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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Updating and Recoding Enamel Microstructure in Mesozoic Mammals: In Search of Discrete Characters for Phylogenetic Reconstruction

The previously unknown enamel microstructure of a variety of Mesozoic and Paleogene mammals ranging from monotremes and docodonts to therians is described and characterized here. The novel information is used to explore the structural diversity of enamel in early mammals and to explore the impact of the new information for systematics. It is presently unclear whether enamel prisms arose several times during mammalian evolution or arose only once with several reversals to prismless structure. At least two undisputed reversions or simplifications are known—in the monotreme clade from Obdurodon to Ornithorhynchus (via Monotrematum?), and (perhaps more than once) within the clade from archaeocete to a variety of odontocete whales. Similarly, both prismatic and nonprismatic enamel is present among docodonts. Seven discrete characters showing enough morphological diversity to be of potential importance in phylogenetic reconstructions may be identified as a more appropriate summary of enamel microstructural diversity among mammaliaforms than the single character “prismatic enamel-present/absent” employed in recent matrices. Inclusion of five of these characters in the matrix of Luo et al. (2002) modifies the original topology by collapsing several nodes involving triconodonts and other nontribosphenic taxa. There is considerable support for prismatic enamel as a synapomorphy of trithelodonts plus Mammaliamorpha, and multituberculates appear to have small or “normal” sized prisms as the ancestral condition, with some (as yet) enigmatic changes to nonprismatic structure in some basal members of the group and the appearance of “gigantoprismatic” structure as an autapomorphic state of less inclusive clades. Other potential qualitative characters and the need for attaining appropriate methods to incorporate quantitative features may be important for future analyses.     

Similar Associations of Tooth Microwear and Morphology Indicate Similar Diet across Marsupial and Placental Mammals

Low-magnification microwear techniques have been used effectively to infer diets within many unrelated mammalian orders, but the extent to which patterns are comparable among such different groups, including long extinct mammal lineages, is unknown. Microwear patterns between ecologically equivalent placental and marsupial mammals are found to be statistically indistinguishable, indicating that microwear can be used to infer diet across the mammals. Microwear data were compared to body size and molar shearing crest length in order to develop a system to distinguish the diet of mammals. Insectivores and carnivores were difficult to distinguish from herbivores using microwear alone, but combining microwear data with body size estimates and tooth morphology provides robust dietary inferences. This approach is a powerful tool for dietary assessment of fossils from extinct lineages and from museum specimens of living species where field study would be difficult owing to the animal’s behavior, habitat, or conservation status.     

Structural Morphology of Molars in Large Mammalian Herbivores: Enamel Content Varies between Tooth Positions

The distribution of dental tissues in mammalian herbivores can be very different from taxon to taxon. While grazers tend to have more elaborated and complexly folded enamel ridges, browsers have less complex enamel ridges which can even be so far reduced that they are completely lost. The gradient in relative enamel content and complexity of structures has so far not been addressed within a single species. However, several studies have noted tooth position specific wear rates in small mammals (rabbits, guinea pigs) which may be related to individual tooth morphology. We investigate whether differentiated enamel content by tooth position is also to be found in large herbivores. We use CT-scanning techniques to quantify relative enamel content in upper and lower molar teeth of 21 large herbivorous mammal species. By using a broad approach and including both perissodactyls and artiodactyls, we address phylogenetic intraspecific differences in relative enamel content. We find that enamel is highly unevenly distributed among molars (upper M1, M2, M3 and lower m1, m2, m3) in most taxa and that relative enamel content is independent of phylogeny. Overall, relative enamel content increases along the molar tooth row and is significantly higher in lower molars compared to upper molars. We relate this differential enamel content to prolonged mineralisation in the posterior tooth positions and suggest a compensatory function of m3 and M3 for functional losses of anterior teeth.