Generic-level relationships within the ant subfamily Dolichoderinae (Hymenoptera: Formicidae)

A cladistic analysis was undertaken to determine relationships among extant genera of the ant subfamily Dolichoderinae. Twenty-one of the twenty-two currently recognized genera within the subfamily were examined using 104 morphological characters. A single fully resolved, most-parsimonious tree was found when a combination of ordered and unordered characters was used. When all characters were coded as unordered, seventy most-parsimonious trees were found. The following results were found with both character coding methods. Leptomyrmex was placed basal to the remainder of the subfamily and the monophyletic sets Dolichoderus + Liometopum + Axinidris + Tapinoma + Technomyrmex, Froggattella + Iridomyrmex + Ochetellus + Papyrius + Philidris + Turneria , and Bothriomyrmex + Dorymyrmex + Forelius + Loweriella were suggested. The genera Linepithema and Doleromyrma showed a small amount of instability in moving between neighbouring sister groups when the character coding method changed. The genera Anillidris and Anonychomyrma were difficult to place as they showed major differences in their positions between the two character coding methods.     

The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants

Abstract. The ant subfamily Pseudomyrmecinae comprises three genera of hyperoptic, arboreal ants, widely distributed in tropical and subtropical regions: Pseudomyrmex (∼200 species, New World), Myrcidris (two species, South America) and Tetraponera (∼100 species, Palaeotropics). The phylogenetic relationships among these ants were investigated using DNA sequence data (∼5.2 kb from 18S rDNA, 28S rDNA, wingless, abdominal-A, and long-wavelength rhodopsin genes) and 144 morphological characters, both separately and in combination. Data were gathered from a representative set of forty-nine pseudomyrmecine species, plus eighteen species from various outgroups. There was substantial agreement among the results obtained from different datasets, and from different methods of phylogenetic inference (parsimony, Bayesian inference). The monophyly of the following groups is strongly supported (100% bootstrap support and 1.00 posterior probability in the molecular dataset): Pseudomyrmecinae, Pseudomyrmex, and Pseudomyrmex + Myrcidris. The status of the genus Tetraponera is less clear: the DNA sequence data indicate that the genus is paraphyletic, but morphological features and a unique insertion in the 28S gene support the monophyly of this taxon. Seven of nine Pseudomyrmex species groups, established previously on the basis of morphology alone, are strongly upheld, but monophyly is rejected for the P. pallens group and the P. viduus group. In the latter case, molecular evidence indicates the existence of two independent clades, associated with the ant-plants Triplaris and Tachigali, respectively, whose convergent morphological features had caused them to be placed erroneously in the same species group. The present results confirm an earlier assertion that obligate associations with domatia-bearing plants have arisen at least twelve times in the subfamily. Molecular and morphological data support the hypothesis of a sister-group relationship between Pseudomyrmecinae and Myrmeciinae (84% parsimony bootstrap, combined dataset), which implies a Cretaceous origin of the stem-group pseudomyrmecines in the southern hemisphere. Pseudomyrmecines appear to have arisen in the Palaeotropics and later dispersed from Africa to South America, where they experienced a pronounced burst of diversification.     

The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): generic revision and relationship to other formicids

Three genera are recognized in the ant subfamily Pseudomyrmecinae: Myrcidris new genus (monotypic; type species M. epicharis , new species, from Amazonas, Brazil); Pseudomyrmex Lund (c. 150 nominal taxa; southern Nearctic and Neotropical regions); and Tetraponera F. Smith (c. 110 nominal taxa; Palaeotropical region). New generic synonymies are: Pseudomyrmex Lund = Ornatinoda Enzmann = Clavanoda Enzmann = Triangulinoda Enzmann = Apendunculata Enzmann = Latinoda Enzmann; and Tetraponera F. Smith = Pachysima Emery = Viticicola Wheeler = Sima Emery (nee Roger) = Parasima Donisthorpe. Autapomorphies are identified for the subfamily and for its constituent genera, indicating the monophyly of all four taxa. Based on available character-state information, Myrcidris is most parsimoniously interpreted as a sister-group to all other Pseudomyrmecinae {Pseudomyrmex + Tetraponera) , although there is almost equally strong support for an alternative grouping: Tetraponera + (Myrcidris + Pseudomyrmex). Phylogenetic relationships of the Pseudomyrmecinae and other ant subfamilies within the 'poneroid complex' ( sensu Taylor, 1978) were assessed by a cladistic analysis of eleven representative ant genera. At the level of subfamily relationships the results suggest that: (1) The Myrmicinae and the Pseudomyrmecinae are sister groups. (2) Within the 'poneroid complex' is an unresolved trichotomy composed of: (i) Pseudomyrmecinae + Myrmicinae, (ii) Myrmeciinae, and (iii) Ponerinae + Leptanillinae + (Cerapachyinae + Dorylinae (s.l.)). (3) The Nothomyrmeciinae are a basal lineage within the 'poneroid complex', rather than being allied with the 'formicoid complex' (Formicinae, Dolichoderinae and Aneuretinae).     

Tales of dracula ants: the evolutionary history of the ant subfamily Amblyoponinae (Hymenoptera: Formicidae)

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.     

中国细蚁亚科系统分类研究（膜翅目：蚁科）

记载中国细蚁亚科Leptanillinae昆虫2属。报道细蚁属Leptanilla Emery 3种：湖南细蚁L. hunanensis Tang, Li et Chen分布于湖南省,台湾细蚁L.taiwanensis Ogata, Terayama et Masuko分布于台湾省,云南细蚁L.yunnanensis sp.nov.分布于云南省。原细蚁属Protanilla Taylor为中国新记录属,在云南省采集并描述该属2新种：单色原细蚁P.concolor sp.nov.和双色原细蚁P.bicolor sp.nov.。编制了细蚁亚科这2属的工蚁分属检索表和中国分布种工蚁分种检索表。     

Higher classification of the ant subfamilies Aneuretinae, Dolichoderinae and Formicinae (Hymenoptera: Formicidae)

Abstract. An analysis of the cladistic relationships among the ant subfamilies Aneuretinae, Dolichoderinae and Formicinae demonstrates the monophyly of these three subfamilies, and places Dolichoderinae and Formicinae as sister groups. This latter result differs from most previous studies which consider Aneuretinae and Dolichoderinae as sister groups.     

Comparative ultrastructure of ant spermatozoa (Formicidae: Hymenoptera)

Mature spermatozoa from spermathecae of founding queens were obtained from 5 species of ants, representing the major subfamilies Myrmicinae (Acromyrmex versicolor, Crematogaster sp.) and Dolichoderinae (Tapinoma sessile, Conomyrma insana, Conomyrma wheeleri). The ultrastructure of ant spermatozoa has many features in common with that of higher insects and is similar to that of other Hymenoptera. Structural similarities to spermatozoa of other Hymenoptera include an acrosome containing an internal rod that extends into the nucleus, two elongate mitochondrial derivatives, a centriolar adjunct, and an axonemal arrangement of 9 + 9 + 2 that includes well-developed coarse, or accessory, tubules. Spermatozoa obtained from A. versicolor, a species that is known to store and utilize viable sperm from this supply for over 10 years, show greater development of the mitochondrial derivatives than do the other species. The most distinctive feature of ant spermatozoa in comparison to other Hymenoptera is the large size of the centriolar adjunct relative to the other organelles. The centriolar adjunct is located posterior to the nucleus, anterior to the mitochondrial derivatives, and opposite the axoneme.     

Revision of the oriental ant genus Cladomyrma,with an outline of the higher classification of the Formicinae (Hymenoptera: Formicidae)

Abstract. The Oriental ant genus Cladomyrma is revised and possible phylogenetic relationships are discussed. Cladomyrma hewitti and hobbyi are newly synonymized with andrei. Cladomyrma cryptata, maschwitzi, mossyna and petalae are described as new species. A lectotype for Cladomyrma hewitti is designated. A synopsis of the classification of Formicinae, based on new morphological characters, is provided. New morphological characters at the generic level are described and the relationships of Cladomyrma within Formicinae are discussed.     

The visual perception of the ant Myrmica ruginodis (Hymenoptera: Formicidae)

Myrmica ruginodis workers are able to distinguish black or white circles from black or white squares, black or white ellipses from black or white rectangles as well as hollow circles or ellipses from hollow squares or rectangles. They can also distinguish differently oriented elements as well as objects containing a various number of elements. These workers are also able to perceive and discriminate transparent cues on a black background and even small luminous spots on a black ceiling. Such visual abilities are in agreement with the species?? eye morphology and system of navigation: the eyes are rather large with a well-developed posterio-dorsal part, and the foragers rely exclusively on cues located above them, neglecting odorous elements as long as visual perception is possible. Probably, they might use cues located in the canopy and the sky to travel in certain circumstances.     

Revision of the dolichoderine ant genus Turneria (Hymenoptera: Formicidae)

Abstract. The Australasian ant genus Turneria is revised. T.bidentata Forel, T.dahlii Forel and T.pacifica Mann are redescribed and a lectotype designated for T.bidentata. T.frenchi Forel is transferred to Stigmacros (comb.n.), and T.butteli Forel is transferred to Iridomyrmex (comb.n.). Three new species, T.arbusta, T.collina and T.postomma, are described from Papua New Guinea. Males of the genus are described for the first time. A key to the workers of recognized species and a distribution map are provided. The group is analysed cladistically and the resulting species relationships are discussed, along with a comment on the use of polymorphic characters.     

Molecular phylogeny of the ant tribe Myrmicini (Hymenoptera: Formicidae)

The interrelationships within ant subfamilies remain elusive, despite the recent establishment of the phylogeny of the major ant lineages. The tribe Myrmicini belongs to the subfamily Myrmicinae, and groups morphologically unspecialized genera. Previous research has struggled with defining Myrmicini, leading to considerable taxonomic instability. Earlier molecular phylogenetic studies have suggested the nonmonophyly of Myrmicini, but were based on limited taxon sampling. We investigated the composition of Myrmicini with phylogenetic analyses of an enlarged set of taxa, using DNA sequences of eight gene fragments taken from 37 representatives of six of the seven genera (Eutetramorium, Huberia, Hylomyrma, Manica, Myrmica, and Pogonomyrmex), and eight outgroups. Our results demonstrate the invalidity of Myrmicini as currently defined. We recovered sister‐group relationships between the genera Myrmica and Manica, and between Pogonomyrmex and Hylomyrma. This study illustrates that to understand the phylogeny of over 6000 myrmicine species, comprehensive taxon sampling and DNA sequencing are an absolute requisite. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 482–495.     

A review of Bahamian ant (Hymenoptera: Formicidae) biogeography

Abstract. The results of recent surveys for ants on Staniel Cay, Exumas (and surrounding small cays) and the northeastern region of Andros are presented. These records are compiled with all previously published collection records of ants from the Bahamas into a single database. A list of all known Bahamian species is presented, in accord with current taxonomy. Distributions within the Bahamas are given, along with the status of each species with reference to its origin (endemic, native, or exotic). At present seventy-five named species of ants are known from the Bahamas, which shares 60% of its ant fauna with Florida and 71% with Cuba. More than one third of the species (37%) are exotics, which appear to exist primarily in areas characterized by human disturbance, and apparently have not invaded the native vegetation to a large degree. Both historical and ecological factors are likely to have played important roles in shaping present-day Bahamian ant species diversity and distribution. The number of species on all major Bahamian islands from which ants have been collected is presented as a function of island area. Most islands have not been very thoroughly surveyed. The species-area relationship for the more thoroughly surveyed islands is relatively flat (z= 0.06). Further collecting in this archipelago will probably lead to the discovery of additional species.     

Exocrine chemistry of the myrmicine ant Zacryptocerus pusillus (Hymenoptera: Formicidae)

Minor workers of the ant Zacryptocerus pusillus have unusual exocrine secretions in both their mandibular and Dufour glands. The mandibular glands contain a 3:1 mixture of 4-heptanone and 4-heptanol, a mixture found only in the related species Z. varians. The Dufour gland contains a mixture of 13 aldehydes from C9 to C18, not previously encountered in ant secretions. The venom glands gave variable results with only nonanal present consistently.     

Population genetics of the black ant Formica lemani (Hymenoptera: Formicidae)

Colony kin structure and spatial population structure were studied in multiple populations of the ant Formica lemani , using allozymes and DNA microsatellites. Average genetic relatedness between nestmate workers varied little between populations ( r  = 0.51–0.76), indicating that the average colony kin structure was relatively simple. Worker genotypes could not be explained with a single breeding pair in all nests, however, and the distribution of relatedness estimates across nests was bimodal, suggesting that single- and multi-queen colonies co-occur. We studied spatial population structure in a successional boreal forest system, which is a mixture of different aged habitats. Newly clear-cut open habitat patches are quickly colonized by F. lemani , where it is able to persist for a limited number of generations. Newly-founded populations showed signs of a founder effect and spatial substructuring, whereas older populations were more homogenous. This suggests that new populations are founded by a limited number of colonizers arriving from more than one source. Genetic differentiation among local populations was minor, indicating strong migration between them. There were, however, indications of both isolation by distance and populations becoming more isolated as habitat patches grew older.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 247–258.     

Communication in the primitive cryptobiotic ant Prionopelta amabilis (Hymenoptera: Formicidae)

Summary The Neotropical ant Prionopelta amabilis, a cryptobiotic species in the phylogenetically primitive tribe Amblyoponini, lives in subterranean habitats, where it preys preferentially on campodeid diplurans and other small arthropods. Here we report that the species employs chemical recruitment and orientation trails during foraging and nest emigrations. The trail pheromone originates in a hitherto unknown basitarsal gland located in the basitarsus of the hindlegs. Trails are laid by a special foot dragging behavior. During the recruitment process the chemical trail signal is complemented by body shaking on the part of the recruiting ant. Foragers frequently wipe the basitarsal gland opening in the hindlegs with the grooming apparatus of the front legs. This latter structure is equipped with unusual glands evidently specialized for this purpose.     

Microsatellites for the Neotropical ant,Camponotus leydigi (Hymenoptera: Formicidae)

Ants (Hymenoptera: Formicidae) are dominant social insects that play important ecological roles in terrestrial ecosystems. Camponotus leydigi (Forel) is widely distributed in the Neotropical region and is frequently found in the Brazilian cerrado savanna interacting with plants and other insects. Field observations indicate that C. leydigi has a polydomous nesting habit, but little is known about the genetic relationship among workers. In this study, we identify the first nine microsatellite loci for C. leydigi that will allow further investigation on its genetic diversity. We used a microsatellite-enriched library method. According to this method, repetitive sequences are captured with (CT)₈ and (GT)₈ biotin-linked probes, with subsequent recovery by streptavidin magnetic-coated beads. We observed that eight loci were polymorphic. The mean (± standard error) observed and expected heterozygosities were 0.55 ± 0.23 and 0.73 ± 0.28, respectively. The rarified allelic richness ranged from 1 to 5.32. The polymorphism contents were similar to diversity estimates found in markers previously developed for other Camponotus ants. These markers will be useful for future studies on population genetics and ecology of Camponotus ants in cerrado, including nesting ecology, colony structure, dispersal and conservation.     

Agonistic relationships among sympatric mediterranean ant species (Hymenoptera: Formicidae)

The potential aggressiveness of nine ant species belonging to the same community was estimated by means of two laboratory approaches: individual confrontations and group confrontations. Interspecific aggressiveness was not directly related to size of species in individual tests, although in group tests mortality of smaller ants increased with increasing size difference between the opponents. Tempo was related to the defense strategy of species (low-tempo species used immobility as a mechanism to avoid attacks, while high-tempo species tended to escape when coming into contact with more aggressive ones), but not with their potential aggressiveness: the most aggressive ants in individual and group tests were both low-tempo and high-tempo species when only these two categories were considered. No specific level of aggression was related to the subfamily to which the species belong (both myrmicines and formicines dominated in individual and group tests) or to the degree of taxonomic similarity between species.     

New exocrine glands in the legionary ants of the genus Leptanilla (Hymenoptera,Formicidae, Leptanillinae)

Summary This paper reports the results of the first detailed study of the morphology of exocrine glands in two species of the ant subfamily Leptanillinae. Workers of Leptanilla escheri and L. japonica possess a large, unpaired sternal gland in the VIIth abdominal sternite, and an hypertrophied poison gland which is surrounded by a massive muscle layer. The sternal gland is absent in the queen of L. japonica, and the poison gland is highly degenerated. The queen is, however, endowed with a series of large, paired, intersegmental tergal and sternal glands, which occur between the IVth through the VIIth segments. The queen also posseses large spiracular plate glands.