to their geographic origin, indicating a recent gene flow among the populations. The molecular data corroborate the morphology based hypothesis that L. austriacus belongs to the Lasius (Lasius s.str.) brunneus group. The invasive species Lasius neglectus forms a sister taxon with L. turcicus, both next related to L. austriacus. Other phylogenetic relationships within the genus Lasius are in accordance with morphological data.First data on the bionomics of L. austriacus are discussed in context with its phylogenetic position. Based on gyne and male morphology, excavations of nests, pitfall trapping and observations in formicaries, we hypothesize that L. austriacus is a mainly hypogaeic, monogynous species with nuptial flight. These are characters of the Lasius brunneus group in general, except the polygynous-polycalic, intranidally copulating L. neglectus. Aggression tests, however, revealed non-aggressive behaviour (antennation) between separated L. austriacus populations, but pronounced interspecific aggression against L. neglectus. This confirms the species status of L. austriacus and indicates a reduced level of intraspecific aggression, similar to L. neglectus.The status of L. austriacus as a native species in Central Europe is confirmed.Received 10 February 2003; revised 24 June 2003; accepted 23 July 2003.     

New ankylosaurian material from the Upper Cretaceous of Transylvania

Ankylosaurian remains from the Transylvanian Basin, Romania, are extremely rare. More than 100 years after the discovery of the first and only better-known assemblage, namely the type material of Struthiosaurus transylvanicus, new ankylosaurian material has been discovered in the Maastrichtian of the Ha?eg Basin, as well as at another locality (Vurp?r), in the Transylvanian Basin, that is described here. The material consists of one tooth in a small jaw fragment (from the Ha?eg Basin) and at least two accummulations of associated, as well as several isolated, postcranial elements (from Vurp?r). No diagnostic elements are preserved that would overlap with the type of S. transylvanicus, so we cannot assign any of the new specimens to this species. The tooth shows marked differences compared to those of other anklyosaurs including S. austriacus and Hungarosaurus in having only six, more or less equally sized, apically pointed cusps separated by deep grooves. The postcranial material from Vurp?r represents at least three different individuals. The humerus is the most diagnostic element among the postcranial remains being most similar both in size and morphology to humeri referred to as Struthiosaurus from different European localities, thus here we refer the humerus and probably associated elements preserved in one assemblage to as cf. Struthiosaurus sp.; the remaining specimens from Vurp?r are retained as Nodosauridae indet. Histological studies have confirmed the adult nature of all sampled bones in the Vurp?r ankylosaur material suggesting that these fully grown animals were of similar size to Struthiosaurus, a small-bodied nodosaurid the ontogenetic status of which, however, has never been investigated histologically. The obviously diminished body size of the Transylvanian ankylosaurs compared to other members of the clade could be explained by insular dwarfism using the same histology-based argument as presented for Magyarosaurus.     

Bone Histology Reveals a High Environmental and Metabolic Plasticity as a Successful Evolutionary Strategy in a Long-Lived Homeostatic Triassic Temnospondyl

Evolutionary stasis (long-term stability of morphology in an evolving lineage) is a pattern for which explanations are usually elusive. The Triassic tetrapod Gerrothorax pulcherrimus, a gill-bearing temnospondyl, survived for 35 million years in the Germanic Basin of Central Europe persisting throughout the dinosaur-dominated Late Triassic period. This evolutionary stasis coincides with the occurrence of this species in a wide range of habitats and environmental conditions. By the combination of palaeoecological and palaeohistological analyses, we found great ecological flexibility in G. pulcherrimus and present substantial evidence of developmental and metabolic plasticity despite the morphological stasis. We conclude that G. pulcherrimus could show the capacity to settle in water bodies too harsh or unpredictable for most other tetrapods. This would have been made possible by a unique life history strategy that involved a wide reaction norm, permitting adjustment to fluctuating conditions such as salinity and level of nutrients. Growth rate, duration of juvenile period, age at maturity, and life span were all subject to broad variation within specimens of G. pulcherrimus in one single lake and in between different lakes. In addition to providing a better understanding of fossil ecosystems, this study shows the potential of such a methodology to encourage palaeobiologists and evolutionary biologists to consider the mechanisms of variation in extant and fossil organisms by using a similar time-scope reference.     

Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well-known. This skeleton represents some of the first associated evidence of cranial and postcranial female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons (Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas (Theropithecus gelada).     

Toxic coral gobies reduce the feeding rate of a corallivorous butterflyfish on Acropora corals

The obligate coral-dwelling gobiid genus Gobiodon inhabits Acropora corals and has developed various physiological, morphological and ethological adaptations towards this life habit. While the advantages of this coral-fish association are well documented for Gobiodon, possible fitness-increasing factors for the host coral are unknown. This study examines the influence of coral-dwelling gobies on the feeding behaviour of obligate corallivorous butterflyfishes. In an aquarium experiment using video observation, the corallivorous butterflyfish Chaetodon austriacus fed significantly less on corals inhabited by two Gobiodon species compared to unoccupied coral colonies of similar size. The more agonistic species G. histrio, which mostly displayed directed movements towards butterflyfishes, decreased butterflyfish bite rate by 62–98 % compared to uninhabited colonies. For Gobiodon sp. 3, which mostly displayed undirected movements in response to visits by C. austriacus, bite rate reduction was 64–68 %. The scale-less skin of Gobiodon spp. is covered by mucus that is toxic and multi-functional by reducing predation as well as affecting parasite attachment. A choice flume experiment suggests that the highly diluted skin mucus of Gobiodon spp. also functions as a corallivore repellent. This study demonstrates that Gobiodon spp. exhibit resource defence against coral-feeding butterflyfishes and also that coral colonies without resident Gobiodon suffer higher predation rates. Although the genus Gobiodon is probably a facultative corallivore, this study shows that by reducing predation on inhabited colonies by other fishes, these obligate coral-dwellers either compensate for their own fitness-decreasing impact on host colonies or live in a mutualistic association with them.     

Morphological evolution through integration: a quantitative study of cranial integration in Homo, Pan, Gorilla and Pongo

Morphological integration refers to coordinated variation among traits that are closely related in development and/or function. Patterns of integration can offer important insight into the structural relationship between phenotypic units, providing a framework to address questions about phenotypic evolvability and constraints. Integrative features of the primate cranium have recently become a popular subject of study. However, an important question that still remains under-investigated is: what is the pattern of cranial shape integration among closely related hominoids? To address this question, we conducted a Procrustes-based geometric morphometrics study to quantify and analyze shape covariation patterns between different cranial regions in Homo, Pan, Gorilla and Pongo. A total of fifty-six 3D landmarks were collected on 407 adult individuals. We then sub-divided the landmarks corresponding to cranial units as outlined in the ‘functional matrix hypothesis.’ Sub-dividing the cranium in this manner allowed us to explore patterns of covariation between the face, basicranium and cranial vault, using the two-block partial least squares approach. Our results suggest that integrated shape changes in the hominoid cranium are complex, but that the overall pattern of integration is similar among human and non-human apes. Thus, despite having very distinct morphologies the way in which the face, basicranium and cranial vault covary is shared among these taxa. These results imply that the pattern of cranial integration among hominoids is conserved.     

Sexual dimorphism of skull shape in a lacertid lizard species (Podarcis spp., Dalmatolacerta sp., Dinarolacerta sp.) revealed by geometric morphometrics

Geometric morphometric techniques were used to examine allometric and non-allometric influences on sexual shape dimorphism (SShD) in the ventral cranium (skull base, palate and upper jaw) of four species of lacertid lizards (Podarcis muralis, Podarcis melisellensis, Dalmatolacerta oxycephala, Dinarolacerta mosorensis). These species differ in body shape, ecology and degree of phylogenetic relatedness. The structures of the ventral cranium that were studied are directly involved in the mechanics of feeding and are connected to the jaw musculature; these structures are potentially subject to both sexual and natural selection. Allometry accounted for a considerable degree of cranial shape variation between the sexes. Allometric shape changes between individuals with smaller cranium size and individuals with larger cranium size are mostly related to changes in the skull base showing pronounced negative allometry. The rostral part, however, either scaled isometrically or showed less pronounced negative allometry than the skull base. Non-allometric intersexual shape variation predominantly involved changes related to the jaw adductor muscle chamber, i.e., changes that are associated with biomechanically relevant traits of the jaw system in females and males. Both allometric and non-allometric shape changes appeared to be species-specific. Our results indicate that natural and sexual selection may be involved in the evolution of SShD.     

New material and revision of hipparions from the upper Miocene locality Ravin des Zouaves-5, RZO of Axios Valley (Macedonia,Greece)

The reconsideration of the hipparions from the locality Ravin des Zouaves-5 (RZO) of the lower Axios Valley provides a better taxonomy, based on the revision of the old known collection (up to 1987), the inclusion of new discoveries and the bibliographic data of the last 30 years related to the study of the hipparions in Greece and the neighboring countries. Based on the cranial material and metapodials, five different taxa have been distinguished. New cranial material confirms the presence of the large-sized Hipparion proboscideum and the small-sized Hipparion macedonicum in the RZO fauna. In the new middle-sized hipparion sample from RZO two species have been identified, Hipparion philippus, and another form similar to Hipparion mediterraneum, which is attributed to Hipparion cf. mediterraneum. Cranial remains of a second small-sized hipparion are also recognized in the RZO fauna, which is attributed to Hipparion sithonis. The relationships of H. mediterraneum and H. proboscideum are discussed; a cladistic analysis suggests that they are sister taxa, sharing a common ancestor with the Hipparion moldavicum group, and a form similar to H. mediterraneum is a possible ancestor of H. proboscideum.     

The reproduction of Chaetodon austriacus,C. fasciatus and C. paucifasciatus (Chaetodontidae,Perciformes) in the Jordan Gulf of Aqaba

Synopsis Aspects of the reproduction of three sympatric and endemic chaetodontids, Chaetodon austriacus, C. fasciatus and C. paucifasciatus, from the Jordan Gulf of Aqaba were investigated. Chaetodon fasciatus had a higher fecundity than the other species which, in turn, had similar fecundities. The major egg release of C. austriacus and C. paucifasciatus began in August, that of C. fasciatus in September. Based on the gonadosomatic index of both sexes, the spawning period of C. austriacus was from July through October, that of C. paucifasciatus from August through October and that of C. fasciatus from September through December. The maturity length of the three chaetodontids is given and reproductive isolation among the sympatric species is discussed.     

Phenetic relationships among four Apodemus species (Rodentia, Muridae) inferred from skull variation

Phenetic relationships among four Apodemus species (A. agrarius, A. epimelas, A. flavicollis and A. sylvaticus) inferred from skull (mandible and cranium) variation were explored using landmark-based geometric morphometrics. Analysis of size variation revealed that mandibles and crania of A. epimelas were the largest, followed by those of A. flavicollis, while A. agrarius and A. sylvaticus had the smallest ones. Phenetic relationships inferred from mandible shape variation better reflected phylogenetic relationships among the analyzed Apodemus species than those inferred from cranial differences. Concerning cranial shape variation, the most differentiated species was A. epimelas, whose ecology clearly differs from the other three species. Thus, differentiation of the mandible provided a pattern fully concordant with the phylogeny, while the cranium differentiation was in agreement with ecology expectations. The most evident shape changes of mandible and cranium involved the angular process and facial region, respectively. We also found that allometry had a significant influence on shape variation and that size-dependent shape variation differed among the analyzed species. Moreover, mandible and cranium are differently influenced by allometric changes. Different phenetic relationships inferred from mandible and cranium shape variation imply that phylogeny, ecology, together with factors related to size differences are all involved in the observed morphological divergence among the analyzed Apodemus species.     

A new species of Theriosuchus (Atoposauridae, Crocodylomorpha) from the Late Jurassic (Kimmeridgian) of Guimarota, Portugal

A new species of an atoposaurid crocodilian, Theriosuchus guimarotae, is described from the Late Jurassic (Kimmeridgian) of Portugal. Theriosuchus guimarotae can be distinguished from other species of Theriosuchus by a lateral surface of squamosal bevelled ventrally; a rounded, caudally projecting and dorsally sculptured caudolateral corner of the squamosal; a premaxillomaxillary suture aligned caudomedially in dorsal aspect; a minimum space between the supratemporal foramina that comprises one third of the total width of the cranial table; a minimum width of the frontal between the orbits that comprises one third of the maximum width of the skull at the orbits; a dentition that comprises only pseudocaniniform and lanceolate-shaped teeth; the presence of an external mandibular fenestra and all vertebral bodies amphicoelous. Its osteology also sheds light on the diagnosis of Theriosuchus within Atoposauridae. The material additionally includes specimens representative of several ontogenetic stages, each of which is discussed here. With its Late Jurassic age, T. guimarotae represents the oldest well-preserved material of Theriosuchus and reveals further knowledge about the palaeobiogeography of the genus in western Europe.     

New remains of Diplocynodon (Crocodylia: Diplocynodontidae) from the Early Miocene of the Iberian Peninsula

We describe crocodylian remains from the Early Miocene (MN4) site of Els Casots (Subirats, Vallès-Penedès Basin, NE of the Iberian Peninsula). Referral to Diplocynodon (Alligatoroidea: Diplocynodontidae) is justified by several cranial and postcranial features, including: (1) the subequal and confluent alveoli of the maxilla (fourth and fifth) and dentary (third and fourth); (2) the position of the foramen aëreum on the quadrate; (3) the small and ventrally reflected medial hemicondyle of the quadrate; (4) the distinct dorsoventral step on the frontal; and (5) the bipartite ventral osteoderms. Multiple morphological features are consistent with an attribution to Diplocynodon ratelii, previously known from the Early Miocene (MN2) of France, and discount an alternative attribution to other species of the genus, including Diplocynodon ungeri from the Middle Miocene (MN5) of Austria. The described material from Els Casots is smaller in size than the French material of D. ratelii, possibly reflecting an earlier ontogenetic stage. The described remains constitute the first report of D. ratelii and the youngest record of Diplocynodon in the Iberian Peninsula, where only Diplocynodon muelleri and Diplocynodon tormis have been previously reported. The presence of Diplocynodon further supports the lacustrine depositional environment previously inferred for Els Casots and also indicates a relatively high temperature.     

Interspecific variation of ontogeny and skull shape among porpoises (Phocoenidae)

All extant members of Phocoenidae (porpoises) have been characterized as pedomorphic based on skeletal characters. To investigate the ontogenetic background for pedomorphosis and assess interspecific differences in ontogeny among phocoenids, samples of the six extant species were compared in terms of development of both epiphyseal and cranial suture fusion. Across all species, full maturity of the vertebral column was rare. Vertebral epiphyseal development did not progress so far in most Phocoena phocoena as in Phocoenoides dalli and Phocoena dioptrica. P. phocoena, Phocoena spinipinnis, Ph. dalli, and P. dioptrica, for which large series were available, were further compared in terms of ontogeny of cranial shape by three‐dimensional geometric morphometrics. Ph. dalli and P. dioptrica generally showed further development of cranial sutures than the other species. Postnatal skull shape development was similar for all species studied; the majority of interspecific shape differences are present at parturition. Smaller species had a higher rate of shape development relative to growth in size than Ph. dalli and P. dioptrica, but they still showed less allometric development due to less postnatal growth. Interspecific shape differences indicate phylogenetic relationships similar to that proposed based on morphology or convergent evolution of the two pelagic species, Ph. dalli and P. dioptrica, under the scenarios suggested by recent molecular studies. A shape trend coinciding with habitat preference was detected; in species with pelagic preference the position and orientation of the foramen magnum aligned the skull with the vertebral column; the rostrum showed less ventral inclination, and the facial region was larger and more concave in lateral aspect. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Skeletal Morphology of Palaeocastor peninsulatus (Rodentia,Castoridae) from the Fort Logan Formation of Montana (early Arikareean): Ontogenetic and Paleoecological Interpretations

Palaeocastorine beavers were diverse members of the burrowing fauna of the Oligocene of North America best known from the Great Plains and the Great Basin. Palaeocastorines are also found in the northern Rocky Mountains. Several specimens of Capacikala, Euhapsis, and Palaeocastor have been reported from Montana within the Cabbage Patch beds fauna. New occurrences of palaeocastorine beavers from the Fort Logan Formation of central Montana show that the fauna there included Palaeocastor fossor, Capacikala sp., Palaeocastor sp., and three specimens of a species previously known only from the John Day Formation of Oregon: Palaeocastor peninsulatus. The three specimens include crania and associated postcrania forming an ontogenetic series from a juvenile to a mature adult. Careful examination of these specimens shows that morphological changes throughout ontogeny are associated with increased burrowing ability in adults. Palaeocastor peninsulatus is a scratch-digger with few cranial adaptations to burrowing susceptible to change with maturity. Dental characters, however, are modified with wear and vary greatly across individuals of similar wear stage. Cranial morphology thus appears to be a reliable source of taxonomic information in palaeocastorines unlike dental material. These new specimens provide evidence for increased chisel-tooth digging in adults compared to juveniles as is observed throughout the evolution of palaeocastorines. This study demonstrates the need for future work focusing on intraspecific variation to help better define species boundaries among palaeocastorines and improve phylogenetic analyses.     

Size, shape, and asymmetry in fossil hominins: the status of the LB1 cranium based on 3D morphometric analyses

The unique set of morphological characteristics of the Liang Bua hominins (Homo floresiensis) has been attributed to explanations as diverse as insular dwarfism and pathological microcephaly. This study examined the relationship between cranial size and shape across a range of hominin and African ape species to test whether or not cranial morphology of LB1 is consistent with the basic pattern of static allometry present in these various taxa. Correlations between size and 3D cranial shape were explored using principal components analysis in shape space and in Procrustes form space. Additionally, patterns of static allometry within both modern humans and Plio-Pleistocene hominins were used to simulate the expected cranial shapes of each group at the size of LB1. These hypothetical specimens were compared to LB1 both visually and statistically. Results of most analyses indicated that LB1 best fits predictions for a small specimen of fossil Homo but not for a small modern human. This was especially true for analyses of neurocranial landmarks. Results from the whole cranium were less clear about the specific affinities of LB1, but, importantly, demonstrated that aspects of facial morphology associated with smaller size converge on modern human morphology. This suggests that facial similarities between LB1 and anatomically modern humans may not be indicative of a close relationship. Landmark data collected from this study were also used to test the degree of cranial asymmetry in LB1. These comparisons indicated that the cranium is fairly asymmetrical, but within the range of asymmetry exhibited by modern humans and all extant African ape species. Compared to other fossil specimens, the degree of asymmetry in LB1 is moderate and readily explained by the taphonomic processes to which all fossils are subject. Taken together, these findings suggest that H. floresiensis was most likely the diminutive descendant of a species of archaic Homo, although the details of this evolutionary history remain obscure.