Ontogenetic and interelemental study of appendicular bones of Caiman latirostris Daudin, 1802 sheds light on osteohistological variability in crocodylians.

The osteohistology of vertebrates provides a reliable source to deduce biological information, particularly regarding growth and development. Although osteohistological studies in Neosuchia (Crocodyliformes, Mesoeucrocodylia) are relatively numerous, the number of species studied within the group is still small. Extant crocodilians are known to exhibit intraspecific variability linked to environmental conditions, habitat, feeding, and other intrapopulation factors. Here, we analyzed the osteohistology of the living South American Caiman latirostris throughout posthatching ontogeny. The histology of several appendicular bones of 13 different-sized captive and wild individuals were examined. Although some thin sections revealed the classic lamellar, parallel-fibered, or woven bone matrices, others showed a variation and a mix between the organization of the bone tissue. These histological differences are likely related to variability in the growth dynamics of caimans. In some bones of the juveniles studied, remnants of embryonic bone were observed. Osteohistological variation related to prevailing environmental conditions is documented. Furthermore, our results show ontogenetic variation in the type of bone tissues deposited throughout the development of C. latirostris. This study offers a broad framework for life history interpretations for C. latirostris and provides insight into the evolutionary history and ontogenetic growth of extinct crocodylian lineages.

Palaeoecological deductions from osteohistology.

Palaeoecological deductions are vital for understanding the evolution and diversification of species within prehistoric environments. This review highlights the multitude of ways in which the microanatomy and microscopic structure of bones enables palaeoecological deductions. The occurrence of growth marks in bones is discussed, and their usefulness in deducing the ontogenetic status and age of individuals is considered, as well as how such marks in bones permit the assessment of the growth dynamics of individuals and species. Here osteohistology is shown to provide insight into the structure of past populations, as well as ecological relationships between individuals. In addition, the response of bones to trauma, disease and moulting is considered. Finally, I explore how osteohistology can give insight into ecomorphological adaptations, such as filter feeding, probe feeding and saltatorial locomotion. Methodological advances in three-dimensional microtomography and synchrotron scanning bodes well for future studies in osteohistology and despite some compromises in terms of tissue identity, circumvents the crucial issue of destructive analyses.

Osteohistology of Dromornis stirtoni (Aves: Dromornithidae) and the biological implications of the bone histology of the Australian mihirung birds.

The late Miocene Dromornis stirtoni is the largest of the giant flightless dromornithid birds. Here, we studied 22 long bones (femora, tibiotarsi, tarsometatarsi) of D. stirtoni to assess its osteohistology to deduce various aspects of its life history. Our results show that D. stirtoni took several years (likely, more than a decade), to reach adult body size, after which its growth rate slowed down, and skeletal maturity occurred. This growth strategy differs from that of its Pleistocene relative, Genyornis newtoni, which experienced faster rates of growth to reach adult body size. We propose that these mihirung birds, separated by millions of years, each responded to the prevailing environmental conditions of the time, by selecting for different growth strategies, with D. stirtoni having an extreme K-selected life history strategy. The presence of medullary bone permitted the identification of female D. stirtoni specimens, and its presence in some bones lacking an OCL layer showed sexual maturity preceded its formation. We postulate that while G. newtoni had a somewhat greater reproductive potential than D. stirtoni, it remained far less than that observed in the extant emu (Dromaius novaehollandiae). Genyornis newtoni survived into the late Pleistocene alongside extant emus and overlapped the arrival of the first humans in Australia, but the former species shortly thereafter became extinct while emus remain prolific.

Bone histology of Neogene angulate tortoises (Testudines: Testudinidae) from South Africa: palaeobiological and skeletochronological implications.

Here we examine the tibial microstructure of modern and fossil angulate tortoises to assess the histology and growth from the late Miocene-early Pliocene, Pleistocene through to modern forms. The cross-sections of all the tibiae sampled revealed highly vascularized, uninterrupted, fibrolamellar bone tissue during early ontogeny, which suggests that early growth was fast. However, later in ontogeny, growth was slower, as indicated by the deposition of parallel-fibred bone tissue in the outer cortex, and even ceased periodically, as indicated by lines of arrested growth. Comparative analyses of the growth rates of the tortoises from different time periods showed that the tortoises from the late Miocene-early Pliocene Langebaanweg locality and from Diepkloof Rock Shelter had relatively slower growth rates under less optimal growth conditions. Additionally, these prehistoric specimens show extensive remodelling, and several generations of secondary osteons further suggest functional and/or metabolic stresses on the skeleton. Palaeoenvironmental reconstructions suggest that it was mostly cooler and drier with seasonal fluctuations in late Miocene-early Pliocene, and it is likely that Chersina responded to these conditions by having a lower rate of growth as compared with their modern counterparts, which thrive in the current prevailing more favourable Mediterranean type of climate.

Osteohistology and palaeobiology of giraffids from the Mio-Pliocene Langebaanweg (South Africa).

The reconstruction of life history traits, such as growth rate, age at maturity and age at death can be estimated from the histological analysis of long bones. Here, we studied 20 long bones (metapodials, tibia and femora) of Sivatherium hendeyi and Giraffa cf. Giraffa jumae recovered from the Miocene-Pliocene locality of Langebaanweg on the West Coast of South Africa. We analysed the long bone histology and growth marks of juvenile and adult specimens of these taxa. Our results show that bone tissue types and vascular canal orientation varies during ontogeny, as well as between the different skeletal elements, and also across single cross sections of bones. Majority of our specimens appear to be still growing, with only an adult metacarpal of S. hendeyi being skeletally mature as indicated by the presence of an outer circumferential layer. We propose that the growth marks preserved in the cortices of the bones studied are most likely related to multiple catastrophic events as opposed to being annual/seasonal.

Comparative morphology and soft tissue histology of the remote-touch bill-tip organ in three ibis species of differing foraging ecology.

Ibises (order: Pelecaniformes, family: Threskiornithidae) are probe-foraging birds that use 'remote-touch' to locate prey items hidden in opaque substrates. This sensory capability allows them to locate their prey using high-frequency vibrations in the substrate in the absence of other sensory cues. Remote-touch is facilitated by a specialised bill-tip organ, comprising high densities of mechanoreceptors (Herbst corpuscles) embedded in numerous foramina in the beak bones. Each foramen and its associated Herbst corpuscles make up a sensory unit, called a 'sensory pit'. These sensory pits are densely clustered in the distal portion of the beak. Previous research has indicated that interspecific differences in the extent of sensory pitting in the beak bones correlate with aquatic habitat use of ibises, and have been suggested to reflect different levels of remote-touch sensitivity. Our study investigates the interspecific differences in the bone and soft tissue histology of the bill-tip organs of three species of southern African ibises from different habitats (mainly terrestrial to mainly aquatic). We analysed the external pitting pattern on the bones, as well as internal structure of the beak using micro-CT scans and soft tissue histological sections of each species' bill-tip organs. The beaks of all three species contain remote-touch bill-tip organs and are described here in detail. Clear interspecific differences are evident between the species' bill-tip organs, both in terms of bone morphology and soft tissue histology. Glossy Ibises, which forage exclusively in wetter substrates, have a greater extent of pitting but lower numbers of Herbst corpuscles in each pit, while species foraging in drier substrates (Hadeda and Sacred Ibises) have more robust beaks, fewer pits and higher densities of Herbst corpuscles. Our data, together with previously published histological descriptions of the bill-tip organs of other remote-touch foraging bird species, indicate that species foraging in drier habitats have more sensitive bill-tip organs (based on their anatomy). The vibrations produced by prey (e.g., burrowing invertebrates) travel poorly in dry substrates compared with wetter ones (i.e., dry soil vs. mud or water), and thus we hypothesise that a more sensitive bill-tip organ may be required to successfully locate prey in dry substrates. Furthermore, our results indicate that the differences in bill-tip organ anatomy between the species reflect complex trade-offs between morphological constraints of beak shape and remote-touch sensitivity requirements, both of which are likely related to each species' foraging behaviour and substrate usage. Our study suggests that structures in the bone of the bill-tip organ could provide valuable osteological correlates for the associated soft tissues, and consequently may provide information on the sensory ecology and habitat usage of the birds in the absence of soft tissues.

Fossorial adaptations in African mole-rats (Bathyergidae) and the unique appendicular phenotype of naked mole-rats.

Life underground has constrained the evolution of subterranean mammals to maximize digging performance. However, the mechanisms modulating morphological change and development of fossorial adaptations in such taxa are still poorly known. We assessed the morpho-functional diversity and early postnatal development of fossorial adaptations (bone superstructures) in the appendicular system of the African mole-rats (Bathyergidae), a highly specialized subterranean rodent family. Although bathyergids can use claws or incisors for digging, all genera presented highly specialized bone superstructures associated with scratch-digging behavior. Surprisingly, Heterocephalus glaber differed substantially from other bathyergids, and from fossorial mammals by possessing a less specialized humerus, tibia and fibula. Our data suggest strong functional and developmental constraints driving the selection of limb specializations in most bathyergids, but more relaxed pressures acting on the limbs of H. glaber. A combination of historical, developmental and ecological factors in Heterocephalus are hypothesized to have played important roles in shaping its appendicular phenotype.

Inter-element variation in the bone histology of Anteosaurus (Dinocephalia, Anteosauridae) from the Tapinocephalus Assemblage Zone of the Karoo Basin of South Africa.

Despite its abundance in the Permian fossil record of South Africa, little is known about the life history of Anteosaurus. Here we examine the bone microstructure of multiple skeletal elements of Anteosaurus from the Tapinocephalus Assemblage Zone of the Karoo Basin. The bone histology of Anteosaurus magnificus reveals that the cortex is composed of highly vascularized, uninterrupted fibrolamellar bone tissue surrounding the inner spongy medullary region. However, the histology of two ribs and a previously described femur of another Anteosaurus taxon revealed an interrupted growth pattern with lines of arrested growth and peripheral rest lines occurring in the compacta, indicating periodic pauses in growth possibly linked to the slowing down of growth during maturity. Given that the fibula of the same individual has well-vascularised fibrolamellar bone tissue without any growth marks in the cortex; this suggests variation in skeletal growth. Based on our histological results, three growth dynamic stages are deduced for the genus Anteosaurus: (i) the earliest growth stage is represented by the predominance of highly vascularized, uninterrupted fibrolamellar bone tissue in the inner cortex, which suggests rapid periosteal bone deposition during early ontogeny; (ii) the next stage of growth shows periodic interruptions in the bone deposition as indicated by the deposition of lines of arrested growth; (iii) the third stage shows the development of lamellar bone tissue with rest lines in the peripheral part of the cortex suggesting a slowing down of growth prior to death. Most of the skeletal elements are characterized by thick bone walls, extensive secondary reconstruction and the complete infilling of the medullary cavity. However, the radius and a previously studied femur have open medullary cavities with struts of bony trabeculae. Based on histologic structures and comparisons with extant taxa, it is likely that Anteosaurus may have been more terrestrial as its osteology point towards terrestriality, but it may have occasionally inhabited ephemeral pools like modern semi-aquatic Hippopotamus.

Bone remodeling in the longest living rodent, the naked mole-rat: Interelement variation and the effects of reproduction.

The pattern of bone remodeling of one of the most peculiar mammals in the world, the naked mole-rat (NMR), was assessed. NMRs are known for their long lifespans among rodents and for having low metabolic rates. We assessed long-term in vivo bone labeling of subordinate individuals, as well as the patterns of bone resorption and bone remodeling in a large sample including reproductive and non-reproductive individuals (n = 70). Over 268 undecalcified thin cross-sections from the midshaft of humerus, ulna, femur and tibia were analyzed with confocal fluorescence and polarized light microscopy. Fluorochrome analysis revealed low osteogenesis, scarce bone resorption and infrequent formation of secondary osteons (Haversian systems) (i.e., slow bone turnover), thus most likely reflecting the low metabolic rates of this species. Secondary osteons occurred regardless of reproductive status. However, considerable differences in the degree of bone remodeling were found between breeders and non-breeders. Pre-reproductive stages (subordinates) exhibited quite stable skeletal homeostasis and bone structure, although the attainment of sexual maturity and beginning of reproductive cycles in female breeders triggered a series of anabolic and catabolic processes that up-regulate bone turnover, most likely associated with the increased metabolic rates of reproduction. Furthermore, bone remodeling was more frequently found in stylopodial elements compared to zeugopodial elements. Despite the limited bone remodeling observed in NMRs, the variation in the pattern of skeletal homeostasis (interelement variation) reported here represents an important aspect to understand the skeletal dynamics of a small mammal with low metabolic rates. Given the relevance of the remodeling process among mammals, this study also permitted the comparison of such process with the well-documented histomorphology of extinct therapsids (i.e., mammalian precursors), thus evidencing that bone remodeling and its endocortical compartmentalization represent ancestral features among the lineage that gave rise to mammals. It is concluded that other factors associated with development (and not uniquely related to biomechanical loading) can also have an important role in the development of bone remodeling.

Long bone histomorphogenesis of the naked mole-rat: Histodiversity and intraspecific variation.

Lacking fur, living in eusocial colonies and having the longest lifespan of any rodent, makes naked mole-rats (NMRs) rather peculiar mammals. Although they exhibit a high degree of polymorphism, skeletal plasticity and are considered a novel model to assess the effects of delayed puberty on the skeletal system, scarce information on their morphogenesis exists. Here, we examined a large ontogenetic sample (n = 76) of subordinate individuals to assess the pattern of bone growth and bone microstructure of fore- and hindlimb bones by using histomorphological techniques. Over 290 undecalcified thin cross-sections from the midshaft of the humerus, ulna, femur, and tibia from pups, juveniles and adults were analyzed with polarized light microscopy. Similar to other fossorial mammals, NMRs exhibited a systematic cortical thickening of their long bones, which clearly indicates a conserved functional adaptation to withstand the mechanical strains imposed during digging, regardless of their chisel-tooth predominance. We describe a high histodiversity of bone matrices and the formation of secondary osteons in NMRs. The bones of pups are extremely thin-walled and grow by periosteal bone formation coupled with considerable expansion of the medullary cavity, a process probably tightly regulated and adapted to optimize the amount of minerals destined for skeletal development, to thus allow the female breeder to produce a higher number of pups, as well as several litters. Subsequent cortical thickening in juveniles involves high amounts of endosteal bone apposition, which contrasts with the bone modeling of other mammals where a periosteal predominance exists. Adults have bone matrices predominantly consisting of parallel-fibered bone and lamellar bone, which indicate intermediate to slow rates of osteogenesis, as well as the development of poorly vascularized lamellar-zonal tissues separated by lines of arrested growth (LAGs) and annuli. These features reflect the low metabolism, low body temperature and slow growth rates reported for this species, as well as indicate a cyclical pattern of osteogenesis. The presence of LAGs in captive individuals was striking and indicates that postnatal osteogenesis and its consequent cortical stratification most likely represents a plesiomorphic thermometabolic strategy among endotherms which has been suggested to be regulated by endogenous rhythms. However, the generalized presence of LAGs in this and other subterranean taxa in the wild, as well as recent investigations on variability of environmental conditions in burrow systems, supports the hypothesis that underground environments experience seasonal fluctuations that may influence the postnatal osteogenesis of animals by limiting the extension of burrow systems during the unfavorable dry seasons and therefore the finding of food resources. Additionally, the intraspecific variation found in the formation of bone tissue matrices and vascularization suggested a high degree of developmental plasticity in NMRs, which may help explaining the polymorphism reported for this species. The results obtained here represent a valuable contribution to understanding the relationship of several aspects involved in the morphogenesis of the skeletal system of a mammal with extraordinary adaptations.

Ontogenetic growth and the development of a unique fibrocartilage entheses in Macropus fuliginosus.

Here we examine the bone histology of the femora and humeri of the Western Grey Kangaroo, Macropus fuliginosus. Our results reveal that bone modelling in response to ontogenetic growth and the development of tuberosities on the femur, and especially in the humerus, lead to a highly complex histology. We propose that the alternating fast and slow rates of bone deposition are seasonal, and are likely correlated with heterothermy related to ecological constraints during the summer months. In females, after the fourth growth mark in the femur, there is a distinctive change to a more lamellar textured bone deposition with sparse vascularisation, directly indicating a slowdown in growth. However, in males, the zones remain woven textured and well vascularised, which is indicative of continued fast growth. Here we also report the novel occurrence of a fibrocartilaginous entheses for the attachment of the m. quadratus femoris to the caudal femoral tuberosity. Using a combination of methodologies, we show that perimeter measurements of growth marks provide a reasonable estimation of the age of kangaroos. Additionally, we observed large individuals that have ceased diaphyseal appositional growth of the femur and the humerus, as well as fusion of the distal epiphyses of both bones, though the proximal epiphyses may remain unfused.

Osteohistology and Life History of the Basal Pygostylian, Confuciusornis sanctus.

More than a thousand specimens of Confuciusornis sanctus have been recovered from the Early Cretaceous Jehol Group of Northeastern China. Here, we investigate the bone microstructure of 33 long bones sampled from 14 C. sanctus specimens in an attempt to assess the life history patterns of this basal pygostylian bird. Analysis of the histology of various skeletal elements (femur, humerus, tibia, radius, and ulna) revealed differences in the histology of their bone walls. Based on the osteohistology, we coded the examined specimens into five histology age classes. We found that histological age was not strictly correlated with body size. The variability in the histology of multiple bones from single skeletons suggests differences in the growth rate of the skeleton in response to allometry, functional demands, and pathology. We show that although fibrolamellar bone is widespread across birds, the extent and duration of this rapid phase of bone deposition is highly variable. Comparisons among Mesozoic birds confirm that early ontogenetic growth was rapid, but that later post-hatching growth was strongly influenced by the ontogenetic age of the individual, body size, and local environment, as well as taxonomy. Our findings indicate that C. sanctus experienced rapid growth from early ontogeny until almost fully grown, and thereafter transitioned to slow, episodic growth (for at least 3-4 years) to reach skeletal maturity. Anat Rec, 303:949-962, 2020. © 2019 American Association for Anatomy.

Sex and Ontogenetic Variation in the Crest of Numida meleagris: Implications for Crested Vertebrates.

Crested vertebrates are known from a wide variety of modern and fossil taxa, however, the actual formation and function of the crest is still debatable. Among modern birds, the globally distributed guinea fowl (Numida meleagris) is characterized by having a cranial bony crest (overlain by keratin), but surprisingly little is known about its development. Here, we studied the crest of 202 wild guinea fowl from the same population, using anatomical measurements as well as 2D-morphometry. Our results show that juveniles have smaller skulls than adults and have smaller, simpler crests that are visible even in very young individuals. Among adults, female skulls are smaller than males, and they have smaller, simpler shaped crests, which permit a discrimination between the sexes of 93% when the keratin is preserved with the bony crest, and of 89% when only the bony crest is available. By extrapolation, these results confirm that the crest can be used as an ontogenetic character, as well as for sex discrimination in the fossil record. Our results also show that the overlying keratin does not always mimic the underlying bony crest, which should be considered when reconstructing extinct crested vertebrates. Anat Rec, 303:1018-1034, 2020. © 2019 American Association for Anatomy.

Long bone histology of Chersina angulata : Interelement variation and life history data.

The current study deduced the growth pattern and lifestyle habits of Chersina angulata based on bone histology and cross-sectional geometry of limb bones. Femora, humeri, and tibiae of seven different-sized individuals representing different ontogenetic stages were assessed to determine the interelement and intraskeletal histological variation within and among the tortoises. The bone histology of adult propodials consists of a highly vascularized, uninterrupted fibrolamellar bone tissue with a woven texture in the perimedullary and midcortical regions suggesting overall fast early growth. However, later in ontogeny, growth was slow and even ceased periodically as suggested by slowly formed parallel-fibered bone tissue and several growth marks in the pericortical region. In juvenile individuals, fibrolamellar bone tissue is restricted to the perimedullary regions of propodials as remnants of bone formed during the earliest stages of ontogeny. The epipodials are characterized by having parallel-fibered bone tissue present in their cortices; however, periodic arrests in growth are recorded at various times. Remnants of fibrolamellar bone tissue formed during early ontogeny occur in the epipodials of only a few individuals. Interelement variation is evident, in terms of variation in the orientation of vascular canals between individuals and within the same diaphyseal cross-sections. Different elements show varying cross-sectional geometry, which appear to be correlated with the fossorial behavior of the species. Our results show that of all the long bones, the tibia is least remodeled during ontogeny and it is therefore the best element for skeletochronology.

Postnatal development of the largest subterranean mammal (Bathyergus suillus): Morphology, osteogenesis, and modularity of the appendicular skeleton.

BACKGROUND: Subterranean mammals show a suite of musculoskeletal adaptations that enables efficient digging. However, little is known about their development. We assessed ontogenetic changes in functionally relevant skeletal traits, and ossification patterns (periosteal and endochondral bone modules) in a truly subterranean scratch-digging rodent, Bathyergus. We studied 52 individuals (202 long bones) from a wild population by using a multiscale approach involving internal and external morphology. RESULTS: Multivariate analysis showed significant morphological changes during ontogeny. A specialized phenotype is expressed perinatally (eg, greater external robustness and developed olecranon, teres major, and deltoid processes), whereas adults presented slender bones with significantly thicker cross-sections. Ossification modules scaled mostly isometrically with body size parameters. Periosteal modules showed high variability and tended to grow faster than endochondral modules. CONCLUSIONS: Scratch-digging adaptations appear at perinatal age and then specialize in subadults. Early development of agonistic and digging behaviors and onset of sexual maturation seems to contribute to its development, although genetic factors also seem to play an important role. Ontogenetic differences are probably a trade-off to counteract weaker cortical bone properties and poor muscle development in juveniles, whereas slender but thicker cortical bones maximize bone resistance during burrow construction without compromising locomotor performance in adults.

A new baby oviraptorid dinosaur (Dinosauria: Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia.

Recent discoveries of new oviraptorosaurs revealed their high diversity from the Cretaceous Period in Asia and North America. Particularly, at the family level, oviraptorids are among the most diverse theropod dinosaurs in the Late Cretaceous of Mongolia and China. A new oviraptorid dinosaur Gobiraptor minutus gen. et sp. nov. from the Upper Cretaceous Nemegt Formation is described here based on a single holotype specimen that includes incomplete cranial and postcranial elements. The most prominent characters of Gobiraptor are its thickened rostrodorsal end of the mandibular symphysis and a rudimentary lingual shelf on each side of the dentary. Each lingual shelf is lined with small occlusal foramina and demarcated by a weakly developed lingual ridge. This mandibular morphology of Gobiraptor is unique among oviraptorids and likely to be linked to a specialized diet that probably included hard materials, such as seeds or bivalves. The osteohistology of the femur of the holotype specimen indicates that the individual was fairly young at the time of its death. Phylogenetic analysis recovers Gobiraptor as a derived oviraptorid close to three taxa from the Ganzhou region in southern China, but rather distantly related to other Nemegt oviraptorids which, as the results of recent studies, are also not closely related to each other. Gobiraptor increases diversity of oviraptorids in the Nemegt Formation and its presence confirms the successful adaptation of oviraptorids to a mesic environment.

Cortical bone adaptation and mineral mobilization in the subterranean mammal Bathyergus suillus (Rodentia: Bathyergidae): effects of age and sex.

The patterns of bone modeling and mineral mobilization (skeletal homeostasis) among mammals other than humans and laboratory rodents are still poorly known. In this study we assessed the pattern of bone formation and bone resorption in the femur of a wild population of Cape dune molerats, Bathyergus suillus (n = 41) (Bathyergidae), a solitary subterranean mammal with a marked extended longevity among rodents, and which also lives in a naturally deficient state of vitamin D. In order to determine ontogenetic and sex effects on histomorphometric parameters of transversal undecalcified bone sections, two-way ANOVA, linear mixed-effects model and regression statistical analyses were performed. During ontogeny, B. suillus increased their cross sectional area, cortical area and cortical thickness, and most importantly, they showed scarce endosteal bone resorption which resulted in a retained medullary cavity size during ontogeny. This resulted in a positively imbalanced bone modeling, where bone formation considerably surpasses bone loss by almost 100-fold in adulthood. This differs markedly from other terrestrial mammals with relatively thin cortical walls. Regarding bone loss and remodeling, three main processes involving intracortical resorption were observed: modeling-related bone loss in early postnatal growth; secondary osteon formation occurring in both sexes; and subendosteal secondary reconstruction observed only in females. The latter is accompanied by females having six-fold more relative bone loss than males, which is evidenced by the development of enlarged resorption cavities (RCs) distributed circumferentially around the medullary cavity. Males have smaller, more circular and randomly distributed RCs. In general, our data indicate no age-related decline in mineral content in B. suillus, and provides strong support for a pattern of sexual dimorphism in skeletal homeostasis, similar to that occurring in humans and other mammals, with females losing more bone throughout aging as compared to males due to reproductive factors. Interestingly as well, despite the high mechanical loads experienced during burrow construction, bone remodeling in B. suillus is kept at very low levels throughout their lifespan, and dense Haversian tissue never forms. This study represents the first comprehensive assessment of skeletal homeostasis in a subterranean mammal, and it enables a better understanding of the complex processes governing the acquisition and maintenance of bone properties in this species with extraordinary fossorial adaptations.

A new Jurassic theropod from China documents a transitional step in the macrostructure of feathers.

Genuine fossils with exquisitely preserved plumage from the Late Jurassic and Early Cretaceous of northeastern China have recently revealed that bird-like theropod dinosaurs had long pennaceous feathers along their hindlimbs and may have used their four wings to glide or fly. Thus, it has been postulated that early bird flight might initially have involved four wings (Xu et al. Nature 421:335-340, 2003; Hu et al. Nature 461:640-643, 2009; Han et al. Nat Commun 5:4382, 2014). Here, we describe Serikornis sungei gen. et sp. nov., a new feathered theropod from the Tiaojishan Fm (Late Jurassic) of Liaoning Province, China. Its skeletal morphology suggests a ground-dwelling ecology with no flying adaptations. Our phylogenetic analysis places Serikornis, together with other Late Jurassic paravians from China, as a basal paravians, outside the Eumaniraptora clade. The tail of Serikornis is covered proximally by filaments and distally by slender rectrices. Thin symmetrical remiges lacking barbules are attached along its forelimbs and elongate hindlimb feathers extend up to its toes, suggesting that hindlimb remiges evolved in ground-dwelling maniraptorans before being co-opted to an arboreal lifestyle or flight.

Novel insight into the origin of the growth dynamics of sauropod dinosaurs.

Sauropod dinosaurs include the largest terrestrial animals and are considered to have uninterrupted rapid rates of growth, which differs from their more basal relatives, which have a slower cyclical growth. Here we examine the bone microstructure of several sauropodomorph dinosaurs, including basal taxa, as well as the more derived sauropods. Although our results agree that the plesiomorphic condition for Sauropodomorpha is cyclical growth dynamics, we found that the hypothesized dichotomy between the growth patterns of basal and more derived sauropodomorphs is not supported. Here, we show that sauropod-like growth dynamics of uninterrupted rapid growth also occurred in some basal sauropodomorphs, and that some basal sauropods retained the plesiomorphic cyclical growth patterns. Among the sauropodomorpha it appears that the basal taxa exploited different growth strategies, but the more derived Eusauropoda successfully utilized rapid, uninterrupted growth strategies.

Long bone histology of the subterranean rodent Bathyergus suillus (Bathyergidae): ontogenetic pattern of cortical bone thickening.

Patterns of bone development in mammals are best known from terrestrial and cursorial groups, but there is a considerable gap in our understanding of how specializations for life underground affect bone growth and development. Likewise, studies of bone microstructure in wild populations are still scarce, and they often include few individuals and tend to be focused on adults. For these reasons, the processes generating bone microstructural variation at intra- and interspecific levels are not fully understood. This study comprehensively examines the bone microstructure of an extant population of Cape dune molerats, Bathyergus suillus (Bathyergidae), the largest subterranean mammal endemic to the Western Cape of South Africa. The aim of this study is to investigate the postnatal bone growth of B. suillus using undecalcified histological sections (n = 197) of the femur, humerus, tibia-fibula, ulna and radius, including males and females belonging to different ontogenetic and reproductive stages (n = 42). Qualitative histological features demonstrate a wide histodiversity with thickening of the cortex mainly resulting from endosteal and periosteal bone depositions, whilst there is scarce endosteal resorption and remodeling throughout ontogeny. This imbalanced bone modeling allows the tissues deposited during ontogeny to remain relatively intact, thus preserving an excellent record of growth. The distribution of the different bone tissues observed in the cortex depends on ontogenetic status, anatomical features (e.g. muscle attachment structures) and location on the bone (e.g. anterior or lateral). The type of bone microstructure and modeling is discussed in relation to digging behavior, reproduction and physiology of this species. This study is the first histological assessment describing the process of cortical thickening in long bones of a fossorial mammal.