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A highly endemic ant fauna is found in the arid regions of southern Africa, including species in the genus Ocymyrmex. This genus of ants has higher species richness in the western arid regions of southern Africa compared to tropical and subtropical parts of the continent. The processes that have produced these patterns of diversity and distribution of arid adapted ants in southern Africa have never been investigated. The diversification of many other taxa in the region has been associated with past climate fluctuations that occurred during the Miocene epoch. In this study, the nature and timing of historical processes that may have led to the diversification within Ocymyrmex were assessed. We hypothesized that past climate oscillations, characterized by long periods of aridification, have driven the current distribution of Ocymyrmex species that resulted in the highest species richness of the genus in the Deserts & xeric shrublands biome in southern Africa. Ninety-four Ocymyrmex worker specimens from Botswana, Kenya, Namibia, South Africa, Tanzania and Zimbabwe, representing 21 currently described species and six morphospecies, were included in a phylogenomic analysis. Phylogenies for the genus, based on next generation sequencing data from ultraconserved elements, were inferred using Maximum Likelihood, and a dating analysis was performed using secondary age estimates as calibration points. A distribution database of Ocymyrmex records was used to assign species ranges, which were then coded according to major biomes in southern Africa and used as input for biogeographical analysis. We explored the phylogenomic relationships of Ocymyrmex and analysed these within a biogeographical and paleoclimatic framework to disentangle the potential processes responsible for diversification in this group. Dating analyses estimated that the crown age of Ocymyrmex dates to the Oligocene, around 32 Ma. Diversification within this group occurred between the mid-Miocene (∼12.5 Ma) and Pleistocene (∼2 Ma). Our biogeographic analyses suggest that Ocymyrmex species originated in the south-western region of southern Africa, which is now part of the Deserts & xeric shrublands biome and diversified into eastern subtropical areas during the Pliocene. Paleoclimatic changes resulting in increased aridity during the Miocene likely drove the diversification of the genus Ocymyrmex. It is most likely that the diversification of grasslands, because of historical climate change, facilitated the diversification of these ants to the eastern parts of southern Africa when open grasslands replaced forests during the early Miocene., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Chalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra-conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first-instar larvae. The phylogeny suggests a transition from smaller soft-bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the "Angiosperm Terrestrial Revolution". Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3-180.5 Ma) and a crown age of 162.2 Ma (153.9-169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history., (© 2023 The Authors. Cladistics published by John Wiley & Sons Ltd on behalf of Willi Hennig Society.)

Bees are the most significant pollinators of flowering plants. This partnership began ca. 120 million years ago, but the uncertainty of how and when bees spread across the planet has greatly obscured investigations of this key mutualism. We present a novel analysis of bee biogeography using extensive new genomic and fossil data to demonstrate that bees originated in Western Gondwana (Africa and South America). Bees likely originated in the Early Cretaceous, shortly before the breakup of Western Gondwana, and the early evolution of any major bee lineage is associated with either the South American or African land masses. Subsequently, bees colonized northern continents via a complex history of vicariance and dispersal. The notable early absences from large landmasses, particularly in Australia and India, have important implications for understanding the assembly of local floras and diverse modes of pollination. How bees spread around the world from their hypothesized Southern Hemisphere origin parallels the histories of numerous flowering plant clades, providing an essential step to studying the evolution of angiosperm pollination syndromes in space and time., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

The order Hymenoptera (wasps, ants, sawflies, and bees) represents one of the most diverse animal lineages, but whether specific key innovations have contributed to its diversification is still unknown. We assembled the largest time-calibrated phylogeny of Hymenoptera to date and investigated the origin and possible correlation of particular morphological and behavioral innovations with diversification in the order: the wasp waist of Apocrita; the stinger of Aculeata; parasitoidism, a specialized form of carnivory; and secondary phytophagy, a reversal to plant-feeding. Here, we show that parasitoidism has been the dominant strategy since the Late Triassic in Hymenoptera, but was not an immediate driver of diversification. Instead, transitions to secondary phytophagy (from parasitoidism) had a major influence on diversification rate in Hymenoptera. Support for the stinger and the wasp waist as key innovations remains equivocal, but these traits may have laid the anatomical and behavioral foundations for adaptations more directly associated with diversification., (© 2023. The Author(s).)

The parasitoid lifestyle is largely regarded as a key innovation that contributed to the evolutionary success and extreme species richness of the order Hymenoptera. Understanding the phylogenetic history of hyperdiverse parasitoid groups is a fundamental step in elucidating the evolution of biological traits linked to parasitoidism. We used a genomic-scale dataset based on ultra-conserved elements and the most comprehensive taxon sampling to date to estimate the evolutionary relationships of Braconidae, the second largest family of Hymenoptera. Based on our results, we propose Braconidae to comprise 41 extant subfamilies, confirmed a number of subfamilial placements and proposed subfamily-level taxonomic changes, notably the restoration of Trachypetinae stat. rev. and Masoninae stat. rev. as subfamilies of Braconidae, confirmation that Apozyx penyai Mason belongs in Braconidae placed in the subfamily Apozyginae and the recognition of Ichneutinae sensu stricto and Proteropinae as non-cyclostome subfamilies robustly supported in a phylogenetic context. The correlation between koinobiosis with endoparasitoidism and idiobiosis with ectoparasitoidism, long thought to be an important aspect in parasitoid life history, was formally tested and confirmed in a phylogenetic framework. Using ancestral reconstruction methods based on both parsimony and maximum likelihood, we suggest that the ancestor of the braconoid complex was a koinobiont endoparasitoid, as was that of the cyclostome sensu lato clade. Our results also provide strong evidence for one transition from endo- to ectoparasitoidism and three reversals back to endoparasitoidism within the cyclostome sensu stricto lineage. Transitions of koino- and idiobiosis were identical to those inferred for endo- versus ectoparasitoidism, except with one additional reversal back to koinobiosis in the small subfamily Rhysipolinae., (Published by Elsevier Inc.)

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.)

Background: Parasitoidism, a specialized life strategy in which a parasite eventually kills its host, is frequently found within the insect order Hymenoptera (wasps, ants and bees). A parasitoid lifestyle is one of two dominant life strategies within the hymenopteran superfamily Cynipoidea, with the other being an unusual plant-feeding behavior known as galling. Less commonly, cynipoid wasps exhibit inquilinism, a strategy where some species have adapted to usurp other species' galls instead of inducing their own. Using a phylogenomic data set of ultraconserved elements from nearly all lineages of Cynipoidea, we here generate a robust phylogenetic framework and timescale to understand cynipoid systematics and the evolution of these life histories., Results: Our reconstructed evolutionary history for Cynipoidea differs considerably from previous hypotheses. Rooting our analyses with non-cynipoid outgroups, the Paraulacini, a group of inquilines, emerged as sister-group to the rest of Cynipoidea, rendering the gall wasp family Cynipidae paraphyletic. The families Ibaliidae and Liopteridae, long considered archaic and early-branching parasitoid lineages, were found nested well within the Cynipoidea as sister-group to the parasitoid Figitidae. Cynipoidea originated in the early Jurassic around 190 Ma. Either inquilinism or parasitoidism is suggested as the ancestral and dominant strategy throughout the early evolution of cynipoids, depending on whether a simple (three states: parasitoidism, inquilinism and galling) or more complex (seven states: parasitoidism, inquilinism and galling split by host use) model is employed., Conclusions: Our study has significant impact on understanding cynipoid evolution and highlights the importance of adequate outgroup sampling. We discuss the evolutionary timescale of the superfamily in relation to their insect hosts and host plants, and outline how phytophagous galling behavior may have evolved from entomophagous, parasitoid cynipoids. Our study has established the framework for further physiological and comparative genomic work between gall-making, inquiline and parasitoid lineages, which could also have significant implications for the evolution of diverse life histories in other Hymenoptera.

Two increasingly popular approaches to reconstruct the Tree of Life involve whole transcriptome sequencing and the target capture of ultraconserved elements (UCEs). Both methods can be used to generate large, multigene datasets for analysis of phylogenetic relationships in non-model organisms. While targeted exon sequencing across divergent lineages is now a standard method, it is still not clear if UCE data can be readily combined with published transcriptomes. In this study, we evaluate the combination of UCEs and transcriptomes in a single analysis using genome-, transcriptome-, and UCE data for 79 bees in the largest and most biologically diverse bee family, Apidae. Using existing tools, we first developed a workflow to assemble phylogenomic data from different sources and produced two large nucleotide matrices of combined data. We then reconstructed the phylogeny of the Apidae using concatenation- and coalescent-based methods, and critically evaluated the resulting phylogenies in the context of previously published genetic, genomic, and morphological data sets. Our estimated phylogenetic trees are robustly supported and largely congruent with previous molecular hypotheses, from deep nodes to shallow species-level phylogenies. Moreover, the combined approach allows us to resolve controversial nodes of the apid Tree of Life, by clarifying the relationships among the genera of orchid bees (Euglossini) and the monophyly of the Centridini. Additionally, we present novel phylogenetic evidence supporting the monophyly of the diverse clade of cleptoparasitic Apidae and the placement of two enigmatic, oil-collecting genera (Ctenoplectra and Tetrapedia). Lastly, we propose a revised classification of the family Apidae that reflects our improved understanding of apid higher-level relationships., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

The evolution of reversed sexual dichromatism and aposematic coloration has long been of interest to both theoreticians and empiricists. Yet despite the potential connections between these phenomena, they have seldom been jointly studied. Large carpenter bees (genus Xylocopa) are a promising group for such comparative investigations as they are a diverse clade in which both aposematism and reversed sexual dichromatism can occur either together or separately. We investigated the evolutionary history of dichromatism and aposematism and a potential correlation of these traits with diversification rates within Xylocopa, using a newly generated phylogeny for 179 Xylocopa species based on ultraconserved elements (UCEs). A monochromatic, inconspicuous ancestor is indicated for the genus, with subsequent convergent evolution of sexual dichromatism and aposematism in multiple lineages. Aposematism is found to covary with reversed sexual dichromatism in many species; however, reversed dichromatism also evolved in non-aposematic species. Bayesian Analysis of Macroevolutionary Models (BAMM) did not show increased diversification in any specific clade in Xylocopa, whereas support from Hidden State Speciation and Extinction (HiSSE) models remained inconclusive regarding an association of increased diversification rates with dichromatism or aposematism. We discuss the evolution of color patterns and diversification in Xylocopa by considering potential drivers of dichromatism and aposematism., (© 2018 The Author(s). Evolution © 2018 The Society for the Study of Evolution.)

Cuticular hydrocarbons (CHCs) are ubiquitous and highly diverse in insects, serving as communication signal and waterproofing agent. Despite their vital function, the causes, mechanisms and constraints on CHC diversification are still poorly understood. Here, we investigated phylogenetic constraints on the evolution of CHC profiles, using a global data set of the species-rich and chemically diverse ant genus Crematogaster. We decomposed CHC profiles into quantitative (relative abundances, chain length) and qualitative traits (presence/absence of CHC classes). A species-level phylogeny was estimated using newly generated and previously published sequences from five nuclear markers. Moreover, we reconstructed a phylogeny for the chemically diverse Crematogaster levior species group using cytochrome oxidase I. Phylogenetic signal was measured for these traits on genus and clade level and within the chemically diverse C. levior group. For most quantitative CHC traits, phylogenetic signal was low and did not differ from random expectation. This was true on the level of genus, clade and species group, indicating that CHC traits are evolutionary labile. In contrast, the presence or absence of alkenes and alkadienes was highly conserved within the C. levior group. Hence, the presence or absence of biosynthetic pathways may be phylogenetically constrained, especially at lower taxonomic levels. Our study shows that CHC composition can evolve rapidly, allowing insects to quickly adapt their chemical profiles to external selection pressures, whereas the presence of biosynthetic pathways appears more constrained. However, our results stress the importance to consider the taxonomic level when investigating phylogenetic constraints., (© 2017 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2017 European Society For Evolutionary Biology.)

The field of sequence based phylogenetic analyses is currently being transformed by novel hybrid-based targeted enrichment methods, such as the use of ultraconserved elements (UCEs). Rather than analyzing relationships among organisms using a small number of genes, these methods now allow us to evaluate relationships with many hundreds to thousands of individual gene loci. However, the inclusion of thousands of loci does not necessarily overcome the long-standing challenge of incongruence among phylogenetic trees derived from different genes or gene regions. One factor that impacts the level of incongruence in phylogenomic data sets is the level of GC bias. GC rich gene regions are prone to higher recombination rates than AT rich regions, driven by a process referred to as "GC biased gene conversion". As a result, high GC content can be negatively associated with phylogenetic accuracy, but the extent to which this impacts incongruence among UCEs is currently unstudied. We investigated the impact of GC content on phylogeny reconstruction using in silico captured UCE data for the corbiculate bees (Hymenoptera: Apidae). The phylogeny of this group has been the subject of extensive study, and incongruence among gene trees is thought to be a source of phylogenetic error. We conducted coalescent- and concatenation-based analyses of 810 individual gene loci from all 13 currently available bee genomes, including 8 corbiculate taxa. Both coalescent- and concatenation-based methods converged on a single topology for the corbiculate tribes. In contrast to concatenation, the coalescent-based methods revealed significant topological conflict at nodes involving the orchid bees (Euglossini) and honeybees (Apini). Partitioning the loci by GC content reveals decreasing support for the inferred topology with increasing GC bias. Based on the results of this study, we report the first evidence that GC biased gene conversion may contribute to topological incongruence in studies based on ultraconserved elements., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Cuticular hydrocarbons (CHCs) cover the cuticles of virtually all insects, serving as a waterproofing agent and as a communication signal. The causes for the high CHC variation between species, and the factors influencing CHC profiles, are scarcely understood. Here, we compare CHC profiles of ant species from seven biogeographic regions, searching for physiological constraints and for climatic and biotic selection pressures. Molecule length constrained CHC composition: long-chain profiles contained fewer linear alkanes, but more hydrocarbons with disruptive features in the molecule. This is probably owing to selection on the physiology to build a semi-fluid cuticular layer, which is necessary for waterproofing and communication. CHC composition also depended on the precipitation in the ants' habitats. Species from wet climates had more alkenes and fewer dimethyl alkanes than those from drier habitats, which can be explained by different waterproofing capacities of these compounds. By contrast, temperature did not affect CHC composition. Mutualistically associated (parabiotic) species possessed profiles highly distinct from non-associated species. Our study is, to our knowledge, the first to show systematic impacts of physiological, climatic and biotic factors on quantitative CHC composition across a global, multi-species dataset. We demonstrate how they jointly shape CHC profiles, and advance our understanding of the evolution of this complex functional trait in insects., (© 2017 The Author(s).)

Acropyga ants are a widespread clade of small subterranean formicines that live in obligate symbiotic associations with root mealybugs. We generated a data set of 944 loci of ultraconserved elements (UCEs) to reconstruct the phylogeny of 41 representatives of 23 Acropyga species using both concatenation and species-tree approaches. We investigated the biogeographic history of the genus through divergence dating analyses and ancestral range reconstructions. We also explored the evolution of the Acropyga-mealybug mutualism using ancestral state reconstruction methods. We recovered a highly supported species phylogeny for Acropyga with both concatenation and species-tree analyses. The age for crown-group Acropyga is estimated to be around 30Ma. The geographic origin of the genus remains uncertain, although phylogenetic affinities within the subfamily Formicinae point to a Paleotropical ancestor. Two main Acropyga lineages are recovered with mutually exclusive distributions in the Old World and New World. Within the Old World clade, a Palearctic and African lineage is suggested as sister to the remaining species. Ancestral state reconstructions indicate that Old World species have diversified mainly in close association with xenococcines from the genus Eumyrmococcus, although present-day associations also involve other mealybug genera. In contrast, New World Acropyga predominantly evolved with Neochavesia until a recent (10-15Ma) switch to rhizoecid mealybug partners (genus Rhizoecus). The striking mandibular variation in Acropyga evolved most likely from a 5-toothed ancestor. Our results provide an initial evolutionary framework for extended investigations of potential co-evolutionary interactions between these ants and their mealybug partners., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Obtaining sequence data from historical museum specimens has been a growing research interest, invigorated by next-generation sequencing methods that allow inputs of highly degraded DNA. We applied a target enrichment and next-generation sequencing protocol to generate ultraconserved elements (UCEs) from 51 large carpenter bee specimens (genus Xylocopa), representing 25 species with specimen ages ranging from 2-121 years. We measured the correlation between specimen age and DNA yield (pre- and post-library preparation DNA concentration) and several UCE sequence capture statistics (raw read count, UCE reads on target, UCE mean contig length and UCE locus count) with linear regression models. We performed piecewise regression to test for specific breakpoints in the relationship of specimen age and DNA yield and sequence capture variables. Additionally, we compared UCE data from newer and older specimens of the same species and reconstructed their phylogeny in order to confirm the validity of our data. We recovered 6-972 UCE loci from samples with pre-library DNA concentrations ranging from 0.06-9.8 ng/μL. All investigated DNA yield and sequence capture variables were significantly but only moderately negatively correlated with specimen age. Specimens of age 20 years or less had significantly higher pre- and post-library concentrations, UCE contig lengths, and locus counts compared to specimens older than 20 years. We found breakpoints in our data indicating a decrease of the initial detrimental effect of specimen age on pre- and post-library DNA concentration and UCE contig length starting around 21-39 years after preservation. Our phylogenetic results confirmed the integrity of our data, giving preliminary insights into relationships within Xylocopa. We consider the effect of additional factors not measured in this study on our age-related sequence capture results, such as DNA fragmentation and preservation method, and discuss the promise of the UCE approach for large-scale projects in insect phylogenomics using museum specimens., Competing Interests: The authors have declared that no competing interests exist.

The classification of the ant subfamily Formicinae is revised to reflect findings from a recent molecular phylogenetic study and complementary morphological investigations. The existing classification is maintained as far as possible, but some tribes and genera are redefined to ensure monophyly. Eleven tribes are recognized, all of which are strongly supported as monophyletic groups: Camponotini, Formicini, Gesomyrmecini, Gigantiopini, Lasiini (= Prenolepidii syn. n.), Melophorini (= Myrmecorhynchini syn. n.; = Notostigmatini syn. n.), Myrmelachistini stat. rev. (= Brachymyrmicini syn. n.), Myrmoteratini, Oecophyllini, Plagiolepidini, and Santschiellini stat. rev. Most of the tribes remain similar in content, but the generic composition of Lasiini, Melophorini, and Plagiolepidini is changed substantially. Species that have been placed in the genus Camponotus belong to three separate lineages. To ensure monophyly of this large, cosmopolitan genus we institute the following changes: Colobopsis and Dinomyrmex, both former subgenera of Camponotus, are elevated to genus level (stat. rev.), and two former genera, Forelophilus and Phasmomyrmex, are demoted to subgenus status (stat. n. and stat. rev., respectively) under Camponotus; two erstwhile subgenera of Phasmomyrmex, Myrmorhachis and Myrmacantha, become junior synonyms (syn. n.) of Camponotus (Phasmomyrmex); and the Camponotus subgenus Myrmogonia becomes a junior synonym (syn. n.) of Colobopsis. Dinomyrmex, represented by a single species from southeast Asia, D. gigas, is quite distinctive, but Camponotus and Colobopsis exhibit more subtle differences, despite being well separated phylogenetically. We identify morphological features of the worker caste that are broadly useful for distinguishing these two genera. Colobopsis species on the islands of New Caledonia and Fiji-regions with few native Camponotus species-tend to exceed these diagnostic bounds, but in this case regionally applicable character differences can be used to distinguish the two clades. Despite confusing similarities in the worker caste Colobopsis and Camponotus retain diagnostic differences in their larvae and pupae.

Background: Ultraconserved elements (UCEs) have been successfully used in phylogenomics for a variety of taxa, but their power in phylogenetic inference has yet to be extensively compared with that of traditional Sanger sequencing data sets. Moreover, UCE data on invertebrates, including insects, are sparse. We compared the phylogenetic informativeness of 959 UCE loci with a multi-locus data set of ten nuclear markers obtained via Sanger sequencing, testing the ability of these two types of data to resolve and date the evolutionary history of the second most species-rich subfamily of ants in the world, the Formicinae., Results: Phylogenetic analyses show that UCEs are superior in resolving ancient and shallow relationships in formicine ants, demonstrated by increased node support and a more resolved phylogeny. Phylogenetic informativeness metrics indicate a twofold improvement relative to the 10-gene data matrix generated from the identical set of taxa. We were able to significantly improve formicine classification based on our comprehensive UCE phylogeny. Our divergence age estimations, using both UCE and Sanger data, indicate that crown-group Formicinae are older (104-117 Ma) than previously suggested. Biogeographic analyses infer that the diversification of the subfamily has occurred on all continents with no particular hub of cladogenesis., Conclusions: We found UCEs to be far superior to the multi-locus data set in estimating formicine relationships. The early history of the clade remains uncertain due to ancient rapid divergence events that are unresolvable even with our genomic-scale data, although this might be largely an effect of several problematic taxa subtended by long branches. Our comparison of divergence ages from both Sanger and UCE data demonstrates the effectiveness of UCEs for dating analyses. This comparative study highlights both the promise and limitations of UCEs for insect phylogenomics, and will prove useful to the growing number of evolutionary biologists considering the transition from Sanger to next-generation sequencing approaches.

We investigated the species-level taxonomy of the Malagasy Crematogaster (Crematogaster) kelleri-group and an additional more distantly related species of the same subgenus. Morphological data from worker, queen and male ants, as well as genetic data from three nuclear genes (long wavelength rhodopsin, arginine kinase and carbomoylphosphate synthase) and one mitochondrial marker (cytochrome oxidase I) led to the recognition of six species. Within the C. kelleri-group, three new species are described: C. hazolava Blaimer sp. n., C. hafahafa Blaimer sp. n. and C. tavaratra Blaimer sp. n. The previously described taxa C. kelleri Forel and C. madagascariensis André are validated by our analysis. Conversely, our data suggests synonymy of C. adrepens Forel (with C. kelleri) and C. gibba Emery (with C. madagascariensis). A more distantly related and phylogenetically isolated species, C. tsisitsilo Blaimer sp. n., is further described. We report high levels of morphological and molecular variation in C. kelleri and illustrate that this variation can be explained partly by geography. Species descriptions, images, distribution maps and identification keys based on worker ants, as well as on queen and male ants where available, are presented for all six species. Our work highlights the elevated species richness of Crematogaster ants throughout Madagascar's humid forests, especially in the far northern tip of the island, and the need to use multiple data sources to ensure clear demarcation of this diversity.

This study unravels the evolution and biogeographic history of the globally distributed ant genus Crematogaster on the basis of a molecular phylogeny, reconstructed from five nuclear protein-coding genes and a total of 3384 bp of sequence data. A particular emphasis is placed on the evolutionary history of these ants in the Malagasy region. Bayesian and likelihood analyses performed on a dataset of 124 Crematogaster ingroup taxa lend strong support for three deeply diverging phylogenetic lineages within the genus: the Orthocrema clade, the Global Crematogaster clade and the Australo-Asian Crematogaster clade. The 15 previous subgenera within Crematogaster are mostly not monophyletic. Divergence dating analyses and ancestral range reconstructions suggest that Crematogaster evolved in South-East Asia in the mid-Eocene (40-45 ma). The three major lineages also originated in this region in the late Oligocene/early Miocene (~24-30 ma). A first dispersal out of S-E Asia by an Orthocrema lineage is supported for 22-30 ma to the Afrotropical region. Successive dispersal events out of S-E Asia began in the early, and continued throughout the late Miocene. The global distribution of Crematogaster was achieved by subsequent colonizations of all major biogeographic regions by the Orthocrema and the Global Crematogaster clade. Molecular dating estimates and ancestral range evolution are discussed in the light of palaeogeographic changes in the S-E Asian region and an evolving ocean circulation system throughout the Eocene, Oligocene and Miocene. Eight dispersal events to/from Madagascar by Crematogaster are supported, with most events occurring in the late Miocene to Pliocene (5.0-9.5 ma). These results suggest that Crematogaster ants possess exceptional dispersal and colonization abilities, and emphasize the need for detailed investigations of traits that have contributed to the global evolutionary success of these ants., (Copyright © 2012 Elsevier Inc. All rights reserved.)

The species-level taxonomy of the subgenus Crematogaster (Orthocrema) in the Malagasy region is evaluated with both morphological data from worker and queen ants, and genetic data from three nuclear markers (long wavelength rhodopsin, arginine kinase and carbamoylphosphate synthase). These two types of data support the existence of six Orthocrema species: Crematogaster madecassa Emery, Crematogaster rasoherinae Forel, Crematogaster telolafysp. n., Crematogaster razanasp. n., Crematogaster volamenasp. n. and Crematogaster mpanjonosp. n.. Two new synonyms of Crematogaster rasoherinae Forel are recognized, Crematogaster rasoherinae brunneola Emery, syn.n. and Crematogaster voeltzkowi Forel, syn. n., as these were not supported as distinct taxa by the data. A neotype is designated for Crematogaster rasoherinae; lectotypes are designated for Crematogaster madecassa, Crematogaster rasoherinae brunneola and Crematogaster voeltzkowi. Species descriptions, images, distribution maps and identification keys based on worker and queen ants are given for all six species. A diagnosis of the subgenus Orthocrema in the Malagasy region is presented for both workers and queens. Within the Malagasy Orthocrema, three distinct phylogenetic lineages are suggested by molecular and morphological data. Newly defined monophyletic species-groups are thus the Crematogaster madecassa-group (Crematogaster madecassa, Crematogaster telolafy and Crematogaster razana) and the Crematogaster volamena-group (Crematogaster volamena and Crematogaster mpanjono); Crematogaster rasoherinae represents an isolated lineage in the Malagasy region and its closest relatives remain unclear. Other interesting biological findings are the presence of an intermediate caste between workers and queens in Crematogaster rasoherinae and Crematogaster madecassa, and unusually large workers in Crematogaster volamena resembling a major caste.