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This study focuses on the genus Dioncomena and its acoustics, particularly the unique songs produced by male Dioncomena that consist of several distinct elements in a fixed sequence, culminating in a coda that typically elicits a response from a receptive female. We also examine the inflated pronotal lobes, which we term prebullae, that are prominently developed in some Dioncomena species but not in others. We discuss the role of prebullae in the context of acoustic communication in Dioncomena and other related Phaneropterini genera that have similar lateral pronotal lobes. We found that prebullae size is correlated with habitat distribution, with larger prebullae occurring in isolated species while aggregation-prone species have smaller or less pronounced prebullae. Using micro-computer tomography we show sexual dimorphism in the 3D geometry of the acoustic tracheae, being larger in the male. Interestingly, the tracheae are coupled by a septum, like in field crickets, which suggests potential cross talk. We define three groups of Dioncomena based on altitude preferences, ecology, color patterns, and songs: the jagoi-, tanneri-, and ornata-groups. We describe the songs of several species, including newly identified species such as D. flavoviridis sp. nov., D. magombera sp. nov., D. ngurumontana sp. nov., D. sanje sp. nov., D. tanneri, D. versicolor sp. nov., and D. zernyi. We also provide information on the nymphs, development time, and mating behavior of various species reared in the laboratory, shedding light on their phenology and adaptations to their habitats.

Accelerating rates of biodiversity loss underscore the need to understand how species achieve resilience—the ability to resist and recover from a/biotic disturbances. Yet, the factors determining the resilience of species remain poorly understood, due to disagreements on its definition and the lack of large-scale analyses. Here, we investigate how the life history of 910 natural populations of animals and plants predicts their intrinsic ability to be resilient. We show that demographic resilience can be achieved through different combinations of compensation, resistance and recovery after a disturbance. We demonstrate that these resilience components are highly correlated with life history traits related to the species’ pace of life and reproductive strategy. Species with longer generation times require longer recovery times post-disturbance, whilst those with greater reproductive capacity have greater resistance and compensation. Our findings highlight the key role of life history traits to understand species resilience, improving our ability to predict how natural populations cope with disturbance regimes.

Trichomonas gallinae, a protozoan parasite infecting a wide range of birds, has been a cause 3 for conservation concern since its recognition as an emerging infectious disease, having had 4 notable impacts on several avian species, including causing widespread mortality in 5 greenfinches (Chloris chloris), and chaffinches (Fringilla coelebs), and hindering the recovery 6 of the endangered pink pigeon (Nesoenas mayeri). Horizontal transmission between birds 7 congregating at feeding sites, such as those provided for conservation and species management 8 purposes, is an important driver in the spread of T. galliane. Supplementary feeding is a key 9 conservation intervention for the European turtle dove (Streptopelia turtur), which is declining 10 across its range, driven at least partially by a loss of natural food resources. Due to the link 11 between T. gallinae transmission and supplementary feeding, we consider the prevalence of 12 this parasite amongst European turtle dove in relation to diet, in the first study to analyse these 13 two factors in the decline of this species together. Using birds sampled from breeding and 14 wintering grounds, the dietary composition of individuals was compared to the presence of T. 15 gallinae, and specific T. gallinae strains. Dietary variation was summarised into two axes using 16 detrended correspondence analysis, neither were associated with the presence of T. gallinae, 17 or any specific strains. The proportion of diet accounted for by cultivated seeds did not affect 18 parasite presence, despite the spread of this parasite being associated with supplementary 19 feeding due to an increase in contact between potentially infected and uninfected birds 20 gathering at high densities at feeding sites. Significant dietary overlap was observed between 21 infected and uninfected individuals in all sites, with almost complete dietary overlap being 22 observed in one breeding site. The level of dietary overlap between individuals infected with 23 specific strains fluctuated between sampling seasons, indicating no consistent relationship 24 between diet and infection status.

Katydids produce sound for signaling and communication by stridulation of the tegmina. Unlike crickets, most katydids are known to sing at ultrasonic frequencies. This has drawn interest in the investigation of the biophysics of ultrasonic sound production, detection, evolution, and ecology (including predator–prey interactions) of these katydids. However, most of these studies are based on species from the Neotropics, while little is known about katydid species from the hyperdiverse region of Southeast Asia. To address this, a concerted effort to document, record, and describe the calling songs of Southeast Asian katydids, especially species that call at ultrasonic frequencies, was made. A study spanning two years (2018–2020) in the Malay Peninsula (Singapore and Malaysia), Borneo (Brunei Darussalam and Sabah), and the Philippines revealed previously unknown calls of 24 katydid species from four subfamilies. The calling songs of Southeast Asian katydid species are highly diversified in terms of time and frequency. Call structure can range from isolated syllables (e.g., Holochlora), continuous trills (e.g., Axylus philippinus), to short pulse-trains (e.g., Euanisous teuthroides) and complex echemes (e.g., Conocephalus spp.), with 87.5% of species having ultrasonic peak frequencies and 12.5% being considered extreme ultrasonic callers (peak frequency >40 kHz). The call spectrum ranges from tonal (e.g., spectral entropy is 6.8 in Casigneta sp. 2) to resonant (entropy is 8.8 in Conocephalus cognatus). Of the 24 species whose calls are described here, we imaged and described the sound-producing structures of 18. This study provides a preliminary overview of the acoustic diversity of katydids in Southeast Asia, and the authors hope to inspire further investigation into the bioacoustics of little-known katydids from these areas. Amassing a database of calling songs and sound-producing organ illustrations from different species is important to address taxonomic impediments while advancing our knowledge about the bioacoustics of Southeast Asian katydids.

Frogs exhibit complex anatomical features of the pelvis, limbs and spine, long assumed to represent specialisations for jumping. Yet frogs employ a wide range of locomotor modes, with several taxa featuring primary locomotor modes other than jumping. Using a combination of techniques (CT imaging and 3D visualization, morphometrics, phylogenetic mapping), this study aims to determine the link between skeletal anatomy and locomotor style, habitat type and phylogenetic history, shedding new light on how functional demands impact morphology. Body and limb measurements for 164 taxa from all the recognised anuran families are extracted from digitally segmented CT scans of whole frog skeletons and analysed using various statistical techniques. We find that the expansion of the sacral diapophyses is the most important variable for predicting locomotor mode, which was more closely correlated with frog morphology than either habitat type or phylogenetic relationships. Predictive analyses suggest that skeletal morphology is a useful indicator of jumping but less so for other locomotor modes, suggesting that there is a wide range of anatomical solutions to performing locomotor styles such as swimming, burrowing or walking.

Although tropical forests differ substantially in form and function, they are often represented as a single biome in global change models, hindering understanding of how different tropical forests will respond to environmental change. The response of the tropical forest biome to environmental change is strongly influenced by forest type. Forest types differ based on functional traits and forest structure, which are readily derived from high resolution airborne remotely sensed data. Whether the spatial resolution of emerging satellite-derived hyperspectral data is sufficient to identify different tropical forest types is unclear. Here, we resample airborne remotely sensed forest data at spatial resolutions relevant to satellite remote sensing (30 m) across two sites in Malaysian Borneo. Using principal component and cluster analysis, we derive and map seven forest types. We find ecologically relevant variations in forest type that correspond to substantial differences in carbon stock, growth, and mortality rate. We find leaf mass per area and canopy phosphorus are critical traits for distinguishing forest type. Our findings highlight the importance of these parameters for accurately mapping tropical forest types using space borne observations.

During the main COVID-19 global pandemic lockdown period of 2020 an impromptu set of pollination ecologists came together via social media and personal contacts to carry out standardised surveys of the flower visits and plants in gardens. The surveys involved 67 rural, suburban and urban gardens, of various sizes, ranging from 61.18° North in Norway to 37.96° South in Australia, resulting in a data set of 25,174 rows, with each row being a unique interaction record for that date/site/plant species, and comprising almost 47,000 visits to flowers, as well as records of flowers that were not visited by pollinators, for over 1,000 species and varieties belonging to more than 460 genera and 96 plant families. The more than 650 species of flower visitors belong to 12 orders of invertebrates and four of vertebrates. In this first publication from the project, we present a brief description of the data and make it freely available for any researchers to use in the future, the only restriction being that they cite this paper in the first instance. The data generated from these global surveys will provide scientific evidence to help us understand the role that private gardens (in urban, rural and suburban areas) can play in conserving insect pollinators and identify management actions to enhance their potential.

The taxonomically diverse terrestrial tetrapod fauna from the late Mississippian East Kirkton Limestone includes the earliest known members of stem Amphibia and stem Amniota. Here we name and describe a new East Kirkton tetrapod with an unusual hindlimb morphology reminiscent of that of several stem- and primitive crown amniotes. It displays a unique ilium with two slender and elongate processes and a 5-digit pes with a long, stout metatarsal IV and a greatly elongate digit IV. The new taxon broadens our knowledge of East Kirkton tetrapods, adding to the remarkable diversity of their hindlimb constructions, functional specializations, locomotory modes, and adaptations to a wide variety of substrates. An unweighted character parsimony analysis places the new taxon in a polytomy alongside some other Carboniferous groups. Conversely, weighted parsimony and Bayesian analyses retrieve it among the earliest diverging stem amniotes, either as the basalmost anthracosaur or within a clade that includes alsoEldeceeonandSilvanerpeton, crownward of an array of chroniosaurs plus anthracosaurs.

Low- and middle-income countries in Southeast and East Asia face a range of challenges related to the rapid pace of urbanisation in the region, the scale of pollution, climate change, loss of ecosystem services and associated difficulties for ecological restoration. Possible pathways towards a more sustainable future lie in the applications of nature-based solutions (NBS). However, there is relatively little literature on the application of NBS in the region, particularly Southeast Asia. In this paper we address this gap by assessing the socio-ecological challenges to the application of NBS in the region – one of the most globally biodiverse. We first provide an overview and background on NBS and its underpinnings in biodiversity and ecosystem services. We then present a typology describing five unique challenges for the application of NBS in the region: (1) Characteristics of urbanisation; (2) Biophysical environmental and climatic context; (3) Environmental risks and challenges for restoration; (4) Human nature relationships and conflicts; and (5) Policy and governance context. Exploiting the opportunities through South-South and North-South collaboration to address the challenges of NBS in Southeast and East Asia needs to be a priority for government, planners and academics. Highlights Most research on Nature-Based Solutions (NBS) is in high income nations with low population growth rates and regulated urban planning.; Urban blue-green infrastructure is being degraded in low and middle-income Southeast and East Asian countries.; Applications of NBS in the region need to address the unique socio-ecological challenges.; We provide an overview of key knowledge gaps to support the implementation of NBS.

Aim: Body size explains most variation in fitness within animal populations and is therefore under constant selection from ecological and reproductive pressures which often promote its evolution in sex-specific directions, leading to sexual size dimorphism (SSD). Multiple hypotheses have been proposed to explain the vast diversity of SSD across species. These hypotheses emphasise (i) the mate competition benefits to larger male size (sexual selection), (ii) the benefits of larger female size for fecundity (fecundity selection), (iii) the simultaneous benefits of niche divergence for males and females to reduce inter-sexual competition for ecological resources (natural selection), and (iv) the underlying impact of geographic variation in climatic pressures expected to shape large-scale patterns of SSD in synergy with the above selection pressures (e.g., intensification of fecundity selection as breeding seasons shorten). Based on a novel, global-scale amphibian dataset, we address the shortage of large-scale, integrative tests of these four hypotheses. Location: Global. Time period: Extant. Major taxa studied: Class Amphibia. Methods: Using a >3,500 species dataset spanning body size, ecological, life history, geographic and climatic data, we performed phylogenetic linear models to address the sexual, fecundity, ecological and climatic hypotheses of SSD. Results: SSD evolution is discordant between anurans and salamanders. Anuran SSD is shaped by climate – male-biased SSD increases with temperature seasonality – and by nesting site. In salamanders, SSD converges across species that occupy the same microhabitat types (‘ecodimorphs’), while reproductive or climatic pressures have no effects on their SSD. These contrasts are associated with latitudinal gradients of SSD in anurans, but not in salamanders. Main conclusions: Amphibian SSD is driven by ecological and climatic pressures, while no roles for sexual or fecundity selection were detected. We show that macroevolutionary processes determined by different forms of selection lead to latitudinal patterns of trait diversity, and the lack of them.

Resilience has emerged as a key concept in ecology and conservation biology to understand and predict ecosystem responses to global change. In its broadest sense, resilience describes the ability of an ecosystem to resist, and recover from, a disturbance. However, the application of such a concept in different subdisciplines of ecology and in different study systems has resulted in a wide disparity of definitions and ways of quantifying resilience.\ud This Special Feature, which spans the Journal of Ecology, Journal of Animal Ecology and Functional Ecology, provides an overview of how ecologists define, quantify, compare and predict resilience across different study systems.\ud The 29 contributions to this Special Feature show the broad range of approaches used by ecologists to study resilience. Almost half of the contributions (48%) study resilience at the community level, with a 30% of them studying resilience at multiple levels of biological organisation. A large proportion of these articles are observational (42%), experimental (14%) or a combination of both (17%) while a 17% utilise theoretical or computational approaches. Although 38%, 21% and 14% of the studies were based solely on plants, animals or micro-organisms, respectively, 17% of them incorporated these multiple trophic levels.\ud Synthesis. A unified ecological understanding of resilience across systems and taxa requires a trans-disciplinary consensus on what resilience actually is and how to best measure it. Here, we provide an overview of how ecologists define, quantify, compare and predict resilience across different ecological systems and subdisciplines, with reference to the diverse approaches used by contributions to this Special Feature. We identify four key recommendations to harmonise future efforts in resilience research: (a) define resilience using existing theoretical frameworks; (b) use common and comparable metrics to measure resilience; (c) clearly contextualise and define the pre- and post-disturbance state of the ecological system and (d) consider explicitly the disturbance type and regime impacting the system.

Through phylogenetic modelling, we previously presented strong support for diversification decline in the three major subclades of dinosaurs (Sakamoto et al . 2016 Proc. Natl Acad. Sci. USA 113 , 5036–5040. ( doi:10.1073/pnas.1521478113 )). Recently, our support for this model has been criticized (Bonsor et al . 2020 R. Soc. Open Sci. 7 , 201195. ( doi:10.1098/rsos.201195 )). Here, we highlight that these criticisms seem to largely stem from a misunderstanding of our study: contrary to Bonsor et al .'s claims, our model accounts for heterogeneity in diversification dynamics, was selected based on deviance information criterion (DIC) scores (not parameter significance), and intercepts were estimated to account for uncertainties in the root age of the phylogenetic tree. We also demonstrate that their new analyses are not comparable to our models: they fit simple, Dinosauria-wide models as a direct comparison to our group-wise models, and their additional trees are subclades that are limited in taxonomic coverage and temporal span, i.e. severely affected by incomplete sampling. We further present results of new analyses on larger, better-sampled trees ( N = 961) of dinosaurs, showing support for the time-quadratic model. Disagreements in how we interpret modelled diversification dynamics are to be expected, but criticisms should be based on sound logic and understanding of the model under discussion.

The braincase anatomy of the Pennsylvanian diadectomorphLimnoscelis dynatisis described in detail, based upon high-resolution X-ray microcomputed tomography. Both supraoccipitals and most of the prootics and opisthotics are preserved. The known portions of the left prootic, opisthotic, and supraoccipital enclose complete sections of the endosseous labyrinth, including the anterior, posterior, and lateral semicircular canals, the vestibule, the cochlear recess, and the canal for the endolymphatic duct. The fossa subarcuata is visible anteromedial to the anterior semicircular canal. The presumed endolymphatic fossae occur in the dorsal wall of the posteromedial portion of the supraoccipital. Both the fossa subarcuata and the fossa endolymphatica lie in the cerebellar portion of the cranial cavity. In order to investigate the phylogenetic position ofL. dynatiswe used a recently published data matrix, including characters of the braincase, and subjected it to maximum parsimony analyses under a variety of character weighting schemes and to a Bayesian analysis.Limnoscelis dynatisemerges as sister taxon toL. paludis, and both species form the sister group to remaining diadectomorphs. Synapsids and diadectomorphs are resolved as sister clades in ∼90% of all the most parsimonious trees from the unweighted analysis, in the single trees from both the reweighted and the implied weights analyses, as well in the Bayesian tree.

Despite the increasing importance of molecular sequence data, morphology still makes an important contribution to resolving the phylogeny of many groups, and is the only source of data for most fossils. Most systematists sample morphological characters as broadly as possible on the principle of total evidence. However, it is not uncommon for sampling to be focused on particular aspects of anatomy, either because characters therein are believed to be more informative, or because preservation biases restrict what is available. Empirically, the optimal trees from partitions of morphological data sets often represent significantly different hypotheses of relationships. Previous work on hard-part versus soft-part characters across animal phyla revealed significant differences in about a half of sampled studies. Similarly, studies of the craniodental versus postcranial characters of vertebrates revealed significantly different trees in about one-third of cases, with the highest rates observed in non-avian dinosaurs. We test whether this is a generality here with a much larger sample of 81 published data matrices across all major dinosaur groups. Using the incongruence length difference test and two variants of the incongruence relationship difference test, we found significant incongruence in about 50% of cases. Incongruence is not uniformly distributed across major dinosaur clades, being highest (63%) in Theropoda and lowest (25%) in Thyreophora. As in previous studies, our partition tests show some sensitivity to matrix dimensions and the amount and distribution of missing entries. Levels of homoplasy and retained synapomorphy are similar between partitions, such that incongruence must partly reflect differences in patterns of homoplasy between partitions, which may itself be a function of modularity and mosaic evolution. Finally, we implement new tests to determine which partition yields trees most similar to those from the entire matrix. Despite no bias across dinosaurs overall, there are striking differences between major groups. The craniodental characters of Ornithischia and the postcranial characters of Saurischia yield trees most similar to the “total evidence” trees derived from the entire matrix. Trees from these same character partitions also tend to be most stratigraphically congruent: a mutual consilience suggesting that those partitions yield more accurate trees. [Dinosauria; homoplasy; partition homogeneity.]

The exponential growth of human population and infrastructure is significantly reducing the amount of ecological resources available for wild animals. We analyzed the effect of human activity on Barbary macaques (Macaca sylvanus), an endangered species restricted to the fragmented forests of Morocco and Algeria, using location data from five social groups inhabiting Ifrane National Park, Morocco. We used a resource selection function to explore the effects of anthropogenic disturbance on macaque habitat selection, using nine natural, social, and anthropogenic disturbance variables as predictors. Forest cover, home range overlap, herding route proximity, and road proximity were all significant predictors of habitat use. Macaques avoided areas used by local shepherds, to reduce the risk of attack by shepherds’ dogs, but approached roads to increase the chances of provisioning by tourists. However, herding route and road use varied seasonally in line with levels of human use, suggesting that macaques may be navigating their environment strategically (in space and time) to balance food acquisition and risk avoidance. The results of this study highlight the importance of assessing human impact on habitat selection in both space and time. Our data on seasonal variations in macaques’ use of roads can help prevent road injuries, a major source of mortality for provisioned macaques, by focusing management efforts by national park workers in time and space. Furthermore, understanding when and where macaques seek provisioning from tourists can help combat provisioning, which negatively impacts macaque health, behavior, and susceptibility to poaching.

Determining the acoustic ecology of extinct or rare species is challenging due to the inability to record their acoustic signals or hearing thresholds. Katydids and their relatives (Orthoptera: Ensifera) offer a model for inferring acoustic ecology of extinct and rare species, due to allometric parameters of their sound production organs. Here, the bioacoustics of the orthopteran Prophalangopsis obscura are investigated. This species is one of only eight remaining members of an ancient family with over 90 extinct species that dominated the acoustic landscape of the Jurassic. The species is known from only a single confirmed specimen–the 150-year-old holotype material housed at the London Natural History Museum. Using Laser-Doppler Vibrometry, 3D surface scanning microscopy, and known scaling relationships, it is shown that P. obscura produces a pure-tone song at a frequency of ~4.7 kHz. This frequency range is distinct but comparable to the calls of Jurassic relatives, suggesting a limitation of early acoustic signals in insects to sonic frequencies (

The long-term accumulation of biodiversity has been punctuated by remarkable evolutionary transitions that allowed organisms to exploit new ecological opportunities. Mesozoic flying reptiles (the pterosaurs), which dominated the skies for more than 150 million years, were the product of one such transition. The ancestors of pterosaurs were small and probably bipedal early archosaurs1, which were certainly well-adapted to terrestrial locomotion. Pterosaurs diverged from dinosaur ancestors in the Early Triassic epoch (around 245 million years ago); however, the first fossils of pterosaurs are dated to 25 million years later, in the Late Triassic epoch. Therefore, in the absence of proto-pterosaur fossils, it is difficult to study how flight first evolved in this group. Here we describe the evolutionary dynamics of the adaptation of pterosaurs to a new method of locomotion. The earliest known pterosaurs took flight and subsequently appear to have become capable and efficient flyers. However, it seems clear that transitioning between forms of locomotion2,3—from terrestrial to volant—challenged early pterosaurs by imposing a high energetic burden, thus requiring flight to provide some offsetting fitness benefits. Using phylogenetic statistical methods and biophysical models combined with information from the fossil record, we detect an evolutionary signal of natural selection that acted to increase flight efficiency over millions of years. Our results show that there was still considerable room for improvement in terms of efficiency after the appearance of flight. However, in the Azhdarchoidea4, a clade that exhibits gigantism, we test the hypothesis that there was a decreased reliance on flight5–7 and find evidence for reduced selection on flight efficiency in this clade. Our approach offers a blueprint to objectively study functional and energetic changes through geological time at a more nuanced level than has previously been possible. Phylogenetic statistical analyses, biophysical models and information from the fossil record show that an evolutionary signal of natural selection acted to increase the flight efficiency of pterosaurs over millions of years.

Spotted wing drosophila, Drosophila suzukii, is a serious invasive pest impacting the production of multiple fruit crops, including soft and stone fruits such as strawberries, raspberries and cherries. Effective control is challenging and reliant on integrated pest management which includes the use of an ever decreasing number of approved insecticides. New means to reduce the impact of this pest that can be integrated into control strategies are urgently required. In many production regions, including the UK, soft fruit are typically grown inside tunnels clad with polyethylene based materials. These can be modified to filter specific wavebands of light. We investigated whether targeted spectral modifications to cladding materials that disrupt insect vision could reduce the incidence of D. suzukii. We present a novel approach that starts from a neuroscientific investigation of insect sensory systems and ends with infield testing of new cladding materials inspired by the biological data. We show D. suzukii are predominantly sensitive to wavelengths below 405 nm (ultraviolet) and above 565 nm (orange & red) and that targeted blocking of lower wavebands (up to 430 nm) using light restricting materials reduces pest populations up to 73% in field trials.

Day-camouflaged leaf-mimic katydids Typophyllum spp. have a remarkable way of evading predators as male and female forewings appear as bite-damaged leaves complete with necrotic spots. As in all other katydids, males produce sound signals to attract females by rubbing their forewings together. The biophysical properties of these special leaf-like forewings remain obscure. Here we study the wing mechanics and resonances of Typophyllum spurioculis, a new species of leaf-mimic katydid with a broad distribution in the Andes from Western Ecuador to the middle Central Cordillera in Colombia. This species performs an unusual laterally directed aposematic display, showing orange spots that simulate eyes at the leg base. At night, males are conspicuous by their loud, audible calling songs, which exhibit two spectral peaks at ca. 7 and 12 kHz. Using micro-scanning laser Doppler vibrometry we find the effective sound radiators of the wings (speculae) vibrate with three modes of vibration, two of which include the frequencies observed in the calling song. Remarkably, this resonance is preserved in the parts of the wings mimicking necrotic leaves, which are in theory not specialised for sound production. The eyespot function is discussed.

Skeletal remains of a new early Miocene (Ottnangian, MN 4 mammal zone) monitor lizard, Varanus mokrensis sp. nov., are described from two karst fissures in the Mokrá-Western Quarry (1/2001 Turtle Joint; 2/2003 Reptile Joint), Czech Republic, providing the first documented example of a European varanid for which osteological data permit a well-supported assignment to the genus Varanus. The new species is morphologically similar to the Recent Indo-Asiatic varanids of the Varanus bengalensis group. It differs from all other Varanus species on the basis of a single autapomorphy and a combination of 11 characters. As a distinguishing feature of V. mokrensis, the parietal and squamosal processes of the postorbitofrontal form a narrowly acute angle. The teeth show distinct, smooth cutting edges along the mesial and distal margins of the apical portion of their crowns. This feature is not observed in most extant Asiatic Varanus species and may represent a plesiomorphic condition. The results of parsimony phylogenetic analyses, with and without character reweighting, reveal poor resolution within Varanus. A Bayesian analysis shows V. mokrensis to be closely related to extant representatives of the Indo-Asiatic Varanus clade, with close affinities to the V. bengalensis species group. The topology of the Bayesian tree supports the hypothesis that Miocene monitors from Mokrá are representatives of a lineage that is ancestral to the well-defined clade of extant African varanids, including the early Miocene V. rusingensis. In addition, our results support a Eurasian origin for the varanid clade. The extant African Varanus species probably originated in the late Oligocene. The radiation of African varanids probably occurred during the late Oligocene to early Miocene time interval. The occurrence of Varanus in the early Miocene of Mokrá-Western Quarry corresponds to the warm phase of the Miocene Climatic Optimum. Remains of a diverse aquatic and heliophobe amphibian fauna at the 2/2003 Reptile Joint site indicate more humid conditions than those at the 1/2001 Turtle Joint site.

The known diversity of tetrapods of the Devonian period has increased markedly in recent decades, but their fossil record consists mostly of tantalizing fragments1–15. The framework for interpreting the morphology and palaeobiology of Devonian tetrapods is dominated by the near complete fossils of Ichthyostega and Acanthostega; the less complete, but partly reconstructable, Ventastega and Tulerpeton have supporting roles2,4,16–34. All four of these genera date to the late Famennian age (about 365–359 million years ago)—they are 10 million years younger than the earliest known tetrapod fragments5,10, and nearly 30 million years younger than the oldest known tetrapod footprints35. Here we describe Parmastega aelidae gen. et sp. nov., a tetrapod from Russia dated to the earliest Famennian age (about 372 million years ago), represented by three-dimensional material that enables the reconstruction of the skull and shoulder girdle. The raised orbits, lateral line canals and weakly ossified postcranial skeleton of P. aelidae suggest a largely aquatic, surface-cruising animal. In Bayesian and parsimony-based phylogenetic analyses, the majority of trees place Parmastega as a sister group to all other tetrapods. Three-dimensionally preserved fossils of Parmastega aelidae, a newly described tetrapod from the earliest Famennian (Late Devonian) of Russia, provide detailed insights into the morphology and palaeobiology of the earliest tetrapods.

Over the last 20 years, a new group of systemic insecticides – the neonicotinoids - has gained prominence in arable systems, and their application globally has risen year on year. Previous modelling studies using long-term data have suggested that neonicotinoid application has had a detrimental impact on bird populations, but these studies were either limited to a single species or neglected to analyse specific exposure pathways in conjunction with observed population trends. Using bird abundance data, neonicotinoid usage records and cropping data for England at a 5x5 km resolution, generalised linear mixed models were used to test for spatio-temporal associations between neonicotinoid use and changes in the populations of 22 farmland bird species between 1994 and 2014, and to determine whether any associations were explained by dietary preferences. We assigned farmland bird species to three categories of dietary exposure to neonicotinoids based on literature data for species diets and neonicotinoid residues present in dietary items. Significant estimates of neonicotinoid-related population change were obtained for 13 of the 22 species (9 positive effects, 4 negative effects). Model estimates for individual species were not collectively explained by dietary risk categories, so dietary exposure to neonicotinoids via ingestion of treated seeds and seedlings could not be confirmed as a causal factor in farmland bird declines. Although it is not possible to infer any generic effect of dietary exposure to neonicotinoids on farmland bird populations, our analysis identifies three species with significant negative estimates that may warrant further research (house sparrow Passer domesticus, skylark Alauda arvensis and red-legged partridge Alectoris rufa). We conclude that there was either no consistent effect of dietary exposure to neonicotinoids on farmland bird populations in England, or that any over-arching effect was not detectable using our study design. The potential for indirect effects of insecticide use on bird populations via reduced food availability was not considered here and should be a focus for future research.

The aim of this study is to investigate the colouration of parrots in an evolutionary context utilising novel metrics for the quantification of colour pattern. Previously, the study of animal colouration has been limited by its inability to produce statistically robust and objective measures of pattern. We propose the use of image correlograms, a method adapted from the field of computer science where it is used in the indexing and retrieval of images for search engines, in order to evaluate the whole colour pattern of parrot species as taken from field guides. This is the largest study of parrot colouration in terms of number of species included in the study (403) and includes the ancestral reconstruction of the original parrot’s colouration as well as how colouration has changed over time. Finally, the information gained on the parrots’ colouration is applied to an issue of pressing concern for those studying the conservation of parrots, how differences in colour pattern relates to conservation status.

Transparency is a greatly advantageous form of camouflage, allowing species to passively avoid detection regardless of the properties of the surface which they occupy. However, it is uncommon and poorly understood in terrestrial species. In one tribe of predacious katydids (Phlugidini), transparency is paired with highly ultrasonic communication for increased predator evasion, yet little is known about the singing capabilities of these species, with only one genus of Phlugidini acoustically well described to date. Here, we describe Speculophlugis hishquten new genus and species of highly transparent crystal katydid species from the Andean cloud forest, discussing the potential use of this species for non-invasive studies of internal anatomy, and analysing its ultrasonic call. Using laser Doppler vibrometry and light microscopy techniques, we found the transparency of the cuticle around the hearing apparatus to be 85-87 % at the wavelength of the laser beam (633 nm), making S. hishquten a candidate for the highest recorded cuticle transparency of any insect. The male song has a fundamental frequency of 50 kHz, matching both the ultrasonic call range and rapid call structure of other Phlugidini species. However, the extent of ultrasonic communication and the level of transparency across the Phlugidini tribe requires more attention.

Summary Larger easily visible animals and plants are negatively affected by agrochemicals used for intensive food production, but we do not understand the general spatial and temporal effects of agrochemicals on the multitudes of bacteria, fungi, and small invertebrate animals that underpin ecosystem productivity. We sequenced the 16S, ITS2, and COI DNA barcode regions from 648 New Zealand vineyard soil samples managed under either conventional or low-agrochemical-input conservation approaches across two regions and three seasons in 1 year and discovered at least 170,000 phylotypes (taxa) with >97% genetic identity. Management approach correlated with a significant 2%–10% difference in the abundances of phylotypes that differed over regions and seasons. Although the data show that agrochemicals do not have a large effect on soil biodiversity on average, the important finding is that the magnitude of impact differs between taxa types and locations, and some taxa most affected also influence the quality of agricultural produce., Graphical abstract, Highlights • Agrochemicals have a small effect on soil bacterial, fungal, and animal biodiversity • The effect differs in space (and less so time) and affects taxa abundances greatest • There is no effect of agrochemicals on taxa richness • The size of the effect differs between taxa but is greatest for eukaryotic taxa, Microorganism; Farming Management; Farming Systems; Relation between Agriculture and Environment; Soil Biology; Beverage

In the cloud forests of the central range of the Colombian Andes, we discovered a species of katydid (Orthoptera: Tettigoniidae) that imitates mosses to an uncanny degree and is exceedingly difficult to detect. The camouflage exhibited by this particular katydid seems quite specific. We discuss the evolutionary consequences of this sort of specialization. Selection to maintain effective disguises can result in reproductive isolation between populations specialized for different microhabitats, which makes it reasonable to speculate that camouflage may increasing diversification rates. Camouflage could also come at the price of elevated extinction risk. This possibility must be considered because although antipredator defenses are often thought of as leading to “escape-and-radiate” dynamics where diversification follows innovation that allows expansion into new niches, recent work has shown unexpected extinction risk associated with some antipredator adaptations. Highly specialized camouflage would seem an ambiguous case because of its obvious benefits, but also potential costs such as inhabiting habitats with low carrying capacities, vulnerability to predators at high densities if predators form search images, or metabolic trade-offs with thermoregulation. Groups such as the Tettigoniidae provide a tantalizing opportunity for their exceptional diversity, wide geographic distribution, and striking array of disguises suggest that many independent evolutionary experiments have already taken place.

© 2017 Elsevier. This manuscript version is made available under the CC-BY-NC-ND 4.0 license: http://creativecommons.org/licenses/by-nc-nd/4.0/, Phylogeography can provide insight into the potential for speciation and identify geographic regions and evolutionary processes associated with species richness and evolutionary endemism. In the marine environment, highly mobile species sometimes show structured patterns of diversity, but the processes isolating populations and promoting differentiation are often unclear. The Delphinidae (oceanic dolphins) are a striking case in point and, in particular, bottlenose dolphins (Tursiops spp.). Understanding the radiation of species in this genus is likely to provide broader inference about the processes that determine patterns of biogeography and speciation, because both fine-scale structure over a range of kilometers and relative panmixia over an oceanic range are known for Tursiops populations. In our study, novel Tursiops spp. sequences from the northwest Indian Ocean (including mitogenomes and two nuDNA loci) are included in a worldwide Tursiops spp. phylogeographic analysis. We discover a new ‘aduncus’ type lineage in the Arabian Sea (off India, Pakistan and Oman) that diverged from the Australasian lineage ∼261 Ka. Effective management of coastal dolphins in the region will need to consider this new lineage as an evolutionarily significant unit. We propose that the establishment of this lineage could have been in response to climate change during the Pleistocene and show data supporting hypotheses for multiple divergence events, including vicariance across the Indo-Pacific barrier and in the northwest Indian Ocean. These data provide valuable transferable inference on the potential mechanisms for population and species differentiation across this geographic range.

Chemical communication plays a pivotal role in shaping sexual and ecological interactions among animals. In lizards, fundamental mechanisms of sexual selection such as female mate choice have rarely been shown to be influenced by quantitative phenotypic traits (e.g., ornaments), while chemical signals have been found to potentially influence multiple forms of sexual and social interactions, including mate choice and territoriality. Chemical signals in lizards are secreted by glands primarily located on the edge of the cloacae (precloacal glands, PG) and thighs (femoral glands), and whose interspecific and interclade number ranges from 0 to > 100. However, elucidating the factors underlying the evolution of such remarkable variation remains an elusive endeavour. Competing hypotheses suggest a dominant role for phylogenetic conservatism (i.e., species within clades share similar numbers of glands) or for natural selection (i.e., their adaptive diversification results in deviating numbers of glands from ancestors). Using the prolific Liolaemus lizard radiation from South America (where PG vary from 0 to 14), we present one of the largest-scale tests of both hypotheses to date. Based on climatic and phylogenetic modelling, we show a clear role for both phylogenetic inertia and adaptation underlying gland variation: (i) solar radiation, net primary productivity, topographic heterogeneity and precipitation range have a significant effect on PG variation, (ii) humid and cold environments tend to concentrate species with a higher number of glands, (iii) there is a strong phylogenetic signal that tends to conserve the number of PG within clades. Collectively, our study confirms that the inertia of niche conservatism can be broken down by the need of species facing different selection regimes to adjust their glands to suit the demands of their specific environments. University of Lincoln PhD scholarship; Research Investment Fund (RIF) Grant - University of Lincoln MJ is fully funded by a University of Lincoln PhD scholarship. DPD thanks financial support provided by a Research Investment Fund (RIF) Grant awarded by the University of Lincoln. The authors thank the insightful critical contribution that the editor and two anonymous referees provided on earlier versions of our work, which helped improve its flow and clarity.

This work was supported by the School of Biology, University of St Andrews, the Scottish Funding Council (MASTS grant reference HR09011) and the Templeton Foundation (grant #60501, 'Putting the Extended Evolutionary Synthesis to the Test'). Coral reefs are a valuable and vulnerable marine ecosystem. The structure of coral reefs influences their health and ability to fulfill ecosystem functions and services. However, monitoring reef corals largely relies on 1D or 2D estimates of coral cover and abundance that overlook change in ecologically significant aspects of the reefs because they do not incorporate vertical or volumetric information. This study explores the relationship between 2D and 3D metrics of coral size. We show that surface area and volume scale consistently with planar area, albeit with morphotype specific conversion parameters. We use a photogrammetric approach using open-source software to estimate the ability of photogrammetry to provide measurement estimates of corals in 3D. Technological developments have made photogrammetry a valid and practical technique for studying coral reefs. We anticipate that these techniques for moving coral research from 2D into 3D will facilitate answering ecological questions by incorporating the 3rd dimension into monitoring. Publisher PDF

Male katydids produce mating calls by stridulation using specialized structures on the forewings. The right wing (RW) bears a scraper connected to a drum-like cell known as the mirror and a left wing (LW) that overlaps the RW and bears a serrated vein on the ventral side, the stridulatory file. Sound is generated with the scraper sweeping across the file, producing vibrations that are amplified by the mirror. Using this sound generator, katydids exploit a range of song carrier frequencies (CF) unsurpassed by any other insect group, with species singing as low as 600 Hz and others as high as 150 kHz. Sound generator size has been shown to scale negatively with CF, but such observations derive from studies based on few species, without phylogenetic control, and/or using only the RW mirror length. We carried out a phylogenetic comparative analysis involving 94 species of katydids to study the relationship between LW and RW components of the sound generator and the CF of the male’s mating call, while taking into account body size and phylogenetic relationships. The results showed that CF negatively scaled with all morphological measures, but was most strongly related to components of the sound generation system (file, LW and RW mirrors). Interestingly, the LW mirror (reduced and non-functional) predicted CF more accurately than the RW mirror, and body size is not a reliable CF predictor. Mathematical models were verified on known species for predicting CF in species for which sound is unknown (e.g. fossils or museum specimens).

Precise and sufficiently detailed morphological taxonomy is vital in biology, for example in the accurate interpretation of ecological and palaeoecological datasets, especially in polar regions, where biodiversity is poor. Testate amoebae on the Antarctic Peninsula (AP) are well-documented and variations in their population size have recently been interpreted as a proxy for microbial productivity changes in response to recent regional climate change. AP testate amoeba assemblages are dominated by a small number of globally ubiquitous taxa. We examine morphological variation in Corythion spp. across the AP, finding clear evidence supporting the presence of two morphospecies. Corythion constricta (Certes 1889) was identified on the AP for the first time and has potentially been previously misidentified. Furthermore, a southerly trend of decreasing average test size in Corythion dubium (Taránek 1881) along the AP suggests adaptive polymorphism, although the precise drivers of this remain unclear, with analysis hindered by limited environmental data. Further work into morphological variation in Corythion is needed elsewhere, alongside molecular analyses, to evaluate the potential for (pseudo)cryptic diversity within the genus. We advocate a parsimonious taxonomical approach that recognises genetic diversity but also examines and develops accurate morphological divisions and descriptions suitable for light microscopy-based ecological and palaeoecological studies. © 2017 The Author(s)

There is increased need to identify sustainable agricultural methods which avoid environmental degradation. Previous studies have focused on the effect of specific agricultural interventions on large organisms, but we have fewer data evaluating how microbes, which are key components of ecosystems, might be affected. Additionally, previous studies have been constrained as they only examined one habitat in an ecosystem and have not gone on to evaluate the effect of agricultural approach on harvested crops. Here we take an ecosystems approach and evaluate the net effect of conventional versus biodynamic management on agricultural ecosystems by quantifying fungal communities in multiple habitats using metagenomics. We go on to measure biodiversity in the crop and key chemical quality parameters in the product consumed by humans. We find that the method of management significantly affects communities in soil, on plant structures, and on the developing crop in subtle but importantly different ways in terms of number, type, and abundance of species. However, management approach has no effect on communities in the final harvested juice, nor on product traits aligned with quality. This shows that while management approach impacts different habitats in the environment in different ways, this does not automatically flow onto the harvested crop.

Male Tettigoniidae emit sound to attract conspecific females. The sound is produced by stridulation. During stridulation the forewings open and close, but it is during the closing stroke that the scraper contacts the file teeth to generate the predominant sound components, which are amplified by adjacent wing cells specialized in sound radiation. The sounds usually exceed the sonic boundary and might occur above 40 kHz, reaching extreme ultrasonic frequencies of 150kHz in some species. Here we test the hypothesis that Tettigoniidae species should prefer microhabitats that favour efficient signal transmission, i.e. that there is a relationship of sound frequency with the vertical distribution of the species (from ground to canopy) at Gorgona National Natural Park, Colombia. We sampled 16 trees and four different altitudinal levels between 1 and 20m above the understory vegetation. We placed collecting blankets separated by vertical distances of 5m, and knocked insects down using the technique known as fogging. We found no correlation between vertical distribution and carrier frequency, but there was a preference for open spaces (below the canopy and above the understory) in species using extreme ultrasound. This is the first quantitative description of the vertical distribution in neotropical species of the family Tettigoniidae and its relationship to the calling song frequency. Rev. Biol. Trop. 62 (Suppl. 1): 289-296. Epub 2014 February 01. Los machos de Tettigoniidae producen sonido frotando las alas anteriores para atraer a sus hembras coespecíficas con fines reproductivos (mecanismo conocido como estridulación). Un borde esclerotizado (o raspador) en el ala derecha es frotado sobre una vena modificada con dientecillos en el ala izquierda. Durante la estridulación las alas abren y cierran, pero es durante el cierre que el raspador contacta los dientecillos de la lima y produce vibraciones que son amplificadas por celdas adyacentes especializadas para radiar sonido. Sonidos que superen los 20 000Hz, se consideran ultrasónicos, en los Tettigoniidae, la mayoría de los cantos superan el umbral sónico y pueden ocurrir a más de 40 000Hz, con casos extremos de hasta 148 000Hz. La esencia de este trabajo es el estudio de esta gama espectacular de frecuencias de la familia Tettigoniidae y su relación con la distribución vertical de los individuos en las especies que habitan en Gorgona. Se muestrearon 16 árboles a cuatro niveles diferentes de altitud entre 1 y ~20m sobre el sotobosque, instalando sabanas colectoras a intervalos de 4-5m. Los insectos se colectaron mediante el método de nebulización térmica. El análisis indica que no hay correlación entre la distribución vertical de las especies y la frecuencia del canto. Sin embargo se observó una preferencia por espacios abiertos en especies que cantan a frecuencias extremas. Este estudio representa la primera descripción cuantitativa de la distribución vertical de Tettigoniidae Neotropicales y su relación con la frecuencia del canto.

Many species are threatened by human activity through processes such as habitat modification, water management, hunting, and introduction of invasive species. These anthropogenic threats must be mitigated as efficiently as possible because both time and money available for mitigation are limited. For example, it is essential to address the type and degree of uncertainties present to derive effective management strategies for managed populations. Decision science provides the tools required to produce effective management strategies that can maximize or minimize the desired objective(s) based on imperfect knowledge, taking into account stochasticity. Of particular importance are questions such as how much of available budgets should be invested in reducing uncertainty and which uncertainties should be reduced. In such instances, decision science can help select efficient environmental management actions that may be subject to stochasticity and imperfect knowledge. Here, we review the use of decision science in environmental management to demonstrate the utility of the decision science framework. Our points are illustrated using examples from the literature. We conclude that collaboration between theoreticians and practitioners is crucial to maximize the benefits of decision science’s rational approach to dealing with uncertainty.

Saccharomyces is one of the best-studied microbial genera but our understanding of the global distributions and evolutionary histories of its members is relatively poor. Recent studies have altered our view of Saccharomyces’ origin, but a lack of sampling from the vast majority of the world precludes a holistic perspective. We evaluate alternate Gondwanan and Far East Asian hypotheses concerning the origin of these yeasts. Being part of Gondwana, and only colonized by humans in the last ~1,000 years, New Zealand represents a unique environment for testing these ideas. Genotyping and ribosomal sequencing of samples from North Island native forest parks identified a widespread population of Saccharomyces. Whole genome sequencing identified the presence of S. arboricola and S. eubayanus in NZ, which is the first report of S. arboricola outside Far East Asia, and also expands S. eubayanus’ known distribution to include the Oceanic region. Phylogenomic approaches place the S. arboricola population as significantly diverged from the only other sequenced Chinese isolate, but indicate that S. eubayanus might be a recent migrant from South America. These data tend to support the Far East Asian origin of the Saccharomyces, but the history of this group is still far from clear.

Studies of the effects of mass extinctions on ancient ecosystems have focused on changes in taxic diversity, morphological disparity, abundance, behaviour and resource availability as key determinants of group survival. Crucially, the contribution of life history traits to survival during terrestrial mass extinctions has not been investigated, despite the critical role of such traits for population viability. We use bone microstructure and body size data to investigate the palaeoecological implications of changes in life history strategies in the therapsid forerunners of mammals before and after the Permo-Triassic Mass Extinction (PTME), the most catastrophic crisis in Phanerozoic history. Our results are consistent with truncated development, shortened life expectancies, elevated mortality rates and higher extinction risks amongst post-extinction species. Various simulations of ecological dynamics indicate that an earlier onset of reproduction leading to shortened generation times could explain the persistence of therapsids in the unpredictable, resource-limited Early Triassic environments and help explain observed body size distributions of some disaster taxa (e.g., Lystrosaurus). Our study accounts for differential survival in mammal ancestors after the PTME and provides a methodological framework for quantifying survival strategies in other vertebrates during major biotic crises.

In Australia, the adventure sport of canyoning occurs predominantly in the protected areas of the Greater Blue Mountains World Heritage Area, 50 km west of Sydney. It involves travelling through narrow, deep gorges using a combination of walking, abseiling, wading, rock scrambling and/ or swimming through the canyon streams. The sport’s popularity is reported to have increased substantially over time, causing concern for the sustainability of these fragile ecosystems. To investigate visitor impacts, benthic macroinvertebrate assemblages and water quality were compared among four canyons subject to high visitation (average 20 – 100 visits per week) with those in four unvisited/low visitation canyons (0 - 5 visits per week). Comparison was also made between impacted and nonimpacted areas within two of the high visitation canyons. Sampling was undertaken over two canyoning seasons (early spring to late autumn). No significant relationship between macroinvertebrate assemblage composition or water quality was observed due to visitation level within and among canyons, although there were inherent differences among canyons. Water quality was consistent with pristine conditions in all locations. Macroinvertebrate assemblages included a range of pollution sensitive taxa (e.g., Ephemeroptera represented by 42.3% of all animals). Although family richness was concentrated within a few orders (84.2% of families were represented by 50% of orders), most families were represented by 2 Recognisable Taxonomic Units, 47.4% of these were represented by a single taxon. We concluded that, at current visitation levels, there is no measurable impact on the predominant animal component of these ecosystems. This finding was contrary to the expectations of the land managers of this area.

Bernard, E.L., Ruta, M., Tarver, J.E., and Benton, M.J. 2010. The fossil record of early tetrapods: Worker effort and the end−Permian mass extinction. Acta Palaeontologica Polonica 55 (2): 229–239. It is important to understand the quality of the fossil record of early tetrapods (Tetrapoda, minus Lissamphibia and Amniota) because of their key role in the transition of vertebrates from water to land, their dominance of terrestrial fau− nas for over 100 million years of the late Palaeozoic and early Mesozoic, and their variable fates during the end−Perm− ian mass extinction. The first description of an early tetrapod dates back to 1824, and since then discoveries have oc− curred at a rather irregular pace, with peaks and troughs corresponding to some of the vicissitudes of human history through the past two centuries. As expected, the record is dominated by the well−sampled sedimentary basins of Europe and North America, but finds from other continents are increasing rapidly. Comparisons of snapshots of knowledge in 1900, 1950, and 2000 show that discovery of new species has changed the shape of the species−level diversification curve, contrary to earlier studies of family−level taxa. There is, however, little evidence that taxon counts relate to re− search effort (as counted by numbers of publications), and there are no biasing effects associated with differential study of different time intervals through the late Palaeozoic and Mesozoic. In fact, levels of effort are apparently not related to geological time, with no evidence that workers have spent more time on more recent parts of the record. In particular, the end−Permian mass extinction was investigated to determine whether diversity changes through that interval might reflect worker effort: it turns out that most records of early tetrapod taxa (when corrected for duration of geological se− ries) occur in the Lower Triassic. Key wor ds: Discovery curve, collector curve, early tetrapods, fossil record, Tetrapoda, Temnospondyli, Lepospondyli, end−Permian mass extinction.

Tree shape analyses are frequently used to infer the location of shifts in diversification rate within the Tree of Life. Many studies have supported a causal relationship between shifts and temporally coincident events such as the evolution of “key innovations”. However, the evidence for such relationships is circumstantial. We investigated patterns of diversification during the early evolution of Amniota from the Carboniferous to the Triassic, subjecting a new supertree to analyses of tree balance in order to infer the timing and location of diversification shifts. We investigated how uneven origination and extinction rates drive diversification shifts and use two case studies (herbivory and an aquatic lifestyle) to examine whether shifts tend to be contemporaneous with evolutionary novelties. Shifts within amniotes tend to occur during periods of elevated extinction, with mass extinctions coinciding with numerous and larger shifts. Diversification shifts occurring in clades that possess evolutionary innovations do not coincide temporally with the appearance of those innovations, but are instead deferred to periods of high extinction rate. We suggest such innovations did not cause increases in the rate of cladogenesis, but allowed clades to survive extinction events. We highlight the importance of examining general patterns of diversification before interpreting specific shifts.

During the late Paleozoic, vascular land plants (tracheophytes) diversified into a remarkable variety of morpho- logical types, ranging from tiny, aphyllous, herbaceous forms to giant leafy trees. Leaf shape is a key determinant of both function and structural diversity of plants, but relatively little is known about the tempo and mode of leaf morphological diversification and its correlation with tracheophyte diversity and abiotic changes during this re- markable macroevolutionary event, the greening of the continents. We use the extensive record of Paleozoic tra- cheophytes from South China to explore models of morphological evolution in early land plants. Our findings suggest that tracheophyte leaf disparity and diversity were decoupled, and that they were under different selec- tive regimes. Two key phases in the evolution of South Chinese tracheophyte leaves can be recognized. In the first phase, from Devonian to Mississippian, taxic diversity increased substantially, as did leaf disparity, at the same time as they acquired novel features in their vascular systems, reproductive organs, and overall architecture. The second phase, through the Carboniferous–Permian transition, saw recovery of wetland communities in South China, associated with a further expansion of morphologies of simple leaves and an offset shift in morphospace occupation by compound leaves. Comparison with Euramerica suggests that the floras from South China were unique in several ways. The Late Devonian radiation of sphenophyllaleans contributed signif- icantly to the expansion of leaf morphospace, such that the evolution of large laminate leaves in this group oc- curred much earlier than those in Euramerica. The Pennsylvanian decrease in taxic richness had little effect on the disparity of compound leaves. Finally, the distribution in morphospace of the Permian pecopterids, gigantopterids, and equisetaleans occurred at the periphery of Carboniferous leaf morphospace.