Identifying the relative importance of urban and non-urban land-use types for potential denitrification derived N2O at a regional scale is critical for quantifying the impacts of human activities on nitrous oxide (N2O) emission under changing environments. In this study we used a regional dataset from China including 197 soil samples and six land-use types to evaluate the main predictors (land use, heavy metals, soil pH, soil moisture, substrate availability, functional and broad microbial abundances) of potential denitrification using multivariate and pathway analysis. Our results provide empirical evidence that soils on farms have the greatest potential denitrifying ability (PDA) (10.92±6.08ng N2O-N·g–1 dry soil·min–1) followed by urban soil (6.80±5.35ng N2O-N·g–1 dry soil·min–1). Our models indicate that land use (low vs. high human activity), followed by total nitrogen (TN) and heavy metals (Cu, Zn, Pb, Cd) was the most important driver of PDA. In addition, our path analysis suggests that at least part of the impacts of land use on potential denitrification were mediated via microbial abundance, soil pH and substrates including TN, dissolved organic carbon and nitrate. This study identifies the main predictors of denitrification at a regional scale which is needed to quantify the impact of human activities on ecosystem functionality under changing conditions.

The Carboniferous tetrapod Crassigyrinus scoticus is an enigmatic animal in terms of its morphology and its phylogenetic position. Crassigyrinus had extremely reduced forelimbs, and was aquatic, perhaps secondarily. Recent phylogenetic analyses tentatively place Crassigyrinus close to the whatcheeriids. Many Carboniferous tetrapods exhibit several characteristics associated with terrestrial locomotion, and much research has focused on how this novel locomotor mode evolved. However, to estimate the selective pressures and constraints during this important time in vertebrate evolution, it is also important to study early tetrapods like Crassigyrinus that either remained aquatic or secondarily became aquatic. We used computed tomographic scanning to search for more data about the skeletal morphology of Crassigyrinus and discovered several elements previously hidden by the matrix. These elements include more ribs, another neural arch, potential evidence of an ossified pubis and maybe of pleurocentra. We also discovered several additional metatarsals with interesting asymmetrical morphology that may have functional implications. Finally, we reclassify what was previously thought to be a left sacral rib as a left fibula and show previously unknown aspects of the morphology of the radius. These discoveries are examined in functional and phylogenetic contexts.

The reconstructed palate of Whatcheeria deltae indicates a skull that was unusually narrow: at least 2.2 times longer than wide if the pterygoids are conservatively placed in the horizontal plane. This maximum width is narrower than any other early tetrapod reconstructed so far. Rotating the pterygoids to produce a vaulted palate would produce an even narrower skull. Primitive palatal features include very narrow interpterygoid vacuities and a vomer, palatine, and ectopterygoid with fang-sized replacement pairs. It is derived in that there is no anterior palatal fenestra and the premaxilla has a substantial palatal shelf – a combination of characters shared only with Proterogyrinus among early tetrapods. There is a possible septomaxilla in one specimen. Whatcheeria differs from and is more derived than Pederpes, its likely sister taxon, in that only the pterygoid is covered with denticles, the vomer, palatine, and ectopterygoid containing labyrinthine teeth only. Reconstructed dental occlusion indicates that the large choana apparently accommodated the large dentary fangs; this would be a unique feature among early tetrapods. The palatal ramus of the pterygoid is longer than the quadrate ramus, which does not have a descending flange. Like Meckel's cartilage in the lower jaw, the palatoquadrate is fully ossified in larger specimens, such that in a posterior view of the skull the pterygoid is mostly hidden from sight by the epipterygoid. The ossified neurocranium consists of the basiparasphenoid and basioccipital; no ossified sphenethmoid has been found. Remains of otic capsules are partial, crushed, and smeared, so no useful morphology is available. The stapes appears to be more columnar and less plate-like than in many other primitive, early tetrapods.

Soil conservation on the Loess Plateau is important not only for local residents but also for reducing sediment downstream in the Yellow River. In this paper, we report a decrease in soil erosion from 2000 to 2010 as a result of the ‘Grain for Green' (GFG) Project. By using the Revised Universal Soil Loss Equation and data on land cover, climate and sediment yield, we found that soil erosion decreased from 6579.55tkm–2yr–1 in 2000 to 1986.66tkm–2yr–1 in 2010. During this period, there was a major land cover change from farmland to grassland in response to the GFG. The area of low vegetation coverage with severe erosion decreased dramatically, whereas the area of high vegetation coverage with slight erosion increased. Our study demonstrates that the reduction in soil erosion on the Loess Plateau contributed to the decrease in the sediment concentration in the Yellow River.

Since the ‘Grain to Green Program' was launched in 1999 in the Loess Plateau, China, the processes and patterns of ecological change have become important. Through the use of the fractional vegetation cover (FVC) index, this study examines the spatial distribution and temporal change of vegetation cover in the Loess Plateau during 2000–2014. Over this period more than 60% of the Loess Plateau has remained with little vegetation cover (FVC <30%). The spatial distribution pattern shows an overall increase from the NW to SE. Temporally, the vegetation cover exhibits a general trend of improvement. In 2000–2005, 2000–2010 and 2000–2014, the percentages of restored vegetation (vegetation with significantly increased FVC, P<0.05) were 2, 21 and 52%, respectively. The rate of vegetation cover restoration was highly variable among different bioclimatic zones. The expansion of restored vegetation was greater in the N of the Loess Plateau than in the south. Both human restoration activities and climatic fluctuation influenced the vegetation cover change. The ‘Grain to Green Program' emphasised vegetation restoration. Regional precipitation also had clear effects on vegetation cover. The results of this study reveal that vegetation change shows a non-linear process in response to climate and ecological restoration measures, and that the change gradually emerges over time. This study highlights the importance of considering the spatiotemporal variability in vegetation cover during the implementation of restoration programs, which could aid decision-making for the effective and sustainable management of large-scale restoration programs.

Specimens of trematopid amphibians from the Asturian (Upper Carboniferous) of Nýřany, Czech Republic, are redescribed as two taxa, namely Mordex calliprepes Steen and Mattauschia (gen. nov) laticeps Fritsch. Mordex calliprepes is represented by a single post-metamorphic specimen and has the diagnostic trematopid characters of the nasal region. Mattauschia laticeps is represented by one adult partial skull and mandible plus some fragments and two small post-metamorphic specimens including the species name-bearer. It has the trematopid-type modified lacrimal and a large but oval naris and appears to be the most primitive trematopid yet described. The stratigraphically sequential large trematopids Mattauschia, Fedexia, Ecolsonia and Acheloma show progressive acquisition of the derived features that characterise the terminal form Acheloma.

Mordex has a combination of primitive and derived characters and its position within the family is less clear. The many ‘branchiosaurs' in the Nýřany assemblage include specimens that could be larvae of both Mordex and Mattauschia but certain attribution is not possible and they are assigned to Olsoniformes incertae sedis. Mordex and Mattauschia appear to be terrestrial exotic elements in the Nýřany tetrapod assemblage, but with possible larvae in the lake assemblage. Representatives of at least four Palaeozoic dissorophoid families were present in late Middle Pennsylvanian/Asturian strata implying diversification of the Dissorophoidea prior to this time.

We describe four newly discovered and three-dimensionally preserved specimens of the discosauriscid seymouriamorph Discosauriscus pulcherrimus from the Lower Permian lacustrine sediments of the Boskovice Basin (Czech Republic). Their excellent preservation revealed new cranial features that allowed us to present a new skull reconstruction of middle-sized specimens of this tetrapod. The new features include: (1) the sculpturing of the anterior wedge-like process of the parasphenoid consisting of irregular ridges, grooves, low elevations and pits; and (2) the presence of a pair of semicircular crests on the posterodorsal surface of the parasphenoidal plate. We discuss the intraspecific variability of several characters, such as the type of dermal cranial ornamentation, and the morphology of the sensory grooves, nasolacrimal canal, dentition on the pterygoids and parasphenoid. The lengths of the skulls of four new specimens range from about 18 to 31mm and represent larval ontogenetic stages.

As the largest industrial and population centre in China, the Pearl River Delta is facing a growing threat of heavy metal pollution from local mining and power industries. This study investigates the distribution and potential health risks of copper (Cu), cadmium (Cd), lead (Pb) and zinc (Zn) in paddy soils and rice at four typical sites. The Nemerow synthetic pollution index (PN) of soils from Fogang, Dabao Mountain, Shaoguan and Lechang were 8.40, 9.10, 4.64 and 10.28, respectively, indicating serious pollution at all four sampling sites. The average concentrations of Cu, Cd, Pb and Zn in rice grains were 2.23, 10.98, 29.84 and 1.62 times their corresponding maximum allowable levels, indicating potential health risks to humans. Cd has greater bioavailability because of its high mobility from soil to roots, and its subsequent transfer to grains. Pb mainly accumulates in roots because of its lower translocation rate from roots to grains. The greatest health risk index for Cd and Pb for adults and children was at the Shaoguan site, probably due to pollution from atmospheric deposition. Cd and Pb had greater health risk indices than Cu and Zn at almost all sites, indicating a major health risk to local people.

The aquatic diplocaulid nectridean Keraterpeton galvani is the commonest taxon represented in the Jarrow Coal assemblage from Kilkenny, Ireland. The Jarrow locality has yielded the earliest known Carboniferous coal-swamp fauna in the fossil record and is, therefore, of importance in understanding the history and diversity of the diplocaulid clade. The morphology of Keraterpeton is described in detail with emphasis on newly observed anatomical features. A reconstruction of the palate includes the presence of interpterygoid vacuities and new morphological details of the pterygoid, parasphenoid and basicranial region. The hyoid apparatus comprising an ossified basibranchial element has not been reported previously in nectrideans. The structure of the scapulocoracoid and primitive nature of the humerus is described and the presence of a five-digit manus confirmed. Previously unrecognised accessory dermal ossifications are present in the pectoral girdle. Keraterpeton longtoni from the Bolsovian in Staffordshire, England, is also described and newly figured. The primitive condition in diplocaulids is defined on the basis of the earliest occurrence at Jarrow and discussed in relation to functional morphology and mode of life. The evolution of the diplocaulid clade is assessed in relation to the revised diagnoses that define the primitive condition in Keraterpeton.

Study of the residues of organochlorine pesticides (OCPs) can assist with the assessment of the contamination level, distribution, sources, transportation and trends in these selected OCPs. We investigated the concentration levels of hexachlorocyclohexanes (HCHs), dichlorodiphenyltrichloroethane (DDT), chlordane, which is currently used, and the never-used aldrin and dieldrin in the Ningbo region – a heavily industrialised and urbanised area in East China. OCPs were found in all soil samples and were predominantly composed of HCHs, DDT and its metabolites. The concentration of DDTs in the soil ranged from 2.2 to 566.6ngg–1, with a mean value of 55.6±94.8ngg–1, while the concentration of HCHs ranged from 2.7 to 28.2ngg–1, with an average of 4.6±2.9ngg–1. Aldrin, dieldrin, endrin and heptachlor were detected at much lower concentrations. The wide distribution of these never-used OCPs indicates that they have been input via long-range atmospheric processes. The results from analysis suggest that HCHs and DDTs were mainly derived from use in historical times.

Soil moisture is a key issue for eco-hydrological research in arid and semi-arid regions, and is primarily concerned with water availability for vegetation. Shallow and deep soil moisture occurs according to the maximum infiltration depth. Soil moisture has three-dimensional characteristics: inter-layer variability, horizontal heterogeneity and temporal variability. Soil moisture is affected by various factors including terrain, soil characteristics, climate and vegetation, and the effects of these change with time (e.g., rainfall patterns) and space (e.g., soil depth). In arid and semi-arid regions, deep soil moisture is of particular importance to vegetation restoration and the evaluation of vegetation sustainability; however, accurate prediction of the spatial distribution of deep soil moisture in the Loess Plateau of China still faces numerous challenges. Therefore, future research should focus on the mechanisms, models and scale effects of soil moisture, particularly for deep soil moisture.

We describe a new specimen of the aïstopod Oestocephalus from Five Points, Ohio, which preserves much of the posterior braincase. The specimen, the largest aïstopod skull described, preserves the postorbital region to the occiput. The posterior braincase has coossified the basioccipital, exoccipitals, and opisthotic. The parasphenoid is rostrally restricted, toothless, and highly vaulted along the cultriform process. The lateral walls of the cultriform process are further reinforced by large longitudinally running, ventral flanges from the parietals. Two large endochondral ventral projections from the basioccipital, previously interpreted as basal tuberosities for hypaxial muscle insertion, are here instead interpreted as articulations for the branchial skeleton. This interpretation is further supported by traces of vasculature that is consistent with what is seen in gill-bearing species. A model for the reorganisation of the basicranial region on the transition from hyomandibula to stapes is proposed, which suggests that gills, or gill-support skeletal elements, might be further distributed along the tetrapod stem than previously thought. These data further support the placement of aïstopods in the tetrapod stem group and require a reconsideration of our understanding of early tetrapod evolution.

In Scotland, the base of the Ballagan Formation has traditionally been placed at the first grey mudstone within a contiguous Late Devonian to Carboniferous succession. This convention places the Devonian–Carboniferous boundary within the Old Red Sandstone (ORS) Kinnesswood Formation. The consequences of this placement are that tetrapods from the Ballagan Formation were dated as late Tournaisian in age and that the ranges of typically Devonian fish found in the Kinnesswood Formation continued into the Carboniferous. The Pease Bay specimen of the fish Remigolepis is from the Kinnesswood Formation. Comparisons with its range in Greenland, calibrated against spores, show it was Famennian in age. Detailed palynological sampling at Burnmouth from the base of the Ballagan Formation proves that the early Tournaisian spore zones (VI and HD plus Cl 1) are present. The Schopfites species that occurs through most of the succession is Schopfites delicatus rather than Schopfites claviger. The latter species defines the late Tournaisian CM spore zone. The first spore assemblage that has been found in Upper ‘ORS' strata underlying the Ballagan Formation (Preston, Whiteadder Water), contains Retispora lepidophyta and is from the early latest Famennian LL spore zone. The spore samples are interbedded with volcaniclastic debris, which shows that the Kelso Volcanic Formation is, in part, early latest Famennian in age. These findings demonstrate that the Ballagan Formation includes most of the Tournaisian with the Devonian–Carboniferous boundary positioned close to the top of the Kinnesswood Formation. The Stage 6 calcrete at Pease Bay can be correlated to the equivalent section at Carham, showing that it represents a time gap equivalent to the latest Famennian glaciation(s). Importantly, some of the recently described Ballagan Formation tetrapods are older than previously dated and now fill the key early part of Romer's Gap.

The lower Mississippian Ballagan Formation of northern Britain is one of only two successions worldwide to yield the earliest known tetrapods with terrestrial capability following the end-Devonian mass extinction event. Studies of the sedimentary environments and habitats in which these beasts lived have been an integral part of a major research project into how, why and under what circumstances this profound step in the evolution of life on Earth occurred. Here, a new palaeogeographic map is constructed from outcrop data integrated with new and archived borehole material. The map shows the extent of a very low-relief coastal wetland developed along the tropical southern continental margin of Laurussia. Coastal floodplains in the Midland Valley and Tweed basins were separated from the marginal marine seaway of the Northumberland–Solway Basin to the south by an archipelago of more elevated areas. A complex mosaic of sedimentary environments was juxtaposed, and included fresh and brackish to saline and hypersaline lakes, a diverse suite of floodplain palaeosols and a persistent fluvial system in the east of the region. The strongly seasonal climate led to the formation of evaporite deposits alternating with flooding events, both meteoric and marine. Storm surges drove marine floods from the SW into both the western Midland Valley and Northumberland–Solway Basin; marine water also flooded into the Tweed Basin and Tayside in the east. The Ballagan Formation is a rare example in the geological record of a tropical, seasonal coastal wetland that contains abundant, small-scale evaporite deposits. The diverse sedimentary environments and palaeosol types indicate a network of different terrestrial and aquatic habitats in which the tetrapods lived.

Specimen-level phylogenetic approaches are widely used in molecular biology for taxonomic and systematic purposes. However, they have been largely ignored in analyses based on morphological traits, where phylogeneticists mostly resort to species-level analyses. Recently, a number of specimen-level studies have been published in vertebrate palaeontology. These studies indicate that specimen-level phylogeny may be a very useful tool for systematic reassessments at low taxonomic levels. Herein, we review the challenges when working with individual organisms as operational taxonomic units in a palaeontological context, and propose guidelines of how best to perform a specimen-level phylogenetic analysis using the maximum parsimony criterion. Given that no single methodology appears to be perfectly suited to resolve relationships among individuals, and that different taxa probably require different approaches to assess their systematics, we advocate the use of a number of methodologies. In particular, we recommend the inclusion of as many specimens and characters as feasible, and the analysis of relationships using an extended implied weighting approach with different downweighting functions. Resulting polytomies should be explored using a posteriori pruning of unstable specimens, and conflicting tree topologies between different iterations of the analysis should be evaluated by a combination of support values such as jackknifing and symmetric resampling. Species delimitation should be consistent among the ingroup and based on a reproducible approach. Although time-consuming and methodologically challenging, specimen-level phylogenetic analysis is a highly useful tool to assess intraspecific variability and provide the basis for a more informed and accurate creation of species-level operational taxonomic units in large-scale systematic studies. It also has the potential to inform us about past speciation processes, morphological trait evolution, and their potential intrinsic and extrinsic drivers in pre-eminent detail.

A comprehensive analysis of early dinosaur relationships raised the possibility that the group may have originated in Laurasia (Northern Hemisphere), rather than Gondwana (Southern Hemisphere) as often thought. However, that study focused solely on morphology and phylogenetic relationships and did not quantitatively evaluate this issue. Here, we investigate dinosaur origins using a novel Bayesian framework uniting tip-dated phylogenetics with dynamic, time-sliced biogeographic methods, which explicitly account for the age and locality of fossils and the changing interconnections of areas through time due to tectonic and eustatic change. Our analysis finds strong support for a Gondwanan origin of Dinosauria, with 99 % probability for South America (83 % for southern South America). Parsimony analysis gives concordant results. Inclusion of time-sliced biogeographic information affects ancestral state reconstructions (e.g., high connectivity between two regions increases uncertainty over which is the ancestral area) and influences tree topology (disfavouring uniting fossil taxa from localities that were widely separated during the relevant time slice). Our approach directly integrates plate tectonics with phylogenetics and divergence dating, and in doing so reaffirms southern South America as the most likely area for the geographic origin of Dinosauria.

Open palates with large interpterygoid vacuities are a diagnostic characteristic of temnospondyl amphibians, the most species-rich group of early tetrapods. Aside from their functional roles, several other aspects of such vacuities, such as their variation and spatial relationships relative to the orbits, have received only scarce attention. The present work examines patterns of shape and size changes in the orbits and vacuities of temnospondyls using a time-calibrated phylogeny of 69 temnospondyl taxa and 13 additional early tetrapod ‘outgroups' (colosteids, an embolomere, ‘microsaurs' and nectrideans). Orbit and vacuity outlines are quantified in a comparative framework using standard eigenshape analyses. In addition, we employ a series of ratios of linear measurements of both orbits and vacuities, and subject them to a phylogenetic principal component analysis in order to evaluate their proportional changes relative to the skull and to one another. Finally, we examine rates of evolutionary change and their associated shifts for shape and size for both structures, and assess the strength and significance of the correlations between these two variables using phylogenetic generalised least squares analyses. Although orbits and vacuities have fairly simple outlines, they both reveal complex models of proportional change across the temnospondyl phylogeny. These changes exhibit strong phylogenetic signal, that is, trait covariance among taxa is predicted by tree topology. We discuss the hypothesis that, early in tetrapod evolution, the functional role of the vacuities was related to the accommodation of the anterior jaw muscles. Only later in evolution did such vacuities serve to accommodate the eye muscles only.

The present study explores the macroevolutionary dynamics of shape changes in the humeri of all major grades and clades of early tetrapods and their fish-like forerunners. Coordinate point eigenshape analysis applied to humeral outlines in extensor view reveals that fish humeri are more disparate than those of most early tetrapod groups and significantly separate from the latter. Our findings indicate sustained changes in humeral shape in the deepest portions of the tetrapod stem group and certain portions of the crown. In the first half of sampled tetrapod history, subclades show larger than expected humeral disparity, suggesting rapid diffusion into morphospace. Later in tetrapod evolution, subclades occupy smaller and non-overlapping morphospace regions. This pattern may reflect in part increasing specialisations in later tetrapod lineages. Bayesian shifts in rates of evolutionary change are distributed discontinuously across the phylogeny, and most of them occur within rather than between major groups. Most shifts with the highest Bayesian posterior probabilities are observed in lepospondyls. Similarly, maximum likelihood analyses of shifts support marked rate accelerations in lepospondyls and in various subclades within that group. In other tetrapod groups, rates either tend to slow down or experience only small increases. Somewhat surprisingly, no shifts are concurrent with structural, functional, or ecological innovations in tetrapod evolution, including the origin of digits, the water–land transition and increasing terrestrialisation. Although counterintuitive, these results are consistent with a model of continual phenotypic innovation that, although decoupled from key evolutionary changes, is possibly triggered by niche segregation in divergent clades and grades of early tetrapods.