Macroevolutionary integration of phenotypes within and across ant worker castes.

Phenotypic traits are often integrated into evolutionary modules: sets of organismal parts that evolve together. In social insect colonies, the concepts of integration and modularity apply to sets of traits both within and among functionally and phenotypically differentiated castes. On macroevolutionary timescales, patterns of integration and modularity within and across castes can be clues to the selective and ecological factors shaping their evolution and diversification. We develop a set of hypotheses describing contrasting patterns of worker integration and apply this framework in a broad (246 species) comparative analysis of major and minor worker evolution in the hyperdiverse ant genus Pheidole. Using geometric morphometrics in a phylogenetic framework, we inferred fast and tightly integrated evolution of mesosoma shape between major and minor workers, but slower and more independent evolution of head shape between the two worker castes. Thus, Pheidole workers are evolving as a mixture of intracaste and intercaste integration and rate heterogeneity. The decoupling of homologous traits across worker castes may represent an important process facilitating the rise of social complexity.

Colonize, radiate, decline: Unraveling the dynamics of island community assembly with Fijian trap-jaw ants.

The study of island community assembly has been fertile ground for developing and testing theoretical ideas in ecology and evolution. The ecoevolutionary trajectory of lineages after colonization has been a particular interest, as this is a key component of understanding community assembly. In this system, existing ideas, such as the taxon cycle, posit that lineages pass through a regular sequence of ecoevolutionary changes after colonization, with lineages shifting toward reduced dispersal ability, increased ecological specialization, and declines in abundance. However, these predictions have historically been difficult to test. Here, we integrate phylogenomics, population genomics, and X-ray microtomography/3D morphometrics, to test hypotheses for whether the ecomorphological diversity of trap-jaw ants (Strumigenys) in the Fijian archipelago is assembled primarily through colonization or postcolonization radiation, and whether species show ecological shifts toward niche specialization, toward upland habitats, and decline in abundance after colonization. We infer that most Fijian endemic Strumigenys evolved in situ from a single colonization and have diversified to fill a large fraction of global morphospace occupied by the genus. Within this adaptive radiation, lineages trend to different degrees toward high elevation, reduced dispersal ability, and demographic decline, and we find no evidence of repeated colonization that displaces the initial radiation. Overall these results are only partially consistent with taxon cycle and associated ideas, while highlighting the potential role of priority effects in assembling island communities.

Genomic and phenomic analysis of island ant community assembly.

Island biodiversity has long fascinated biologists as it typically presents tractable systems for unpicking the eco-evolutionary processes driving community assembly. In general, two recurring themes are of central theoretical interest. First, immigration, diversification, and extinction typically depend on island geographical properties (e.g., area, isolation, and age). Second, predictable ecological and evolutionary trajectories readily occur after colonization, such as the evolution of adaptive trait syndromes, trends toward specialization, adaptive radiation, and eventual ecological decline. Hypotheses such as the taxon cycle draw on several of these themes to posit particular constraints on colonization and subsequent eco-evolutionary dynamics. However, it has been challenging to examine these integrated dynamics with traditional methods. Here, we combine phylogenomics, population genomics and phenomics, to unravel community assembly dynamics among Pheidole (Hymenoptera, Formicidae) ants in the isolated Fijian archipelago. We uphold basic island biogeographic predictions that isolated islands accumulate diversity primarily through in situ evolution rather than dispersal, and population genomic support for taxon cycle predictions that endemic species have decreased dispersal ability and demography relative to regionally widespread taxa. However, rather than trending toward island syndromes, ecomorphological diversification in Fiji was intense, filling much of the genus-level global morphospace. Furthermore, while most endemic species exhibit demographic decline and reduced dispersal, we show that the archipelago is not an evolutionary dead-end. Rather, several endemic species show signatures of population and range expansion, including a successful colonization to the Cook islands. These results shed light on the processes shaping island biotas and refine our understanding of island biogeographic theory.

An ant genus-group (Prenolepis) illuminates the biogeography and drivers of insect diversification in the Indo-Pacific.

The Malay Archipelago and the tropical South Pacific (hereafter the Indo-Pacific region) are considered biodiversity hotspots, yet a general understanding of the origins and diversification of species-rich groups in the region remains elusive. We aimed to test hypotheses for the evolutionary processes driving insect species diversity in the Indo-Pacific using a higher-level and comprehensive phylogenetic hypothesis for an ant clade consisting of seven genera. We estimated divergence times and reconstructed the biogeographical history of ant species in the Prenolepis genus-group (Formicidae: Formicinae: Lasiini). We used a fossil-calibrated phylogeny to infer ancestral geographical ranges utilizing a biogeographic model that includes founder-event speciation. Ancestral state reconstructions of the ants' ecological preferences, and diversification rates were estimated for selected Indo-Pacific clades. Overall, we report that faunal interchange between Asia and Australia has occurred since at least 20-25 Ma, and early dispersal to the Fijian Basin happened during the early and mid-Miocene (ca. 10-20 Ma). Differences in diversification rates across Indo-Pacific clades may be related to ecological preference breadth, which in turn may have facilitated geographical range expansions. Ancient dispersal routes suggested by our results agree with the palaeogeography of the region. For this particular group of ants, the rapid orogenesis in New Guinea and possibly subsequent ecological shifts may have promoted their rapid diversification and widespread distribution across the Indo-Pacific.

Revision and Microtomography of the Pheidole knowlesi Group, an Endemic Ant Radiation in Fiji (Hymenoptera, Formicidae, Myrmicinae)Myrmicinae).

The Fijian islands, a remote archipelago in the southwestern Pacific, are home to a number of spectacular endemic radiations of plants and animals. Unlike most Pacific archipelagos, these evolutionary radiations extend to social insects, including ants. One of the most dramatic examples of ant radiation in Fiji has occurred in the hyperdiverse genus Pheidole. Most of the 17 native Fijian Pheidole belong to one of two species groups that descended from a single colonization, yet have evolved dramatically contrasting morphologies: the spinescent P. roosevelti species group, and the more morphologically conservative P. knowlesi species group. Here we revise the knowlesi group, in light of recent phylogenetic results, and enhanced with modern methods of X-ray microtomography. We recognize six species belonging to this group, including two of which we describe as new: Pheidole caldwelli Mann, Pheidole kava sp. n., Pheidole knowlesi Mann, P. ululevu sp. n., P. vatu Mann, and P. wilsoni Mann. Detailed measurements and descriptions, identification keys, and high-resolution images for queens, major and minor workers are provided. In addition, we include highly detailed 3D surface reconstructions for all available castes.

Inordinate Spinescence: Taxonomic Revision and Microtomography of the Pheidole cervicornis Species Group (Hymenoptera, Formicidae).

The ant genus Pheidole-for all of its hyperdiversity and global ubiquity-is remarkably conservative with regard to morphological disparity. A striking exception to this constrained morphology is the spinescent morphotype, which has evolved multiple times across distantly related lineages of Indoaustralian Pheidole. The Pheidole cervicornis group contains perhaps the most extraordinary spinescent forms of all Pheidole. Here we present a taxonomic revision of the P. cervicornis group, and use microtomographic scanning technology to investigate the internal anatomy of the thoracic spines. Our findings suggest the pronotal spines of Pheidole majors, are possibly skeletomuscular adaptations for supporting their disproportionately large heads. The 'head support hypothesis' is an alternative to the mechanical defense hypothesis most often used to explain spinescence in ants. The P. cervicornis group is known only from New Guinea and is represented by the following four species, including two described here as new: P. barumtaun Donisthorpe, P. drogon sp. nov., P. cervicornis Emery, and P. viserion sp. nov. The group is most readily identified by the minor worker caste, which has extremely long pronotal spines and strongly bifurcating propodeal spines. The major and minor workers of all species are illustrated with specimen photographs, with the exception of the major worker of P. cervicornis, which is not known.

Revision of the ant genus Proceratium Roger (Hymenoptera, Proceratiinae) in Fiji.

The Fiji archipelago harbours a surprisingly diverse and endemic ant fauna, despite its isolated and remote location in the South Pacific. The ant genus Proceratium is present on Fiji with three endemic species, of which Proceratiumoceanicum De Andrade, 2003 and Proceratiumrelictum Mann, 1921 were previously known. In this study we describe the third species: Proceratiumvinaka sp. n. All three species are members of the widespread and species-rich Proceratiumsilaceum clade. In order to integrate the new species into the current taxonomic system, we present an illustrated identification key to the worker caste of the three Fijian species. In addition, we provide a detailed description of Proceratiumvinaka, as well as species accounts for the other two species, which include diagnoses, taxonomic discussions, specimen photographs, and a distribution map.

Global phylogenetic structure of the hyperdiverse ant genus Pheidole reveals the repeated evolution of macroecological patterns.

Adaptive radiations are of particular interest owing to what they reveal about the ecological and evolutionary regulation of biodiversity. This applies to localized island radiations such as Darwin's finches, and also to rapid radiations occurring on a global scale. Here we analyse the macroevolution and macroecology of Pheidole, a famously hyperdiverse and ecologically dominant ant genus. We generate and analyse four novel datasets: (i) a robust global phylogeny including 285 Pheidole species, (ii) a global database on regional Pheidole richness in 365 political areas summarizing over 97 000 individual records from more than 6500 studies, (iii) a global database of Pheidole richness from 3796 local communities and (iv) a database of Pheidole body sizes across species. Analysis of the potential climate drivers of richness revealed that the patterns are statistically very similar across different biogeographic regions, with both regional and local richness associated with the same coefficients of temperature and precipitation. This similarity occurs even though phylogenetic analysis shows that Pheidole reached dominance in communities through serial localized radiations into different biomes within different continents and islands. Pheidole body size distributions have likewise converged across geographical regions. We propose these cases of convergence indicate that the global radiation of Pheidole is structured by deterministic factors regulating diversification and diversity.

Breaking out of biogeographical modules: range expansion and taxon cycles in the hyperdiverse ant genus Pheidole.

AIM: We sought to reconstruct the biogeographical structure and dynamics of a hyperdiverse ant genus, Pheidole, and to test several predictions of the taxon cycle hypothesis. Using large datasets on Pheidole geographical distributions and phylogeny, we (1) inferred patterns of biogeographical modularity (clusters of areas with similar faunal composition), (2) tested whether species in open habitats are more likely to be expanding their range beyond module boundaries, and (3) tested whether there is a bias of lineage flow from high- to low-diversity areas. LOCATION: The Old World. METHODS: We compiled and jointly analysed a comprehensive database of Pheidole geographical distributions, the ecological affinities of different species, and a multilocus phylogeny of the Old World radiation. We used network modularity methods to infer biogeographical structure in the genus and comparative methods to evaluate the hypotheses. RESULTS: The network analysis identified eight biogeographical modules, and a suite of species with anomalous ranges that are statistically more likely to occur in open habitat, supporting the hypothesis that open habitats promote range expansion. Phylogenetic analysis shows evidence for a cascade pattern of colonization from Asia to New Guinea to the Pacific, but no 'upstream' colonization in the reverse direction. MAIN CONCLUSIONS: The distributions of Pheidole lineages in the Old World are highly modular, with modules generally corresponding to biogeographical regions inferred in other groups of organisms. However, some lineages have expanded their ranges across module boundaries, and these species are more likely to be adapted to open habitats rather than interior forest. In addition, there is a cascade pattern of dispersal from higher to lower diversity areas during these range expansions. Our findings are consistent with the taxon cycle hypothesis, although they do not rule out alternative interpretations.

Molecular phylogeny of Indo-Pacific carpenter ants (Hymenoptera: Formicidae, Camponotus) reveals waves of dispersal and colonization from diverse source areas.

Ants that resemble Camponotus maculatus (Fabricius, 1782) present an opportunity to test the hypothesis that the origin of the Pacific island fauna was primarily New Guinea, the Philippines, and the Indo-Malay archipelago (collectively known as Malesia). We sequenced two mitochondrial and four nuclear markers from 146 specimens from Pacific islands, Australia, and Malesia. We also added 211 specimens representing a larger worldwide sample and performed a series of phylogenetic analyses and ancestral area reconstructions. Results indicate that the Pacific members of this group comprise several robust clades that have distinctly different biogeographical histories, and they suggest an important role for Australia as a source of Pacific colonizations. Malesian areas were recovered mostly in derived positions, and one lineage appears to be Neotropical. Phylogenetic hypotheses indicate that the orange, pan-Pacific form commonly identified as C. chloroticus Emery 1897 actually consists of two distantly related lineages. Also, the lineage on Hawai'i, which has been called C. variegatus (Smith, 1858), appears to be closely related to C. tortuganus Emery, 1895 in Florida and other lineages in the New World. In Micronesia and Polynesia the C. chloroticus-like species support predictions of the taxon-cycle hypothesis and could be candidates for human-mediated dispersal.

Introduced Pheidole of the world: taxonomy, biology and distribution.

The objective of this study is to provide a detailed taxonomic resource for identifying and studying ants in the genus Pheidole that have established beyond their native ranges. There is an increasing need for systematists to study taxa of specific concern to 21(st) century environmental, food security and public health challenges. Systematics has an important role to play in both the theoretical and applied disciplines of invasion biology. Few invaders impact terrestrial ecosystems more than ants. Among the world's 100 worst invasive species is the cosmopolitan and highly destructive Pheidole megacephala (Fabricius). Accurate identification of Pheidole megacephala is imperative for the success of screening, management and eradication programs designed to protect native ecosystems from the impacts of this destructive species. However, accurate identification of Pheidole species is difficult because of their taxonomic diversity, dimorphic worker caste and lack of taxonomic resources. Illustrated keys are included, along with the taxonomic history, taxonomic diagnoses, biological notes and risk statements for the 14 most invasive members of the genus. Global distribution maps based on over 14,000 specimen and literature records are presented for each species. These results of this work will facilitate identification of pest species, determination of climatic and habitat requirements, discovery of pest origins, horizon scanning and assessment of invasion pathways. The following new synonym is proposed, with the senior synonym listed first and the junior synonyms in parentheses: Pheidole indica Mayr (= Pheidole teneriffana Forel, and its synonyms Pheidole taina Aguayo and Pheidole voeltzkowii Forel). Pheidole navigans Forel, stat. rev., stat. n. is removed from synonymy and elevated to species rank. It is proposed that records of Pheidole moerens Forel outside of the Mesoamerica and the Caribbean refer instead to Pheidole navigans or other heterospecific taxa in the Pheidole flavens species complex. We propose that the names Pheidole anastasii Emery and Pheidole floridana Emery have been widely misapplied to North American outdoor records of Pheidole bilimeki Mayr. It is suggested that the synonymy of Pheidole lauta Wheeler be transferred from Pheidole floridana Emery to Pheidole bilimeki Mayr.

Pristomyrmex tsujii sp. n. and P. mandibularis Mann (Hymenoptera, Formicidae) from Fiji.

Pristomyrmex tsujii sp. n., an endemic species of the Fiji islands, is described from the worker, ergatoid queen, alate queen and male castes. The alate queen and male castes of Pristomyrmex mandibularis Mann are also described for the first time. The ergatoid queens for both species appear to be morphologically intermediate between the worker and alate queen castes. Pristomyrmex tsujii is readily distinguished from Pristomyrmex mandibularis by the lack of well-developed propodeal spines. Although both species occur across the Fijian archipelago, they are rarely encountered and workers are most often collected from sifted litter. The descriptions are illustrated with specimen photographs, line drawings and a distribution map.

Checklist of the ants (Hymenoptera, Formicidae) of the Solomon Islands and a new survey of Makira Island.

The intent of this paper is to facilitate future research of the Solomon Islands ant fauna by providing the first comprehensively researched species inventory in over 75 years. The species list presented here includes the names of all ant species recorded from the islands that are available in the literature together with specimen records from several museum collections and new records from our 2008 Makira field expedition. All the names of described species presented are valid in accordance with the most recent Formicidae classification. In total, the checklist is composed of 237 species and subspecies (including 30 morphospecies) in 59 genera representing nine subfamilies. We report that the recent field expedition added 67 new species records to Makira and 28 new species records to the Solomon Islands. Our research recovered species occurrence records for 32 individual islands and five island groups. The five islands with the highest number of recorded species are: Makira (142 spp.), Guadalcanal (107 spp.), Malaita (70 spp.), Santa Isabel (68 spp.), and Rennell (66 spp.). Based on our results, we discuss the taxonomic composition of the archipelago's ant fauna, which islands are most in need of additional sampling, and the importance of establishing biodiversity baselines before environmental threats such as the invasive ant Wasmannia auropunctata cause irrevocable harm to the native biodiversity.

Revisiting the ants of Melanesia and the taxon cycle: historical and human-mediated invasions of a tropical archipelago.

Understanding the historical evolution of biotas and the dynamics of contemporary human-mediated species introductions are two central tasks of biology. One hypothesis may address both-the taxon cycle. Taxon cycles are phases of range expansion and contraction coupled to ecological and evolutionary niche shifts. These historical invasion processes resemble human-mediated invasions in pattern and possibly mechanism, but both the existence of historical cycles and the roles of recent introductions are in question. We return to the system that originally inspired the taxon cycle-Melanesian ants-and perform novel tests of the hypothesis. We analyze (i) the habitat distributions of Fiji's entire ant fauna (183 species), (ii) ecological shifts associated with the in situ radiation of Fijian Pheidole in a phylogenetic context, and (iii) the ecological structure of a massive exotic ant invasion of the archipelago. Our analyses indicate lineages shift toward primary habitats, higher elevation, rarity, and ecological specialization with increasing level of endemism, consistent with taxon cycle predictions. The marginal habitats that historically formed a dispersal conduit in the Pacific are now mostly replaced by human-modified habitats dominated by a colonization pulse of exotic species. We propose this may represent the first phase of an incipient global cycle of human-mediated colonization, ecological shifts, and diversification.

Biogeography and morphological evolution in a Pacific island ant radiation.

While insular radiations are documented for many terrestrial arthropods, few examples are known for eusocial insects. This study seeks to ascertain whether the spinescence observed among Fijian Pheidole ants was inherited from an oversea ancestor or is evidence of ecological release from interspecific competitors. We broaden our understanding of morphological convergence, insular radiation and Pacific biogeography by testing three hypotheses proposed previously for the Fijian Pheidole roosevelti group: (i) the group is monophyletic; (ii) spinescence is a plesiomorphic trait inherited from an overseas ancestor; and (iii) the group is closely related to spinescent New Guinean relatives. The analysis included the fragments of two mitochondrial genes (COI, cytb) and two nuclear genes (H3, EF1α-F2) from 66 taxa, including all members of the roosevelti group, representatives from the spinescent subgenus Pheidolacanthinus, Fijian congeners and widespread Pacific congeners. Our results yield new insights into the biogeographic history of Fiji, reveal a fascinating example of convergent evolution and serve as a novel example of ecological release occurring within an insular eusocial insect lineage. These findings recover the history of a presumably unremarkable ant species that colonized a remote oceanic archipelago in the Miocene (17-10 Ma) and radiated across the emerging islands into niche-space occupied elsewhere in the Pacific by distantly related spinescent congeners. We propose the radiation of Fijian Pheidole into spinescent morphotypes was the consequence of ecological opportunities afforded by the absence of competing ant lineages with conspicuous epigaeic foraging strategies.