The Phylogenetic Relationships of Eucynodontia (Amniota: Synapsida)

The phylogeny of Eucynodontia is an important topic in vertebrate paleontology and is the foundation for understanding the origin of mammals. However, consensus on the phylogeny of Eucynodontia remains elusive. To clarify their interrelationships, a cladistic analysis, based on 145 characters and 31 species, and intergrating most prior works, was performed. The monophyly of Eucynodontia is confirmed, although the results slightly differ from those of previous analyses with respect to the composition of both Cynognathia and Probainognathia. This is also the first numerical cladistic analysis to recover a monophyletic Traversodontidae. Brasilodon is the plesiomorphic sister taxon of Mammalia, although it is younger than the oldest mammals and is specialized in some characters. A monophyletic Prozostrodontia, including tritheledontids, tritylodontids, and mammals, is well supported by many characters. Pruning highly incomplete taxa generally has little effect on the inferred pattern of relationships among the more complete taxa, although exceptions sometimes occur when basal fragmentary taxa are removed. Taxon sampling of the current data matrix shows that taxon sampling was poor in some previous studies, implying that their results are not reliable. Two major unresolved questions in cynodont phylogenetics are whether tritylodontids are more closely related to mammals or to traversodontids, and whether tritylodontids or tritheledontids are closer to mammals. Analyses of possible synapomorphies support a relatively close relationship between mammals and tritylodontids, to the exclusion of traversodontids, but do not clearly indicate whether or not tritheledontids are closer to mammals than are tritylodontids.     

Origin and relationships of the Ictidosauria to non-mammalian cynodonts and mammals

Abstract

Ictidosaurian genera are allocated to two families, Tritheledontidae and Therioherpetidae. This paper provides a diagnosis for Ictidosauria. The previously named family Brasilodontidae is shown to be a junior synonym of a family, Therioherpetidae. It is concluded that Ictidosauria originated from Late Permian procynosuchid non-mammalian cynodonts rather than from Middle Triassic probainognathid non-mammalian cynodonts. The structure of the skull and jaws of a derived traversodontid Ischignathus sudamericanus from the early Late Triassic of Argentina supports an earlier view that tritylodontids are more closely related to traversodontid than probainognathid non-mammalian cynodonts. Tritylodontids should not be included in Ictidosauria, nor should they considered to be a sister group to mammaliaforms.     

MOLAR OCCLUSION IN LATE TRIASSIC MAMMALS

1. A new genus and species of late Triassic mammal, Megazostrodon rudnerae, from Lesotho in southern Africa is described. The molars are similar to those of the British Eozostrodon parvus except that they are slightly larger and the upper molars have a large external cingulum supporting well-developed cusps. 2. Molar occlusion is discussed in two groups of late Triassic mammals: Eoxostrodon and the closely related Megazostrodon on one the hand and the unnamed primitive symmetrodonts on the other. It is shown that in Eoxostrodon the upper and lower molars did not have matching occlusal surfaces upon eruption but that wear produced matching occlusal surfaces. These surfaces are confined to the internal surface of the upper molars and the external surface of the lower molars and form a series of wide-angled triangles. The main cusp of an upper molar occluded between the main and posterior subsidiary cusp of the lower molar and the main cusp of the lower molar occluded between the main and anterior subsidiary cusp of the upper molar, 3. It is shown that the molars of Docodon and HaIdanodon were possibly derived from those of a primitive mammal such as Eozostrodon. The transition involved the development on the upper molars of an internal extension which, as it increased in size, established contact with the dorsal surfaces of two adjacent lower molars. The process involved is fundamentally different from that leading to tribosphenic molars. 4. In Megaxostrodon the main cusp of the upper molars occluded between the posterior and anterior subsidiary cusps of two adjacent lower molars, i.e. more posteriorly than in Eozostrodon. Primitive Rhaetic symmetrodonts were derived from mammals which had this type of occlusion and which were also closely related to Eoxostrodon and Megaxostrodon. The transition involved a rotation of the subsidiary cusps of the upper molars externally and those of the lower molars internally. This rotation increased the shearing surfaces between occluding upper and lower molars. Cusp rotation was carried further in the acute-angled symmetrodonts (Peralestes and Spalacotherium) and pantotheres. It appears that marked cusp rotation was coupled with the acquisition of transverse movements of the lower jaw during mastication. Transverse movement was apparently not possible in cynodonts, in Eoxostrodon (and related forms) and in Docodon. 5. The evolution of therian molars involves cusp rotation as originally proposed by the Cope—Osborn theory. Criticisms of the Cope—Osborn theory are re-evaluated in light of the new late Triassic material. 6. In Rhaetic symmetrodonts, molar wear produces matching occlusal facets, but the amount of attrition necessary to produce these facets was considerably less than in Eoxostrodon. In acute-angled symmetrodonts and in pantotheres, the molars erupt with more precise occlusal surfaces and attrition was not necessary to produce matching surfaces. 7. On the basis of the structure of the molar teeth it was concluded that Eozostrodon, Megazostrodon and Erythrotherium were closely related to the Rhaetic symmetrodonts. Slightly different occlusal relationships between upper and lower molars indicated that in these early mammals constant occlusal relations were being established. 8. Primitive cynodonts, such as Thrinaxodon, are characterized by alternate tooth replacement; there is a total lack of a constant occlusal relationship between upper and lower postcanine teeth. In Thrinaxodon individual postcanines were replaced several times. The crown structures of successive generations of postcanines were different so that a freshly erupted postcanine tooth had a crown structure quite distinct from the tooth which it replaced. It has been shown that the crown structure of one of the generations of postcanine teeth of Thrinaxodon is almost identical to that of Eozostrodon except that Thrinaxodon postcanines have a single root, On the basis of this similarity and the over-all structure of the primitive cynodont skull, it was concluded that Rhaetic mammals (excluding ictidosaurs and haramyids) could be derived from primitive cynodonts. 9. All the orders of Jurassic mammals (with the possible exception of multituber-culates) were probably derived from late Triassic mammals. The apparent close relationship of late Triassic mammals is evidence of a monophyletic origin of this class.     

Callitrichids as phyletic dwarfs,and the place of the callitrichidae in platyrrhini

Cebuella, Callithrix, Leontopithecus, andSaguinus share five distinguishing features. All of these features are best interpreted as derived character states within Platyrrhini, and these animals are phyletic dwarfs. These derived traits may form a single complex that evolved as a result of dwarfing. Two changes in the dentition are shown to be correlated with dwarfing in mammals. These four platyrrhine genera may or may not form a monophyletic group. It is suggested thatCallimico is an “incipient dwarf platyrrhine.” Causes of dwarfing in mammals are discussed.     

Utilization of food resources by small and medium‐sized mammals in the Monte Desert biome,Argentina

Theoretical models of species coexistence between desert mammals have generally been based on a combination of food and microhabitat selection by granivorous rodents. Although these models are applicable in various deserts of the world, they cannot explain resource use by mammals in Neotropical deserts. The present study examines diet composition in a mammal assemblage in the Monte desert, Argentina. The results show that two main strategies are used by these mammals: medium‐sized species (hystricognath rodents: Dolichotis patagonum, Lagostomus maximus, Microcavia australis and Galea musteloides; and an exotic lagomorph: Lepus europaeus) are herbivores, whereas small‐sized species (a marsupial: Thylamys pusillus; and sigmodontine rodents: Graomys griseoflavus, Akodon molinae, Calomys musculinus, Eligmodontia typus) are omnivorous. Small mammals also show a tendency towards granivory (C. musculinus), insectivory (A. molinae and T. pusillus) and folivory (G. griseoflavus).     

Composition des faunes de mammifères quaternaires en Chine selon un gradient Nord-Sud

During Pleistocene Epoch, quite a number of meridionale mammals inhabited north China, such as Stegodon, Macaca, Hystrix, Dicerorhinus, Bubalus and several kinds of other probscideans. On the contrary, some boreal mammals once appeared in south China, such as Megaloceros and Ursus arctos etc. The frequent appearance of warm-adapted mammals in North China can be explained by two alternative assumptions: one is that those mammals were of palearctic origin; the other is the climate-driven northward dispersal. According to the fossil evidences, it seems that the northward dispersal events were much more frequent than the southward ones during Pleistocene in China. It maybe means that during Pleistocene, the temperature fluctuated very frequently and the boundary between the Paleartic and Oriental Regions were always keeping on shifting. But it's still not clear how many episodes or phases of such dispersal events had occurred. The Qinling Mountains weren't responsible for the faunal differentiation between the North and the South, neither was the Yangtze River; because the former is not big enough, and the latter cannot be regarded as an efficient barrier for those animals, which are capable of flying and swimming. Maybe the climate zonation is the root cause of zoogeographical changes.     

The origin and evolution of genomic imprinting and viviparity in mammals

Genomic imprinting is widespread in eutherian mammals. Marsupial mammals also have genomic imprinting, but in fewer loci. It has long been thought that genomic imprinting is somehow related to placentation and/or viviparity in mammals, although neither is restricted to mammals. Most imprinted genes are expressed in the placenta. There is no evidence for genomic imprinting in the egg-laying monotreme mammals, despite their short-lived placenta that transfers nutrients from mother to embryo. Post natal genomic imprinting also occurs, especially in the brain. However, little attention has been paid to the primary source of nutrition in the neonate in all mammals, the mammary gland. Differentially methylated regions (DMRs) play an important role as imprinting control centres in each imprinted region which usually comprises both paternally and maternally expressed genes (PEGs and MEGs). The DMR is established in the male or female germline (the gDMR). Comprehensive comparative genome studies demonstrated that two imprinted regions, PEG10 and IGF2-H19, are conserved in both marsupials and eutherians and that PEG10 and H19 DMRs emerged in the therian ancestor at least 160 Ma, indicating the ancestral origin of genomic imprinting during therian mammal evolution. Importantly, these regions are known to be deeply involved in placental and embryonic growth. It appears that most maternal gDMRs are always associated with imprinting in eutherian mammals, but emerged at differing times during mammalian evolution. Thus, genomic imprinting could evolve from a defence mechanism against transposable elements that depended on DNA methylation established in germ cells.     

Review of negative effects of introduced rodents on small mammals on islands

In this first comprehensive review of negative effects of introduced rodents on insular small mammals, the focal species Rattus rattus, R. norvegicus, R. exulans and Mus musculus are implicated in at least 11 extinctions. Furthermore, removal experiments, eradication campaigns and control programmes provide evidence for negative effects on extant populations. While data are currently insufficient for meaningful generalisation with regard to the most threatening rodents, the most threatened small mammals, and the true extent of the problem, it is interesting that R. rattus is implicated in the majority of impacts. This may be explained by its extensive distribution and ecological plasticity. I conclude with methodological recommendations to guide data collection for impact quantification and the study of impact mechanism. This information should facilitate the prioritisation and justification of eradication campaigns, control programmes and biosecurity measures while ensuring that much-needed attention is paid to the conservation of insular small mammals.     

A comprehensive morphological analysis of talpid moles (Mammalia) phylogenetic relationships

Some talpid moles show one of the most specialized suites of morphological characters seen among small mammals. Fossorial and more generalized shrew‐looking moles inhabit both North America and Eurasia but these land masses share none of the same genera. One of the central questions of mole evolution has been that of how many times specialized fossorial habits evolved. We investigated the origin of mole characters with a maximum parsimony analysis of 157 characters, mostly craniodental and postcranial, of representatives of all 17 living mole genera and three shrews and one hedgehog as outgroups. The result was one most‐parsimonious tree and its most novel aspect was the position of a Japanese shrew mole clade (Urotrichus, Dymecodon), which branched off after Uropsilus and was not closely related to the American shrew mole (Neurotrichus). The desmans (Galemys and Desmana) were the next clade in the tree, followed by Neurotrichus. We confirmed the monophyly of the Eurasian fossorial mole clade Talpini (Euroscaptor, Parascaptor, Mogera, Scaptochirus and Talpa). Condylura, the star‐nosed mole from North America, was sister group to a clade consisting of the Talpini plus Scaptonyx and the Scalopini (Scalopus, Scapanus, Parascalops, and Scapanulus). Based on our results and on the assumption that moles originated in Eurasia, it is most parsimonious to infer one migration from Eurasia to North America and two back‐migrations to Eurasia. It is ambiguous if Talpini and Scalopini evolved their full fossorial habits independently or not. © The Willi Hennig Society 2006.     

Identification and expression of an elastase homologue in <Emphasis Type="Italic">Branchiostoma belcheri</Emphasis> with implications to the origin of vertebrate pancreas

Elastases have been identified in a variety of organisms ranging from bacteria to insects to mammals, yet little is known to date about them in amphioxus, a model animal for insights into the origin and evolution of vertebrates. In this study we demonstrate the presence of an elastase homologue, named BbElas, in Branchiostoma belcheri. The recombinant BbElas hydrolyses the elastase specific substrate N-succinyl-Ala-Ala-Ala p-nitroanilide, which can be inhibited by the serine proteinase inhibitor PMSF, the elastase-specific inhibitor elastatinal and the cysteine proteinase inhibitor PCMB. Phylogenetic analysis shows that BbElas represents the archetype of vertebrate elastases, hinting at the clues that the different isoforms of vertebrate elastases are originated from an ancestral gene like BbElas. Our results also suggest that the mid-gut in amphioxus is to homologous vertebrate pancreas, a novel proposal which deserves further study.     

The end of an 80-million year experiment: a review of evidence describing the impact of introduced rodents on New Zealand’s ‘mammal-free’ invertebrate fauna

Since separating from its super-continental origin 80 million years ago, New Zealand has effectively been isolated from the impacts of terrestrial mammals. The arrival of Polynesians in 13th C heralded the end of this era, with the introduction of kiore, (Rattus exulans, or Pacific rat), which had far-reaching effects on plant regeneration, survival of small ground vertebrates, larger invertebrates, and seabird breeding colonies. This paper reviews the evidence available from raptor nest sites and Quaternary beetle fossils to summarise extinctions thought to be caused by kiore in New Zealand. It also utilises invertebrate comparisons between islands with and without rats, or where rats have been eradicated, in order to document the impacts of rats (R. exulans, R. norvegicus) on invertebrate abundance, body mass, and the behavioural responses of some large New Zealand insects to the presence of rats. The role of a ‘mammal-free’ evolutionary history is discussed.     

Population dynamics of three Neotropical small mammals: Time series models and the role of delayed density-dependence in population irruptions

It is widely believed that only precipitation levels (through increased primary production) determine irruptions of small mammals in semi-arid areas of western South America. Nevertheless, density-dependent factors may also drive population fluctuations. To test statistically these putative effects we analysed 11 years of population records on three sympatric species of small mammals at two different habitat types in north central Chile. We applied the classical diagnostic tools of time series analysis (the autocorrelation function: ACF) to the observed time series of three neotropical small mammals. We also used simple linear autoregressive time series models to reconstruct the endogenous dynamics of these populations. The analysis strongly suggests that population fluctuations of the three species have an important density-dependent component, with the most irruptive species (Phyllotis darwini, Waterhouse 1837) displaying stronger second order population feedbacks than the other two (Akodon olivaceus, Waterhouse 1837 and Thylamys elegans, Waterhouse 1839). The latter two species showed direct density-dependent feedbacks. We hypothesize that the frequent population outbreaks of P. darwini (and perhaps of other species) in semi-arid regions of western South America, may be the result of population-level (direct density- dependence) and community-level processes (delayed density-dependence), interacting with exogenous perturbations (rainfall and associated primary production).     

Evolution of the mandibular symphysis in Notharctinae (Adapidae,Primates)

The Eocene Notharctinae provide a record of increasing fusion of the mandibular symphysis. The two sympatric genera,Notharctus andSmilodectes, differed through time in two respects.Notharctus increased in body size and evolved a partially fused mandibular symphysis.Smilodectes changed little in body size and retained an unfused symphysis. Similarities in molar morphology between these two genera and extant leaf-eating mammals suggest thatNotharctus andSmilodectes were specialized for folivory, a dietary regime correlated with partial symphyseal fusion in many extant mammals. It is concluded that the presence and the extant of symphyseal fusion is a function of body size, diet, and jaw mechanics, complicated by lineagespecific factors that vary among higher mammalian taxa.     

Locking Yourself Out: Diversity Among Dentally Zalambdodont Therian Mammals

We review the evolution of dental zalambdodonty across therian mammals. Among zalambdodonts, there is little or no occlusion between the protocone and talonid basin and one of the central cusps of the upper molars (metacone or paracone) and the talonid basin of the lower molars are lost or reduced. Over two dozen genera of therian mammals show zalambdodont molars, including tenrecids, chrysochlorids, Solenodon, the marsupial mole Notoryctes, the extinct placentals Apternodus, Oligoryctes, Parapternodus, and Koniaryctes, and the extinct marsupials Necrolestes and Yalkaparidon. The chiropteran Harpiocephalus is nearly zalambdodont. Transformation series provided by paleontological or ontogenetic data, plus occlusal relationships, can be used to determine the homology of molar cusps in zalambdodont taxa. Zalambdodont dental specialization does not appear to have led to ecological specialization. With the important exception of golden moles and tenrecs, dentally zalambdodont taxa are less speciose than their nonzalambdodont sister taxa.     

Calcitonin receptor-stimulating peptide: Its evolutionary and functional relationship with calcitonin/calcitonin gene-related peptide based on gene structure

This review focuses on the evolutionary and functional relationship of calcitonin receptor-stimulating peptide (CRSP) with calcitonin (CT)/calcitonin gene-related peptide (CGRP) in mammals. CRSP shows high sequence identity with CGRP, but distinct biological properties. CRSP genes (CRSPs) have been identified in mammals such as pigs and dogs of the Laurasiatheria, but not in primates and rodents of the Euarchontoglires or in non-placental mammals. CRSPs have genomic organizations highly similar to those of CT/CGRP genes (CT/CGRPs), which are located along with CGRPs in a locus between CYP2R1 and INSC, while the other members of the CGRP superfamily, adrenomedullin and amylin, show genomic organizations and locations distinct from CT, CGRP, and CRSP. Thus, we categorized these three peptides into the CT/CGRP/CRSP family. Non-placental mammals having one and placental mammals having multiple CT/CGRP/CRSP family genes suggests that multiplicity of CT/CGRP started at an early stage of mammalian evolution. In the placental mammals, Laurasiatheria generally possesses multiple CRSPs and only one CT/CGRP, while Euarchontoglires possesses CT/CGRP and CGRPβ but no CRSP, indicating an increase in the diversity and multiplicity of this family of genes in mammalian evolution. Phylogenetic analysis suggests that some CRSPs have been generated very recently in mammalian evolution. Taken together, the increase in the number and complexity of the CT/CGRP/CRSP family genes may have due to evolutionary pressure to facilitate adaptation during mammalian evolution. In this regard, it is important to elucidate the physiological roles of CT, CGRP and CRSP from the viewpoint of the CT/CGRP/CRSP family even in Euarchontoglires.     

Genetic Heterogeneity in Wild Isolates of Cellular Slime Mold Social Groups

This study addresses the issues of spatial distribution, dispersal, and genetic heterogeneity in social groups of the cellular slime molds (CSMs). The CSMs are soil amoebae with an unusual life cycle that consists of alternating solitary and social phases. Because the social phase involves division of labor with what appears to be an extreme form of “altruism”, the CSMs raise interesting evolutionary questions regarding the origin and maintenance of sociality. Knowledge of the genetic structure of social groups in the wild is necessary for answering these questions. We confirm that CSMs are widespread in undisturbed forest soil from South India. They are dispersed over long distances via the dung of a variety of large mammals. Consistent with this mode of dispersal, most social groups in the two species examined for detailed study, Dictyostelium giganteum and Dictyostelium purpureum, are multi-clonal.     

Genome‐wide adaptive evolution to underground stresses in subterranean mammals: Hypoxia adaption,immunity promotion,and sensory specialization

Life underground has provided remarkable examples of adaptive evolution in subterranean mammals; however, genome‐wide adaptive evolution to underground stresses still needs further research. There are approximately 250 species of subterranean mammals across three suborders and six families. These species not only inhabit hypoxic and dark burrows but also exhibit evolved adaptation to hypoxia, cancer resistance, and specialized sensory systems, making them an excellent model of evolution. The adaptive evolution of subterranean mammals has attracted great attention and needs further study. In the present study, phylogenetic analysis of 5,853 single‐copy orthologous gene families of five subterranean mammals (Nannospalax galili, Heterocephalus glaber, Fukomys damarensis, Condylura cristata, and Chrysochloris asiatica) showed that they formed fou distinct clusters. This result is consistent with the traditional systematics of these species. Furthermore, comparison of the high‐quality genomes of these five subterranean mammalian species led to the identification of the genomic signatures of adaptive evolution. Our results show that the five subterranean mammalian did not share positively selected genes but had similar functional enrichment categories, including hypoxia tolerance, immunity promotion, and sensory specialization, which adapted to the environment of underground stresses. Moreover, variations in soil hardness, climate, and lifestyles have resulted in different molecular mechanisms of adaptation to the hypoxic environment and different degrees of visual degradation. These results provide insights into the genome‐wide adaptive evolution to underground stresses in subterranean mammals, with special focus on the characteristics of hypoxia adaption, immunity promotion, and sensory specialization response to the life underground.     

New Studies of Enamel Microstructure in Mesozoic Mammals: A Review of Enamel Prisms as a Mammalian Synapomorphy

Characters from enamel microstructure have not been used in recent phylogenetic analyses of Mesozoic Mammalia. Reasons are that enamel characters have been perceived as (A) variable without regard to systematic position of taxa, (B) inconsistently reported within the literature, and (C) simply scored as either prismatic or not prismatic in earlier mammals. Our work on Mesozoic mammals such as Sinoconodon, Gobiconodon, Triconodontidae, Docodon, Laolestes, and others suggests that synapsid columnar enamel (SCE) structure was easily transformed into plesiomorphic prismatic enamel (PPE) and that PPE may be described with at least five independent character states. Two PPE characters—a flat, open prism sheath and a planar prism seam—were present in the cynodont Pachygenelus and in several Jurassic and Cretaceous mammals. We propose that appearance of a prism sheath transforms SCE into PPE and that reduction and loss of a prism sheath reverse PPE into SCE, in both phylogeny and ontogeny. We further propose that no amniote vertebrates other than the trithelodontid cynodont, Pachygenelus, plus Mammalia have ever evolved an ameloblastic Tomes process capable of secreting PPE and that the genetic potential to secrete PPE is a synapomorphy of Pachygenelus plus Mammalia, whether or not all lineages of the clade have expressed that potential.