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Mesopelagic fishes play a central role in marine food webs linking primary consumers to top predators. In this study, measures of direct calorimetry were used to analyse the energy density (ED) of 34 mesopelagic species collected at 12 stations in the equatorial and tropical Atlantic. Mean ED ranged from 2.7 kJ g -1 wet weight (WW) for the lanterfish Lampanyctus nobilis to 8.7 kJ g -1 WW for the lanterfish Benthosema glaciale. This study includes species of the orders Myctophiformes and Stomiiformes and represents migrants, non-migrants and partial-migrants species. The majority of the species were grouped into the medium-energy quality category (ED from 4 to 6 kJ g -1 ); Myctophiformes showed higher energetic values than the Stomiiformes. For the different species, the ED values were discussed in relation to spawning period and energy allocation strategies for reproduction and growth and feeding and migratory behaviour, as well as the ecoregion of study. These values will be useful for future assessment of energetic transfer between trophic levels and energetic modelling of Atlantic ecosystems., (© 2023 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

The latest Permian mass extinction (LPME) was triggered by magmatism of the Siberian Traps Large Igneous Province (STLIP), which left an extensive record of sedimentary Hg anomalies at Northern Hemisphere and tropical sites. Here, we present Hg records from terrestrial sites in southern Pangea, nearly antipodal to contemporaneous STLIP activity, providing insights into the global distribution of volcanogenic Hg during this event and its environmental processing. These profiles (two from Karoo Basin, South Africa; two from Sydney Basin, Australia) exhibit significant Hg enrichments within the uppermost Permian extinction interval as well as positive Δ 199 Hg excursions (to ~0.3‰), providing evidence of long-distance atmospheric transfer of volcanogenic Hg. These results demonstrate the far-reaching effects of the Siberian Traps as well as refine stratigraphic placement of the LPME interval in the Karoo Basin at a temporal resolution of ~10 5 years based on global isochronism of volcanogenic Hg anomalies., (© 2023. The Author(s).)

All modern mammals are descendants of the paraphyletic non-mammaliaform Synapsida, colloquially referred to as the "mammal-like reptiles." It has long been assumed that these mammalian ancestors were essentially reptile-like in their morphology, biology, and behavior, i.e., they had a small brain, displayed simple behavior, and their sensory organs were unrefined compared to those of modern mammals. Recent works have, however, revealed that neurological, sensory, and behavioral traits previously considered typically mammalian, such as whiskers, enhanced olfaction, nocturnality, parental care, and complex social interactions evolved before the origin of Mammaliaformes, among the early-diverging "mammal-like reptiles." In contrast, an enlarged brain did not evolve immediately after the origin of mammaliaforms. As such, in terms of paleoneurology, the last "mammal-like reptiles" were not significantly different from the earliest mammaliaforms. The abundant data and literature published in the last 10 years no longer supports the "three pulses" scenario of synapsid brain evolution proposed by Rowe and colleagues in 2011, but supports the new "outside-in" model of Rodrigues and colleagues proposed in 2018, instead. As Mesozoic reptiles were becoming the dominant taxa within terrestrial ecosystems, synapsids gradually adapted to smaller body sizes and nocturnality. This resulted in a sensory revolution in synapsids as olfaction, audition, and somatosensation compensated for the loss of visual cues. This altered sensory input is aligned with changes in the brain, the most significant of which was an increase in relative brain size., (Copyright © 2023 Elsevier B.V. All rights reserved.)

A new species of Nemestrinidae, Atriadops irwini Barraclough sp. nov., is described and illustrated from Sakaramy in the far north of Madagascar at about the same latitude as the Comoros Islands. The species was collected in lowland tropical forest by Dr Michael Irwin. Atriadops Wandolleck, 1897, is a small genus of Nemestrinidae in the subfamily Trichopisideinae. Six named species are distributed across the Australasian, Oriental, Afrotropical and Neotropical Regions. Atriadops irwini sp. nov. is thus only the seventh species in the genus (the first new species described in almost 70 years) and the second valid species from the Afrotropical Region. This is the first record of Atriadops from Madagascar, where the new species is the only endemic nemestrinid. Nycterimyia capensis Bezzi is the only other nemestrinid recorded from Madagascar, but it also occurs in South Africa (where it was first described) and Kenya. The world fauna of Atriadops is discussed and all known larval host records are listed.

A longstanding question in evolutionary biology is how natural selection and environmental pressures shape the mitochondrial genomic architectures of organisms. Mitochondria play a pivotal role in cellular respiration and aerobic metabolism, making their genomes functionally highly constrained. Evaluating selective pressures on mitochondrial genes can provide functional and ecological insights into the evolution of organisms. Collembola (springtails) are an ancient hexapod group that includes the oldest terrestrial arthropods in the fossil record, and that are closely associated with soil environments. Of interest is the diversity of habitat stratification preferences (life forms) exhibited by different species within the group. To understand whether signals of positive selection are linked to the evolution of life forms, we analysed 32 published Collembola mitogenomes in a phylomitogenomic framework. We found no evidence that signatures of selection are correlated with the evolution of novel life forms, but rather that mutations have accumulated as a function of time. Our results highlight the importance of nuclear-mitochondrial interactions in the evolution of collembolan life forms and that mitochondrial genomic data should be interpreted with caution, as complex selection signals may complicate evolutionary inferences., (© 2022. The Author(s).)

The sternum is a stabilizing element in the axial skeleton of most tetrapods, closely linked with the function of the pectoral girdle of the appendicular skeleton. Modern mammals have a distinctive sternum characterized by multiple ossified segments, the origins of which are poorly understood. Although the evolution of the pectoral girdle has been extensively studied in early members of the mammalian total group (Synapsida), only limited data exist for the sternum. Ancestrally, synapsids exhibit a single sternal element and previously the earliest report of a segmental sternum in non-mammalian synapsids was in the Middle Triassic cynodont Diademodon tetragonus. Here, we describe the well-preserved sternum of a gorgonopsian, a group of sabre-toothed synapsids from the Permian. It represents an ossified, multipartite element resembling the mammalian condition. This discovery pulls back the origin of the distinctive "mammalian" sternum to the base of Theriodontia, significantly extending the temporal range of this morphology. Through a review of sternal morphology across Synapsida, we reconstruct the evolutionary history of this structure. Furthermore, we explore its role in the evolution of mammalian posture, gait, and ventilation through progressive regionalization of the postcranium as well as the posteriorization of musculature associated with mammalian breathing., (© 2022. The Author(s).)

The endemic Afrotropical genus Dinapsis is revised, and seven new species are described and illustrated: D.bicolor van Noort & Shaw, sp. nov. , D.gamka van Noort & Shaw, sp. nov. , D.igneus van Noort & Shaw, sp. nov. , D.spinitibia van Noort & Shaw, sp. nov. , D.taita van Noort & Shaw, sp. nov. , D.tricolor Shaw & van Noort, sp. nov. , D.zulu Shaw & van Noort, sp. nov. The distribution of the Central African Republican species D.centralis Shaw & van Noort, 2009 is extended to include Cameroon, Kenya and Uganda. Dinapsisturneri Waterston, 1922, previously only known from the poorly preserved holotype female, is redescribed based on newly collected specimens. The distribution of this Western Cape species is extended to include the Eastern and Northern Cape provinces of South Africa. Four distinct species groups within the genus are proposed and diagnosed. An illustrated identification key to all described species of Dinapsis is provided. Online interactive Lucid keys to Afrotropical megalyrid genera and Dinapsis species are available at http://www.waspweb.org., (Simon van Noort, Scott Richard Shaw, Robert S. Copeland.)

A new species of Setanodosa, S. jacquesi sp. nov. is described from the Western Cape (South Africa). It differs from other species of the genus by its unique shocking red pigmentation, the number of vesicles in the post antennal organ, and the number of clavate tenent hairs on the tibiotarsi. A comparative table of the world Setanodosa and a key of Brachystomellidae species known from South Africa are provided. DNA barcoding results are provided for several Brachystomellidae species from South Africa, Australia and the sub-Antarctic to support our findings. It shows that a species provisionally identified as Brachystomella cf. platensis is unambiguously present in both South Africa and Australia.

Afronympha gen. nov. is described for the single known species A. eminpashai sp. nov. collected in the Democratic Republic of the Congo and Uganda. This new genus is similar to some extent to Asecodes, but differs from this and other genera of Entedoninae in having a produced and medially notched anterior margin of the clypeus, and robust mandibles. Similarities and differences between Afronympha and other genera are discussed. The type species, A. eminpashai sp. nov., was collected in degraded mid-altitude rainforest in Kibale National Park, western Uganda and disturbed areas of lowland rainforest in the Ituri region of eastern Democratic Republic of the Congo, suggesting its ecological plasticity under circumstances of habitat transformation.

Middle Miocene foraminifera from the northern Namibian outer continental shelf are indicators of a period prior to the initiation of the Benguela Upwelling System (BUS). This study provides an update to the occurrence and taxonomy of Miocene foraminifera from the continental margin of Namibia. The taxonomy of 51 benthic and 12 planktic foraminiferal species from the northern Namibian shelf are discussed, their stratigraphic significance given, and their ecological preferences and regional distribution summarised within this study. The identification of extinct planktic foraminifera provided key stratigraphic control for the middle Miocene strata of this region. The taxa identified in this study provide a distinct and different assemblage to the overlying younger strata. Many of the species recorded in this study have not been identified in the region and are reported for the first time from the middle Miocene on the southwestern continental shelf of Africa, off Namibia. A total of 47 species are identified and discussed for the first time from this region. Nineteen species recorded in this study are extinct and eleven taxa reported here have previously only been reported on the genus level on the southwestern shelf of South Africa. Seven benthic species (Amphicoryna scalaris, Marginulina obesa, Glandulina laevigata, Globocassidulina subglobosa, Uvigerina peregrina, Sphaeroidina bulloides and Melonis affinis) and two planktic species (Globigerina bulloides and Orbulina universa) did not disappear from the regional stratigraphy and continued to occur in Plio-Pleistocene to Recent sediments along the southwestern continental shelf of Africa.

Olive lace bugs (Hemiptera: Tingidae) are small sap-sucking insects that feed on wild and cultivated Olea europaea . The diversity of olive lace bug species in South Africa, the most important olive producer on the continent, has been incompletely surveyed. Adult specimens were collected in the Western Cape province for morphological and DNA-based species identification, and sequencing of complete mitogenomes. Cysteochila lineata , Plerochila australis , Neoplerochila paliatseasi and Neoplerochila sp. were found at 12 sites. Intra- and interspecific genetic divergences and phylogenetic clustering in 30 species in 18 genera of Tingidae using new and publicly available DNA barcodes showed high levels of congruity between taxonomic and genetic data. The phylogenetic position of the four species found in South Africa was inferred using new and available mitogenomes of Tingidae. Notably, olive lace bugs formed a cluster of closely related species. However, Cysteochila was non-monophyletic as C. lineata was recovered as a sister species to P. australis whereas Cysteochila chiniana , the other representative of the genus, was grouped with Trachypeplus jacobsoni and Tingis cardui in a different cluster. This result suggests that feeding on O. europaea may have a common origin in Tingidae and warrants future research on potential evolutionary adaptations of olive lace bugs to this plant host.

Earth's largest biotic crisis occurred during the Permo-Triassic Transition (PTT). On land, this event witnessed a turnover from synapsid- to archosauromorph-dominated assemblages and a restructuring of terrestrial ecosystems. However, understanding extinction patterns has been limited by a lack of high-precision fossil occurrence data to resolve events on submillion-year timescales. We analyzed a unique database of 588 fossil tetrapod specimens from South Africa's Karoo Basin, spanning ∼4 My, and 13 stratigraphic bin intervals averaging 300,000 y each. Using sample-standardized methods, we characterized faunal assemblage dynamics during the PTT. High regional extinction rates occurred through a protracted interval of ∼1 Ma, initially co-occurring with low origination rates. This resulted in declining diversity up to the acme of extinction near the Daptocephalus - Lystrosaurus declivis Assemblage Zone boundary. Regional origination rates increased abruptly above this boundary, co-occurring with high extinction rates to drive rapid turnover and an assemblage of short-lived species symptomatic of ecosystem instability. The "disaster taxon" Lystrosaurus shows a long-term trend of increasing abundance initiated in the latest Permian. Lystrosaurus comprised 54% of all specimens by the onset of mass extinction and 70% in the extinction aftermath. This early Lystrosaurus abundance suggests its expansion was facilitated by environmental changes rather than by ecological opportunity following the extinctions of other species as commonly assumed for disaster taxa. Our findings conservatively place the Karoo extinction interval closer in time, but not coeval with, the more rapid marine event and reveal key differences between the PTT extinctions on land and in the oceans., Competing Interests: The authors declare no competing interest.

Trait-environment interactions have contributed to the remarkable plant radiations in the Cape Floristic Region (CFR) of southern Africa. Whether such interactions have also resulted in the diversification of the invertebrate fauna, independently of direct associations with plants is, however, not clear. One candidate where this may be the case is the unusually diverse Collembola genus Seira. Including 89 species in the CFR, many of which are localised habitat specialists, this genus includes many species inhabiting the warm, dry fynbos shrubland-a habitat atypical of usually desiccation-sensitive Collembola. Here, we investigate whether desiccation tolerance may have contributed to the considerable diversity of Seira in the CFR. First, we demonstrate, by measuring vapour pressure deficits (VPD) of the species' microhabitats (fynbos shrubland and moister Afrotemperate Forests), that the fynbos shrublands are dry environments (mean ± S.E. maximum VPD 5.2 ± 0.1 kPa) compared with the Afrotemperate Forest patches (0.3 ± 0.02 kPa) during the summer activity period of Seira. Then we show that Seira species living in these shrublands are more desiccation tolerant (mean ± S.E. survival time at 76% relative humidity: 74.3 ± 3.3 h) than their congeners in the cooler, moister Afrotemperate Forests (34.3 ± 2.8 h), and compared with Collembola species globally (3.7 ± 0.2 h). These results, and a previous demonstration of pronounced thermal tolerance in the fynbos shrubland species, suggest that the diversity of Seira in the CFR is at least partly due to pronounced desiccation and thermal tolerance, which has enabled species in the genus to exploit the hot and dry habitats of the CFR.

Recent technical advances combined with novel computational approaches have promised the acceleration of our understanding of the tree of life. However, when it comes to hyperdiverse and poorly known groups of invertebrates, studies are still scarce. As published phylogenies will be rarely challenged by future taxonomists, careful attention must be paid to potential analytical bias. We present the first molecular phylogenetic hypothesis for the family Chalcididae, a group of parasitoid wasps, with a representative sampling (144 ingroups and seven outgroups) that covers all described subfamilies and tribes, and 82% of the known genera. Analyses of 538 Ultra-Conserved Elements (UCEs) with supermatrix (RAxML and IQTREE) and gene tree reconciliation approaches (ASTRAL, ASTRID) resulted in highly supported topologies in overall agreement with morphology but reveal conflicting topologies for some of the deepest nodes. To resolve these conflicts, we explored the phylogenetic tree space with clustering and gene genealogy interrogation methods, analyzed marker and taxon properties that could bias inferences and performed a thorough morphological analysis (130 characters encoded for 40 taxa representative of the diversity). This joint analysis reveals that UCEs enable attainment of resolution between ancestry and convergent/divergent evolution when morphology is not informative enough, but also shows that a systematic exploration of bias with different analytical methods and a careful analysis of morphological features is required to prevent publication of artifactual results. We highlight a GC content bias for maximum-likelihood approaches, an artifactual mid-point rooting of the ASTRAL tree and a deleterious effect of high percentage of missing data (>85% missing UCEs) on gene tree reconciliation methods. Based on the results we propose a new classification of the family into eight subfamilies and ten tribes that lay the foundation for future studies on the evolutionary history of Chalcididae., (© The Willi Hennig Society 2020.)

The Afrotropical banchine fauna comprises 12 genera: Apophua Morley, Atropha Kriechbaumer, Cryptopimpla Taschenberg, Exetastes Gravenhorst, Glyptopimpla Morley, Himertosoma Schmiedeknecht, Lissonota Gravenhorst, Sjostedtiella Szépligeti, Spilopimpla Cameron, Syzeuctus Förster, Tetractenion Seyrig, and Tossinola Viktorov. A well-illustrated revised key to the genera using high definition images is provided, and the endemic Afrotropical genus Tetractenion is revised, previously represented by two described species. Four new species are described: T.ibayaensis sp. nov. , T.pascali sp. nov. , T.pseudolutea sp. nov. , and T.rosei sp. nov. The first species-level identification key is provided for this rare genus. Based on morphological attributes the hypothesis is presented that the species in this genus are probably nocturnal. All images and online interactive Lucid keys are available at: www.waspweb.org and the associated underlying data is made available as Suppl. materials 1, 2 LIF3 files to this paper for inter-exchange with other key production software., (Terry Reynolds Berry, Simon van Noort.)

Archosauria (birds, crocodilians and their extinct relatives) form a major part of terrestrial ecosystems today, with over 10 000 living species, and came to dominate the land for most of the Mesozoic (over 150 Myr) after radiating following the Permian-Triassic extinction. The archosaur skull has been essential to this diversification, itself diversified into myriad forms. The archosauriform Euparkeria capensis from the Middle Triassic (Anisian) of South Africa has been of great interest since its initial description in 1913, because its anatomy shed light on the origins and early evolution of crown Archosauria and potentially approached that of the archosaur common ancestor. Euparkeria has been widely used as an outgroup in phylogenetic analyses and when investigating patterns of trait evolution among archosaurs. Although described monographically in 1965, subsequent years have seen great advances in the understanding of early archosaurs and in imaging techniques. Here, the cranium and mandible of Euparkeria are fully redescribed and documented using all fossil material and computed tomographic data. Details previously unclear are fully described, including vomerine dentition, the epiptergoid, number of premaxillary teeth and palatal arrangement. A new diagnosis and cranial and braincase reconstruction is provided, and an anatomical network analysis is performed on the skull of Euparkeria and compared with other amniotes. The modular composition of the cranium suggests a flexible skull well adapted to feeding on agile food, but with a clear tendency towards more carnivorous behaviour, placing the taxon at the interface between ancestral diapsid and crown archosaur ecomorphology, corresponding to increases in brain size, visual sensitivity, upright locomotion and metabolism around this point in archosauriform evolution. The skull of Euparkeria epitomizes a major evolutionary transition, and places crown archosaur morphology in an evolutionary context., Competing Interests: The authors declare no competing financial or non-financial interests., (© 2020 The Authors.)

Although best known for its extraordinary radiations of endemic plant species, the South African fynbos is home to a great diversity of phytophagous insects, including butterflies in the genus Chrysoritis (Lepidoptera: Lycaenidae). These butterflies are remarkably uniform morphologically; nevertheless, they comprise 43 currently accepted species and 68 currently valid taxonomic names. While many species have highly restricted, dot-like distributions, others are widespread. Here, we investigate the phylogenetic and biogeographic history underlying their diversification by analyzing molecular markers from 406 representatives of all described species throughout their respective ranges. We recover monophyletic clades for both C. chrysaor and C. thysbe species-groups, and identify a set of lineages that fall between them. The estimated age of divergence for the genus is 32 Mya, and we document significantly rapid diversification of the thysbe species-group in the Pleistocene (~2 Mya). Using ancestral geographic range reconstruction, we show that West Fynbos is the most likely region of origin for the radiation of the thysbe species-group. The colonization of this region occurred 9 Mya and appears to have been followed by a long period of relative stasis before a recent increase in diversification. Thus, the thysbe radiation does not appear to have resulted from the colonization of new biogeographic areas. Rather, the impact of species interactions (with ants and plants), the appearance of key innovations, and/or the opening of new ecological niche space in the region might explain the sudden burst of speciation that occurred in this group 2 Mya. The biogeographic model suggests two different diversification processes with few historical cross-colonisations, one in eastern South Africa for the C. chrysaor group and the other in western South Africa for the remaining taxa. Distributional range assessments and ecological niche models for each species show important niche overlap, and in a few cases, complete overlap. However, these shared traits are not explained by phylogenetic history. Chrysoritis taxa frequently fly in sympatry and gene tree reticulation appears to be widespread at the species level, suggesting that several episodes of range shifts might have led to secondary sympatries, allowing limited gene flow that challenges species delimitation efforts. In addition, the unusually high diversification rate for the thysbe clade of 1.35 [0.91-1.81] lineages per million years also suggests the possibility of taxonomic oversplitting. The phylogeny presented here provides a framework for a taxonomic revision of the genus. We highlight cases of potential synonymy both in allopatry and sympatry, and stress the importance of dedicated studies to assess potential pre- and post-zygotic barriers giving rise to species delimitations of the thysbe group., (Copyright © 2020 Elsevier Inc. All rights reserved.)

As global climates change, alien species are anticipated to have a growing advantage relative to their indigenous counterparts, mediated through consistent trait differences between the groups. These insights have largely been developed based on interspecific comparisons using multiple species examined from different locations. Whether such consistent physiological trait differences are present within assemblages is not well understood, especially for animals. Yet, it is at the assemblage level that interactions play out. Here, we examine whether physiological trait differences observed at the interspecific level are also applicable to assemblages. We focus on the Collembola, an important component of the soil fauna characterized by invasions globally, and five traits related to fitness: critical thermal maximum, minimum and range, desiccation resistance and egg development rate. We test the predictions that the alien component of a local assemblage has greater basal physiological tolerances or higher rates, and more pronounced phenotypic plasticity than the indigenous component. Basal critical thermal maximum, thermal tolerance range, desiccation resistance, optimum temperature for egg development, the rate of development at that optimum and the upper temperature limiting egg hatching success are all significantly higher, on average, for the alien than the indigenous components of the assemblage. Outcomes for critical thermal minimum are variable. No significant differences in phenotypic plasticity exist between the alien and indigenous components of the assemblage. These results are consistent with previous interspecific studies investigating basal thermal tolerance limits and development rates and their phenotypic plasticity, in arthropods, but are inconsistent with results from previous work on desiccation resistance. Thus, for the Collembola, the anticipated advantage of alien over indigenous species under warming and drying is likely to be manifest in local assemblages, globally., (© The Author(s) 2020. Published by Oxford University Press and the Society for Experimental Biology.)

Length-weight relationships (LWRs) were estimated for 36 mesopelagic fish species collected from the equatorial and tropical Atlantic encompassing several oceanographic regions: oligotrophic, equatorial, Cape Blanc, Cape Verde and the Canary Islands. The sample was composed of myctophids (25 species), gonostomatids (5), sternoptychids (3), stomiids (2) and phosichthyids (1). The species were clustered according to body shape: "short-deep" (sternoptychids), "elongate" (gonostomatids, stomiids and some phosichthyids) and "fusiform" (myctophids and some phosichthyids). Three types of weight and LWRs were considered: wet weight (WW), eviscerated wet weight (eWW) and eviscerated dry weight (eDW). The study demonstrated that most species present a positive allometric growth, independent of the weight used. However, the allometric value varied in 40-50% of species depending on the type of weight considered. Significant variations linked to fish morphology were found in the relationship between the slope and intercept of the LWR equation. Significant differences were also noted in the water content linked to fish body shape. Based on the distributions of several species we compare their fitness between oceanographic regions using the relative condition factor (K rel ). Except for Diaphus brachycephalus (oligotrophic vs. equatorial waters) and Lampanyctus alatus (equatorial, Cape Blanc, Cape Verde and the Canary Islands), no regional significant differences were observed in the species analysed., (© 2020 The Fisheries Society of the British Isles.)

The Pedra de Fogo Formation in the Parnaíba Basin of northeastern Brazil hosts a recently discovered lacustrine fauna and provides the only known record of the Captorhinidae in South America. Here, new captorhinid remains from this unit are described. Two partial mandibles, including one formerly ascribed to the genus Captorhinus , are here referred to Captorhinikos sp. a genus previously described from North America. The natural mould of a large mandible probably represents a new taxon within the captorhinid subclade Moradisaurinae, and a small skull roof is regarded as Captorhinidae indet. Captorhinids are generally considered to have been herbivores or omnivores. The Pedra de Fogo captorhinids likely played an important ecological role as primary consumers in the palaeoenvironment of this geological unit, which is also known for its extensive record of petrified forests. The new finds reinforce the close relationships between the continental faunas of palaeotropical western Gondwana and palaeoequatorial North America during the Cisuralian., Competing Interests: Claudia A. Marsicano is an Academic Editor for PeerJ., (© 2020 Cisneros et al.)

Tropical forest invertebrates, such as the parasitoid wasp family Ichneumonidae, are poorly known. This work reports some of the first results of an extensive survey implemented in Kibale National Park, Uganda. A total of 456 individuals was caught of the subfamily Rhyssinae Morley, 1913, which in the Afrotropical region was previously known from only 30 specimens. Here, the six species found at the site are described and the Afrotropical Rhyssinae are reviewed. Two new species, Epirhyssa johanna Hopkins, sp. nov. and E. quagga sp. nov. , are described and a key, diagnostic characters, and descriptions for all 13 known Afrotropical species are provided, including the first description of the male of Epirhyssa overlaeti Seyrig, 1937. Epirhyssa gavinbroadi Rousse & van Noort, 2014, syn. nov. is proposed to be a synonym of E. uelensis Benoit, 1951. Extensive sampling with Malaise traps gave an unprecedented sample size, and the method is recommended for other poorly known tropical areas., (Tapani Hopkins, Heikki Roininen, Simon van Noort, Gavin R. Broad, Kari Kaunisto, Ilari E. Sääksjärvi.)

Thermal performance curves (TPCs) are intended to approximate the relationship between temperature and fitness, and are commonly integrated into species distributional models for understanding climate change responses. However, TPCs may vary across traits because selection and environmental sensitivity (plasticity) differ across traits or because the timing and duration of the temperature exposure, here termed time scale, may alter trait variation. Yet, the extent to which TPCs vary temporally and across traits is rarely considered in assessments of climate change responses. Using a common garden approach, we estimated TPCs for standard metabolic rate (SMR), and activity in Drosophila melanogaster at three test temperatures (16, 25 and 30°C), using flies from each of six developmental temperatures (16, 18, 20, 25, 28 and 30°C). We examined the effects of time scale of temperature exposure (minutes/hours versus days/weeks) in altering TPC shape and position, and commonly used descriptors of the TPC: thermal optimum ( T opt ), thermal limits ( T min and T max ) and thermal breadth ( T br ). In addition, we collated previously published estimates of TPCs for fecundity and egg-to-adult viability in D. melanogaster We found that the descriptors of the TPCs varied across traits (egg-to-adult viability, SMR, activity and fecundity), but variation in TPCs within these traits was small across studies when measured at the same time scales. The time scale at which traits were measured contributed to greater variation in TPCs than the observed variance across traits, although the relative importance of time scale differed depending on the trait (activity versus fecundity). Variation in the TPC across traits and time scales suggests that TPCs using single traits may not be an accurate predictor of fitness and thermal adaptation across environments., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

The Afrotropical species of Glenosema are reviewed. Seventeen species are recognized, four previously described species, G. niloticum (Kieffer), G. durum Barbosa, Kawada Azevedo, G. denteatum Lanes Azevedo and G. elevatum Lanes Azevedo, and thirteen species described and illustrated as new: G. qaqambile sp. nov., G. deki sp. nov., G. ebazileyo sp. nov., G. tyaba sp. nov., G. umphanda sp. nov., G. banzi sp. nov., G. bubanzi sp. nov., G. ahlukileyo sp. nov., G. icephe sp. nov., G. mxinwa sp. nov., G. lingana sp. nov., G. uphahla sp. nov. and G. tyheli sp. nov. A key to the Afrotropical species is presented. We highlight potential character suites that may be phylogenetically significant and consider evolutionary scenarios that may have led to the present distribution pattern associated with semi-arid areas on the African continent.

Bactrocera biguttula is an African olive fruit fly that does not attack cultivated olives but rather develops in the fruits of wild species of Olea and Noronhia. The complete mitochondrial genome of an individual specimen was characterized in comparison to other Bactrocera. The phylogenetic relationships of B. biguttula with other Dacini were investigated, with special focus on B. oleae, an agricultural pest known to attack cultivated and wild olives. The sequence had a total length of 15,829 bp, and included the typical features of insect mitogenomes, similarly to the other Bactrocera analysed. Start codons included ATG, ATC, ATT, and TCG (in COI). The majority of stop codons (TAA) were fully encoded, whereas in some cases only TA or T were present. The complete sequence was biased towards A + T, with a positive AT-skew and a negative GC-skew. The predicted cloverleaf structure of tRNA Ser1 showed absence of the DHU arm, a common feature in insects and other Metazoans. Phylogenetic reconstruction showed that B. biguttula and B. oleae are sister species, having diverged from a common ancestor < 10 Myr ago. This result warrants future genomic comparisons between these two closely related species for investigating the specific adaptations to the different hosts., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Wild and cultivated olives harbor and share a diversity of insects, some of which are considered agricultural pests, such as the olive fruit fly. The assemblage of olive-associated parasitoids and seed wasps is rich and specialized in sub-Saharan Africa, with native species possibly coevolving with their hosts. Although historical entomological surveys reported on the diversity of olive wasp species in the Western Cape Province of South Africa, no comprehensive study has been performed in the region in the molecular era. In this study, a dual approach combining morphological and DNA-based methods was used for the identification of adult specimens reared from olive fruits. Four species of Braconidae and six species of Chalcidoidea were identified, and DNA barcoding methodologies were used to investigate conspecificity among individuals, based on randomly selected representative specimens. Morphological identifications were congruent with DNA data, as NJ and ML trees correctly placed the sequences for each species either at the genus or species level, depending on the available taxa coverage, and genetic distances strongly supported conspecificity. No clear evidence of cryptic diversity was found. Overall seed infestation and parasitism rates were higher in wild olives compared to cultivated olives, and highest for Eupelmus spermophilus and Utetes africanus . These results can be used for early DNA-based detection of wasp larvae in olives and to further investigate the biology and ecology of these species.

A new Afrotropical species of Polynema Haliday, 1833 (Hymenoptera: Mymaridae), Polynema (Polynema) sagittaria van Noort & Triapitsyn, sp. n. , is described and illustrated based on specimens collected in the Cape Floral region in south-western South Africa. This species is morphologically closely related to the recently described Polynema (Polynema) dikobraz Triapitsyn, 2017 from Madagascar, both species possessing enlarged spine-like microtrichia on the fore wing disc that are unique among all the known world fairyflies. This new species belongs to the informal dikobraz species-group of the nominate subgenus of Polynema , which previously was only known from Madagascar. In addition, P.sagittaria has the ovipositor extending ventrally under the mesosoma to well in front of the head, in a bow-like curve, and housed in a narrow, anterior elongation of the metasoma, the basal sac. Occurrence and possible significance of such a bizarre ovipositor in other Mymaridae is discussed. All images and online keys are available on www.waspweb.org.

Chalcidoidea are a megadiverse group of mostly parasitoid wasps of major ecological and economical importance that are omnipresent in almost all extant terrestrial habitats. The timing and pattern of chalcidoid diversification is so far poorly understood and has left many important questions on the evolutionary history of Chalcidoidea unanswered. In this study, we infer the early divergence events within Chalcidoidea and address the question of whether or not ancestral chalcidoids were small egg parasitoids. We also trace the evolution of some key traits: jumping ability, development of enlarged hind femora, and associations with figs. Our phylogenetic inference is based on the analysis of 3,239 single-copy genes across 48 chalcidoid wasps and outgroups representatives. We applied an innovative a posteriori evaluation approach to molecular clock-dating based on nine carefully validated fossils, resulting in the first molecular clock-based estimation of deep Chalcidoidea divergence times. Our results suggest a late Jurassic origin of Chalcidoidea, with a first divergence of morphologically and biologically distinct groups in the early to mid Cretaceous, between 129 and 81 million years ago (mya). Diversification of most extant lineages happened rapidly after the Cretaceous in the early Paleogene, between 75 and 53 mya. The inferred Chalcidoidea tree suggests a transition from ancestral minute egg parasitoids to larger-bodied parasitoids of other host stages during the early history of chalcidoid evolution. The ability to jump evolved independently at least three times, namely in Eupelmidae, Encyrtidae, and Tanaostigmatidae. Furthermore, the large-bodied strongly sclerotized species with enlarged hind femora in Chalcididae and Leucospidae are not closely related. Finally, the close association of some chalcidoid wasps with figs, either as pollinators, or as inquilines/gallers or as parasitoids, likely evolved at least twice independently: in the Eocene, giving rise to fig pollinators, and in the Oligocene or Miocene, resulting in non-pollinating fig-wasps, including gallers and parasitoids. The origins of very speciose lineages (e.g., Mymaridae, Eulophidae, Pteromalinae) are evenly spread across the period of chalcidoid evolution from early Cretaceous to the late Eocene. Several shifts in biology and morphology (e.g., in host exploitation, body shape and size, life history), each followed by rapid radiations, have likely enabled the evolutionary success of Chalcidoidea., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Two new species of therocephalian therapsids are described from the upper Permian Teekloof Formation of the Karoo Basin, South Africa. They include two specimens of a whaitsiid, Microwhaitsia mendrezi gen. et sp. nov., and a single, small whaitsioid Ophidostoma tatarinovi gen. et sp. nov., which preserves a combination of primitive and apomorphic features. A phylogenetic analysis of 56 therapsid taxa and 136 craniodental and postcranial characters places the new taxa within the monophyletic sister group of baurioids-Whaitsioidea-with Microwhaitsia as a basal whaitsiid and Ophidostoma as an aberrant whaitsioid just outside the hofmeyriid+whaitsiid subclade. The new records support that whaitsioids were diverse during the early-late Permian (Wuchiapingian) and that the dichotomy between whaitsiid-line and baurioid-line eutherocephalians was established early on. The oldest Gondwanan whaitsiid Microwhaitsia and additional records from the lower strata of the Teekloof Formation suggest that whaitsioids had diversified by the early Wuchiapingian and no later than Pristerognathus Assemblage Zone times. Prior extinction estimates based on species counts are reflected in an analysis of origination/extinction rates, which imply increasing faunal turnover from Guadalupian to Lopingian (late Permian) times. The new records support a growing body of evidence that some key Lopingian synapsid clades originated near or prior to the Guadalupian-Lopingian boundary ca. 260-259 million years ago, but only radiated following the end-Guadalupian extinction of dinocephalians and basal therocephalian predators (long-fuse model). Ongoing collecting in older portions of the Teekloof Formation (e.g., Pristerognathus Assemblage Zone) will shed further light on early eutherocephalians during this murky but critical time in their evolutionary diversification., Competing Interests: The authors declare there are no competing interests.

Major transformations in brain size and proportions, such as the enlargement of the brain during the evolution of birds, are accompanied by profound modifications to the skull roof. However, the hypothesis of concerted evolution of shape between brain and skull roof over major phylogenetic transitions, and in particular of an ontogenetic relationship between specific regions of the brain and the skull roof, has never been formally tested. We performed 3D morphometric analyses to examine the deep history of brain and skull-roof morphology in Reptilia, focusing on changes during the well-documented transition from early reptiles through archosauromorphs, including nonavian dinosaurs, to birds. Non-avialan taxa cluster tightly together in morphospace, whereas Archaeopteryx and crown birds occupy a separate region. There is a one-to-one correspondence between the forebrain and frontal bone and the midbrain and parietal bone. Furthermore, the position of the forebrain-midbrain boundary correlates significantly with the position of the frontoparietal suture across the phylogenetic breadth of Reptilia and during the ontogeny of individual taxa. Conservation of position and identity in the skull roof is apparent, and there is no support for previous hypotheses that the avian parietal is a transformed postparietal. The correlation and apparent developmental link between regions of the brain and bony skull elements are likely to be ancestral to Tetrapoda and may be fundamental to all of Osteichthyes, coeval with the origin of the dermatocranium.

Late Permian Karoo Basin tectonics in South Africa are reflected as two fining-upward megacycles in the Balfour and upper Teekloof formations. Foreland tectonics are used to explain the cyclic nature and distribution of sedimentation, caused by phases of loading and unloading in the southern source areas adjacent to the basin. New data supports this model, and identifies potential climatic effects on the tectonic regime. Diachronous second-order subaerial unconformities (SU) are identified at the base and top of the Balfour Formation. One third-order SU identified coincides with a faunal turnover which could be related to the Permo-Triassic mass extinction (PTME). The SU are traced, for the first time, to the western portion of the basin (upper Teekloof Formation). Their age determinations support the foreland basin model as they coincide with dated paroxysms. A condensed distal (northern) stratigraphic record is additional support for this tectonic regime because orogenic loading and unloading throughout the basin was not equally distributed, nor was it in-phase. This resulted in more frequent non-deposition with increased distance from the tectonically active source. Refining basin dynamics allows us to distinguish between tectonic and climatic effects and how they have influenced ancient ecosystems and sedimentation through time.

The only true living endothermic vertebrates are birds and mammals, which produce and regulate their internal temperature quite independently from their surroundings. For mammal ancestors, anatomical clues suggest that endothermy originated during the Permian or Triassic. Here we investigate the origin of mammalian thermoregulation by analysing apatite stable oxygen isotope compositions (δ 18 O p ) of some of their Permo-Triassic therapsid relatives. Comparing of the δ 18 O p values of therapsid bone and tooth apatites to those of co-existing non-therapsid tetrapods, demonstrates different body temperatures and thermoregulatory strategies. It is proposed that cynodonts and dicynodonts independently acquired constant elevated thermometabolism, respectively within the Eucynodontia and Lystrosauridae + Kannemeyeriiformes clades. We conclude that mammalian endothermy originated in the Epicynodontia during the middle-late Permian. Major global climatic and environmental fluctuations were the most likely selective pressures on the success of such elevated thermometabolism.

The relationship between dinosaurs and other reptiles is well established, but the sequence of acquisition of dinosaurian features has been obscured by the scarcity of fossils with transitional morphologies. The closest extinct relatives of dinosaurs either have highly derived morphologies or are known from poorly preserved or incomplete material. Here we describe one of the stratigraphically lowest and phylogenetically earliest members of the avian stem lineage (Avemetatarsalia), Teleocrater rhadinus gen. et sp. nov., from the Middle Triassic epoch. The anatomy of T. rhadinus provides key information that unites several enigmatic taxa from across Pangaea into a previously unrecognized clade, Aphanosauria. This clade is the sister taxon of Ornithodira (pterosaurs and birds) and shortens the ghost lineage inferred at the base of Avemetatarsalia. We demonstrate that several anatomical features long thought to characterize Dinosauria and dinosauriforms evolved much earlier, soon after the bird-crocodylian split, and that the earliest avemetatarsalians retained the crocodylian-like ankle morphology and hindlimb proportions of stem archosaurs and early pseudosuchians. Early avemetatarsalians were substantially more species-rich, widely geographically distributed and morphologically diverse than previously recognized. Moreover, several early dinosauromorphs that were previously used as models to understand dinosaur origins may represent specialized forms rather than the ancestral avemetatarsalian morphology.

Elucidating patterns of population structure for species with complex life histories, and disentangling the processes driving such patterns, remains a significant analytical challenge. Humpback whale (Megaptera novaeangliae) populations display complex genetic structures that have not been fully resolved at all spatial scales. We generated a data set of nuclear markers for 3575 samples spanning the seven breeding stocks and substocks found in the South Atlantic and western and northern Indian Oceans. For the total sample, and males and females separately, we assessed genetic diversity, tested for genetic differentiation between putative populations and isolation by distance, estimated the number of genetic clusters without a priori population information and estimated rates of gene flow using maximum-likelihood and Bayesian approaches. At the ocean basin scale, structure is governed by geographical distance (IBD P < 0.05) and female fidelity to breeding areas, in line with current understanding of the drivers of broadscale population structure. Consistent with previous studies, the Arabian Sea breeding stock was highly genetically differentiated (F ST 0.034-0.161; P < 0.01 for all comparisons). However, the breeding stock boundary between west South Africa and east Africa was more porous than expected based on genetic differentiation, cluster and geneflow analyses. Instances of male fidelity to breeding areas and relatively high rates of dispersal for females were also observed between the three substocks in the western Indian Ocean. The relationships between demographic units and current management boundaries may have ramifications for assessments of the status and continued protections of populations still in recovery from commercial whaling., (© 2017 John Wiley & Sons Ltd.)

A revision of Afrotropical species of Dyscritobaeus Perkins is presented with re-description of the four known species (D. bicolor O'Connor et Ashe, D. comitans Perkins, D. parvipennis (Dodd) and D. sulawensis Mineo, O'Connor et Ashe) and description of six new species (D. antananarivensis Tortorici et Caleca sp. nov., D. flavus Tortorici et Caleca sp. nov., D. kilimanjarensis Tortorici et Caleca sp. nov., D. madagascarensis Tortorici et Caleca sp. nov., D. ndokii Tortorici et Caleca sp. nov. and D. tanzaniensis Tortorici et Caleca sp. nov.). Dyscritobaeus cerosus is considered to be a junior synonym of D. comitans, D. hannibal is considered to be a junior synonym of D. sulawensis and D. maputanus is a junior synonym of D. parvipennis; so far these species are the only three Dyscritobaeus species that are widely distributed across four zoogeographical regions: Afrotropical, Australian, Oriental and Palearctic. Dyscritobaeus comitans and D. ndokii Tortorici et Caleca sp. nov. belong to the comitans-group, the other eight species belong to the orientalis-group. Dyscritobaeus antananarivensis Tortorici et Caleca sp. nov., D. madagascarensis Tortorici et Caleca sp. nov. and D. sulawensis are morphologically similar to D. orientalis bearing the specillum on T2; D. bicolor, D. flavus Tortorici et Caleca sp. nov., D. kilimanjarensis Tortorici et Caleca sp. nov., are morphologically similar to D. parvipennis; D. tanzaniensis Tortorici et Caleca sp. nov. shares the lack of a protruding metascutellum together with D. aspinosus Mineo, O'Connor et Ashe.Dyscritobaeus species are sexually dimorphic, particularly in the following features: the anterior and posterior fringes of the fore wing are longer in the female than in the male; the odontoid process, when present in females, is less developed or absent in the corresponding males; the specillum, when present in females, is less defined and smaller or absent in the corresponding males; the sculpture of the head is more evident in males than in females; and the first and second tergites are frequently lighter in males than in females.

The turtle shell is a complex structure that currently serves a largely protective function in this iconically slow-moving group [1]. Developmental [2, 3] and fossil [4-7] data indicate that one of the first steps toward the shelled body plan was broadening of the ribs (approximately 50 my before the completed shell [5]). Broadened ribs alone provide little protection [8] and confer significant locomotory [9, 10] and respiratory [9, 11] costs. They increase thoracic rigidity [8], which decreases speed of locomotion due to shortened stride length [10], and they inhibit effective costal ventilation [9, 11]. New fossil material of the oldest hypothesized stem turtle, Eunotosaurus africanus [12] (260 mya) [13, 14] from the Karoo Basin of South Africa, indicates the initiation of rib broadening was an adaptive response to fossoriality. Similar to extant fossorial taxa [8], the broad ribs of Eunotosaurus provide an intrinsically stable base on which to operate a powerful forelimb digging mechanism. Numerous fossorial correlates [15-17] are expressed throughout Eunotosaurus' skeleton. Most of these features are widely distributed along the turtle stem and into the crown clade, indicating the common ancestor of Eunotosaurus and modern turtles possessed a body plan significantly influenced by digging. The adaptations related to fossoriality likely facilitated movement of stem turtles into aquatic environments early in the groups' evolutionary history, and this ecology may have played an important role in stem turtles surviving the Permian/Triassic extinction event., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Since its discovery, Euparkeria capensis has been a key taxon for understanding the early evolution of archosaurs. The braincase of Euparkeria was described based on a single specimen, but much uncertainty remained. For the first time, all available braincase material of Euparkeria is re-examined using micro-computed tomography scanning. Contrary to previous work, the parabasisphenoid does not form the posterior border of the fenestra ovalis in lateral view, but it does bear a dorsal projection that forms the anteroventral half of the fenestra. No bone pneumatization was found, but the lateral depression of the parabasisphenoid may have been pneumatic. We propose that the lateral depression likely corresponds to the anterior tympanic recess present in crown archosaurs. The presence of a laterosphenoid is confirmed for Euparkeria. It largely conforms to the crocodilian condition, but shows some features which make it more similar to the avemetatarsalian laterosphenoid. The cochlea of Euparkeria is elongated, forming a deep cochlear recess. In comparison with other basal archosauromorphs, the metotic foramen is much enlarged and regionalized into vagus and recessus scalae tympani areas, indicating an increase in its pressure-relief mechanism. The anterior semicircular canal is extended and corresponds to an enlarged floccular fossa. These aspects of the braincase morphology may be related to the development of a more upright posture and active lifestyle. They also indicate further adaptations of the hearing system of Euparkeria to terrestriality.

An updated checklist of Dryinidae, Embolemidae and Sclerogibbidae from Central African Republic and Uganda is presented. The following new species of Dryinidae are described: from Central African Republic: Anteon dzanganum sp. n. (Anteoninae); from Uganda: Anteon granulatum sp. n., Anteon kibalense sp. n., Anteon makererense sp. n., Anteon mubfs sp. n. (Anteoninae); Bocchus kibalensis sp. n. (Bocchinae); Dryinus kibalus sp. n. (Dryininae); Gonatopus kanyawarus sp. n. (Gonatopodinae). The following species have been recorded for the first time from Central African Republic: Embolemidae: Ampulicomorpha madecassa Olmi, 1999a; Embolemus capensis Olmi, 1997; Dryinidae: Aphelopus mediocarinatus (Benoit, 1951d), Aphelopus testaceus Olmi, 1991, Aphelopus wittei Benoit, 1951c (Aphelopinae); Anteon cautum Olmi, 1994a, Anteon evertsi Olmi, 1989, Anteon gutturnium (Benoit, 1951b), Anteon inflatrix Benoit, 1951b, Anteon kivuanum (Benoit, 1951c), Anteon semajanna Olmi, Copeland & Guglielmino, 2015, Anteon zairense Benoit, 1951d (Anteoninae); Pseudodryinus townesi (Olmi, 1984) (Dryininae); Echthrodelphax tauricus Ponomarenko, 1970, Gonatopus camerounensis Olmi, 2011, Gonatopus kolyadai Olmi, 2007b, Neodryinus antiquus Benoit, 1954, Neodryinus tussaci Olmi, 2004b (Gonatopodinae); Sclerogibbidae: Probethylus callani Richards, 1939b; Sclerogibba algerica Benoit, 1963, Sclerogibba rapax Olmi, 2005a. The following species have been recorded for the first time from Uganda: Embolemidae: Ampulicomorpha magna Olmi, 1996; Dryinidae: Anteon cautum Olmi, 1994a, Anteon fisheri Olmi, 2003, Anteon hoyoi Olmi, 1984, Anteon kivuanum (Benoit, 1951c), Anteon townesi Olmi, 1984, Anteon zairense Benoit, 1951d (Anteoninae); Bocchus bini Olmi, 1984 (Bocchinae); Dryinus saussurei (Ceballos, 1936) (Dryininae); Echthrodelphax migratorius Benoit, 1954, Neodryinus tussaci Olmi, 2004b (Gonatopodinae). The following further species has been recorded for the first time from Mali: Sclerogibba algerica Benoit, 1963 (Sclerogibbidae); from Ivory Coast: Adryinus oweni Olmi, 1984 (Gonatopodinae); from Cameroon and South Africa: Gonatopus operosus Olmi, 1993 (Gonatopodinae); from Democratic Republic of the Congo and Zambia: Neodryinus antiquus Benoit, 1954 (Gonatopodinae); from South Africa: Anteon striatum Olmi, 2005b (Anteoninae). Including the above new records, 23 species of Dryinidae (previously six), two species of Embolemidae (previously none) and three species of Sclerogibbidae (previously two) are now known from Central African Republic. For Uganda, 39 species of Dryinidae (previously 23), one species of Embolemidae (previously none) and four species of Sclerogibbidae (previously four) are now known. Additional new faunistic records are provided for Cameroon, Democratic Republic of the Congo, Ivory Coast, Mali, South Africa and Zambia.

Background: Afrotremex Pasteels, 1951 is a rare genus of wasps endemic to the Afrotropical region, containing 6 species represented by 14 specimens. Specimens were previously only recorded from central Africa: Cameroon, Congo, Democratic Republic of Congo, Gabon and Uganda., New Information: Here we record two additional specimens housed in the Natural History Museum in London (BMNH), one of which is a male of A. xylophagus Goulet, 2014 collected in Ghana (previously Gold Coast). This record extends the known distribution of the genus into west Africa, and represents the second known male specimen for the genus. The other BMNH specimen is a female paratype of A. violaceus Pasteels, 1951 collected in the Democratic Republic of Congo. We provide high quality photographs of these additional two specimens. Images of all six known species are openly available online on WaspWeb. In addition we have developed interactive online Lucid Matrix and Lucid Phoenix identification keys to the species, which are openly available on WaspWeb at: http://www.waspweb.org/Siricoidea/Siricidae/Keys/index.htm.

Terrestrial vertebrates are first known to colonize high-latitude regions during the middle Permian (Guadalupian) about 270 million years ago, following the Pennsylvanian Gondwanan continental glaciation. However, despite over 150 years of study in these areas, the biogeographic origins of these rich communities of land-dwelling vertebrates remain obscure. Here we report on a new early Permian continental tetrapod fauna from South America in tropical Western Gondwana that sheds new light on patterns of tetrapod distribution. Northeastern Brazil hosted an extensive lacustrine system inhabited by a unique community of temnospondyl amphibians and reptiles that considerably expand the known temporal and geographic ranges of key subgroups. Our findings demonstrate that tetrapod groups common in later Permian and Triassic temperate communities were already present in tropical Gondwana by the early Permian (Cisuralian). This new fauna constitutes a new biogeographic province with North American affinities and clearly demonstrates that tetrapod dispersal into Gondwana was already underway at the beginning of the Permian.

Cetaceans, occupying the top levels in marine food chains, are vulnerable to elevated levels of potentially toxic trace elements, such as aluminium (Al), mercury (Hg) and nickel (Ni). Negative effects associated with these toxic metals include infection by opportunistic microbial invaders. To corroborate the link between the presence of cutaneous fungal invaders and trace element levels, skin samples from 40 stranded false killer whales (FKWs) were analysed using culture techniques and inductively coupled plasma-mass spectroscopy. Twenty-two skin samples yielded 18 clinically relevant fungal species. While evidence for bioaccumulation of Hg in the skin of the FKWs was observed, a strong link was found to exist between the occurrence of opportunistic fungal invaders and higher Al : Se and Al : Zn ratios. This study provides indications that elevated levels of some toxic metals, such as Al, contribute to immunotoxicity rendering FKWs susceptible to colonization by cutaneous opportunistic fungal invaders., (© 2015 Society for Applied Microbiology and John Wiley & Sons Ltd.)

There is compelling evidence that more diverse ecosystems deliver greater benefits to people, and these ecosystem services have become a key argument for biodiversity conservation. However, it is unclear how much biodiversity is needed to deliver ecosystem services in a cost-effective way. Here we show that, while the contribution of wild bees to crop production is significant, service delivery is restricted to a limited subset of all known bee species. Across crops, years and biogeographical regions, crop-visiting wild bee communities are dominated by a small number of common species, and threatened species are rarely observed on crops. Dominant crop pollinators persist under agricultural expansion and many are easily enhanced by simple conservation measures, suggesting that cost-effective management strategies to promote crop pollination should target a different set of species than management strategies to promote threatened bees. Conserving the biological diversity of bees therefore requires more than just ecosystem-service-based arguments.

The Afrotropical Cynipoidea are represented by 306 described species and 54 genera in four families: Cynipidae, Figitidae, Liopteridae and Ibaliidae, the latter represented by a single introduced species. Seven of these genera are only represented by undescribed species in the region. Seven new genus-level synonymies, one genus resurrected from synonymy, 54 new combinations, one combination reinstated, and one new replacement name are presented. We provide identification keys to the families, subfamilies and genera of cynipoid wasps occurring in the Afrotropical region (Africa south of the Sahara, including Madagascar and southern Arabian Peninsula). Online interactive Lucid Phoenix and Lucid matrix keys are available at: http://www.waspweb.org/Cynipoidea/Keys/index.htm. An overview of the biology and checklists of species for each genus are provided. This paper constitutes the first contributory chapter to the book on Afrotropical Hymenoptera.

The revision of the Afrotropical Ophioninae is updated, based on the examination of about 800-900 individuals in the South African and European museum collections. A robust interactive matrix key was built to provide quick and reliable identifications. The key is available online at http://www.waspweb.org. Two new species are described: Dicamptusmaxipol sp. n. and Enicospilusgauldetmitchellorum sp. n. Numerous new distribution and biological records are provided, and noticeable morphological intra-specific variations are detailed. Enicospilusbatus Gauld & Mitchell, syn. n. is considered as a junior synonym of Enicospilusluebberti (Enderlein).

Three new Afrotropical Lycorina species are described: L. horstmanni sp. nov., L. jacksonfive sp. nov. and L. riftensis sp. nov. The description of L. globiceps is expanded to include the large variability of the colour pattern. New distribution records are provided for L. fici and L. globiceps. An illustrated dichotomous key, and an online interactive matrix key available on www.waspweb.org, are provided for the identification of the seven known Afrotropical Lycorininae species.

The cynipoid subfamily Figitinae is poorly represented in the Afrotropical region with two genera (Figites Latreille and Xyalophora Kieffer) and six species currently known. Here we record an additional two genera (Neralsia Cameron and Lonchidia Thomson) for the region and describe three new species: Neralsiahaddocki sp. n.; Xyalophoratedjoansi sp. n.; Xyalophoratintini sp. n. Benoit's species described in 1956 are synonymized under Figitesaciculatus (Benoit, 1956): Figiteseffossus syn. n.; Figitesfavonius syn. n.; Figitesfurvus syn. n.; Figitesfraudator syn. n. Identification keys to the figitine genera and species occurring in the Afrotropical region are provided. Online interactive Lucid Phoenix and Lucid matrix keys are available at: http://www.waspweb.org/Cynipoidea/Keys/index.htm.

Background: Temnospondyls are one of the earliest radiations of limbed vertebrates. Skeletal remains of more than 190 genera have been identified from late Paleozoic and early Mesozoic rocks. Paleozoic temnospondyls comprise mainly small to medium sized forms of diverse habits ranging from fully aquatic to fully terrestrial. Accordingly, their ichnological record includes tracks described from many Laurasian localities. Mesozoic temnospondyls, in contrast, include mostly medium to large aquatic or semi-aquatic forms. Exceedingly few fossil tracks or trackways have been attributed to Mesozoic temnospondyls, and as a consequence very little is known of their locomotor capabilities on land., Methodology/principal Findings: We report a ca. 200 Ma trackway, Episcopopus ventrosus, from Lesotho, southern Africa that was made by a 3.5 m-long animal. This relatively long trackway records the trackmaker dragging its body along a wet substrate using only the tips of its digits, which in the manus left characteristic drag marks. Based on detailed mapping, casting, and laser scanning of the best-preserved part of the trackway, we identified synapomorphies (e.g., tetradactyl manus, pentadactyl pes) and symplesiomorphies (e.g., absence of claws) in the Episcopopus trackway that indicate a temnospondyl trackmaker., Conclusions/significance: Our analysis shows that the Episcopopus trackmaker progressed with a sprawling posture, using a lateral-sequence walk. Its forelimbs were the major propulsive elements and there was little lateral bending of the trunk. We suggest this locomotor style, which differs dramatically from the hindlimb-driven locomotion of salamanders and other extant terrestrial tetrapods can be explained by the forwardly shifted center of mass resulting from the relatively large heads and heavily pectoral girdles of temnospondyls.

The Afrotropical cremastine fauna (Hymenoptera: Ichneumonidae) is characterized by the near absence of the genus Cremastus, with a single species reported from Madagascar. The fauna is also characterized by the presence of several endemic genera. Among the latter is the exceptional monotypic genus Belesica. We describe two new species from these extremely rare Afrotropical genera, namely Belesica madiba sp. nov. and Cremastus tutui sp. nov.. Both are only known from South Africa.

The genus Scelio is a cosmopolitan and speciose group of solitary parasitoids of the eggs of short-horned grasshoppers (Orthoptera: Acrididae). A number of these hosts are important pests, including plague locusts of the genus Schistocerca. Species of Scelio are recognized as potentially important biological control agents, but this possibility has yet to be fully realized, in part because the species-level taxonomy is still incompletely developed. The species of the pulchripennis group have been recently revised. As a continuation of this effort, here we revise the Afrotropical species of Scelio, excluding the pulchripennis species group. Sixty two (62) species are treated, 48 of which are new. Species are classified into the following species groups: ernstii (12 species, 9 new), howardi (23 species, 19 new), ipomeae (6 species, 5 new), irwini (4 species, 3 new), simoni (3 new species) and walkeri (12 species, 9 new). Keys to species groups and to the species within each group are provided. New species described are: S. albatus Yoder, sp. n., S. aphares Yoder, sp. n., S. apospastos Yoder, sp. n., S. ardelio Yoder, sp. n., S. aurantium Yoder, sp. n., S. balo Valerio & Yoder, sp. n., S. bayanga Yoder, sp. n., S. bubulo Yoder, sp. n., S. cano Yoder, sp. n., S. clypeatus Yoder, sp. n., S. concavus Yoder, sp. n., S. copelandi Yoder, sp. n., S. crepo Yoder, sp. n., S. destico Yoder, sp. n., S. dupondi Yoder, sp. n., S. effervesco Yoder, sp. n., S. erugatus Yoder, sp. n., S. exophthalmus Yoder, sp. n., S. fremo Valerio & Yoder, sp. n., S. gemo Yoder, sp. n., S. grunnio Yoder, sp. n., S. harinhalai Yoder, sp. n., S. igland Yoder, sp. n., S. impostor Yoder, sp. n., S. irwini Yoder, sp. n., S. janseni Yoder, sp. n., S. latro Yoder, sp. n., S. memorabilis Yoder, sp. n., S. modulus Yoder, sp. n., S. mutio Yoder, sp. n., S. ntchisii Yoder, sp. n., S. parkeri Yoder, sp. n., S. phaeoprora Yoder, sp. n., S. pilosilatus Yoder, sp. n., S. pipilo Yoder, sp. n., S. quasiclypeatus Yoder, sp. n., S. retifrons Yoder, sp. n., S. ructo Yoder, sp. n., S. scomma Yoder, sp. n., S. simoni Yoder, sp. n., S. simonolus Yoder, sp. n., S. somaliensis Yoder, sp. n., S. susurro Yoder, sp. n., S. tono Yoder, sp. n., S. transtrum Yoder, sp. n., S. tritus Yoder, sp. n., S. ululo Yoder, sp. n., S. vannoorti Valerio & Yoder, sp. n. The following species are redescribed: S. afer Kieffer, S. chapmani Nixon, S. howardi Crawford, S. ipomeae Risbec, stat. n., S. mauritanicus Risbec, S. philippinensis Ashmead, S. remaudierei Ferrière, S. striatus Priesner,S. taylori Nixon, and S. zolotarevskyi Ferrière. The genus Lepidoscelio Kieffer is treated as a junior synonym of Scelio Latreille, syn. n.; its type species, Lepidoscelio fuscipennis Kieffer, 1905 is transferred to Scelio, renamed Scelio obscuripennis Johnson, nom. n. (preoccupied by Scelio fuscipennis Ashmead, 1887), and redescribed. The following additional species are transferred from Lepidoscelio to Scelio: S. cayennensis (Risbec), comb. n., S. insularis Ashmead, rev. comb., S. luteus (Cameron), comb. n., S. thoracicus Ashmead, rev. comb. Lectotypes are designated for S. africanus Risbec, S. ipomeae Risbec, S. mauritanicus Risbec, S. remaudierei Ferrière, S. sudanensis Ferrière, and S. zolotarevskyi Ferrière. Scelio gaudens Nixon is a junior synonym of Scelio striatus Priesner, syn. n.; Scelio africanus Risbec and Scelio clarus Fouts are both junior synonyms of Scelio afer Kieffer, syn. n.; Scelio sudanensis Ferrière and Scelio cheops Nixon are both junior synonyms of Scelio zolotarevskyi Ferrière, syn. n.; Scelio cahirensis Priesner is a junior synonym of Scelio mauritanicus Risbec, syn. n. The name Scelio chapmanni Nixon is an incorrect original spelling, requiring an emendation to S. chapmani. Digital versions of the identification keys are available at http://www.waspweb.org/Platygastroidea/Keys/index.htm.

We revise the 10 genera and 23 species of the tribe Phaeogenini (Ichneumonidae: Ichneumoninae) known to occur in the Afrotropical region. We describe the following 13 new taxa: Kibalus Rousse, van Noort & Diller, gen. n.; K. toro Rousse, van Noort & Diller, sp. n.; K. mubfs Rousse & van Noort, sp. n.; Arearia oxymoron Rousse & van Noort, sp. n.; Chauvinia nyanga Rousse & van Noort, sp. n.; Dicaelotus asantesana Rousse & van Noort, sp. n.; D. hoerikwaggoensis Rousse & van Noort, sp. n.; D. tablemountainensis Rousse & van Noort, sp. n.; Heterischnus mfongosi Rousse & van Noort, sp. n.; H. mkomazi Rousse & van Noort, sp. n.; Lusius flummox Rousse & van Noort, sp. n.; Tycherus amatola Rousse & van Noort, sp. n.; and T. nardousberg Rousse & van Noort, sp. n. New distribution records: Heterischnus africanus (Heinrich, 1936) from South Africa, Tanzania and Uganda; H. krausi Schönitzer, 1999 from Rwanda; Lusius tenuissimus (Heinrich, 1938) from Democratic Republic of Congo, Malawi, Nigeria, South Africa, and Zimbabwe. A doubtful record of Aethecerus foveolatus Gregor, 1940 from Sao Tome is additionaly reported in the appendix. We provide illustrated diagnoses and identification notes. Online interactive dichotomous and matrix Lucid keys to genera and species are available at http://www.waspweb.org.

Chalcidoidea (Hymenoptera) is extremely diverse with an estimated 500 000 species. We present the first phylogenetic analysis of the superfamily based on both morphological and molecular data. A web-based, systematics workbench mx was used to score 945 character states illustrated by 648 figures for 233 morphological characters for a total of 66 645 observations for 300 taxa. The matrix covers 22 chalcidoid families recognized herein and includes 268 genera within 78 of 83 subfamilies. Morphological data were analysed alone and in combination with molecular data from ribosomal 18S (2105 bp) and 28S D2-D5 expansion regions (1812 bp). Analyses were analysed alone and in combined datasets using implied-weights parsimony and likelihood. Proposed changes in higher classification resulting from the analyses include: (i) recognition of Eriaporidae, revised status; (ii) recognition of Cynipencyrtidae, revised status; (iii) recognition of Azotidae, revised status; (iv) inclusion of Sycophaginae in Agaonidae, revised status; (v) reclassification of Aphelinidae to include Aphelininae, Calesinae, Coccophaginae, Eretmocerinae and Eriaphytinae; (vi) inclusion of Cratominae and Panstenoninae within Pteromalinae (Pteromalidae), new synonymy; (vii) inclusion of Epichrysomallinae in Pteromalidae, revised status. At a higher level, Chalcidoidea was monophyletic, with Mymaridae the sister group of Rotoitidae plus the remaining Chalcidoidea. A eulophid lineage was recovered that included Aphelinidae, Azotidae, Eulophidae, Signiphoridae, Tetracampidae and Trichogrammatidae. Eucharitidae and Perilampidae were monophyletic if Eutrichosomatinae (Pteromalidae) was included, and Eupelmidae was monophyletic if Oodera (Pteromalidae: Cleonyminae) was included. Likelihood recovered a clade of Eupelmidae + (Tanaostigmatidae + (Cynipencyrtus + Encyrtidae). Support for other lineages and their impact on the classification of Chalcidoidea is discussed. Several life-history traits are mapped onto the new phylogeny., (© 2012 Her Majesty the Queen in Right of Canada Cladistics © The Willi Hennig Society 2013 Reproduced with the permission of the Minister of Agriculture and Agri-Food Canada.)