Rise of dinosaurs reveals major body-size transitions are driven by passive processes of trait evolution

A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252-201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian-Middle Jurassic. Maximum-likelihood analyses indicate that Cope's rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope's rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle-early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely-accepted 'rule' that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates.     

Septomaxilla of nonmammalian synapsids: soft-tissue correlates and a new functional interpretation. DELETE

The function of the septomaxilla of nonmammalian synapsids has long been problematic. Distinctive features of this bone, including a prominent intranarial process and a septomaxillary canal and foramen, are characteristic of pelycosaurs and nonmammalian therapsids, but are lost in their mammalian descendants. Numerous contradictory reconstructions have been proposed for the soft anatomy associated with the septomaxilla of nonmammalian synapsids. This review supports the following conclusions: 1) No particular correlation exists between the septomaxilla and the vomeronasal organ (VNO), and the most likely location for the VNO is on the dorsal surface of the palatal process of the vomer; 2) The most likely occupant of the septomaxillary canal is the nasolacrimal duct, which opened either anterior or medial to the intranarial process, near the opening of the VNO duct; and 3) The occupant of the septomaxillary foramen remains uncertain. These conclusions suggest that the functional significance of the septomaxilla in the nonmammalian synapsids is tied to that of the nasolacrimal duct. The association of this duct and the VNO in these animals resembles the condition in Recent amphibians and lepidosaurs, in which the nasolacrimal duct supplies orbital fluids to the VNO, apparently to enhance vomeronasal function. The peculiar shape of the synapsid septomaxilla may have served to collect vomeronasal odor molecules. The changes of the septomaxilla in early mammals, and its nearly complete loss in extant mammals, are probably correlated with a dissociation of the nasolacrimal duct and VNO, and functional changes in both structures.     

Long bone histology of Ophiacodon reveals the geologically earliest occurrence of fibrolamellar bone in the mammalian stem lineage

Shared histological characteristics have been observed in the bone matrix and vascularity between Ophiacodontidae and the later therapsids (Synapsida). Historically, this coincidence has been explained as simply a reflection of the presumed aquatic lifestyle of Ophiacodon or even a sign of immaturity. Here we show, by histologically sampling an ontogenetic series of Ophiacodon humeri, as well as additional material, the existence of fibrolamellar bone (FLB) in the postcranial bones of a pelycosaur. Our findings have reaffirmed what previous studies first described as fast growing tissue, and by proxy, have disproven that the highly vascularized cortex is simply a reflection of young age. This tissue demonstrates the classic histological characteristics of true FLB. The cortex consists of primary osteons in a woven bone matrix and remains highly vascularized throughout ontogeny, providing evidence for fast skeletal growth. Overall, the FLB tissue we have described in Ophiacodon is more advanced or “mammal-like” in terms of the osteonal development, bone matrix, and skeletal growth than what has been described thus far for any other pelycosaur taxon. With regards to the histological record, our results remain inconclusive as to the preferred ecology of Ophiacodon due to a similar cortical vascularity pattern exhibited by other carnivorous pelycosaurs. Our findings have set the evolutionary origins of FLB and high skeletal growth rates back approximately 20 million years to the Early Permian, and by phylogenetic extension perhaps the Late Carboniferous.     

鄂尔多斯高原生态过渡带的判定及生物群区特征

 本文用Holdridge生命地带方法对鄂尔多斯高原的生物群区进行了分类研究,研制了用于生态过渡带判定的计算机模型,模拟结果与实际情况基本吻合。鄂尔多斯高原的生物群区可分为草原、荒漠灌丛和荒漠3大类,草原群区以本氏针茅(Stipa bungeana)群落为主,荒漠灌丛群区以冷蒿(Artemisia frigida),狭叶锦鸡儿(Caragana stenophylla)群落为主,荒漠群区以红砂(Reaumuria soongorica)群落为主。鄂尔多斯高原的地带性植被在传统上分为典型草原、荒漠草原和草原化荒漠3部分,在典型草原的边界划分上,传统划分与本文结果一致。而在荒漠草原和草原化荒漠的边界方面,模拟结果与传统划分有一定的差异。荒漠灌丛和荒漠草原以及荒漠和草原化荒漠在生物群落特征上具有一定的相似性,荒漠灌丛和荒漠在生态特征上较荒漠草原和草原化荒漠更为旱化,在地理位置上更为靠西部。该计算机模型结合了地理空间分析,能准确地确定生态过渡带的位置和宽度,为研究鄂尔多斯高原生态过渡带的特征提供了较可靠的技术手段。     

Natural environment and thermal behaviour of Dimetrodon limbatus

This paper examines the body temperature variation of Dimetrodon during the different seasons of the year. The effect of the sail of Dimetrodon on its body temperature is also evaluated. It is shown that the sail of pelycosaurs provided an advantage to the reptile by warming it up quicker in the morning in cold environments. This would be a benefit, allowing Dimetrodon to prey on large reptiles, above 55kg, in the early morning while they were sluggish. From the results presented a climate similar to that of March for Cyprus could be representative of that of Permian period.     

Does a General Temperature-Dependent Q10 Model of Soil Respiration Exist at Biome and Global Scale?

Soil respiration (SR) is commonly modeled by a Q10 (an indicator of temperature sensitivity)function in ecosystem models. Q10is usually treated as a constant of 2 in these models, although Q10 value of SR often decreases with increasing temperatures. It remains unclear whether a general temperaturedependent Q10 model of SR exists at biome and global scale. In this paper, we have compiled the long-term Q10 data of 38 SR studies ranging from the Boreal, Temperate, to Tropical/Subtropical biome on four continents.Our analysis indicated that the general temperature-dependent biome Q10 models of SR existed, especially in the Boreal and Temperate biomes. A single-exponential model was better than a simple linear model in fitting the average Q10 values at the biome scale. Average soil temperature is a better predictor of Q10 value than average air temperature in these models, especially in the Boreal biome. Soil temperature alone could explain about 50% of the Q10 variations in both the Boreal and Temperate biome single-exponential Q10 model. Q10 value of SR decreased with increasing soil temperature but at quite different rates among the three biome Q10 models. The k values (Q10 decay rate constants) were 0.09, 0.07, and 0.02/℃ in the Boreal, Temperate, and Tropical/Subtropical biome, respectively, suggesting that Q10 value is the most sensitive to soil temperature change in the Boreal biome, the second in the Temperate biome, and the least sensitive in the Tropical/Subtropical biome. This also indirectly confirms that acclimation of SR in many soil warming experiments probably occurs. The k value in the "global" single-exponential Q10 model which combined both the Boreal and Temperate biome data set was 0.08/℃. However, the global general temperature-dependent Q10model developed using the data sets of the three biomes is not adequate for predicting Q10 values of SR globally.The existence of the general temperature-dependent Q10 models of SR in the Boreal and Temperate biome has important implications for modeling SR, especially in the Boreal biome. More detail model runs are needed to exactly evaluate the impact of using a fixed Q10 vs a temperature-dependent Q10 on SR estimate in ecosystem models (e.g., TEM, Biome-BGC, and PnET).     

Molecular phylogenetics reveals a pattern of biome conservatism in New World anchovies (family Engraulidae)

Evolutionary transitions between marine and freshwater biomes are relatively rare events, yielding a widespread pattern of biome conservatism among aquatic organisms. We investigated biome transitions in anchovies (Engraulidae), a globally distributed clade of economically important fishes. Most anchovy species are near-shore marine fishes, but several exclusively freshwater species are known from tropical rivers of South America and were previously thought to be the product of six or more independent freshwater invasions. We generated a comprehensive molecular phylogeny for Engraulidae, including representatives from 15 of 17 currently recognized genera. Our data support previous hypotheses of higher-level relationships within Engraulidae, but show that most New World genera are not monophyletic and in need of revision. Ancestral character reconstruction reveals that New World freshwater anchovies are the product of a single marine to freshwater transition, supporting a pattern of biome conservatism. We argue that competition is the principal mechanism that regulates aquatic biome transitions on a continental scale.     

The evolution of the dicynodont feeding system

The skull structure of dicynodonts may be regarded as a complex adaptation towards herbivorous feeding. The present work examines how and why this adaptation may have evolved. A cladogram of the dicynodonts is presented and from it a sequence of hypothetical ancestral forms is inferred. The jaw musculature of dicynodonts and other therapsids is described and in particular the early dicynodont Eodicynodon oosthuizeni is described in detail. This information is used to draw up a sequence of ancestral stages whose basic skull anatomy, jaw muscle organization and masticatory properties are described. Differences in masticatory properties between these stages are pinpointed and an explanation to account for the development of these differences is advanced. It is concluded that the changes in skull organization seen during the evolution of dicynodonts are consistent with the hypothesis that a propalinal jaw action was being improved by selection, and that this was required to permit dicynodonts to be efficient herbivores.     

Evolution of the mammalian middle ear.

The structure and evolution of the mandible, suspensorium, and stapes of mammal-like reptiles and early mammals are examined in an attempt to determine how, why, and when in phylogeny the precursors of the mammalian tympanic bone, malleus, and incus (postdentary jaw elements and quadrate) came to function in the reception of air-borne sound. The following conclusions are reached: It is possible that at no stage in mammalian phylogeny was there a middle ear similar to that of "typical" living reptiles, with a postquadrate tympanic membrane contracted by an extrastapes. The aquamosal sulcus of cynodonts and other therapsids, usually thought to have housed a long external acoustic meatus, possibly held a depressor mandibulae muscle. In therapsids an air-filled chamber (recessus mandibularis of Westoll) extended deep to the reflected lamina and into the depression (external fossa) on the outer aspect of the angular element. A similar chamber was present in sphenacodontids but pterygoideus musculature occupied the small external fossa. The thin tissues superficial to the recessus mandibularis served as eardrum. Primitively, vibrations reached the stapes mainly via the anterior hyoid cornu, but in dicynodonts, therocephalians, and cynodants vibrations passed mainly or exclusively from mandible to quadrate to stapes and the reflected lamina was a component of the eardrum. In the therapsid phase of mammalian phylogeny, auditory adaptation was an important aspect of jaw evolution. Auditory efficiency, and sensitivity to higher sound frequencies were enhanced by diminution and loosening of the postdentary elements and quadrate, along with transference of musculature from postdentary elements to the dentary. These changes were made possible by associated modifications, including posterior expansion of the dentary. Establishment of a dentary-squamosal articulation permitted continuation of these trends, leading to the definitive mammalian condition, with no major change in auditory mechanism except that in most mammals (not monotremes) the angular, as tympanic, eventually bcame a non-vibrating structure.     

New therapsid specimens and the origin of the secondary hard and soft palate of mammals

A newly prepared palate of the moschorhinid therocephalian Promoschorhynchus platyrhinus from the Upper Permian region of South Africa was used for reconsidering the initial evolutionary development of the secondary palate of mammals. The very distinctive choanal crests are considered to be indicative of strong choanal folds that were probably already fused anteriorly. This gingival bridge probably served as the basis for the outgrowing bony processes of the hard palate that, in therapsids, developed independently at least three times. The soft palate ( velum palatinum ) is considered to be a remnant of the choanal folds that were muscularized from behind. It has been assumed that, in therapsids, a ventral portion of the medial pterygoid musculature shifted its insertion onto the ventral side of the pterygoid; this portion is supposed to have differentiated into the mm. tensor tympani and tensor veli palatini . Investigation of serial sections of extant marsupials confirm the view that the levator of the velum is derived from the upper constrictor of the pharynx. The choanal folds and the secondary palate are discussed within the wider framework of the evolutionary biology of mammalian forerunners. It is suggested that the formation of a sealed mouth cavity is not only related to the improved passage of air, but also to sucking in neonate hatchlings. The importance of the secondary palate of mammals to a number of resulting basic adaptations (dentition, olfactory system, etc.) of mammals is discussed.     

Local and regional-scale responses of ant diversity to a semiarid biome transition

The locations of biome transitions and ecotones are frequently defined by the rapid shift from one form of dominant vegetation to another. The composition of animal taxa is predicted to shift in parallel with that of dominant plants and species diversity is predicted to he greater in transitional zones than in adjacent areas. We asked whether ant species diversity and composition supported these predictions across a biome transition between shortgrass steppe and Chihuahuan desert vegetation. Neither species richness nor diversity was highest at the biome transition region as a whole, or within habitats in the biome transition. The biome transition region was not intermediate in ant species composition or in the representation of different faunal complexes. The community similarity between matched habitats shared between the biome transition zone and adjacent regions was less than that between distinct habitats occurring within regions. A zoogeographic transition for ants may occur to the north of the phytogeographic transition and may be coincident with the northern limits of monsoonal precipitation patterns. In contrast, the phytogeographic transition may be related to less extreme climatic variation within the monsoonal region occurring further south.     

Does a General Temperature-Dependent Q10 Model of Soil Respiration Exist at Biome and Global Scale？

Soil respiration （SR） is commonly modeled by a Q10 （an indicator of temperature sensitivity） function in ecosystem models. Q10 is usually treated as a constant of 2 in these models, although Q10 value of SR often decreases with increasing temperatures. It remains unclear whether a general temperature- dependent Q10 model of SR exists at biome and global scale. In this paper, we have compiled the long-term Q10 data of 38 SR studies ranging from the Boreal, Temperate, to Tropical/Sublropical biome on four continents. Our analysis indicated that the general temperature-dependent biome Q10 models of SR existed, especially in the Boreal and Temperate biomes. A single-exponential model was better than a simple linear model in fitting the average Q10 values at the biome scale. Average soil temperature is a better predictor of Q10 value than average air temperature in these models, especially in the Boreal biome. Soil temperature alone could explain about 50% of the Q10 variations in both the Boreal and Temperate biome single-exponential Q10 model. Q10 value of SR decreased with increasing soil temperature but at quite different rates among the three biome Q10 models. The k values （Q10 decay rate constants） were 0.09, 0.07, and 0.02/℃ in the Boreal, Temperate, and Tropical/Subtropical biome, respectively, suggesting that Q10 value is the most sensitive to soil temperature change in the Boreal biome, the second in the Temperate biome, and the least sensitive in the Tropical/ Subtropical biome. This also indirectly confirms that acclimation of SR in many soil warming experiments probably occurs. The k value in the ＂global＂ single-exponential Q10 model which combined both the Boreal and Temperate biome data set was 0.08/℃. However, the global general temperature-dependent Q10 model developed using the data sets of the three biomes is not adequate for predicting Q10 values of SR globally. The existence of the general temperature-dependent Q10 models of SR in the Boreal and Temperate biome has important implications for modeling SR, especially in the Boreal biome. More detail model runs are needed to exactly evaluate the impact of using a fixed Q10 vs a temperature-dependent Q10 on SR estimate in ecosystem models （e.g., TEM, Biome-BGC, and PnET）.     

The origins and evolution of the North American grassland biome: the story from the hoofed mammals

The North American grassland biome first appeared around 18 Ma in the mid Miocene. The familiar story of the Neogene evolution of this biome is of the replacement of ungulates (hoofed mammals) having a primarily browsing diet by the more derived grazing ungulates. However, new data show a more complicated pattern of faunal succession. There was a maximum taxonomic diversity of ungulates at 16–14 Ma, including a large number of grazers, and the subsequent decline in overall diversity was largely due to the decline of the browsers, with little corresponding increase in the grazers. Additionally the mid Miocene faunas (∼18–12 Ma) contained a much greater number of browsers than any comparable present-day habitat. We discuss possible explanations for these non-analogous grassland faunas, including the possibility that the primary productivity of the vegetation was greater in the early to middle Miocene than it is today. One possible explanation for increased primary productivity is higher Miocene levels of atmospheric carbon dioxide than in the present day. The proposed difference in vegetational productivity also may explain why horses radiated as the main grazers in North America, in contrast to the radiation of antelope in the Plio–Pleistocene African grasslands.     

The genetic structure of the mountain forest butterfly Erebia euryale unravels the late Pleistocene and postglacial history of the mountain coniferous forest biome in Europe

The distribution of the mountain coniferous forest biome in Europe throughout time is not sufficiently understood. One character species of this habitat type is the large ringlet, Erebia euryale well reflecting the extension of this biome today, and the genetic differentiation of this species among and within mountain systems may unravel the late Pleistocene history of this habitat type. We therefore analysed the allozyme pattern of 381 E. euryale individuals from 11 populations in four different European mountain systems (Pyrenees, Alps, Carpathians, Rila). All loci analysed were polymorphic. The mean F(ST) over all samples was high (20%). Furthermore, the mean genetic distance among samples was quite high (0.049). We found four different groups well supported by cluster analyses, bootstraps and hierarchical variance analyses: Pyrenees, western Alps, eastern Alps and southeastern Europe (Carpathians and Rila). The genetic diversity of the populations was highest in the southeastern European group and stepwise decreased westwards. Interestingly, the populations from Bulgaria and Romania were almost identical; therefore, we assume that they were not separated by the Danube Valley, at least during the last ice age. On the contrary, the differentiation among the three western Alps populations was considerable. For all these reasons, we assume that (i) the most important refugial area for the coniferous mountain forest biome in Europe has been located in southeastern Europe including at least parts of the Carpathians and the Bulgarian mountains; (ii) important refugial areas for this biome existed at the southeastern edge of the Alps; (iii) fragments of this habitat types survived along the southwestern Alps, but in a more scattered distribution; and (iv) relatively small relicts have persisted somewhere at the foothills of the Pyrenees.     

A biome classification of China based on plant functional types and the BIOME3 model

A biome classification for China was established based on plant functional types (PFTs) using the BIOME3 model to include 16 biomes. In the eastern part of China, the PFTs of trees determine mostly the physiognomy of landscape. Biomes range from boreal deciduous coniferous forest/woodland, boreal mixed forest/woodland, temperate mixed forest, temperate broad-leaved deciduous forest, warm-temperate broad-leaved evergreen/mixed forest, warm-temperate/cool-temperate evergreen coniferous forest, xeric woodland/scrub, to tropical seasonal and rain forest, and tropical deciduous forest from north to south. In the northern and western part of China, grass is the dominant PFT. From northeast to west and southwest the biomes range from moist savannas, tall grassland, short grassland, dry savannas, arid shrubland/steppe, desert, to alpine tundra/ice/polar desert. Comparisons between the classification introduced here and the four classifications which were established over the past two decades, i.e. the vegetation classification, the vegetation division, the physical ecoregion, and the initial biome classification have showed that the different aims of biome classifications have resulted in different biome schemes each with its own unique characteristics and disadvantages for global change study. The new biome classification relies not only on climatic variables, but also on soil factor, vegetation functional variables, ecophysiological parameters and competition among the PFTs. It is a comprehensive classification that using multivariables better expresses the vegetation distribution and can be compared with world biome classifications. It can be easily used in the response study of Chinese biomes to global change, regionally and globally.     

Assessing the vulnerability of species richness to anthropogenic climate change in a biodiversity hotspot

Aim To compare theoretical approaches towards estimating risks of plant species loss to anthropogenic climate change impacts in a biodiversity hotspot, and to develop a practical method to detect signs of climate change impacts on natural populations. Location The Fynbos biome of South Africa, within the Cape Floristic Kingdom. Methods Bioclimatic modelling was used to identify environmental limits for vegetation at both biome and species scale. For the biome as a whole, and for 330 species of the endemic family Proteaceae, tolerance limits were determined for five temperature and water availability‐related parameters assumed critical for plant survival. Climate scenarios for 2050 generated by the general circulation models HadCM2 and CSM were interpolated for the region. Geographic Information Systems‐based methods were used to map current and future modelled ranges of the biome and 330 selected species. In the biome‐based approach, predictions of biome areal loss were overlayed with species richness data for the family Proteaceae to estimate extinction risk. In the species‐based approach, predictions of range dislocation (no overlap between current range and future projected range) were used as an indicator of extinction risk. A method of identifying local populations imminently threatened by climate change‐induced mortality is also described. Results A loss of Fynbos biome area of between 51% and 65% is projected by 2050 (depending on the climate scenario used), and roughly 10% of the endemic Proteaceae have ranges restricted to the area lost. Species range projections suggest that a third could suffer complete range dislocation by 2050, and only 5% could retain more than two thirds of their range. Projected changes to individual species ranges could be sufficient to detect climate change impacts within ten years. Main conclusions The biome‐level approach appears to underestimate the risk of species diversity loss from climate change impacts in the Fynbos Biome because many narrow range endemics suffer range dislocation throughout the biome, and not only in areas identified as biome contractions. We suggest that targeted vulnerable species could be monitored both for early warning signs of climate change and as empirical tests of predictions.     

Generation of isogenic pluripotent stem cells differing exclusively at two early onset Parkinson point mutations

Patient-specific induced pluripotent stem cells (iPSCs) derived from somatic cells provide a unique tool for the study of human disease, as well as a promising source for cell replacement therapies. One crucial limitation has been the inability to perform experiments under genetically defined conditions. This is particularly relevant for late age onset disorders in which in vitro phenotypes are predicted to be subtle and susceptible to significant effects of genetic background variations. By combining zinc finger nuclease (ZFN)-mediated genome editing and iPSC technology, we provide a generally applicable solution to this problem, generating sets of isogenic disease and control human pluripotent stem cells that differ exclusively at either of two susceptibility variants for Parkinson's disease by modifying the underlying point mutations in the α-synuclein gene. The robust capability to genetically correct disease-causing point mutations in patient-derived hiPSCs represents significant progress for basic biomedical research and an advance toward hiPSC-based cell replacement therapies.     

The skull of Morganucodon

Morganucodon is a triconodont (atherian) mammal from the Lower Jurassic. Two species are described: M. oehleri from China and M. watsoni from Wales. The skull in M. walsoni is 26 mm long; M. oehleri is slightly larger. The dentition is differentiated functionally into incisors, canines, premolars and molars. The pineal foramen is closed. The prefrontals, postfrontals and postorbitals are lost. Septomaxilla, quadratojugal, tabular and pterygoid flanges are retained. The bony external nares are unpaired. The nasal cavity had the mammalian complement of turbinals. The posterior palate has ridges and troughs similar to those in tritylodonts, triconodonts and multituberculates. The alisphenoid ascending process is narrow and is not in contact with the anterior lamina of the petrosal, lying lateral to it. There is a cavum epiptericum, as in late therapsids. The anterior lamina forms the lateral braincase wall, perforated by the foramina pseudovale and pseudorotundum. There is a squamosal-dentary articulation, but the reptilian jaw joint is retained. The ear resembles that in later therapsids, with the tympanum in the lower jaw. The small quadrate was moveable, buttressed medially be a large stapes. Sound conduction from the tympanum was via articular, quadrate and stapes. The systematic position of Morganucodon is discussed.     

中国东北样带生物群区及其对全球气候变化响应的初步探讨

 本文应用室内模拟与野外实地调查相结合的方法,对中国东北森林草原样带上生物群区及其过渡带的位置与面积进行了判定,并预测了未来全球变化条件下生物群区及过渡带可能的变化趋势。结果表明：中国东北样带内的生物群区和过渡带可由基于气候数据和Ho1dridge方法的数量模型较确切地判定出,与实际调查的结果吻合较好。在未来气候变化下,过渡带面积呈扩大化趋势;森林区对降水量的变化反应敏感;草原区地理分布位置变动很大,但其相对面积变化较小;荒漠灌丛对全球变化的反应最为剧烈。