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The discovery in the Netherlands in a shipping container of the ant Monomorium sahlbergi Emery, 1898, a species similar to the invasive pharaoh ant M. pharaonis (Linnaeus, 1758), led to a quest to better define the distribution of this species, which was initially obscure due to uncertain specimen identifications. Here it is shown that M. sahlbergi, like M. pharaonis, is found worldwide, almost certainly as a result of introductions. Including quarantine interceptions, this species is recorded from seven global biogeographic regions, but its established outdoor distribution is currently limited to the tropics and subtropics. Monomorium dichroum Forel, 1902 is here presented as a junior synonym of M. sahlbergisyn. nov. based on morphometric and CO1 analyses.

Species delimitation offered by DNA-based approaches can provide important insights into the natural history and diversity of species, but the cogency of such processes is limited without multigene phylogenies. Recent attempts to barcode various Solenopsidini ant taxa (Hymenoptera: Formicidae: Myrmicinae), including the thief ant Solenopsis saudiensis Sharaf & Aldawood, 2011 described from the Kingdom of Saudi Arabia (KSA), were precipitated by the unexpected existence of a closely related species, the Nearctic S. abdita Thompson, 1989 within the S. molesta species complex native to Florida. This finding left the species status of the former uncertain. Here, we investigated the taxonomy and phylogeny of these two species to determine whether or not S. abdita represents a new global tramp species. We inferred a phylogeny of the two species using DNA sequence data from four nuclear genes (Abd-A, EF1α-F1, EF1α-F2, and Wingless) and one mitochondrial gene (COI) sampled from populations in Florida, Guatemala, Hawaii, and Saudi Arabia. Both species clustered into one distinct and robust clade. The taxonomy of S. saudiensis was re‐examined using morphometrics. A reassessment of the morphological characters used to diagnose the worker and queen castes were consistent with molecular evidence. Based on combined morphological and molecular evidences S. saudiensis is declared as a junior synonym of S. abdita (syn. nov.). In addition, our findings indicate that S. abdita is a novel global tramp species which has a far wider distribution than previously thought and has established itself in many new habitats and different geographic realms.

The present study represents a taxonomic revision of the Pheidole sikorae species group from Madagascar. Forty-four members of this group are recognised and described, and an illustrated identification key to this group is also presented. One species is raised to species level: P. litigiosa Forel, 1892 stat. nov.Pheidole veteratrix angustinoda Forel, 1892 syn. nov. is proposed as a junior synonym of Pheidole veteratrix Forel, 1891. Worker castes are also described and lectotypes designated for P. litigiosa Forel, 1892, P. sikorae Forel, 1891, and P. veteratrix Forel, 1891. The following 41 new species are described: P. alinasp. nov., P. ambohimangasp. nov., P. analavelonasp. nov., P. andohahelasp. nov., P. anomalasp. nov., P. anosyennesp. nov., P. antranohofasp. nov., P. beankasp. nov., P. befotakasp. nov., P. dasossp. nov., P. flavominutasp. nov., P. gracilissp. nov., P. habokasp. nov., P. havoanasp. nov., P. hazosp. nov., P. itremosp. nov., P. joffrevillesp. nov., P. kelysp. nov., P. lavasoasp. nov., P. mahamavosp. nov., P. maintysp. nov., P. mamiratrasp. nov., P. mananteninasp. nov., P. masoandrosp. nov., P. mavohavoanasp. nov., P. midongysp. nov., P. mikrossp. nov., P. mivorysp. nov., P. nitidobrunasp. nov., P. parvulasp. nov., P. parvulogibbasp. nov., P. reniranosp. nov., P. savasp. nov., P. sofiasp. nov., P. sparsasp. nov., P. tamponysp. nov., P. trichotossp. nov., P. tsaravonianasp. nov., P. vadumsp. nov., P. volontanysp. nov., and P. vonysp. nov. At present, there are 109 valid species and subspecies of Pheidole known from Madagascar, but this number is expected to increase with upcoming taxonomic revisions of the species groups not revised in this study.

Termites are major decomposers of organic matter in terrestrial ecosystems and the second most diverse lineage of social insects. The Kalotermitidae, the second-largest termite family, are widely distributed across tropical and subtropical ecosystems, where they typically live in small colonies confined to single wood items inhabited by individuals with no foraging abilities. How the Kalotermitidae have acquired their global distribution patterns remains unresolved. Similarly, it is unclear whether foraging is ancestral to Kalotermitidae or was secondarily acquired in a few species. These questions can be addressed in a phylogenetic framework. We inferred time-calibrated phylogenetic trees of Kalotermitidae using mitochondrial genomes and nuclear ribosomal RNA genes of ∼120 species, about 27% of kalotermitid diversity, including representatives of 22 of the 23 kalotermitid genera. We found that extant kalotermitids shared a common ancestor 81 Mya (72–91 Mya 95% HPD), indicating that a few disjunctions among early-diverging kalotermitid lineages may predate Gondwana breakup. However, most of the ∼40 disjunctions among biogeographic realms were dated at less than 50 Mya, indicating that transoceanic dispersals, and more recently human-mediated dispersals, have been the major drivers of the global distribution of Kalotermitidae. Our phylogeny also revealed that the capacity to forage is often found in early-diverging kalotermitid lineages, implying that the ancestors of Kalotermitidae were able to forage among multiple wood pieces. Our phylogenetic estimates provide a platform for a critical taxonomic revision of the family and for future comparative analyses of Kalotermitidae.

Forests, and the vertebrate species within them, are irreplaceable sources of food and nutrition for millions of people living in areas of high biodiversity. Unfortunately, many of these forests are being cleared for agriculture, and many animals are threatened with extinction from unsustainable hunting. Forest clearing and the hunting of threatened species are untenable solutions to long‐term food insecurity and adequate nutrition, jeopardizing these species' survival, the healthy functioning of ecosystems, and the cultural identities of local people. Working with communities to develop culturally appropriate ways for people to obtain sustainable and legal sources of food from forests outside of protected areas is a key component of improving both conservation and food security. We tested the feasibility, suitability, and viability of farming an abundant and traditionally eaten forest insect, Zanna tenebrosa (locally known as sakondry), in rural communities whose food security relies heavily on the hunting of threatened vertebrates. We found that the insect is high in macro‐ and micronutrients, and can be cheaply, easily, and sustainably cultivated to sustainably diversify forest food systems without increasing habitat loss. Given the range of Z. tenebrosa, which covers a broad swath of central Africa, increasing production of this native insect may support multipronged agroecological approaches to promoting food security, adequate nutrition, and wildlife conservation.

The present study represents an introduction to the revision of Pheidole Westwood, 1839 from Madagascar. Sixteen species groups are established, of which eleven are revised below, and illustrated identification keys to species groups and species of groups revised in this monograph are presented. Two species are raised to species level: Pheidole petax Forel, 1895 stat. nov., and P. scabrata Forel, 1895 stat. nov. We also redescribe worker castes and designate lectotypes for P. annemariae Forel, 1918, P. nemoralis Forel, 1892, P. petax Forel, 1895, P. ensifera Forel, 1897, P. longispinosa Forel, 1891, and P. scabrata Forel, 1895. The following 46 new species are described: Pheidole aelloeasp. nov., P. alasp. nov., P. andapasp. nov., P. ankeranasp. nov., P. avaratrasp. nov., P. bemarahaensissp. nov., P. bemarivoensissp. nov., P. binarasp. nov., P. boriborasp. nov., P. brevipilosasp. nov., P. curvistriatasp. nov., P. diakritossp. nov., P. ehazoarasp. nov., P. ferrugineasp. nov., P. fisakasp. nov., P. fitaratasp. nov., P. glabrasp. nov., P. goavanasp. nov., P. lamperossp. nov., P. longipilosasp. nov., P. luteasp. nov., P. madinikasp. nov., P. mahaboensissp. nov., P. maizinasp. nov., P. makaensissp. nov., P. makirovanasp. nov., P. manantenensissp. nov., P. mantadiasp. nov., P. marieannaesp. nov., P. masoalasp. nov., P. mavesatrasp. nov., P. miramilasp. nov., P. moramanaensissp. nov., P. navoatrensissp. nov., P. ocypodeasp. nov., P. parvioculasp. nov., P. podargeasp. nov., P. praegrandissp. nov., P. ranohirensissp. nov., P. rugocephalasp. nov., P. rugofitaratasp. nov., P. typhlossp. nov., P. vatovavensissp. nov., P. voasarasp. nov., P. vohemarensissp. nov., and P. zavamanirasp. nov. At present, there are 69 valid species and subspecies of Pheidole known from Madagascar, but this number is expected to increase significantly with upcoming taxonomic revisions of the species groups not revised in this study.

Ants, an ecologically successful and numerically dominant group of animals, play key ecological roles as soil engineers, predators, nutrient recyclers, and regulators of plant growth and reproduction in most terrestrial ecosystems. Further, ants are widely used as bioindicators of the ecological impact of land use. We gathered information of ant species in the Atlantic Forest of South America. The ATLANTIC ANTS data set-which is part of the ATLANTIC SERIES data papers-is a compilation of ant records from collections (18,713 records), unpublished data (29,651 records), and published sources (106,910 records; 1059 references), including papers, theses, dissertations, and book chapters published from 1886 to 2020. In total, the data set contains 153,818 ant records from 7,636 study locations in the Atlantic Forest, representing 10 subfamilies, 99 genera, 1,114 ant species identified with updated taxonomic certainty, and 2,235 morphospecies codes. Our data set reflects the heterogeneity in ant records, which include ants sampled at the beginning of the taxonomic history of myrmecology (the nineteenth and twentieth centuries) and more recent ant surveys designed to address specific questions in ecology and biology. The data set can be used by researchers to develop strategies to deal with different macroecological and regional-wide questions, focusing on assemblages, species occurrences and distribution patterns. Furthermore, the data can be used to assess the consequences of changes in land use in the Atlantic Forest on different ecological processes. No copyright restrictions apply to the use of this data set, but we request that authors cite this data paper when using these data in publications or teaching events.

Background Madagascar is famous for its extremely rich biodiversity; the island harbors predominantly endemic and threatened communities meriting special attention from biodiversity scientists. Continuing ongoing efforts to inventory the Malagasy ant fauna, we revise the species currently placed in the myrmicine genus Aphaenogaster Mayr. One species described from Madagascar, Aphaenogaster friederichsi Forel, is synonymized with the Palearctic A. subterranea Latreille syn. nov. This species is considered neither native to Madagascar nor established in the region. This revision focuses on the balance of species in the A. swammerdami group which are all endemic to Madagascar. Methods The diversity of the Malagasy Aphaenogaster fauna was assessed via application of multiple lines of evidence involving quantitative morphometric, qualitative morphological, and DNA sequence data. (1) Morphometric investigation was based on hypothesis-free Nest Centroid clustering (NC-clustering) combined with PArtitioning based on Recursive Thresholding (PART) to estimate the number of morphological clusters and determine the most probable boundaries between them. This protocol provides a repeatable and testable approach to find patterns in continuous morphometric data. Species boundaries and the reliability of morphological clusters recognized by these exploratory analyses were tested via confirmatory Linear Discriminant Analysis (LDA). (2) Qualitative, external morphological characteristics (e.g., shape, coloration patterns, setae number) were subjectively evaluated in order to create a priori grouping hypotheses, and confirm and improve species delimitation. (3) Species delimitation analyses based on mitochondrial DNA sequences from cytochrome oxidase I (COI) gene fragments were carried out to test the putative species previously delimited by morphological and morphometric analyses. Results Five species can be inferred based on the integrated evaluation of multiple lines of evidence; of these, three are new to science: Aphaenogaster bressleri sp. n., A. gonacantha (Emery, 1899), A. makay sp. n., A. sahafina sp. n., and A. swammerdami Forel, 1886. In addition, three new synonymies were found for A. swammerdami Forel, 1886 (A. swammerdami clara Santschi, 1915 syn. n., A. swammerdami curta Forel, 1891 syn. n. and A. swammerdami spinipes Santschi, 1911 syn. n.). Descriptions and redefinitions for each taxon and an identification key for their worker castes using qualitative traits and morphometric data are given. Geographic maps depicting species distributions and biological information regarding nesting habits for the species are also provided.

Evolutionary innovations underlie the rise of diversity and complexity—the 2 long-term trends in the history of life. How does natural selection redesign multiple interacting parts to achieve a new emergent function? We investigated the evolution of a biomechanical innovation, the latch-spring mechanism of trap-jaw ants, to address 2 outstanding evolutionary problems: how form and function change in a system during the evolution of new complex traits, and whether such innovations and the diversity they beget are repeatable in time and space. Using a new phylogenetic reconstruction of 470 species, and X-ray microtomography and high-speed videography of representative taxa, we found the trap-jaw mechanism evolved independently 7 to 10 times in a single ant genus (Strumigenys), resulting in the repeated evolution of diverse forms on different continents. The trap mechanism facilitates a 6 to 7 order of magnitude greater mandible acceleration relative to simpler ancestors, currently the fastest recorded acceleration of a resettable animal movement. We found that most morphological diversification occurred after evolution of latch-spring mechanisms, which evolved via minor realignments of mouthpart structures. This finding, whereby incremental changes in form lead to a change of function, followed by large morphological reorganization around the new function, provides a model for understanding the evolution of complex biomechanical traits, as well as insights into why such innovations often happen repeatedly., The mousetrap-like mandibles of trap-jaw ants are a functional innovation capable of record-breaking acceleration. This study uses phylogenomics, X-ray microtomography and high-speed videography to reveal that they evolved multiple times around the world through a pathway of incremental steps.

Madagascar, one of the top megadiversity regions, hosts one of the highest numbers of endemic and threatened organisms on earth. One of the most spectacular examples of ant radiation on the island has occurred in the hyperdiverse genusPheidole. To this date, there are 117 described MadagascanPheidoledivided into 16 species-groups, and 97% of them are endemic to the island. Only two of these species-groups contain widely distributed invasive species in addition to native, endemic taxa:megacephala, andfervensspecies-groups. Here we revise thefervensspecies-group and discuss updated distribution records of its introduced members on Madagascar. We recognize six species belonging to this group, including five new to science:Pheidole ampangabesp. nov.,P.arivosp. nov.,P.comosasp. nov.,P.indicaMayr,P.mamirapiratrasp. nov., andP.menasp. nov. Detailed descriptions are supplemented with measurements, diagnoses, identification key, high-resolution images for major and minor worker, and comments on distribution and biology.

The ant fauna of the State of Qatar is reviewed based on collected material and literature. In total, 23 species, belonging to 12 genera and four subfamilies, were reported from the country. Given that Messor arenarius diabolus Santschi, 1938 is endemic in Egypt, and it was recorded by mistake in Qatar, we excluded it from the country faunal list. Six genera are recorded for the first time from the country, Cardiocondyla Emery, 1896, Crematogaster Lund, 1831, Lepisiota Santschi, 1926, Monomorium Mayr, 1855, Paratrechina Motschoulsky, 1863, and Pheidole Westwood, 1839. Fourteen species are recorded for the first time from Qatar, namely, Camponotus oasium Forel, 1890, Cardiocondyla emeryiForel, 1881, Cataglyphis livida (Andre, 1881), Cataglyphis arenariaFinzi, 1940, Cataglyphis nigra (Andre, 1881), Lepisiota bipartita (Smith, 1861), Lepisiota gracilicornis (Forel, 1892), Monomorium abeillei Andre, 1881, Monomorium areniphilumSantschi, 1911, Monomorium subopacum (Smith, 1858), Monomorium venustum (Smith, 1858), Pheidole sinaiticaMayr, 1862. Tapinoma melanocephalum (Fabricius, 1793), and Tapinoma simrothiKrausse, 1911. Preliminary analysis of the zoogeography of Qatar's ants reveals the remarkable predominance of Palearctic faunal elements (13 species/ ∼57%), followed by minor proportions of Afrotropical (4 species/ ∼17%) and Indomalayan (1 species/ ∼4%) species, while the five invasive species (∼22%) reflect human impacts on the country. Monomorium tumaireCollingwood & Agosti, 1996 is known as endemic in the Arabian Peninsula. No species are considered endemic to Qatar. Ecological and biological notes on species are provided when available.

The present study represents a taxonomic revision of the P. bessonii species-group from Madagascar. Eighteen members of this group are recognized and described, and an illustrated identification key to this group is also presented. One name is raised to species level: P. decollata Forel, 1892 stat. nov. We also redescribe worker castes and designate lectotypes for P. bessonii Forel, 1891, P. decollata Forel, 1892, P. grallatrix Emery, 1899, P. madecassa Forel, 1892, and P. oswaldi 1891. The following 13 new species are described: Pheidole antsahabe sp. nov., Pheidole atsirakambiaty sp. nov., Pheidole clara sp. nov., Pheidole flammea sp. nov., Pheidole flavodepressa sp. nov., Pheidole mantadioflava sp. nov., Pheidole maro sp. nov., Pheidole ovalinoda sp. nov., Pheidole similis sp. nov., Pheidole tenebrovulgaris sp. nov., Pheidole uranus sp. nov., Pheidole voreios sp. nov., Pheidole zirafy sp. nov.

The genusCarebarais revised for the Malagasy region, and based on the examination of over 10,000 specimens, twenty-three species are recognized. Twenty one of these are described as new (C.barasp. n.,C.berivelosp. n.,C.betsisp. n.,C.creoleisp. n.,C.demetersp. n.,C.dotasp. n.,C.haintenysp. n.,C.hiragasysp. n.,C.jajobysp. n.,C.kabosysp. n.,C.lovasp. n.,C.mahafalysp. n.,C.malagasysp. n.,C.omasisp. n.,C.placidasp. n.,C.raberisp. n.,C.salegisp. n.,C.sampisp. n.,C.tanasp. n.,C.tananasp. n.,C.vazimbasp. n.), and two are redescribed,C.grandidieriForel (=C.voeltzkowiForeln. syn.) andC.nosindamboForel. A lectotype is designated forC.nosindambo. C.creoleisp. n.is known only from Mauritius and Seychelles,C.grandidieriForel is distributed in Comoros, Madagascar and Mayotte, and the other twenty-one species are endemic to Madagascar. Most of theCarebaraspecies recorded in this work are endemic to a specific habitat (ecoregion), but some of them (C.barasp. n.,C.grandidieriForel,C.jajobysp. n.,C.kabosysp. n., andC.nosindamboForel) are widespread within Madagascar across all major habitats. The worker caste ofCarebaracan be differentiated from other genera in the Myrmicinae subfamily by the presence of the following combination of characters: antennae of eight to eleven segments, with a two-segmented club; anterior clypeal margin without central isolated seta (rarely present in some species or specimens), and usually with four distinct setae; mandibles with four to seven teeth (except in one species from Ghana -C.crigensiswith three teeth); and palp formula 2,2 or 1,2. We report that almost allCarebaraspecies found in the Malagasy region have intermediates (distinct forms) in the major worker subcaste, with the largest major workers showing remnants of queen flight sclerites and ocelli. The widespread presence of intermediates in the major worker subcaste expands the morphological boundaries ofCarebara. We present an overview of the natural history ofCarebarain the Malagasy region, an illustrated key for the identification of the known Malagasy species ofCarebara, as well as high-resolution images and distribution maps. Unique identifiers are used for all specimens studied, including type material, and the raw data that forms the basis of this study are available on www.antweb.org (open access).

Ants evolved about 140 million years ago and have diversified into more than 15,000 species with tremendous ecological and morphological diversity, yet evolution of the gene regulatory networks (GRNs) underlying this diversification remains poorly understood. Wing polyphenism, the ability of a single genome to produce either winged or wingless castes during development in response to environmental cues, is a nearly universal feature of ants. The underlying wing GRN is evolutionarily labile in worker castes of phylogenetically derived species: it is conserved in winged castes but interrupted at different points in wingless castes of different species. However, it remains unknown whether the wing GRN is interrupted in wingless castes of species from early branching lineages, and if so, whether it is interrupted at similar locations in worker castes of derived species. We therefore used in situ hybridization to assay the expression of nine genes in the wing GRN in wing imaginal discs of larvae from two species from the early branching ('basal') genus Mystrium. These species possess two castes each: Mystrium rogeri has winged queens and wingless workers, and M. oberthueri has wingless queens and wingless workers. In contrast to derived species, we found no evidence of interruption points in the wing GRN kernel of wingless castes. Our finding supports: (1) a "phylogenetic ladder model" of wing GRN evolution, where interruption points move further upstream in the wing GRN as ant lineages become more derived; and (2) that evolutionary lability of the GRN underlying wing polyphenism originated later during ant evolution.

The ant genus Camponotus (Mayr, 1861) is one of the most abundant and species rich ant genera in the Malagasy zoogeographical region. Although this group is commonly encountered, its taxonomy is far from complete. Here, we clarify the taxonomy of the Malagasy-endemic Camponotus subgenus Myrmopytia (Emery, 1920). Species delimitation was based on traditional morphological characters and multivariate morphometric analyses, including exploratory Nest Centroid clustering and confirmatory cross-validated Linear Discriminant Analysis. Four species are recognized: Camponotus imitator (Forel, 1891), Camponotus jodina sp. n., Camponotus karaha sp. n., and Camponotus longicollis sp. n. All four species appear to mimic co-occurring Aphaenogaster species. A diagnosis of the subgenus Myrmopytia, species descriptions, an identification key based on minor and major subcastes of workers, and the known geographical distribution of each species are provided.

Madagascar has a long history of using Orthoptera as food and feed. Our understanding of the biological diversity of this resource, its contemporary use, and its future potentials in Madagascar is extremely limited. The present study contributes basic knowledge of the biological diversity and local uses of edible Orthoptera in Malagasy food cultures. Data was collected with key informants in 47 localities covering most of the ecoregions of Madagascar and corresponding to 12 of the 19 ethnic groups. Orthoptera are consumed throughout Madagascar. We report 37 edible Orthoptera species, of which 28 are new species records of edible Orthoptera in Madagascar and 24 are new species records of edible Orthoptera in the world. Most species are endemic and occur in farming zones. Children are the primary collectors and consumers of edible Orthoptera. The insects are eaten both as snacks and main meals. Edible Orthoptera are primarily collected casually and marketing is rare, with the notable exceptions of the large cricket Brachytrupes membranaceus colosseus and during locust outbreaks (e.g., Locusta migratoria). The use of Orthoptera as feed seems rare. Further investigations of cultural and personal preferences are required to assess the future potential roles of Orthoptera in Malagasy food habits.

Here we reveal the diversity of the next fragment of the Malagasy elements of the ant genus Nesomyrmex using a combination of advanced exploratory analyses on quantitative morphological data. The diversity of the Nesomyrmex madecassus species-group was assessed via hypothesis-free nest centroid clustering combined with recursive partitioning to estimate the number of clusters and determine the most probable boundaries between them. This combination of methods provides a highly automated species delineation protocol based on continuous morphometric data, and thereby it obviates the need of subjective interpretation of morphological patterns. Delimitations of clusters recognized by these exploratory analyses were tested via confirmatory Linear Discriminant Analysis (LDA). Our results suggest the existence of four morphologically distinct species, Nesomyrmex flavus sp. n., Nesomyrmex gibber, Nesomyrmex madecassus and Nesomyrmex nitidus sp. n.; all are described here and an identification key for their worker castes using morphometric data is given. Two members of the newly outlined madecasus species-group, Nesomyrmex flavus sp. n. and Nesomyrmex nitidus sp. n., represent true cryptic species. Geographic maps depicting species distributions and elevational information for the sites where populations of particular species were collected are also provided.

The ants in the subfamily Amblyoponinae are an old, relictual group with an unusual suite of morphological and behavioural features. Adult workers pierce the integument of their larvae to imbibe haemolymph, earning them the vernacular name ‘dracula ants’. We investigate the phylogeny of this group with a data set based on 54 ingroup taxa, 23 outgroups and 11 nuclear gene fragments (7.4 kb). We find that the genus Opamyrma has been misplaced in this subfamily: it is a member of the leptanilline clade and sister to all other extant Leptanillinae. Transfer of Opamyrma to Leptanillinae renders the Amblyoponinae monophyletic. The enigmatic Afrotropical genus Apomyrma is sister to all other amblyoponines, and the latter cleave into two distinct and well-supported clades, here termed POA and XMMAS. The POA clade, containing Prionopelta, Onychomyrmex and Amblyopone, is well resolved internally, and its structure supports synonymy of the genus Concoctio under Prionopelta (syn.n.). The XMMAS clade comprises two well-supported groups: (i) a predominantly Neotropical clade, for which we resurrect the genus name Fulakora (stat.r., stat.n.), with junior synonyms Paraprionopelta (syn.n.) and Ericapelta (syn.n.); and (ii) the remaining taxa, or ‘core XMMAS’, which are manifested in our study as a poorly resolved bush of about a dozen lineages, suggesting rapid radiation at the time of their origin. Most of these XMMAS lineages have been assigned to the catch-all genus Stigmatomma, but the more distinctive elements have been treated as separate genera (Xymmer, Mystrium, Myopopone and Adetomyrma). Resolution of basal relationships in the core XMMAS clade and reconfiguration of ‘Stigmatomma’ to restore monophyly of all named genera will require more extensive genetic data and additional morphological analysis. However, the genus Bannapone can be synonymized under Stigmatomma (syn.n.) because it is embedded within a clade that contains S. denticulatum, the type species of Stigmatomma. Divergence dating analysis indicates that crown Amblyoponinae arose in the mid-Cretaceous, about 107 Ma (95% highest probability density: 93–121 Ma). The POA and XMMAS clades have estimated crown ages of 47 and 73 Ma, respectively. The initial burst of diversification in the core XMMAS clade occurred in the Late Paleocene/Early Eocene (50–60 Ma). Ancestral range reconstruction suggests that amblyoponines originated in the Afrotropics, and dispersed to the Indo-Malayan region and to the New World. During none of these dispersal events did the ants break out of their cryptobiotic lifestyle.

Most ponerine ants have flying queens that monopolise sexual reproduction, but in a minority of species, workers can also mate and lay fertilised eggs. Such ‘gamergates' reproduce in addition to queens in some species but have replaced queens entirely in other species. The occurrence of a functional spermatheca in workers often appears associated with a slight difference in body size relative to winged queens, as is the case in Euponera sikorae (Forel) studied here. Eight colonies (19.9±8.7 workers and 0–2 dealate queens) were collected in humid forests of Madagascar. A mated queen reproduced in most colonies, but ovarian dissections indicated that 1–3 workers were also inseminated, and one of these laid eggs if the original founding queen was missing or no longer fecund. Exoskeleton remains (including 29 heads) of a staphylinid beetle were found in one nest of E. sikorae, pointing to specialised predation. We review the occurrence of non-flying reproductives (gamergates and ergatoid queens) in Odontomach...

The Malagasy Camponotus edmondi species group is revised based on both qualitative morphological traits and multivariate analysis of continuous morphometric data. To minimize the effect of the scaling properties of diverse traits due to worker caste polymorphism, and to achieve the desired near-linearity of data, morphometric analyses were done only on minor workers. The majority of traits exhibit broken scaling on head size, dividing Camponotus workers into two discrete subcastes, minors and majors. This broken scaling prevents the application of algorithms that uses linear combination of data to the entire dataset, hence only minor workers were analyzed statistically. The elimination of major workers resulted in linearity and the data meet required assumptions. However, morphometric ratios for the subsets of minor and major workers were used in species descriptions and redefinitions. Prior species hypotheses and the goodness of clusters were tested on raw data by confirmatory linear discriminant analysis. Due to the small sample size available for some species, a factor known to reduce statistical reliability, hypotheses generated by exploratory analyses were tested with extreme care and species delimitations were inferred via the combined evidence of both qualitative (morphology and biology) and quantitative data. Altogether, fifteen species are recognized, of which 11 are new to science: Camponotus alamaina sp. n., Camponotus androy sp. n., Camponotus bevohitra sp. n., Camponotus galoko sp. n., Camponotus matsilo sp. n., Camponotus mifaka sp. n., Camponotus orombe sp. n., Camponotus tafo sp. n., Camponotus tratra sp. n., Camponotus varatra sp. n., and Camponotus zavo sp. n. Four species are redescribed: Camponotus echinoploides Forel, Camponotus edmondi André, Camponotus ethicus Forel, and Camponotus robustus Roger. Camponotus edmondi ernesti Forel, syn. n. is synonymized under Camponotus edmondi. This revision also includes an identification key to species for both minor and major castes, information on geographic distribution and biology, taxonomic discussions, and descriptions of intraspecific variation. Traditional taxonomy and multivariate morphometric analysis are independent sources of information which, in combination, allow more precise species delimitation. Moreover, quantitative characters included in identification keys improve accuracy of determination in difficult cases.

Summary A basic expectation of evolution by natural selection is that species morphologies will adapt to their ecological niche. In social organisms, this may include selective pressure from the social environment. Many non-ant parasites of ant colonies are known to mimic the morphology of their host species, often in striking fashion [ 1 , 2 ], indicating there is selection on parasite morphology to match the host (Batesian and/or Wasmannian mimicry [ 3 ]). However, ants that parasitize other ant societies are usually closely related to their hosts (Emery’s rule) [ 4 , 5 , 6 , 7 , 8 ] and expected to be similar due to common ancestry, making any kind of mimicry difficult to detect [ 9 ]. Here, we investigate the diversification of the hyperdiverse ant genus Pheidole in Madagascar, including the evolution of 13 putative social parasite species within a broader radiation of over 100 ant species on the island. We find that the parasitic species are monophyletic and that their associated hosts are spread across the Malagasy Pheidole radiation. This provides an opportunity to test for selection on morphological similarity and divergence between parasites and hosts. Using X-ray microtomography and both linear measurements and three-dimensional (3D) geometric morphometrics, we show that ant social parasite worker morphologies feature a mix of “host-matching” and “parasitic” traits, where the former converge on the host phenotype and the latter diverge from typical Pheidole phenotypes to match a common parasitic syndrome. This finding highlights the role of social context in shaping the evolution of phenotypes and raises questions about the role of morphological sensing in nestmate recognition.

This open-access special issue features 12 full articles representing emerging trends from the international DNAbarcoding community. Several articles highlight how DNA-based techniques are elucidating the species diversity,biogeography, and conservation status of Africas biodiversity. Another prominent theme is the movementtowards big biodiversity data using high-throughput, individual-based DNA barcoding methods, which preservevoucher specimens and abundance data, as well as bulk sample-based metabarcoding. Methodological developments are enhancing the detection of specific species and whole communities using environmental DNA(eDNA) barcoding and metabarcoding. Data are also expanding in terms of genetic coverage; in this issue, a newdatabase is established for a secondary fungalDNAbarcode marker, and multi-kingdom, multi-marker biodiversitysurveys are gaining traction. DNA barcode sequence data, often combined with complementary markers or taxonomic information, are increasingly contributing to large-scale phylogenetic projects, with implications for understanding evolutionary history, community structure, and conservation priorities. Fil: Adamowicz, Sarah J.. University of Guelph; Canadá Fil: Boatwright, James S.. University of The Western Cape; Sudáfrica Fil: Chain, Frédéric. University of Massachusetts; Estados Unidos Fil: Fisher, Brian L.. California Academy Of Sciences.; Estados Unidos Fil: Hogg, Ian D.. Polar Knowledge Canada; Canadá Fil: Leese, Florian. Universitat Essen; Alemania Fil: Lijtmaer, Dario Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Mwale, Monica. South African National Biodiversity Institute; Sudáfrica Fil: Naaum, Amanda M.. The Queens University of Belfast; Irlanda Fil: Pochon, Xavier. University of Auckland; Nueva Zelanda Fil: Schubert, Dirk W.. University of Guelph; Canadá Fil: Wilson, John James. National Museums Liverpool; Reino Unido Fil: Wood, Susanna. Cawthron Institute; Nueva Zelanda Fil: Xu, Jianping. Mcmaster University; Canadá Fil: Xu, Sen. University of Texas at Arlington; Estados Unidos Fil: Zhou, Xin. China Agricultural University; China Fil: Van Der Bank, Michelle. University of Johannesburg; Sudáfrica

Blaimer, Bonnie B., Ward, Philip S., Schultz, Ted R., Fisher, Brian L., Brady, Seán G. (2018): Paleotropical Diversification Dominates the Evolution of the Hyperdiverse Ant Tribe Crematogastrini (Hymenoptera: Formicidae). Insect Systematics and Diversity (AIFB) 2 (5), No. 3: 1-14, DOI: 10.1093/isd/ixy013, URL: http://dx.doi.org/10.1093/isd/ixy013

The Camponotus subgenus Mayria is revised based on the analysis of both qualitative morphological characters and morphometric traits. The multivariate analysis combined the Nest Centroid (NC)-clustering method and Partitioning Algorithm based on Recursive Thresholding (PART) function to generate species hypotheses based on 19 continuous morphological traits of minor workers. The proposed species hypotheses were confirmed by cumulative Linear Discriminant Analysis (LDA). Morphometric ratios for the subsets of minor and major workers were used in species descriptions and redefinitions. In this study, 14 species are recognized, of which seven are newly described: C. lamosy sp. n., C. liandia sp. n., C. mainty sp. n., C. manabo sp. n., C. raina sp. n., C. sada sp. n., C. tanosy sp. n. Four species are redescribed: C. christi Forel, C. dromedarius Forel, C. lubbocki Forel, and C. repens Forel. The following are raised to species and redescribed: C. foersteri Forel stat. n., C. maculiventris Emery stat. n., and C. pulcher stat. n. The following are synonymized under C. foersteri: Camponotus cambouei Forel, syn. n.; C. christi ambustus Forel, syn. n.; C. christi ferrugineus Emery, syn. n. and C. pictipes Forel, syn. n. The following are synonymized under C. lubbocki: Camponotus lubbocki christoides Forel, syn. n. and C. lubbocki rectus Forel, syn. n. Also included in this revision are an illustrated species identification key, taxonomic discussions, images, and distribution maps for each species superimposed with the ecoregions of Madagascar.

In this study we revise the taxonomy of the genus Prionopelta for the Malagasy region, treating seven species, six of which are newly described (Prionopelta laurae sp. n., Prionopelta seychelles sp. n., Prionopelta subtilis sp. n., Prionopelta talos sp. n., Prionopelta vampira sp. n., Prionopelta xerosilva sp. n.), and one redescribed (Prionopelta descarpentriesi Santschi). One species, Prionopelta seychelles, is restricted to Seychelles, while the six remaining species treated are endemic to Madagascar.

Ant communities in tropical forests may be governed by varying assembly mechanisms, depending on the particular habitat investigated. We compared phylogenetic diversity and structure across two forest biomes (dry and humid) and two vertical layers (arboreal and terricolous) in ant communities in Madagascar, and assessed the infl uence of invasive species on this community structure. We estimated phylogenetic signal and correlated evolution for habitat and several functional traits and tested for conservatism in relevant functional and habitat traits. Ancestral states were reconstructed to illuminate the evolution of habitat traits. All analyses utilized phylogenies estimated from newly generated data from three nuclear markers for 290 Malagasy ant taxa. Dry forests, although lower in species richness, were found to support equally high lineage diversity as humid forests. In contrast, phylogenetic diversity was much lower in arboreal than in terricolous communities. We observed signifi cant phylogenetic clustering in the combined humid forest and in the arboreal – humid, arboreal – dry and terricolous – humid communities, whereas the combined dry forest community was overdispersed. Among ant communities in Madagascar, overdispersion and competition therefore may be more prevalent in dry forest, and habitat fi ltering may be more dominant in humid forest. Excluding invasive ant species had little overall eff ect on community structure. All investigated traits showed low to intermediate conservatism; strong support for correlated evolution was found for increased eye size and an arboreal lifestyle. Habitat transitions from humid to dry and from terricolous to arboreal occurred more frequently, and ancestors of most lineages were predicted to be terricolous or humid-forest adapted. We conclude that most Malagasy ant clades fi rst colonized humid forests and subsequently transitioned into dry forests, indicating that previous hypotheses on the evolution of Madagascar ’ s hyperdiverse biota may not apply to ants and other arthropods.

Knowledge of the internal phylogeny and evolutionary history of ants (Formicidae), the world’s most species-rich clade of eusocial organisms, has dramatically improved since the advent of molecular phylogenetics. A number of relationships at the subfamily level, however, remain uncertain. Key unresolved issues include placement of the root of the ant tree of life and the relationships among the so-called poneroid subfamilies. Here we assemble a new data set to attempt a resolution of these two problems and carry out divergence dating, focusing on the age of the root node of crown Formicidae. For the phylogenetic analyses we included data from 110 ant species, including the key species Martialis heureka. We focused taxon sampling on non-formicoid lineages of ants to gain insight about deep nodes in the ant phylogeny. For divergence dating we retained a subset of 62 extant taxa and 42 fossils in order to approximate diversified sampling in the context of the fossilized birth-death process. We sequenced 11 nuclear gene fragments for a total of ~7.5 kb and investigated the DNA sequence data for the presence of among-taxon compositional heterogeneity, a property known to mislead phylogenetic inference, and for its potential to affect the rooting of the ant phylogeny. We found sequences of the Leptanillinae and several outgroup taxa to be rich in adenine and thymine (51% average AT content) compared to the remaining ants (45% average). To investigate whether this heterogeneity could bias phylogenetic inference we performed outgroup removal experiments, analysis of compositionally homogeneous sites, and a simulation study. We found that compositional heterogeneity indeed appears to affect the placement of the root of the ant tree but has limited impact on more recent nodes. We put forward a novel hypothesis regarding the rooting of the ant phylogeny, in which Martialis and the Leptanillinae together constitute a clade that is sister to all other ants. After correcting for compositional heterogeneity this emerges as the best-supported hypothesis of relationships at deep nodes in the ant tree. The results of our divergence dating under the fossilized birth-death process and diversified sampling suggest that the crown Formicidae originated during the Albian or Aptian ages of the Lower Cretaceous (103–124 Ma). In addition, we found support for monophyletic poneroids comprising the subfamilies Agroecomyrmecinae, Amblyoponinae, Apomyrminae, Paraponerinae, Ponerinae, and Proceratiinae, and well-supported relationships among these subfamilies except for the placement of Proceratiinae and (Amblyoponinae + Apomyrminae). Our phylogeny also highlights the non-monophyly of several ant genera, including Protanilla and Leptanilla in the Leptanillinae, Proceratium in the Proceratiinae, and Cryptopone, Euponera, and Mesoponera within the Ponerinae.

Knowledge of the internal phylogeny and evolutionary history of ants (Formicidae), the world’s most species-rich clade of eusocial organisms, has dramatically improved since the advent of molecular phylogenetics. A number of relationships at the subfamily level, however, remain uncertain. Key unresolved issues include placement of the root of the ant tree of life and the relationships among the so-called poneroid subfamilies. Here we assemble a new data set to attempt a resolution of these two problems and carry out divergence dating, focusing on the age of the root node of crown Formicidae. For the phylogenetic analyses we included data from 110 ant species, including the key species Martialis heureka. We focused taxon sampling on non-formicoid lineages of ants to gain insight about deep nodes in the ant phylogeny. For divergence dating we retained a subset of 62 extant taxa and 42 fossils in order to approximate diversified sampling in the context of the fossilized birth-death process. We sequenced 11 nuclear gene fragments for a total of ∼ 7.5 kb and investigated the DNA sequence data for the presence of among-taxon compositional heterogeneity, a property known to mislead phylogenetic inference, and for its potential to affect the rooting of the ant phylogeny. We found sequences of the Leptanillinae and several outgroup taxa to be rich in adenine and thymine (51% average AT content) compared to the remaining ants (45% average). To investigate whether this heterogeneity could bias phylogenetic inference we performed outgroup removal experiments, analysis of compositionally homogeneous sites, and a simulation study. We found that compositional heterogeneity indeed appears to affect the placement of the root of the ant tree but has limited impact on more recent nodes. Our findings have implications for outgroup choice in phylogenetics, which should be made not only on the basis of close relationship to the ingroup, but should also take into account sequence divergence and other properties relative to the ingroup. We put forward a hypothesis regarding the rooting of the ant phylogeny, in which Martialis and the Leptanillinae together constitute a clade that is sister to all other ants. After correcting for compositional heterogeneity this emerges as the best-supported hypothesis of relationships at deep nodes in the ant tree. The results of our divergence dating under the fossilized birth-death process and diversified sampling suggest that the crown Formicidae originated during the Albian or Aptian ages of the Lower Cretaceous (103–124 Ma). In addition, we found support for monophyletic poneroids comprising the subfamilies Agroecomyrmecinae, Amblyoponinae, Apomyrminae, Paraponerinae, Ponerinae, and Proceratiinae, and well-supported relationships among these subfamilies except for the placement of Proceratiinae and (Amblyoponinae + Apomyrminae). Our phylogeny also highlights the non-monophyly of several ant genera, including Protanilla and Leptanilla in the Leptanillinae, Proceratium in the Proceratiinae, and Cryptopone, Euponera, and Mesoponera within the Ponerinae.

Mutualisms between ants and sap-sucking insects generally involve clear benefits for both partners: the ants provide protection in exchange for honeydew. However, a single ant genus associates with armoured scale insects (Diaspididae) that do not excrete honeydew. We studied three colonies ofMelissotarsus emeryiants from two localities in Mozambique. Vast numbers of the diaspididMorganella conspicuaoccupied galleries dug by the ants under the bark of living trees. Unlike free-livingM. conspicuaand other diaspidids,M. conspicualiving with ants are known to lack shields, likely because they gain protection against enemies and desiccation. Nevertheless, we documented the occurrence of rare individuals with shields inside ant galleries, indicating that their glands continue to secrete wax and proteins as building material. This is likely to constitute a significant portion of the ants’ diet, in addition to diaspidid exuviae and excretions from the Malpighian tubules. Indeed,Melissotarsusworkers cannot walk outside the galleries due to modified middle legs, forcing them to obtain all nourishment within the tree.Melissotarsusfounding queens, however, must locate a suitable host tree while flying, and acquire diaspidid crawlers. This mutualism involves ants that are highly specialised to chew through living wood, and diaspidids that can also live freely outside the bark. It is extremely widespread in Africa and Madagascar, recorded from 20 tree families, and harmful effects on plant hosts require rapid study.

The Malagasy endemic ant Malagidris sofina (Bolton and Fisher 2014) nests on cliff faces in natural rock alcoves or clay banks. Colonies have single ergatoid queens and reproduce by fission. Each nest has a funnel-shaped entrance that projects horizontally from the cliff face. We examine three hypotheses for the function of the funnels—water exclusion, gas exchange and defense. Entrance funnels are relatively impermeable and divert water from nests, but simple tubes would achieve the same result. Consistent with the gas exchange hypothesis, projected funnel entrances likely increase gas exchange rates over sixfold compared to simple tubes and may increase air flow within the nest. Gas exchange may explain the recurrent evolution of funnel entrances in several ant lineages, especially among cliff dwelling species. We outline M. sofina defense responses to conspecifics and co-occurring ant species, and find no support for a defense role of entrance funnels. Workers display little aggression but respond to several species with an original form of nest defense––cliff jumping—in which workers drop off the cliff face while clinging to invaders and then return to their nest. M. sofina is a restricted range species under threat of extinction by habitat destruction. Its novel lifestyle underscores the urgency of exploration and conservation in a tropical biodiversity hotspot.

This study investigates the evolutionary history of a hyperdiverse clade, the ant subfamily Myrmicinae (Hymenoptera: Formicidae), based on analyses of a data matrix comprising 251 species and 11 nuclear gene fragments. Under both maximum likelihood and Bayesian methods of inference, we recover a robust phylogeny that reveals six major clades of Myrmicinae, here treated as newly defined tribes and occur- ring as a pectinate series: Myrmicini, Pogonomyrmecini trib.n., Stenammini, Solenop- sidini, Attini and Crematogastrini. Because we condense the former 25 myrmicine tribes into a new six-tribe scheme, membership in some tribes is now notably different, espe- cially regarding Attini. We demonstrate that the monotypic genus Ankylomyrma is nei- ther in the Myrmicinae nor even a member of the more inclusive formicoid clade - rather it is a poneroid ant, sister to the genus Tatuidris (Agroecomyrmecinae). Several species-rich myrmicine genera are shown to be nonmonophyletic, including Pogono- myrmex, Aphaenogaster, Messor, Monomorium, Pheidole, Temnothorax andTetramor- ium. We propose a number of generic synonymies to partially alleviate these problems (senior synonym listed first): Pheidole = Anisopheidole syn.n. = Machomyrma syn.n.; Temnothorax = Chalepoxenus syn.n. = Myrmoxenus syn.n. = Protomognathus syn.n.; Tetramorium = Rhoptromyrmex syn.n. = Anergates syn.n. = Teleutomyrmex syn.n. The genus Veromessor stat.r. is resurrected for the New World species previously placed in Messor; Syllophopsis stat.r. is resurrected from synonymy under Monomorium to contain the species in the hildebrandti group;Trichomyrmex stat.r. is resurrected from synonymy underMonomorium to contain the species in the scabriceps- anddestruc- tor-groups; and the monotypic genus Epelysidris stat.r. is reinstated for Monomorium brocha. Bayesian divergence dating indicates that the crown group Myrmicinae origi- nated about 98.6 Ma (95% highest probability density 87.9-109.6 Ma) but the six major clades are considerably younger, with age estimates ranging from 52.3 to 71.1 Ma. Although these and other suprageneric taxa arose mostly in the middle Eocene or earlier, a number of prominent, species-rich genera, such as Pheidole, Cephalotes, Strumigenys, Crematogaster andTetramorium, have estimated crown group origins in the late Eocene or Oligocene. Most myrmicine species diversity resides in the two sister clades, Attini and Crematogastrini, which are estimated to have originated and diversified extensively in the Neotropics and Paleotropics, respectively. The newly circumscribed Myrmicini is Holarctic in distribution, and ancestral range estimation suggests a Nearctic origin. The Pogonomyrmecini and Solenopsidini are reconstructed as being Neotropical in origin

Fisher and Robertson (Insect Soc 46: 78-83, 1999) discovered the production of silk-like secretions emerging from slit-shaped openings along the anterior margin of the ventral hypostoma of Melissotarsus ant workers. The current histological study describes a hitherto unknown hypostomal gland from which this silk-like substance originates. In addi- tion, this study describes a new basitarsal gland in the three pairs of legs of Melissotarsus workers.

Recent survey work in the Seychelles has revealed two new species of Paraparatrechina that are here described: P. illusio sp. n. and P. luminella sp. n. A revised key to the workers of Paraparatrechina for the Afrotropical and Malagasy regions is provided. The taxonomy of the hypogaeic weissi species-group is also reviewed in light of recent field collections. The species P. sordida is revived from synonymy and given new status (as a full species) and a discussion of the morphologically peculiar species-group is provided. With the description of the two species and the removal of another species from weissi synonomy there are now 16 Paraparatrechina species known from the Afrotropical and Malagasy regions.

The ant genus Paratrechina is reexamined based on the discovery of two new species from Madagascar (P. ankarana sp. n. and P. antsingy sp. n.). Paratrechina kohli, a species known from central Africa, is transferred to Paratrechina from Prenolepis based on a new morphological interpretation of the genus and an updated morphological diagnosis of the genus is provided. This means that other than the widespread P. longicornis, whose origins remain uncertain, all Paratrechina are restricted either to the Afrotropical or Malagasy regions. It would also appear that of the five Paratrechina species now known, three are from dry forest habitats. With this reexamination of the genus, the possible origins of P. longicornis are discussed. A key to the genera of the Prenolepis genus-group is provided, as is a key to the workers of Paratrechina. In addition, we also designate a lectotype for Paratrechina kohli.

In this study we synonymise the genus Decamorium Forel under Tetramorium Mayr, revise the new T. decem species group by providing a diagnosis of the group, an illustrated identification key to species level, and worker-based species descriptions for all five species, which include diagnoses, discussions, images, and distribution maps. The following species are revised in this study: T. decem Forel, comb. r., T. raptor sp. n., T. uelense Santschi, comb. r., T. ultor Forel, comb. r., stat. r. & stat. n., and T. venator sp. n. In addition, we also designate lectotypes for T. decem, T. uelense, and T. ultor.

Nest emigrations are perilous for social insect colonies. Outside their nests, adults and brood are exposed to dangers. The behavioural mechanisms of emigration are thus likely to be under strong selective pressures. 2. Most studies on emigration have focused on monogynous species where survival of the queen is paramount, but emigration processes are largely unknown for species having several queens per colony. 3. In colonies of Mystrium oberthueri Forel, members of the morphological queen caste are as numerous as workers, although only a few of them mate and reproduce (polygyny). All queens perform intranidal tasks, such as brood care. Accordingly, we expected them to participate actively in emigration and to be less well protected. 4. Using four colonies, we studied the dynamics of 16 emigrations with a special focus on individual behavioural profile. 5. Workers were more involved than non-reproductive queens in recruitment and brood transport. Reproductive queens and young ants preferentially walked directly to the new nest without carrying brood. A chemical trail was probably used. The physiological status of individuals had more impact on their behavioural profile than their morphological caste. 6. This highly organized emigration process may underpin dependent colony foundation, as both involve the coordinated movement of nestmates.

In this study we revise the taxonomy of the Tetramorium tortuosum species group members encountered in the Afrotropical region, which we have placed in its own subgroup: the T. capillosum species complex. We re-describe the two previously known species T. capillosum Bolton and T. tabarum Bolton, and describe the new species T. hecate sp. n. The geographic distribution of the three species appears to be restricted to the equatorial rainforests of Central Africa. We provide a diagnosis of the T. capillosum species complex, an illustrated identification key to species level, and worker-based species descriptions, which include diagnoses, discussions, high-quality montage images, and distribution maps. Furthermore, we discuss biogeography and composition of the globally distributed T. tortuosum group.

The species-level taxonomy of the ant genus Meranoplus F. Smith from Madagascar is revised. Two new species, M. cryptomys sp. n. and sylvarius sp. n. are described from workers and queens; M. mayri Forel, 1910, and M. radamae Forel, 1891, are redescribed, and queens and males for these two species are described for the first time. The first diagnosis of Meranoplus males for any biogeographic region is provided based on Malagasy species. Illustrated keys to all known Malagasy castes and species are presented. Diagnoses are given for two species groups: the M. mayri group and the M. nanus group. The diagnosis of the M. nanus species group from Bolton (1981) is thereby expanded with six new characters. Two species are known from the M. mayri species group and seven described species are known for the M. nanus species group, including the two new species described herein. The mouthparts, genitalia, and all castes, where known, of Malagasy Meranoplus are illustrated.

The animal fauna of the Socotra Archipelago is influenced by three biogeographical regions, the Afrotropical, the Oriental and the Palaearctic. Consequently, the Archipelago shares faunal elements of these three regions in addition to unique endemic taxa. The ant fauna of Socotra Island was studied and is reviewed based on literature and newly collected material. In total, 28 species, belonging to 10 genera and four subfamilies, were collected from the main island. Eighteen of these (64%) are successful invasive species, seven are native (25%), and three are considered endemic (11%), Cardiocondyla longiceps Seifert, Monomorium elghazalyi sp. nov. and Monomorium nimihil Collingwood et al. Two genera are recorded for the first time from the island, Hypoponera Santschi, and Syllophopsis Santschi. Ten species are recorded for the first time, Cardiocondyla mauritanica Forel, Cardiocondyla minutior Forel, Monomorium atomum Forel, Monomorium dichroum Forel, Monomorium exiguum Forel, Pheidole pallidula (N...

Defining species limits and describing species of ants are important to identify taxa and habitats with elevated diversity in areas of high conservation priority such as the Malagasy region. The Pachycondyla wasmannii-group is revised in the Malagasy region where eight species are recognized, four of which are new: P. masoala sp. n., P. planicornis sp. n., P. tavaratra sp. n., and P. vazimba sp. n. Four species have been previously described: P. cambouei Forel, P. comorensis (André), P. perroti Forel, and P. wasmannii Forel. Pachycondyla perroti admista Forel is newly synonymized under P. perroti. Pachycondyla cambouei is widespread in eastern Madagascar, morphologically variable, and divided into seven morphotypes. An identification key to species and distribution maps are provided for the genus in the Malagasy region. All species are known only from Madagascar except P. wasmannii, which occurs also on Anjouan of the Comoros Islands.

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Constanta, Romania Univ Mississippi, Oxford, MS USA Royal Belgian Inst Nat Sci, Brussels, Belgium Univ New Mexico, Albuquerque, NM 87131 USA Inst Nacl de Pesquisas da Amazonia, Manaus, Amazonas, Brazil Univ Stellenbosch, Matieland, South Africa Univ Fed Minas Gerais, Belo Horizonte, MG, Brazil CIBIO Ctr Invest Biodiversidade & Recursos Genet, Vairao, Portugal Museum Hist Nat, Geneva, Switzerland Univ Fed Mato Grosso, Cuiaba, Brazil Univ Austral Chile, Valdivia, Chile Univ Malaysia, Sarawak, Malaysia Indian Inst Sci, Bangalore, Karnataka, India Scottish Assoc Marine Sci, Oban, Argyll, Scotland Sam Houston State Univ, Huntsville, TX 77340 USA Museo Nacl Ciencias Nat, Madrid, Spain Sorbonne Univ, CR2P, Museum Natl Hist Nat, Paris, France Drexel Univ, Acad Nat Sci, Philadelphia, PA 19104 USA Univ Richmond, Richmond, VA 23173 USA Ctr Pesquisas Cacau, Itabuna, Brazil Univ Torino, Turin, Italy Soc Hist Nat Alcide dOrbigny, Aubiere, France Wolfden Sci Consulting, Murcia, Spain Univ Fed Rio de Janeiro, Macae, Brazil Univ Tennessee, Knoxville, TN USA Inst Humboldt, Villa De Leyva, Colombia Escuela Politec Nacl, Quito, Ecuador Calif Acad Sci, San Francisco, CA 94118 USA Museum Dept Hist Nat Var, Toulon, France Nat Environm & Wildlife Soc, Angul, India Univ Kansas, Lawrence, KS 66045 USA Kyoto Univ, Kyoto, Japan Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivadavia, Buenos Aires, DF, Argentina Univ Fed Paraiba, Joao Pessoa, Paraiba, Brazil Univ Fed Parana, Curitiba, Parana, Brazil Univ Fed Rio de Janeiro, Rio De Janeiro, Brazil Nat Hist Museum Narodini Museum, Prague, Czech Republic Nat Hist Museum Zimbabwe, Bulawayo, Zimbabwe Ateneo Manila Univ, Quezon City, Philippines Pontificia Univ Javeriana, Bogota, Colombia RACINE, St Maugan, France Univ Paris 06, Sorbonne Univ Paris, ISYEB, Paris, France Ocean Univ China, Qingdao, Peoples R China Museo Nacl Hist Nat Paraguay, San Lorenzo, Paraguay Univ Carabobo, Valencia, Venezuela Natl Ctr Biol Sci, Bengaluru, India Univ Nebraska, Lincoln, NE USA CENAK Ctr Nat Kunde, Hamburg, Germany Ohio Hist Connect, Columbus, OH USA Univ Fed Amazonas, Manaus, Amazonas, Brazil Univ Malaga, Malaga, Spain Chinese Acad Sci, Qingdao, Shandong, Peoples R China Benemerita Univ Autonoma Puebla, Puebla, Mexico Natl Polytech Inst, Ctr Res & Adv Studies, Irapuato, Mexico Field Museum Nat Hist, Chicago, IL 60605 USA Univ Texas El Paso, El Paso, TX 79968 USA Univ Fed Lavras, Lavras, Brazil Nat Hist Museum, Vienna, Austria La Sierra Univ, Riverside, CA USA Univ San Francisco Quito, Quito, Ecuador Univ Hong Kong, Hong Kong, Peoples R China CUNY, New York, NY 10021 USA CAS, Inst Vertebrate Biol, Brno, Czech Republic Sorbonne Univ, MECADEV, Museum Natl Hist Nat, Paris, France Univ Estadual Paulista, Rio Claro, Brazil Zool Staatssammlung Munchen, Munich, Germany Univ Michigan, Dearborn, MI 48128 USA Smithsonian Inst, Natl Museum Nat Hist, Washington, DC USA Helmholtz Ctr Environm Res, Leipzig, Germany Santa Rosa Jr Coll, Santa Rosa, CA USA George Washington Univ, Washington, DC 20037 USA Inst Teknol Bandung, Bandung, Indonesia Earlham Coll, Richmond, IN USA Senckenberg Forschungsinst & Nat Museum, Frankfurt, Germany Univ Estadual Londrina, Londrina, Parana, Brazil Univ Koblenz Landau, Koblenz, Germany Nat Hist Museum, London, England Ditsong Natl Museum Nat Hist, Pretoria, South Africa Bombay Nat Hist Soc, Bombay, Maharashtra, India Philipps Univ Marburg, Marburg, Germany Hess Landesmuseum, Darmstadt, Germany Smithsonian Conservat Biol Inst, Washington, DC USA Univ Pannonia, Keszthely, Hungary Staatliches Museum Nat Kunde, Stuttgart, Germany UFT, Tocantins, Portugal Museum Nat Kunde, Berlin, Germany Nat Hist Riksmuseet, Stockholm, Sweden Hikarigaoka, Munakata, Japan Inst Agr & Environm Sci, Tartu, Estonia Univ Los Andes, Merida, Venezuela Bat Jungle, Monteverde, Costa Rica Texas A&M Univ, College Stn, TX USA Yale Univ, New Haven, CT USA Univ Estadual Paulista, Sao Jose Do Rio Preto, Brazil Museo Nacl Hist Nat, Montevideo, Uruguay Consejo Nacl Invest Cient & Tecn, Fdn Miguel Lillo, San Miguel De Tucuman, Argentina Roger Tory Peterson Inst Nat Hist, Jamestown, VA USA Univ Fed Reconcavo Bahia, Cruz Das Almas, Brazil South China Normal Univ, Guangzhou 510631, Guangdong, Peoples R China Museu Ciencias Tecnol PUCRS, Porto Alegre, RS, Brazil Univ Fed Tocantins, Porto Nacl, Brazil Univ Toronto, Scarborough, ON, Canada Univ Nacl Colombia, Inst Ciencias Nat, Bogota, Colombia Natl Museum Philippines, Manila, Philippines NOVA Univ Lisbon, Caparica, Portugal Royal Saskatchewan Museum, Regina, SK, Canada Deutsch Zentrum Marine Biodiversitatsforsch, Hamburg, Germany Ctr Invest Biol Noroeste, La Paz, Mexico Naturalis Biodivers Ctr, Leiden, Netherlands Univ Douala, Douala, Cameroon Vet & Farmaceut Univ Brno, Brno, Czech Republic Univ Swaziland, Kwaluseni, Eswatini Fundacao Oswaldo Cruz, Rio De Janeiro, Brazil Univ Claude Bernard, Lyon, France Museum Vertebrate Zool, Berkeley, CA USA Smithsonian Inst, Washington, DC 20560 USA Nat Hist Museum & Inst, Chiba, Japan Senckenberg Biodiversitat & Klima Forschunsgzentr, Frankfurt, Germany Marquette Univ, Milwaukee, WI 53233 USA Univ Firenze, Museo Storia Nat, Florence, Italy Univ Fed Bahia, Salvador, BA, Brazil Fed Univ ABC, Sao Bernardo, Brazil Univ Fed Goias, Goiania, Go, Brazil Univ Alaska Museum, Fairbanks, AK USA Egerton Univ, Egerton, Kenya Museo Hist Nat, Lima, Peru IKIAM Univ Reg Amazon, Tena, Ecuador Univ Hyogo, Sanda, Japan Inst Rech Dev, Paris, France Univ Sao Paulo, Piracicaba, Brazil Niedersachs Landesbetrieb Wasserwirtschaft Kusten, Hannover, Germany Univ Complutense Madrid, Madrid, Spain Consejo Nacl Invest Cient & Tecn, Ctr Nacl Patagon, Puerto Madryn, Argentina Roosevelt Univ, Coll Profess Studies, Chicago, IL 60605 USA Western Kentucky Univ, Bowling Green, KY 42101 USA Univ Naples Federico II, Naples, Italy Univ Fed Acre, Rio Branco, Acre, Brazil Univ Fed Santa Catarina, Florianopolis, SC, Brazil Inst Bio & Geociencias Noroeste Argentino, Salta, Argentina Inst Venezolano Invest Cient, Caracas, Venezuela Umweltbundesamt, Vienna, Austria Penclen, Plumelec, France CNRS MNHN UPMC, Ctr Rech Paleobiodivers & Paleoenvironm, Paris, France Staatl Museum Nat Kunde, Stuttgart, Germany Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivada, Buenos Aires, DF, Argentina Ctr Biol Gest Populat INRA, Montferrier Sur Lez, France Loyola Univ Chicago, Chicago, IL USA Pontificia Univ Catolica Rio do Sul, Porto Alegre, RS, Brazil Durban Museum Nat Sci, Durban, South Africa Univ Estado Amazonas, Manaus, Amazonas, Brazil Brigham Young Univ, Provo, UT 84602 USA Museo Hist Nat La Salle, Caracas, Venezuela Univ Vet Med Hannover, Hannover, Germany Australian Museum, Sydney, NSW, Australia Texas Tech Univ, Lubbock, TX 79409 USA Francis Marion Univ, Florence, SC USA Univ Washington, Seattle, WA 98195 USA Univ Connecticut, Groton, CT USA Fed Univ ABC, Santo Andre, SP, Brazil Univ Fed Uberlandia, Uberlandia, MG, Brazil Fed Univ Para, Belem, Para, Brazil Yale Peabody Museum, New Haven, CT USA Univ Texas Arlington, Arlington, TX 76019 USA Senckenberg Deutsch Entomol Inst, Muncheberg, Germany CSIRO, Natl Res Collect, Canberra, ACT, Australia Univ Venda, Thohoyandou, South Africa Univ Ulsan, Ulsan, South Korea Univ Estadual Sao Paulo, Botucatu, SP, Brazil Senckenberg Forsch Inst & Nat Museum, Frankfurt, Germany Ocean Univ China, Inst Marine Biodivers & Evolut, Qingdao, Shandong, Peoples R China Univ Fed Para, Altamira, Brazil Univ Estadual Paulista, Assis, Brazil UPMC, Ctr Rech Paleobiodiversite & Paleoenvironm, CNRS, MNHN, Paris, France Tech Univ Kaiserslautern, Kaiserslautern, Germany Charles Univ Prague, Dept Zool, Prague, Czech Republic Univ Nacl Expt los Llanos Occident Ezequiel Zamor, Guanare, Venezuela Long Isl Univ, Brooklyn, NY USA Univ Ryukyus, Okinawa, Japan Pontificia Univ Catolica Ecuador, Museo Zool, Escuela Ciencias Biol, Quito, Ecuador Indiana Univ Penn, 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Evolutionary novelty can arise through the recombination of ancestral characters. In ants, colonies occasionally produce anomalous individuals that are mosaics of queen and worker characters. These intermediate individuals may be the first step in the evolution of new castes such as wingless queens and soldiers. While the morphological features of intercastes support this hypothesis, their rarity has meant their behaviour is unknown. In this study we carried out a behavioural study of 32 intercastes of Mystrium rogeri to assess whether they are a burden for their colonies, and whether their unusual morphologies are associated with consistent behavioural profiles and new functions. In addition, we measured dominance interactions within colonies, reproductive behaviours during dyadic encounters with males and aggressive interactions during dyadic encounters with heterospecific workers. We found that intercastes were active in colonies and that their behavioural profiles did not differ from those of virgin queens and workers. However, some intercastes were dominant, had developed ovaries and attracted males. One intercaste successfully mated and laid diploid eggs, demonstrating her reproductive potential. Moreover, we found that intercastes were as capable of fighting as workers because of their large size and could thus be useful for colony defence. Our results show that the production of intercastes in M. rogeri inflicts few costs but can yield benefits. Accordingly, certain ecological circumstances could select for increased frequency of intercaste production, ultimately resulting in the evolution of new reproductive and defensive castes.

The monotypic ant genus Xymmer Santschi, 1914 was established for X. muticus (Santschi, 1914) which has so far been known only from the West African subregion. The genus is easily distinguishable from the other amblyoponine genera by the anteromedian part of clypeus which is produced as a short rectangular lobe. Additional undescribed species had been found exclusively from Madagascar and Africa until recently. However, in March, 2015, a Xymmer colony was found in northern Central Vietnam. In the present paper Xymmer phungi sp. nov. is described as the first Oriental species of the genus. Although the worker of X. phungi is morphologically quite similar to that of X. muticus , the former is distinguishable from the latter by the following two characteristics of the worker: clypeal lobe 1/6 times as long as broad (vs. 1/3 times as long as broad in X. muticus ); anterior margin of the lobe weakly concave (vs. almost straight in X. muticus ). Xymmer phungi is well distinguished from Ethiopian Xymmer spp. for which 28S sequences are available from GenBank. Our observations suggest that the species feeds on geophilids (Chilopoda: Geophilomorpha). An updated key to Vietnamese genera of the subfamily Amblyoponinae is also provided.

Here we describe the diversity of the Malagasy Nesomyrmex brevicornis-group as the final installment of series describing the complete Malagasy Nesomyrmex fauna. In the current paper introduce the Nesomyrmex brevicornis-group, a newly outlined species group. We split this group from one of the four former groups, Nesomyrmex madecassus-group (sensu Csősz and Fisher 2015). The relatively small number of individuals available within the Nesomyrmex brevicornis-group relative to the large number of OTUs hampers the use of sophisticated statistical techniques to evaluate the morphological data. Instead, we assess the diversity of the Nesomyrmex brevicornis-group via a conventional morphology-based approach relying upon non-numeric characters (shape, sculpture, color etc.) recorded by simple eye inspection. Morphometric data are used as secondary data to confirm the most probable boundaries between species, but without employing statistical analyses. We rejected prior species hypotheses imposed by the primary approach if a non-overlapping range of a single index calculated from continuous numeric traits was found between two hypothesized species. Our methods render every species in this revisionary work separable via both salient features and morphometric traits. Our approach reveals the existence of nine morphologically distinct species, Nesomyrmex brevicornis sp. n., Nesomyrmex brunneus sp. n., Nesomyrmex cingulatus sp. n., Nesomyrmex edentates sp. n., Nesomyrmex flavigaster sp. n., Nesomyrmex longiceps sp. n., Nesomyrmex minutus sp. n., Nesomyrmex punctaticeps sp. n., and Nesomyrmex sellaris sp. n.. Geographic maps showing the distribution of each species are provided, along with an updated key to workers of Malagasy Nesomyrmex species groups that includes the newly outlined Nesomyrmex brevicornis-group.

Madagascar is one of the world’s greatest biodiversity hotspots, meriting special attention from biodiversity scientists. It is an excellent testing ground for novel techniques in taxonomy that aim to increase classification objectivity and yield greater taxonomic resolving power. Here we reveal the diversity of a unique and largely unexplored fragment of the Malagasy ant fauna using an advanced combination of exploratory analyses on quantitative morphological data allowing for increased objectivity in taxonomic workflow. The diversity of the Nesomyrmex sikorai species-group was assessed via hypothesis-free nest-centroid-clustering combined with recursive partitioning to estimate the number of morphological clusters and determine the most probable boundaries between them. This combination of methods provides a highly automated and objective species delineation protocol based on continuous morphometric data. Delimitations of clusters recognized by these exploratory analyses were tested via confirmatory Linear Discriminant Analysis (LDA) and Multivariate Ratio Analysis (MRA). The final species hypotheses are corroborated by many qualitative characters, and the recognized species exhibit different spatial distributions and occupy different ecological regions. We describe and redescribe eight morphologically distinct species including six new species: Nesomyrmex excelsior sp. n., N. modestus sp. n., N. reticulatus sp. n., N. retusispinosus (Forel, 1892), N. rugosus sp. n., N. sikorai (Emery, 1896), N. striatus sp. n., and N. tamatavensis sp. n. An identification key for their worker castes using morphometric data is provided.

Background.Applying quantitative morphological approaches in systematics research is a promising way to discover cryptic biological diversity. Information obtained through twenty-first century science poses new challenges to taxonomy by offering the possibility of increased objectivity in independent and automated hypothesis formation. In recent years a number of promising new algorithmic approaches have been developed to recognize morphological diversity among insects based on multivariate morphometric analyses. These algorithms objectively delimit components in the data by automatically assigning objects into clusters.Method.In this paper, hypotheses on the diversity of the MalagasyNesomyrmex angulatusgroup are formulated via a highly automated protocol involving a fusion of two algorithms, (1) Nest Centroid clustering (NC clustering) and (2) Partitioning Algorithm based on Recursive Thresholding (PART). Both algorithms assign samples into clusters, making the class assignment results of different algorithms readily inferable. The results were tested by confirmatory cross-validated Linear Discriminant Analysis (LOOCV-LDA).Results.Here we reveal the diversity of a unique and largely unexplored fragment of the Malagasy ant fauna using NC-PART-clustering on continuous morphological data, an approach that brings increased objectivity to taxonomy. We describe eight morphologically distinct species, including seven new species:Nesomyrmex angulatus(Mayr, 1862),N. bidentatussp. n.,N. clypeatussp. n.,N. deviussp. n.,N. exiguussp. n.,N. fragilissp. n.,N. gracilissp. n., andN. hirtellussp. n.. An identification key for their worker castes using morphometric data is provided.Conclusions.Combining the dimensionality reduction feature of NC clustering with the assignment of samples into clusters by PART advances the automatization of morphometry-based alpha taxonomy.

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

Five main conclusions arise from this review of the responses of species to anthropogenic disturbance in Madagascar: First, species’ reactions to anthropogenic disturbance are generally negative, but remain poorly known. Our knowledge is patchy among and within higher taxonomic groups; we are still largely gathering case studies. Second, taxonomic groups vary considerably in which proximate factors are most important. Third, several groups show differing responses within different ecoregions. Whether these differences are consistent across groups requires further testing. Fourth, related species often have divergent reactions to disturbance, even within lower taxonomic groupings (families or genera). Thus, we cannot rely on phylogenetic relatedness or even ecological similarity to infer similarity in responses. Finally, disturbance typically reduces species diversity (especially of native and/or endemic species), but also causes species turnover, typically with forest specialists replaced by grassland generalists, and endemics replaced by non-endemics (including invasives). Given these knowledge gaps, we stress the urgency of applied studies that assess species’ ecology, behaviour and health across disturbance gradients, including purely anthropogenic landscapes. Remaining natural vegetation and protected areas will be unable to preserve Madagascar’s biodiversity under the impact of climatic change; we must understand responses of plants and animals to disturbance in order to create buffer zones and corridors combining secondary, degraded and natural habitats.