The Inner Ear of Two Late Cretaceous Multituberculate Mammals, and Its Implications for Multituberculate Hearing

The inner ear of the Late Cretaceous multituberculates Nemegtbaatar gobiensis and Chulsan-baatar vulgaris is described from serial sections and enlarged models. The size and proportions of the inner ear as a whole are as expected for extant small mammals. The lengths of the cochlea (Nemegtbaatar gobiensis, 3.0 mm, Chulsanbaatar vulgaris, 2.0 mm) are comparable to those of other multituberculates, when ratios of length of the cochlea to skull length are calculated. The vestibule is not as expanded in the two taxa as in Lambdopsalis, ?Meniscoessus, and ?Catopsalis; the estimated volume for Nemegtbaatar gobiensis is 9 mm³. A slightly laterally curved, anteriomedially directed cochlea, relatively robust ear ossicles, and the estimations of the area of the tympanic membrane and stapedial footplate in Chulsanbaatar suggest high-frequency hearing but a relatively low sensitivity to low-decibel sounds. The semicircular canals of Nemegtbaatar and Chulsanbaatar are fully developed; the size of the anterior, posterior, and lateral canals and their angles and proportions are comparable to those of extant mammals of similar size. The anterior semicircular canal of Nemegtbaatar forms a smooth half-circle and thus is more derived than the angular canal of Ornithorhynchus. The notable differences between the ratio of the width of the lateral semicircular canal to skull length and the size of the vestibule in Nemegtbaatar and the Paleocene multituberculate Lambdopsalis bulla are probably related to different modes of life.     

Shell Morphology of the Late Cretaceous Planktonic Foraminifera and Its Value in Modern Taxonomy

Planktonic foraminifers in the Late Cretaceous developed by the way of biological progress. Their taxonomic diversity increased, the shell morphology became more complicated, the number of individuals in the population grew, and the territorial dispersal of the group was active. This process had an intermittently-continuous pattern, as there was alternation of poly- and oligotaxic stages of development of the group. Each of these stages was characterized by the predominant morphotype of shell structure, a certain type of life strategy (k and r strategists), as well as by a certain type of evolutionary process (gradualism or punctualism). All these events are reflected in the systematics of the group, which is regularly updated.     

A Systematic Study on Tooth Enamel Microstructures of Lambdopsalis bulla (Multituberculate,Mammalia) - Implications for Multituberculate Biology and Phylogeny

Tooth enamel microstructure is a reliable and widely used indicator of dietary interpretations and data for phylogenetic reconstruction, if all levels of variability are investigated. It is usually difficult to have a thorough examination at all levels of enamel structures for any mammals, especially for the early mammals, which are commonly represented by sparse specimens. Because of the random preservation of specimens, enamel microstructures from different teeth in various species are often compared. There are few examples that convincingly show intraspecific variation of tooth enamel microstructure in full dentition of a species, including multituberculates. Here we present a systematic survey of tooth enamel microstructures of Lambdopsalis bulla, a taeniolabidoid multituberculate from the Late Paleocene Nomogen Formation, Inner Mongolia. We examined enamel structures at all hierarchical levels. The samples are treated differently in section orientations and acid preparation and examined using different imaging methods. The results show that, except for preparation artifacts, the crystallites, enamel types, Schmelzmuster and dentition types of Lambdopsalis are relatively consistent in all permanent teeth, but the prism type, including the prism shape, size and density, may vary in different portions of a single tooth or among different teeth of an individual animal. The most common Schmelzmuster of the permanent teeth in Lambdopsalis is a combination of radial enamel in the inner and middle layers, aprismatic enamel in the outer layer, and irregular decussations in tooth crown area with great curvature. The prism seam is another comparably stable characteristic that may be a useful feature for multituberculate taxonomy. The systematic documentation of enamel structures in Lambdopsalis may be generalized for the enamel microstructure study, and thus for taxonomy and phylogenetic reconstruction, of multituberculates and even informative for the enamel study of other early mammals.     

Phylogeny and Systematics of Multituberculate Mammals

We present a synopsis of high-rank multituberculate systematicsand a manually generated cladogram illustrating multituberculate interrelationships. We divide the Multituberculata into the paraphyletic suborder 'Plagiaulacida', an apparently monophyletic suborder Cimolodonta, and one family incertae sedis. Within 'Plagiaulacida' we recognise three informal lines: paulchoffatiid (three families), plagiaulacid (three families) and allodontid (two families and the genus Glirodon). The Cimolodonta are divided into an informal Paracimexomys group; three superfamilies: Ptilodontoidea, Djadochtatherioidea (new), and Taeniolabidoidea (restricted to Taeniolabididae); and five families (superfamily incertae sedis): Eucosmodontidae, Microcosmodontidae, Cimolodontidae, Boffiidae, and Kogaionidae; and some genera incertae sedis. New characters used in our analysis are (1) a tendency of molar cusps to coalesce; and (2) ornamentation of grooves, pits, and ridges on the molars. We argue that the Ptilodontoidea, and less certainly also the Cimolodontidae and Boffiidae, might have originated from amongthe plagiaulacid line, a possible intermediate link being the Paracimexomys group. The remaining Cimolodonta might have originated from unknown members of the Paracimexomys group with separated molar cusps and smooth enamel. The origin of two types of prismatic enamel and a relationship between them are stumbling blocks in understanding the origin of the Cimolodonta; we conclude that microprismatic enamel made its appearance only once. Revised diagnoses of high-rank multituberculate taxa, including lists of all known genera, are given.     

Tooth Enamel Microstructure of Living and Extinct Hyracoids Reveals Unique Enamel Types Among Mammals

Among medium- to large-sized terrestrial ‘ungulates,’ there is often a relationship between increasing body size, correlated changes in diet, and increased complexity of the enamel microstructures [notably the development of Hunter-Schreger bands (HSB)]. An exhaustive survey of the enamel microstructures of living and extinct Hyracoidea demonstrates, however, that the Schmelzmuster within this order of mammals is generally one-layered and formed by radial enamel despite a large range of body sizes and dietary adaptations; HSB are remarkably absent. Radial enamel is characteristic of early diverging hyracoids, as well as more derived members of the extinct families Geniohyidae and Pliohyracidae, and the extant Procaviidae. Only some large ‘Saghatheriidae,’ and all members of the family Titanohyracidae, developed a more complex enamel microstructure (i.e., with prisms decussating), a unique condition among Mammalia that we name ‘bundled enamel’ (BE). This structure is reminiscent to some degree of both the ‘Pyrotherium enamel’ and the ‘3D enamel’ of proboscideans. Hyracoids with BE represented a major component of the diversity of mid- to large-sized herbivores during the Paleogene in Africa. Like HSB, which are developed by most other ‘ungulates,’ the BE is regarded as a device for resisting propagation of cracks during mastication. Hyracoids never developed however the ‘modified radial enamel’ that is characteristic of most large and hypsodont perissodactyls and artiodactyls that entered Africa during the Miocene.     

On the Scaling of Tooth Size in Mammals

We must establish the allometric regularities of functionalscalingin interspecific, "mouse-to elephant" plots in orderto provide criteria for the recognition of special adaptationsunrelated to the requirements of size. The qualitative literaturesuggests that postcanine tooth areas of herbivorous mammalsshould increase with positive allometry in such plots. Thispositive allometry might reflect the demands of metabolism orthe ecological strategies of large vs. small hervivores embodiedin Levins' concept of environmental grain. Plots of postcaninearea vs. body size display the expected postive allometry inall groups studied: hystricomorph rodents, suine artiodactyls(pigs, peccaries, and hippos), cervoid artiodactyls (deer, s.l),and four groups of primates considered separately (lemuroids,ceboids, cercopithecoids, and great apes). Sketchy data foraustralopithecines also indicate positive allometry and therelatively larger cheek teeth of robust forms may only reflecttheir larger body size and not the dietary differences so oftenadvocated. Phyletic dwarfs of large herbivores display negativeallometry (relatively larger cheek teeth in dwarfs) in oppositionto the interspecific trend.     

Molecular Decay of the Tooth Gene Enamelin (ENAM) Mirrors the Loss of Enamel in the Fossil Record of Placental Mammals

Vestigial structures occur at both the anatomical and molecular levels, but studies documenting the co-occurrence of morphological degeneration in the fossil record and molecular decay in the genome are rare. Here, we use morphology, the fossil record, and phylogenetics to predict the occurrence of “molecular fossils” of the enamelin (ENAM) gene in four different orders of placental mammals (Tubulidentata, Pholidota, Cetacea, Xenarthra) with toothless and/or enamelless taxa. Our results support the “molecular fossil” hypothesis and demonstrate the occurrence of frameshift mutations and/or stop codons in all toothless and enamelless taxa. We then use a novel method based on selection intensity estimates for codons (ω) to calculate the timing of iterated enamel loss in the fossil record of aardvarks and pangolins, and further show that the molecular evolutionary history of ENAM predicts the occurrence of enamel in basal representatives of Xenarthra (sloths, anteaters, armadillos) even though frameshift mutations are ubiquitous in ENAM sequences of living xenarthrans. The molecular decay of ENAM parallels the morphological degeneration of enamel in the fossil record of placental mammals and provides manifest evidence for the predictive power of Darwin''s theory.     

Taxonomy,morphology and phylogeny of Late Cretaceous spirulid coleoids (Cephalopoda) from Greenland and Canada

Abstract: Groenlandibelus rosenkrantzi from the Maastrichtian of Greenland has long been thought to constitute an early representative of spirulid coleoids. This study shows that this view must be reassessed, at least in part. A re‐investigation of the types and of material recorded subsequently has revealed that none of these specimens is conspecific with the holotype of G. rosenkrantzi. Cyrtobelus birkelundae gen. nov, sp. nov. differs from the type of G. rosenkrantzi in having lower chambers and in lacking an apically elongated sheath. The longiconic phragmocone of G. rosenkrantzi has more features in common with the presumed spirulid genus Naefia. A specimen described in detail by J. A. Jeletzky in the mid 1960s as ‘G. rosenkrantzi’ is designated holotype of C. birkelundae sp. nov., which means that internal phragmocone features are still unknown in G. rosenkrantzi. Cyrtobelus hornbyense gen. nov, sp. nov. from the Campanian of western Canada constitutes the first record of early spirulids from the northeast Pacific, being based on seventeen extraordinarily well‐preserved phragmocones. This species differs from C. birkelundae sp. nov. only in the width of the siphuncular tube. The presence of a caecum, a nacre‐less conotheca that represents the continuation of the protoconch conotheca, conothecal flaps that anchor the mural parts of the septa, and a thin investment‐like sheath are characters shared only with Recent Spirula. In particular, the unusual protoconch architecture of Cyrtobelus gen. nov. challenges a phylogenetic origin within bactritoid‐like coleoids.     

Seed Size, Fruit Size, and Dispersal Systems in Angiosperms from the Early Cretaceous to the Late Tertiary

Fossil data from 25 angiosperm floras from the Early Cretaceous ( approximately 124 million years ago) to the Pliocene ( approximately 2 million years ago) were compiled to estimate sizes of seeds and fruits and the relative proportion of two different seed-dispersal systems by animals and by wind. The results suggest that, first, seed and fruit sizes were generally small during most of the Cretaceous, in agreement with previous suggestions, but the trend of increasing sizes started before the Cretaceous-Tertiary boundary; second, there was a decrease in both seed and fruit sizes during late Eocene and Oligocene, reaching a level that has continued to the Late Tertiary; third, the fraction of animal dispersal was, in contrast to previous suggestions, rather high also during the Cretaceous but increased drastically in the Early Tertiary and declined congruently with the declining seed and fruit sizes from the late Eocene; and fourth, the fraction of wind dispersal showed a bimodal pattern, being high in the Late Cretaceous and in the Oligocene-Miocene but with a drop in between. We find that the observed trends are only weakly related to the availability of animal fruit dispersers. Instead, the trends are congruent with a climate-driven change in environmental conditions for recruitment, where larger seeds are favored by closed forest vegetation. The prevalence of semiopen, dry, and probably herbivore-disturbed vegetation during the Cretaceous, the development of closed multistratal forests in the Eocene, and the later development of a more open vegetation and grasslands starting in the Oligocene-Miocene, are reflected in the distribution of angiosperm seed and fruit sizes and in the dispersal systems.     

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.     

Giants and Bizarres: Body Size of Some Southern South American Cretaceous Dinosaurs

Body masses of some South American dinosaurs are estimated. The sauropod Argentinosaurus huinculensis reached 73 tonnes, and therefore, is the largest of all land animals whose mass has been rigorously obtained. Another sauropod, Antarctosaurus giganteus, was the second largest, at nearly 69 tonnes, while Antarctosaurus wichmannianus reached 34 tonnes. A third sauropod, the bizarre-looking Amargasaurus cazaui, was much smaller, with a body mass of only 2.5 tonnes. Among theropods, the body mass of the strangely looking, horned Carnotaurus sastrei, was volumetrically estimated at 1.5 tonnes, while allometric equations on limb measurements yielded overestimations. Moreover, the holotype specimen of Giganotosaurus carolinii (MUCPv-CH-1) was about as large as the average-sized Tyrannosaurus rex, and only marginally smaller than “Sue”, the largest specimen. However, a new dentary of Giganotosaurus (MUCPv-95) is 8% longer than that of the holotype. Assuming geometric similarity, that individual must have had a body mass above 8 tonnes and hence must have been the largest theropod ever found.     

Genetic Patterns of Paternity and Testes Size in Mammals

Background

Testes size is used as a proxy of male intrasexual competition, with larger testes indicative of greater competition. It has been shown that in some taxa, social mating systems reflect variance in testes size, but results are not consistent, and instead it has been suggested that genetic patterns of mating may reflect testes size. However, there are different measures of genetic patterns of mating. Multiple paternity rates are the most widely used measure but are limited to species that produce multi-offspring litters, so, at least for group living species, other measures such as loss of paternity to males outside the social group (extra group paternity) or the proportion of offspring sired by the dominant male (alpha paternity) might be appropriate. This study examines the relationship between testes size and three genetic patterns of mating: multiple paternity, extragroup paternity and alpha paternity.

Methodology/Principal Findings

Using data from mammals, phylogenetically corrected general linear models demonstrate that both multiple paternity and alpha paternity, but not extra group paternity, relate to testes size. Testes size is greater in species with high multiple paternity rates, whereas the converse is found for alpha paternity. Additionally, length of mating season, ovulation mode and litter size significantly influenced testes size in one model.

Conclusions/Significance

These results demonstrate that patterns of mating (multiple paternity and alpha paternity rates) determined by genetic analysis can provide reliable indicators of male postcopulatory intrasexual competition (testes size), and that other variables (length of mating season, ovulation mode, litter size) may also be important.     

不同食性哺乳动物及人的牙釉质微结构对比研究

根据动物食性将搜集于西藏、新疆、江苏和吉林4省和自治区16种哺乳动物的581颗牙齿分为食草、杂食和食肉3类,设计了录像-截图-测量-定标4步微米量级牙釉质结构测量方法,测量得到牙釉质的颊面厚度和舌面厚度比、臼齿外轮廓平均牙釉质厚度比等牙釉质分布规律,观测到不同哺乳动物釉柱的组装直径均在100 μm 左右,但柱间晶体的宽度不同,发现3类哺乳动物牙釉柱与釉牙本质界夹角的规律:人和食肉动物狗的釉柱角度最大,接近90°,杂食动物猪釉柱角度约为70°,而食草动物釉柱角度在54°～68°之间.研究表明,不同食性哺乳动物牙齿外在和内在结构都和其生物力学功能密切相关,牙釉质的异型结构是使得釉质具有优异的力学性能的优化结构模式.     

Phylogeny,Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary

Background

Rhabdodontid ornithopod dinosaurs are characteristic elements of Late Cretaceous European vertebrate faunas and were previously collected from lower Campanian to Maastrichtian continental deposits. Phylogenetic analyses have placed rhabdodontids among basal ornithopods as the sister taxon to the clade consisting of Tenontosaurus, Dryosaurus, Camptosaurus, and Iguanodon. Recent studies considered Zalmoxes, the best known representative of the clade, to be significantly smaller than closely related ornithopods such as Tenontosaurus, Camptosaurus, or Rhabdodon, and concluded that it was probably an island dwarf that inhabited the Maastrichtian Haţeg Island.

Methodology/Principal Findings

Rhabdodontid remains from the Santonian of western Hungary provide evidence for a new, small-bodied form, which we assign to Mochlodon vorosi n. sp. The new species is most similar to the early Campanian M. suessi from Austria, and the close affinities of the two species is further supported by the results of a global phylogenetic analysis of ornithischian dinosaurs. Bone histological studies of representatives of all rhabdodontids indicate a similar adult body length of 1.6–1.8 m in the Hungarian and Austrian species, 2.4–2.5 m in the subadults of both Zalmoxes robustus and Z. shqiperorum and a much larger, 5–6 m adult body length in Rhabdodon. Phylogenetic mapping of femoral lengths onto the results of the phylogenetic analysis suggests a femoral length of around 340 mm as the ancestral state for Rhabdodontidae, close to the adult femoral lengths known for Zalmoxes (320–333 mm).

Conclusions/Significance

Our analysis of body size evolution does not support the hypothesis of autapomorhic nanism for Zalmoxes. However, Rhabdodon is reconstructed as having undergone autapomorphic giantism and the reconstructed small femoral length (245 mm) of Mochlodon is consistent with a reduction in size relative to the ancestral rhabdodontid condition. Our results imply a pre-Santonian divergence between western and eastern rhabdodontid lineages within the western Tethyan archipelago.     

Scale Effects between Body Size and Limb Design in Quadrupedal Mammals

Recently the metabolic cost of swinging the limbs has been found to be much greater than previously thought, raising the possibility that limb rotational inertia influences the energetics of locomotion. Larger mammals have a lower mass-specific cost of transport than smaller mammals. The scaling of the mass-specific cost of transport is partly explained by decreasing stride frequency with increasing body size; however, it is unknown if limb rotational inertia also influences the mass-specific cost of transport. Limb length and inertial properties – limb mass, center of mass (COM) position, moment of inertia, radius of gyration, and natural frequency – were measured in 44 species of terrestrial mammals, spanning eight taxonomic orders. Limb length increases disproportionately with body mass via positive allometry (length ∝ body mass^0.40); the positive allometry of limb length may help explain the scaling of the metabolic cost of transport. When scaled against body mass, forelimb inertial properties, apart from mass, scale with positive allometry. Fore- and hindlimb mass scale according to geometric similarity (limb mass ∝ body mass^1.0), as do the remaining hindlimb inertial properties. The positive allometry of limb length is largely the result of absolute differences in limb inertial properties between mammalian subgroups. Though likely detrimental to locomotor costs in large mammals, scale effects in limb inertial properties appear to be concomitant with scale effects in sensorimotor control and locomotor ability in terrestrial mammals. Across mammals, the forelimb''s potential for angular acceleration scales according to geometric similarity, whereas the hindlimb''s potential for angular acceleration scales with positive allometry.     

Mammals of south-east Asian islands and their Late Pleistocene environments

Aim The environments that existed in south‐east Asian islands during the last glacial are poorly known, limiting our understanding of mammalian biogeography in the region. The objective of this research is to investigate the ecological characteristics of mammal faunas on small islands, and to see whether the habitat requirements of the species in those faunas can be used to deduct the vegetation types that existed on islands before becoming isolated by rising sea levels. Location The maps presented here cover the small islands of tropical south‐east Asia, including the Burmese, Thai and Cambodian islands in the north, the islands off the coast of west Sumatra in the west, the islands around Java in the south, and the islands off the east coast of Borneo in the east, including the Philippine islands of Palawan and those in the Sulu Archipelago. Methods The presence records of mammal species on 215 small islands in the region were compiled, and the habitat requirements for each of these species was assessed (species that had probably been introduced by humans were excluded from the analysis). For each island location (longitude and latitude), maximum altitude of the island, total area, depth to nearest land, distance to nearest island, and distance to nearest mainland were assessed. Geographical and statistical analyses were used to investigate patterns of mammalian habitat requirements. Results The geographical analysis showed that forest‐dependent species, i.e. species that are only found in primary forest (lowland and mountainous), appear to be concentrated on islands off west Sumatra, in the Lingga and Riau Archipelagos, around Palawan, and around Bunguran Island; they are absent mostly from the islands of the Java Sea, those off the east coast of eastern Borneo, from most islands in the Sunda Strait, several islands in the northern South China Sea, and from all islands off the west coast of the Malay/Thai Peninsula and in the Gulf of Thailand. Species that generally occur outside primary forest, that is those in secondary forest, gardens, plantations and open areas mostly occurred on islands where the forest‐dependent species were absent. The statistical analysis showed that latitude and size of islands were important factors that determined the absence and presence of forest‐dependent species on small islands. Main conclusions The data suggest that during the last glacial there were several areas in the Sundaic region that remained forest covered: west of Sumatra, north‐west of Borneo, the Malacca Straits and around Palawan. Other areas may have been covered by more open vegetation types like tree savanna, or open deciduous forest: on and to the east of the Malay/Thai Peninsula, the Java Sea area, including the Sunda Strait, and eastern Borneo.     

Some Morphogenetic Factors in Taxonomy of Dinoflagellates

The formative ability of ectoplasm in situ and in isolation from the living cell is shown to be an autodynamic morphogenetic factor creating sui generis membrane structure in species. Heteromorphic division and formation of specific aberrants, which differ from the parent cell by characters which have been used to separate genera, may be caused by molecular changes in ectoplasm as a primary factor. The subpellicular vacuom reticulum in Gyrodinium and other genera originates possibly from the upset metabolism of protoplasm. It is purely phenotypic. Oxygen deficiency induces in Prorocentrum endogenous cysts and dwarf-cell formation, a plasmolysis-like effect. The morphological norm in dinoflagellates is maintained also by the angle of nuclear fission and ensuing cytoplasm disposition. It is constant in sea and culture in some species, almost constant in sea but multivariable in culture in other species. The static form in some species is related to a single method of division (Exuviaella marina). Polymorphy of Exuviaella, Prorocentrum and other genera is induced by the presence of many types of divisions, which also favours an abundance of aberrants. “Osmomorphoses” originate in critical salinity-temperature conditions owing to flexibility of the pellicle in Gymnodinioideae; it is prevented by the membrane rigidity in the thecate forms. Flexibility of membrane in juvenile cells favours exogenous division. Rigid membranes of adult cells inhibit exogenous, favour endogenous fission. “Cyclomorphoses”, “phagomorphoses” (feeding habit) and different types of aberrants are useful to complete classification in modern and fossil dinoflagellates since morphogenesis rules work regardless of time and space.