A molecular phylogeny of <Emphasis Type="Italic">Dorylus</Emphasis> army ants provides evidence for multiple evolutionary transitions in foraging niche

Background  

Army ants are the prime arthropod predators in tropical forests, with huge colonies and an evolutionary derived nomadic life style. Five of the six recognized subgenera of Old World Dorylus army ants forage in the soil, whereas some species of the sixth subgenus (Anomma) forage in the leaf-litter and some as conspicuous swarm raiders on the forest floor and in the lower vegetation (the infamous driver ants). Here we use a combination of nuclear and mitochondrial DNA sequences to reconstruct the phylogeny of the Dorylus s.l. army ants and to infer the evolutionary transitions in foraging niche and associated morphological adaptations.     

Subterranean species of the ant genus Crematogaster in Asia (Hymenoptera: Formicidae)

Three Crematogaster ant species, C. (Orthocrema) javanica Menozzi, C. (O.) myops Forel and C. (O.) masukoi sp. nov., share reduced compound eyes which characterizes them among Asian species of the subgenus Orthocrema. The new species is described based on material from Borneo. It can be distinguished from C. javanica and C. myops by its smooth surface of clypeus and acutely produced subpetiolar process. Reduced compound eyes and yellowish body suggest that these three species are subterranean.     

Both female castes contribute to colony emigration in the polygynous ant Mystrium oberthueri

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From the Western Alps across Central Europe: Postglacial recolonisation of the tufa stream specialist Rhyacophila pubescens (Insecta,Trichoptera)

Background

Hybridization can have complex effects on evolutionary dynamics in ants because of the combination of haplodiploid sex-determination and eusociality. While hybrid non-reproductive workers have been found in a range of species, examples of gene-flow via hybrid queens and males are rare. We studied hybridization in East African army ants (Dorylus subgenus Anomma) using morphology, mitochondrial DNA sequences, and nuclear microsatellites.

Results

While the mitochondrial phylogeny had a strong geographic signal, different species were not recovered as monophyletic. At our main study site at Kakamega Forest, a mitochondrial haplotype was shared between a "Dorylus molestus-like" and a "Dorylus wilverthi-like" form. This pattern is best explained by introgression following hybridization between D. molestus and D. wilverthi. Microsatellite data from workers showed that the two morphological forms correspond to two distinct genetic clusters, with a significant proportion of individuals being classified as hybrids.

Conclusions

We conclude that hybridization and gene-flow between the two army ant species D. molestus and D. wilverthi has occurred, and that mating between the two forms continues to regularly produce hybrid workers. Hybridization is particularly surprising in army ants because workers have control over which males are allowed to mate with a young virgin queen inside the colony.     

Evolution of a soldier caste specialized to lay unfertilized eggs in the ant genus Crematogaster (subgenus Orthocrema)

Among social Hymenoptera, only some ant genera have more than one morphological kind of non-reproductive adults. Individuals that are bigger than ordinary workers can function for defence and/or food storage. In Crematogaster (Orthocrema) smithi from Arizona, a third caste exists in addition to winged queens and workers; it is intermediate in size, weight and morphology, and individuals lay many unfertilized eggs that are mostly eaten by larvae (Heinze et al., 1995, 1999). We studied another three species belonging to the subgenus Orthocrema: Crematogaster pygmaea from Brazil, Crematogaster biroi and Crematogaster schimmeri from Taiwan. Using scanning electron microscopy and ovarian dissections, we show that ‘intermediates’ are a patchwork of queen-like and worker-like traits, just as in C. smithi; importantly the combinations differ across species. ‘Intermediates’ are numerically few in the colonies, and in C. pygmaea they are produced seasonally. Using histology we confirmed the lack of a spermatheca, thus they are not ergatoid queens. Based on the similarity of their mosaic phenotypes with those in other ant lineages, we suggest that Orthocrema ‘intermediates’ are a soldier caste with a specialized trophic function. This soldier caste has been reported in other Orthocrema species from Madagascar, Guinea and Costa Rica, suggesting that it is widespread in this subgenus.     

Systematics and biogeography of the ant genus Crematogaster Lund subgenus Orthocrema Santschi in Asia (Hymenoptera: Formicidae)

The subgenus Orthocrema of the ant genus Crematogaster is a well defined group and diverse in the tropical Asia. Its systematics has remained poorly understood because of a lack of modern revisionary work. Crematogaster (Orthocrema) is revised for the Asian region, and 27 species including ten new species are recognized. Five species groups: the C. baduvi group (4 spp.); the C. binghamii group (3 spp.); the C. biroi group (10 spp.); the C. moatensis group (1 sp.); the C. quadriruga group (9 spp.) are established based on worker caste morphology. A key to Asian species of the subgenus Orthocrema based on the worker caste is given. Phylogenetic relationships of Asian Orthocrema are analyzed. The analysis revealed that the C. baduvi‐, C. binghamii‐, and C. biroi groups are monophyletic, and that the allopatric distribution patterns of closely related species imply that Asian Orthocrema is composed of relatively young taxa. There have been at least three west‐to‐east dispersal events across Wallace's line in the C. baduvi‐, C. quadriruga‐ and C. biroi groups. © 2016 The Linnean Society of London     

The generic and species relationships of the reputed endemic Malagasy genus Pseudocossyphus (family Turdidae)

The generic and specific relationships of the Malagasy rock-thrushes, variously placed in the genus Monticola or as an endemic genus Pseudocossyphus, are examined using morphological and molecular characters. On the basis of mitochondrial DNA cytochrome b sequence data it was concluded that all Malagasy rock-thrushes fall between Asian Monticola and a species group of Monticola occurring in numerous Old World areas. It is recommended that all Malagasy species be placed in the genus Monticola. Relationships between various named forms of Malagasy rock-thrushes indicate: M. imerinus is a distinct species, M. bensoni is a synonym of M. sharpei, the subspecies M. sharpei erythronotus should be elevated to a full species, and that no geographic forms of M. sharpei (including interioris and salomonseni) should be recognized on the basis of current evidence. These taxonomic conclusions have important implications for conservation projects currently underway on Madagascar.     

Global domination by crazy ants: phylogenomics reveals biogeographical history and invasive species relationships in the genus Nylanderia (Hymenoptera: Formicidae)

Nylanderia (Emery) is one of the world's most diverse ant genera, with 123 described species worldwide and hundreds more undescribed. Fifteen globetrotting or invasive species have widespread distributions and are often encountered outside their native ranges. A molecular approach to understanding the evolutionary history and to revision of Nylanderia taxonomy is needed because historical efforts based on morphology have proven insufficient to define major lineages and delimit species boundaries, especially where adventive species are concerned. To address these problems, we generated the first genus-wide genomic dataset of Nylanderia using ultraconserved elements (UCEs) to resolve the phylogeny of major lineages, determine the age and origin of the genus, and describe global biogeographical patterns. Sampling from seven biogeographical regions revealed a Southeast Asian origin of Nylanderia in the mid-Eocene and four distinct biogeographical clades in the Nearctic, the Neotropics, the Afrotropics/Malagasy region, and Australasia. The Nearctic and Neotropical clades are distantly related, indicating two separate dispersal events to the Americas between the late Oligocene and early Miocene. We also addressed the problem of misidentification that has characterized species-level taxonomy in Nylanderia as a result of limited morphological variation in the worker caste by evaluating the integrity of species boundaries in six of the most widespread Nylanderia species. We sampled across ranges of species in the N. bourbonica complex (N. bourbonica (Forel) + N. vaga (Forel)), the N. fulva complex (N. fulva (Mayr) + N. pubens (Forel)), and the N. guatemalensis complex (N. guatemalensis (Forel) + N. steinheili (Forel)) to clarify their phylogenetic placement. Deep splits within these complexes suggest that some species names – specifically N. bourbonica and N. guatemalensis – each are applied to multiple cryptic species. In exhaustively sampling Nylanderia diversity in the West Indies, a ‘hot spot’ for invasive taxa, we found five adventive species among 22 in the region; many remain morphologically indistinguishable from one another, despite being distantly related. We stress that overcoming the taxonomic impediment through the use of molecular phylogeny and revisionary study is essential for conservation and invasive species management.     

Congruence of Microsatellite and Mitochondrial DNA Variation in Acrobat Ants (Crematogaster Subgenus Decacrema,Formicidae: Myrmicinae) Inhabiting Macaranga (Euphorbiaceae) Myrmecophytes

A previously reported mitochondrial DNA (mtDNA) phylogeny of Crematogaster (subgenus Decacrema) ants inhabiting Macaranga myrmecophytes indicated that the partners diversified synchronously and their specific association has been maintained for 20 million years. However, the mtDNA clades did not exactly match morphological species, probably owing to introgressive hybridization among younger species. In this study, we determined the congruence between nuclear simple sequence repeat (SSR, also called microsatellite) genotyping and mtDNA phylogeny to confirm the suitability of the mtDNA phylogeny for inferring the evolutionary history of Decacrema ants. Analyses of ant samples from Lambir Hills National park, northeastern Borneo, showed overall congruence between the SSR and mtDNA groupings, indicating that mtDNA markers are useful for delimiting species, at least at the local level. We also found overall high host-plant specificity of the SSR genotypes of Decacrema ants, consistent with the specificity based on the mtDNA phylogeny. Further, we detected cryptic genetic assemblages exhibiting high specificity toward particular plant species within a single mtDNA clade. This finding, which may be evidence for rapid ecological and genetic differentiation following a host shift, is a new insight into the previously suggested long-term codiversification of Decacrema ants and Macaranga plants.     

Incipient morphological castes in <Emphasis Type="Italic">Polistes gallicus</Emphasis> (Vespidae,Hymenoptera)

The occurrence of a pre-imaginal caste determination represents a sort of “point of no return” to eusociality. In some social insect taxa, including Polistes species, the occurrence of distinct queen and worker castes is still debated. Before this report, no clear morphological differences between reproductive and non-reproductive individuals were known. Here we show that several pre-imaginal morphological differences of the Van der Vecht organ occur between foundresses (queens) and workers. Geometric morphometrics revealed that queens are characterized by shape deformations of this organ, which is responsible for a typical allometric growth of the secretory area. This organ is predicted to be larger in foundresses compared to workers because its secretion is involved in defense against ants, in nestmate recognition and in preventing workers from challenging for direct reproduction. The results presented here indicate the existence of an incipient morphological caste determination Polistes gallicus and suggest that this species may have passed the “point of no return” for eusociality.     

中国多刺蚁属驼背亚属系统分类研究（膜翅目：蚁科）

记述了在云南省南部西双版纳自然保护区热带雨林区域采集到的多刺蚁属Polyrhachis Smith驼背亚属Cyrtomyrma Forel 5新种：方肩多刺蚁P. (C.) cornihumera sp. Nov.,齿肩多刺蚁P. (C.) dentihumera sp. Nov.,圆肩多刺蚁P. (C.) orbihumera sp. Nov.,圆顶多刺蚁P. (C.) rotoccipita sp. Nov.,短胸多刺蚁P. (C.) brevicorpa sp. Nov.。目前为止在中国和中南半岛记载该亚属13种,其中中国已知9种。提供了中国和中南半岛已知13种的检索表。     

Two new myrmecophytic associations from the Malay Peninsula: Ants of the genusCladomyrma (Formicidae,Camponotinae) as partners ofSaraca thaipingensis (Caesalpiniaceae) andCrypteronia griffithii (Crypteroniaceae)

Summary In Peninsular Malaysia the treesSaraca thaipingensis (Caesalpiniaceae) andCrypteronia griffithii (Crypteroniaceae) are inhabited by ants. In the vicinity of Gombak, near Kuala Lumpur, the hollow internodes of youngSaraca thaipingensis plants are colonized mainly by twoCladomyrma species. In larger trees aCrematogaster sp. is also found.Crypteronia griffithii is inhabited by a third species ofCladomyrma. None of these species is conspecific with any of the threeCladomyrma taxa so far described. The colonies are founded by single mated queens, which have a conspicuous, sphecid wasp-like behaviour when searching for host plants and nest sites. They chew holes into the plant internodes and hollow them out to provide nest sites. Coccids and pseudococcids are cultivated within the internodes. The homopterans are not carried by queens on their nuptial flights. They apparently find their way by themselves into the cavities or are perhaps carried there by the worker ants. TheCladomyrma ants onCrypteronia are not aggressive, in contrast to those onSaraca thaipingensis.The relationship ofCrypteronia with ants seems to be obligatory, whereasSaraca was only partly colonized byCladomyrma. The interaction ofSaraca withCrematogaster sp. is loose and facultative, since theCrematogaster sp. also lives on other three species. Our studies have now revealed fourCladomyrma spp. which are regularly associated with plants. The genus therefore seems to have an entirely myrmecophytic way of life.     

Polyphagy and Obligate Myrmecophily in the Butterfly Hallonympha paucipuncta (Lepidoptera: Riodinidae) in the Neotropical Cerrado Savanna

This study analyzes host plant use and obligate myrmecophily in Hallonympha paucipuncta, an endemic butterfly of the Cerrado. Larvae of H. paucipuncta are polyphagous, using at least 19 species of plants in 10 families. All larvae found were being tended by Crematogaster ants. Spatial distribution of larvae and tending ants were strongly aggregated, suggesting an influence of ants on oviposition and/or larval survival. Implications of obligate myrmecophily in the evolution of polyphagy in Riodinidae are discussed.     

Disruption of parasitism of the California red scale (Homoptera: Diaspididae) by three ant species (Hymenoptera: Formicidae)

The relative influence of the southern fire ant, Solenopsis xyloni McCook, Argentine ant, Linepithema humile (Mayr), and native gray ant, Formica aerata (Francoeur), on parasitism of California red scale, Aonidiella aurantii Maskell, was studied in the laboratory for two parasitoids, Comperiella bifasciata Howard and Aphytis melinus DeBach. All three ant species reduced percentage parasitism by C. bifasciata and both percentage parasitism and host mutilation by A. melinus. Southern fire ant was the least disruptive and native gray ant the most disruptive. Southern fire ant removed 12% of scale from the lemons, presumably to feed on them, while the other ant species did not exhibit significant removal of scale compared to the controls. Percentage parasitism of California red scale exhibited by C. bifasciata was more than 2-fold the level exhibited by A. melinus. Percentage mutilation of California red scale, including probing and host feeding, was nearly 5-fold higher for A. melinus than C. bifasciata. Because A. melinus required a longer total host examination + oviposition period in the absence of ants than C. bifasciata and because oviposition occurs as the last act in a sequence of behaviors, disruption by the ants had a more significant negative effect on oviposition by A. melinus.     

Characterization of microsatellite markers for plant‐ants of the genus Crematogaster subgenus Decacrema

We present primer sequences for five polymorphic microsatellite loci in ants of the genus Crematogaster subgenus Decacrema that live in obligate symbiosis with host plants of the euphorb genus Macaranga. Microsatellite loci were isolated with a highly efficient method of enrichment. The number of alleles ranged from 10 to 18 for Crematogaster morphospecies 2, for which these markers were developed, with an observed heterozygosity ranging from 0.6578 to 0.9474. The markers were designed for the study of the population as well as of the social structure of colonies.     

A phylogeny of Pacific fiddler crabs of the subgenus Minuca (Crustacea,Brachyura, Ocypodidae: Uca) with the description of a new species from a tropical gulf in Pacific Costa Rica

The phylogeny of all Pacific fiddler crab representatives of the subgenus Minuca Bott, 1954 (sensu Beinlich and von Hagen, 2006) is reconstructed. For the molecular analysis, Cox1 mitochondrial and 28S ribosomal nuclear DNA sequences were used. According to these data, same transisthmian sister species relationships are confirmed and a new species of the genus Uca Leach, 1814, Uca osa sp. n., is described from Golfo Dulce, a tropical gulf in Pacific Costa Rica. Morphological as well as molecular data confirm distinctness of this species compared with all other members of the subgenus Minuca, to which it belongs. Distinctive morphological traits are presented to distinguish Uca osa sp. n. from its congeners in the Eastern Pacific.     

Molecular phylogeny of the subgenus Holothuria (Selenkothuria) Deichmann, 1958 (Holothuroidea: Aspidochirotida)

The subgenus Selenkothuria comprises 12 species of tropical shallow water sea cucumbers that share morphological features, such as rods in the body wall and tube feet, modified tentacles for suspension feeding, and cryptic colours. The taxonomic status of this taxon has been controversial, but currently it is accepted as a subgenus of the genus Holothuria. Phylogenetic analyses of mitochondrial genes [cytochrome c oxidase subunit 1 (COI), 16S RNA] of ten species of Selenkothuria and related subgenera showed the polyphyly of this subgenus; monophyly was rejected by a likelihood ratio test. A geographical split divides the species of this subgenus into three different groups: one Indo‐West‐Pacific (IWP) group and two American groups. The IWP group is more closely related to Holothuria (Semperothuria) cinerascens and to other subgenera such as Roweothuria, Holothuria, and Vaneyothuria, whereas the two American groups are more closely related to each other and to some species of the subgenus Halodeima. These results suggest multiple parallel originations and diversification of ossicle morphology within the subgenus Selenkothuria. The current scheme of subgenera for the genus Holothuria is not supported, suggesting the need for a new classification. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 109–120.     

The cephalic labial gland secretions of two socially parasitic bumblebees Bombus hyperboreus (Alpinobombus) and Bombus inexspectatus (Thoracobombus) question their inquiline strategy

Social parasitic Hymenopterans have evolved morphological, chemical, and behavioral adaptations to overcome the sophisticated recognition and defense systems of their social host to invade host nests and exploit their worker force. In bumblebees, social parasitism appeared in at least 3 subgenera independently: in the subgenus Psithyrus consisting entirely of parasitic species, in the subgenus Alpinobombus with Bombus hyperboreus, and in the subgenus Thoracobombus with B. inexspectatus. Cuckoo bumblebee males utilize species‐specific cephalic labial gland secretions for mating purposes that can impact their inquiline strategy. We performed cephalic labial gland secretions in B. hyperboreus, B. inexspectatus and their hosts. Males of both parasitic species exhibited high species specific levels of cephalic gland secretions, including different main compounds. Our results showed no chemical mimicry in the cephalic gland secretions between inquilines and their host and we did not identify the repellent compounds already known in other cuckoo bumblebees.