Epipubic bones in eutherian mammals from the late Cretaceous of Mongolia

An important transformation in the evolution of mammals was the loss of the epipubic bones. These are elements projecting anteriorly from the pelvic girdle into the abdominal region in a variety of Mesozoic mammals, related tritylodonts, marsupials and monotremes but not in living eutherian (placental) mammals. Here we describe a new eutherian from the Late Cretaceous period of Mongolia, and report the first record of epipubic bones in two distinct eutherian lineages. The presence of epipubic bones and other primitive features suggests that these groups occupy a basal position in the Eutheria. It has been argued that the epipubic bones support the pouch in living mammals, but epipubic bones have since been related to locomotion and suspension of the litter mass of several attached, lactating offspring. The loss of the epipubic bones in eutherians can be related to the evolution of prolonged gestation, which would not require prolonged external attachment of altricial young. Thus the occurrence of epipubic bones in two Cretaceous eutherians suggests that the dramatic modifications connected with typical placental reproduction may have been later events in the evolution of the Eutheria.     

A new symmetrodont mammal from China and its implications for mammalian evolution

A new symmetrodont mammal has been discovered in the Mesozoic era (Late Jurassic or Early Cretaceous period) of Liaoning Province, China. Archaic therian mammals, including symmetrodonts, are extinct relatives of the living marsupial and placental therians. However, these archaic therians have been mostly documented by fragmentary fossils. This newfossil taxon, represented by a nearly complete postcranial skeleton and a partial skull with dentition, is the best-preserved symmetrodont mammal yet discovered. It provides a new insight into the relationships of the major lineages of mammals and the evolution of the mammalian skeleton. Our analysis suggests that this new taxon represents a part of the early therian radiation before the divergence of living marsupials and placentals; that therians and multituberculates are more closely related to each other than either group is to other mammalian lineages; that archaic therians lacked the more parasagittal posture of the forelimb of most living therian mammals; and that archaic therians, such as symmetrodonts, retained the primitive feature of a finger-like promontorium (possibly with a straight cochlea) of the non-therian mammals. The fully coiled cochlea evolved later in more derived therian mammals, and is therefore convergent to the partially coiled cochlea of monotremes.     

The interphotoreceptor retinoid binding protein gene in therian mammals: implications for higher level relationships and evidence for loss of function in the marsupial mole

The subclass Theria of Mammalia includes marsupials (infraclass Metatheria) and placentals (infraclass Eutheria). Within each group, interordinal relationships remain unclear. One limitation of many studies is incomplete ordinal representation. Here, we analyze DNA sequences for part of exon 1 of the interphotoreceptor retinoid binding protein gene, including 10 that are newly reported, for representatives of all therian orders. Among placentals, the most robust clades are Cetartiodactyla, Paenungulata, and an expanded African clade that includes paenungulates, tubulidentates, and macroscelideans. Anagalida, Archonta, Altungulata, Hyracoidea + Perissodactyla, Ungulata, and the "flying primate" hypothesis are rejected by statistical tests. Among marsupials, the most robust clade includes all orders except Didelphimorphia. The phylogenetic placement of the monito del monte and the marsupial mole remains unclear. However, the marsupial mole sequence contains three frameshift indels and numerous stop codons in all three reading frames. Given that the interphotoreceptor retinoid binding protein gene is a single-copy gene that functions in the visual cycle and that the marsupial mole is blind with degenerate eyes, this finding suggests that phenotypic degeneration of the eyes is accompanied by parallel changes at the molecular level as a result of relaxed selective constraints.     

The development of the external features of the platypus (Ornithorhynchus anatinus)

The present study describes the post-hatching development of the external features of the platypus. Specimens range in age from the day of hatching through to 6 months old, and provide the first comprehensive view of the developmental sequence of these features. Various features, those specific to the platypus, those specific to monotremes and those shared with marsupials and eutherians, are described. Features specific to the platypus, including the bill and webbing of the forefeet, are seen to develop precociously. Many features show similarities to marsupials, although marsupials show differential development both in timing and in morphology. The developmental progression is related to the age, in days, although the exact age of the specimens is unclear, and relies on ages given to the specimens at the time of collection, sometimes as long as 70 years ago. Despite this, the progression of development of these features suggests that the ageing given is relatively accurate.     

The origin of the dog-like borhyaenoid marsupials of South America

Dog-like marsupials (superfamily Borhyaenoidea) were the largest predacious mammals during the Tertiary period in South America. They are critical to our understanding of marsupial origin, phylogeny and palaeobiogeography because they have been related to various marsupial lineages of several continents: didelphoids (mainly New World, but also Europe, Asia and Africa), pediomyid, stagodontids (North America), dasyuroids (Australia) and deltatheroidans (predominantly Asian). These relationships, based mainly on dental morphology, have been discussed and rejected several times. Here we report the discovery of exceptionally well preserved skulls and skeletons, referrable to the didelphoid Andinodelphys, which shed new light on the phylogenetic and palaeobiogeographic origin of dog-like marsupials. The skulls of Mayulestes (boryhyaenoid), Andinodelphys and Pucadelphys (didelphoids) from the early Palaeocene epoch of Bolivia are the oldest known for American marsupials. Comparison of their basicranial anatomy suggests that dog-like marsupials are closely related to an early didelphimorphian radiation in South America, rather than to Asiatic (deltatheroidan), North American (stagodontid), or Australian (dasyuroid) lineages.     

Aspects of masticatory form and function in common tree shrews, Tupaia glis

Tree shrews have relatively primitive tribosphenic molars that are apparently similar to those of basal eutherians; thus, these animals have been used as a model to describe mastication in early mammals. In this study the gross morphology of the bony skull, joints, dentition, and muscles of mastication are related to potential jaw movements and cuspal relationships. Potential for complex mandibular movements is indicated by a mobile mandibular symphysis, shallow mandibular fossa that is large compared to its resident condyle, and relatively loose temporomandibular joint ligaments. Abrasive tooth wear is noticeable, and is most marked at the first molars and buccal aspects of the upper cheek teeth distal to P2. Muscle morphology is basically similar to that previously described for Tupaia minor and Ptilocercus lowii. However, in T. glis, an intraorbital part of deep temporalis has the potential for inducing lingual translation of its dentary, and the large medial pterygoid has extended its origin anteriorly to the floor of the orbit, which would enhance protrusion. The importance of the tongue and hyoid muscles during mastication is suggested by broadly expanded anterior bellies of digastrics, which may assist mylohyoids in tensing the floor of the mouth during forceful tongue actions, and by preliminary electromyography, which suggests that masticatory muscles alone cannot fully account for jaw movements in this species.     

A New Hadrosauriform Dinosaur from the Early Cretaceous of Gansu Province, China

A new basal hadrosauriform dinosaur, Xuwulong yueluni gen. et sp. nov., is reported based on an articulated specimen including a complete cranium, almost complete axial skeleton, and complete left pelvic girdle. The specimen is recovered from the Early Cretaceous Yujingzi Basin in the Jiuquan area, Gansu Province of northwestern China. Xuwulong is among the most basal members of badrosauriform dinosaurs, and distinguisbed by its unique lower jaw in having a V-sbaped rostral end of the dentary in lateral view and a mandibular articular surface which levels with mid-height of the dentary ramus, as well as an enlarged and downturned prepubic process. Including Xuwulong, three basal hadrosauriforms have been discovered from the Early Cretaceous Xinminpu Group in the Mazongshan area, making it an important area to study the origin and early evolution of duck-billed dinosaurs.     

Tarsier-like locomotor specializations in the oligocene primate afrotarsius

Tarsiers and extinct tarsier-like primates have played a central role in views of primate phylogeny and evolution for more than a century. Because of the importance of tarsiers in so many primatological problems, there has been particular interest in questions about the origin of tarsier specializations and the biogeography of early tarsioid radiations. We report on a new fossil of rare Afrotarsius that shows near identity to modern Tarsius in unique specializations of the leg, which provides information about the locomotor behavior and clarifies the phylogenetic position of this previously controversial primate. These specializations constitute evidence that Afrotarsius is a tarsiid, closely related to extant Tarsius; hence, it is now excluded from being a generalized sister taxon to Anthropoidea.     

Special challenges of maintaining wild animals in captivity in Australia and New Zealand: prevention of infectious and parasitic diseases

The authors review infectious and parasitic diseases of eutherians (excluding marine mammals), marsupials, reptiles and birds in Australia and New Zealand. The diseases discussed are those which are common and/or more likely to affect humans. The emphasis is placed on the type of animal affected, the epidemiology, and hence the control or prevention measures necessary. Clinical signs, pathology and treatment are not considered in any detail.     

Marsupial light chains: complexity and conservation of lambda in the opossum Monodelphis domestica

The Ig lambda chains in the South American opossum, Monodelphis domestica, were analyzed at the expressed cDNA and genomic organization level, the first described for a nonplacental mammal. The V lambda segment repertoire in the opossum was found to be comprised of at least three diverse V lambda families. Each of these families appears to be related to distinct V lambda families present in placental mammals, suggesting the divergence of these genes before the separation of metatherians and eutherians more than 100 million years ago. Based on framework and constant region sequences from full-length cDNAs and intron sequences from genomic clones, it appears that there are multiple functional J lambda-C lambda pairs in the opossum locus. The opossum J lambda-C lambda sequences are phylogenetically clustered, suggesting that these gene duplications are more recent and species specific. Sequence analysis of a large set of functional, expressed V lambda-J lambda recombinations is consistent with an unbiased, highly diverse lambda light chain repertoire in the adult opossum. Overall, the complexity of the Ig lambda locus appears to be greater than that found in the Ig heavy chain locus in the opossum, and light chains are therefore likely to contribute significantly to Ig diversity in this species.     

A new pitheciin primate from the middle Miocene of Argentina

We report here a new fossil primate from the middle Miocene of Argentina. The material consists of isolated teeth, mandibular fragments, and a talus. The fossils were collected in the Collón Cura formation at Ca?adón del Tordillo in Neuquén Province. An age of 15.71 +/- 0.07 Ma has been reported for the Pilcaniyeu Ignimbrite, which lies just below the paleosols in which the fossils were found. This material is thus the youngest occurrence of fossil primates in Argentina (hitherto documented in the Santacrucian and older land mammal ages) but still is older than the middle Miocene platyrrhine primates from La Venta, Colombia, in particular the pitheciins Nuciruptor and Cebupithecia. The material is recognized as a new genus and species of Pitheciinae, Propithecia neuquenensis. The mesiodistally compressed, high-crowned incisors are specialized and similar to species in the tribe Pithecini and to the nonpitheciin Soriacebus (early Miocene, Patagonia). We rule out a phylogenetic relationship to the latter because of differences in molar morphology. Propithecia does, however, fit well into the pattern of pitheciin evolution, being more derived than the middle Miocene pitheciin Nuciruptor but not as much as another middle Miocene taxon, Cebupithecia. As such, this makes Propithecia the oldest taxon that can be confidently placed within this modern New World monkey subfamily. By analogy with the molar structures and diets of extant platyrrhines, Propithecia has a molar structure consistent with a variety of low-fiber diets ranging from fruit and gum to seeds. Its incisors suggest seed-eating in much the same way as extant pitheciins, like Pithecia. The talus resembles that of Callicebus, suggesting arboreal quadrupedal locomotion.     

Evolution of thyroid hormone binding by transthyretins in birds and mammals

Transthyretin, a protein synthesized and secreted by the choroid plexus and liver, binds thyroid hormones in extracellular compartments. This binding prevents accumulation of thyroid hormones in the lipids of membranes, establishing extracellular thyroid hormone pools for the distribution of the hormones throughout the body and brain. The N-termini of the transthyretin subunits are longer and more hydrophobic in chicken than in eutherian transthyretins. Here, we show that this is a general structural feature of avian transthyretins. Systematic changes of protein structure during evolution result from selection pressure leading to changes in function. The evolution of transthyretin function, namely, the binding of thyroid hormones, was studied in nine vertebrate species. The affinity of thyroxine binding to transthyretin is lowest in avians (mean Kd of about 30 nm), intermediate in metatherians (mean Kd of about 17 nm) and highest in eutherians (mean Kd of about 11 nm). The affinity for 3,5,3'-triiodothyronine shows an opposite trend, being four times higher for avian transthyretins than for mammalian transthyretins.     

A revised systematic scheme for the Eurasian Miocene fossil Hominidae

The current systematics of the European hominids has recently been reexamined in light of new and important fossil discoveries. It has been argued by Begun (1992a) that the European dryopithecines (including Graecopithecus from Greece) represent one of the earliest members of the Hominidae. Andrews (1992) and Dean & Delson (1992) agree that Graecopithecus should probably be placed within the Homininae, but that Dryopithecus should be excluded from this subfamily and retained within the Dryopithecinae. De Bonis & Koufos (1993, 1994) suggest that Graecopithecus shares a sister-group relationship with hominins. Moyà-Solà & K?hler (1993, 1995), however, argue that Dryopithecus and Graecopithecus both represent primitive members of the Ponginae. A phylogenetic and taxonomic revision of these taxa presented here supports the allocation of Graecopithecus to Homininae. It is also shown that the apparent disagreement concerning Dryopithecus is a result of the presence of two distinct genera within the fossil samples examined. It is concluded that the Hungarian hominid should be retained within the genus Dryopithecus, and the subfamily Dryopithecinae; the Spanish hominids are allocated to the resurrected genus Hispanopithecus Villalta & Crusafont-Pairó, 1944 which is likely to be a primitive member of the Ponginae.     

Evidence for a clade of nematodes, arthropods and other moulting animals

The arthropods constitute the most diverse animal group, but, despite their rich fossil record and a century of study, their phylogenetic relationships remain unclear. Taxa previously proposed to be sister groups to the arthropods include Annelida, Onychophora, Tardigrada and others, but hypotheses of phylogenetic relationships have been conflicting. For example, onychophorans, like arthropods, moult periodically, have an arthropod arrangement of haemocoel, and have been related to arthropods in morphological and mitochondrial DNA sequence analyses. Like annelids, they possess segmental nephridia and muscles that are a combination of smooth and obliquely striated fibres. Our phylogenetic analysis of 18S ribosomal DNA sequences indicates a close relationship between arthropods, nematodes and all other moulting phyla. The results suggest that ecdysis (moulting) arose once and support the idea of a new clade, Ecdysozoa, containing moulting animals: arthropods, tardigrades, onychophorans, nematodes, nematomorphs, kinorhynchs and priapulids. No support is found for a clade of segmented animals, the Articulata, uniting annelids with arthropods. The hypothesis that nematodes are related to arthropods has important implications for developmental genetic studies using as model systems the nematode Caenorhabditis elegans and the arthropod Drosophila melanogaster, which are generally held to be phylogenetically distant from each other.     

A tribosphenic mammal from the Mesozoic of Australia

A small, well-preserved dentary of a tribosphenic mammal with the most posterior premolar and all three molars in place has been found in Aptian (Early Cretaceous) rocks of southeastern Australia. In most respects, dental and mandibular anatomy of the specimen is similar to that of primitive placental mammals. With the possible exception of a single tooth reported as Eocene in age, terrestrial placentals are otherwise unknown in Australia until the Pliocene. This possible Australian placental is similar in age to Prokennalestes from the late Aptian/early Albian Khoboor Beds of Mongolia, the oldest currently accepted member of the infraclass Placentalia.     

Microsporidia are related to Fungi: evidence from the largest subunit of RNA polymerase II and other proteins

We have determined complete gene sequences encoding the largest subunit of the RNA polymerase II (RBP1) from two Microsporidia, Vairimorpha necatrix and Nosema locustae. Phylogenetic analyses of these and other RPB1 sequences strongly support the notion that Microsporidia are not early-diverging eukaryotes but instead are specifically related to Fungi. Our reexamination of elongation factors EF-1alpha and EF-2 sequence data that had previously been taken as support for an early (Archezoan) divergence of these amitochondriate protists show such support to be weak and likely caused by artifacts in phylogenetic analyses. These EF data sets are, in fact, not inconsistent with a Microsporidia + Fungi relationship. In addition, we show that none of these proteins strongly support a deep divergence of Parabasalia and Metamonada, the other amitochondriate protist groups currently thought to compose early branches. Thus, the phylogenetic placement among eukaryotes for these protist taxa is in need of further critical examination.     

Patterns of postnatal development in skulls of lynxes, genus Lynx (Mammalia: Carnivora)

Studies on ossification patterns and other ontogenetic events associated with postnatal cranial growth of wild felids are scarce. An analysis of developmental processes undergone by several cranial structures (presphenoidal and sphenooccipital synchondroses, temporal and sagittal crests, and deciduous and permanent teeth) during postnatal growth has been conducted on a sample of 336 specimens belonging to the four Recent species of lynxes (Lynx pardinus, Lynx lynx, Lynx rufus, and Lynx canadensis). Age has been estimated based on tooth replacement, skull size, and by counting the annual lines of cementum growth. Comparison of the results obtained for each of the four species reveal (1) a single pattern for both tooth replacement and ossification of the sphenooccipital synchondrosis, (2) two ossification patterns for the presphenoidal synchondrosis, (3) a common pattern for development of temporal ridges and sagittal crest showing different degrees of morphological expression, and (4) evidence suggesting the involvement of a heterochronic process, neoteny, in the morphological differentiation of several populations and species of the genus Lynx. These data also support the hypothesis that processes involved in the replacement of carnassials are based on functional requirements.