The internal phylogeny of ants (Hymenoptera: Formicidae)

Abstract. The higher phylogeny of the Formicidae was analysed using 68 characters and 19 taxa: the 14 currently recognized ant subfamilies plus 5 potentially critical infrasubfamilial taxa. The results justified the recognition of 3 additional subfamilies: Aenictogitoninae Ashmead (new status), Apomyrminae Dlussky & Fedoseeva (new status), and Leptanilloidinae Bolton (new subfamily). A second analysis on these better delimited 17 subfamilies resulted in 24 equally most parsimonious trees. All trees showed a basal division of extant Formicidae into two groups, the first containing (Myrmicinae, Pseudomyrmecinae, Nothomyrmeciinae, Myrmeciinae, Formicinae, Dolichoderinae, Aneuretinae) and the second the remaining subfamilies. Clades appearing within these groups included the Cerapachyinae plus 'army ants', the Nothomyrmeciinae plus Myrmeciinae, the 'formicoid' subfamilies (Aneuretinae + Dolichoderinae + Formicinae), and the Old World army ants (Aenictinae + Aenictogitoninae + Doryline), but relationships within the last two groups were not resolved, and the relative positions of the Apomyrminae, Leptanillinae and Ponerinae remained ambiguous. Moreover, a bootstrap analysis produced a consensus tree in which all branches were represented in proportions much lower than 95%. A reconstruction of the ground plan of the Formicidae indicated that the most specialized of all recent ants are the members of the subfamily Dorylinae and the least specialized ones are the monotypic Apomyrminae.     

Miocene ants (Hymenoptera,Formicidae) from Crimea

Ant imprints, new and previously known, from Middle Miocene deposits near Kerch (Crimean Peninsula, Russia) are described or redescribed. A new Myrmicinae species, Solenopsis atavinus sp. nov., is described based on a wingless female. A winged female similar to the earlier described Dolichoderus tauricus Dlussky, 1981 is found: due to the excellent preservation of this specimen, the new specimen and the holotype of D. tauricus are redescribed and can be reclassified as Ponerites tauricus (Dlussky, 1981), comb. nov. Oecophylla taurica sp. nov. is described based on a partly preserved imprint of a female thorax with a forewing, the venation of which allows it to be positively identified as a weaver ant. Two other species are transferred from the formal genus Camponotites to Oecophylla based on forewing venation: O. kraussei (Dlussky et Rasnitsyn, 1999), comb. nov. (Early Eocene, United States) and O. macroptera (Dlussky, 1981), comb. nov. (Middle Miocene, Stavropol, Russia). One of the studied forewing imprints is similar in venation to Paraphaenogaster microphthalmus Dlussky, 1981, described from the Middle Miocene of Vishnevaya Balka (Stavropol province, Russia) and so is attributed to this species. Dolichoderus tavridus sp. nov. is described based on a forewing.     

Observer ants: Storers of foraging area information inFormica rufa L. (Formicidae,Hymenoptera)

Summary The structure of a foraging area in Formica s. str. is maintained by specific foraging roads and by experienced ants familiar with the foraging area, namely foragers and observer ants which used to be foragers. After all the foragers have been removed, observer ants of a F. rufa nest experimental column resume foraging, rapidly restoring the initial structure of the foraging area. For a nest population to lose completely information of the foraging area of a column, not only foragers but also observer ants must be removed and direct access from the nest to experimental road blocked.

---

Les fourmis observatrices: des stockeuses d'informations sur l'aire d'affouragement chez Formica rufa L.

Resume La structure de l'aire d'affouragement chezFormica rufa est maintenue grâce à des pistes d'affouragement spécifiques et des Fourmis expérimentées qui sont familiarisées avec l'aire d'affouragement: des fourrageuses et des observatrices qui sont habituellement des fourrageuses. Lorsque toutes les fourrageuses sont enlevées, les observatrices d'un nid expérimental deFormica rufa reprennent l'affouragement et rétablissent rapidement la structure initiale de l'aire d'affouragement. Pour que la population d'un nid perde toute information sur l'aire d'affouragement d'une colonne, il faut que non seulement les fourrageuses mais aussi les observatrices soient enlevées et que l'accès du nid à la piste soit empêché.

     

Morphological phylogeny of army ants and other dorylomorphs (Hymenoptera: Formicidae)

Abstract. The dorylomorph group of ants comprises the three subfamilies of army ants (Aenictinae, Dorylinae, Ecitoninae) together with the subfamilies Aenictogitoninae, Cerapachyinae, and Leptanilloidinae. We describe new morphological characters and synthesize data from the literature in order to present the first hypothesis of phylogenetic relationships among all dorylomorph genera. These data include the first available character information from the newly discovered male caste of Leptanilloidinae. We used ant taxa from Leptanillinae, Myrmeciinae, and the poneromorph (Ponerinae sensu lato) subfamilies Amblyoponinae, Ectatomminae, and Paraponerinae as outgroups. We scored a total of 126 characters from twenty-two terminal taxa and used these data to conduct maximum parsimony and bootstrap analyses. The single most-parsimonious tree and bootstrap results support a single origin of army ants. The Old World army ant genus Dorylus forms a monophyletic group with the enigmatic genus Aenictogiton, which is currently known only from males; the second Old World army ant genus Aenictus is sister to this clade. This result generates the prediction that females of Aenictogiton, when discovered, will be observed to possess the army ant syndrome of behavioural and reproductive traits. The monophyly of the New World army ants (Ecitoninae) is supported very strongly, and within this group the genera Eciton, Nomamyrmex, and Labidus form a robust clade. The monophyly of Leptanilloidinae is also upheld. The subfamily Cerapachyinae appears paraphyletic, although this conclusion is not supported by strong bootstrap results. Relationships among genera of Cerapachyinae similarly are not resolved robustly, although parsimony results suggest clades consisting of (Acanthostichus + Cylindromyrmex) and (Cerapachys + Sphinctomyrmex). We tested for the effect of incompletely known taxa by conducting a secondary analysis in which the two genera containing ∼50% missing character data (Aenictogiton and Asphinctanilloides) were removed. The strict consensus of the seventeen most-parsimonious trees from this secondary analysis is poorly resolved outside the army ants and contains no clades conflicting with the primary analysis. The position of Leptanilla shifts from forming the sister group to Leptanilloidinae (without high bootstrap support) in the primary analysis, to falling within a polytomy at the base of the root of the dorylomorphs when incompletely known taxa are removed. This instability suggests that the placement of Leptanilla within the dorylomorphs in our primary analysis may be spurious.     

Molecular markers allow sibling species identification in red wood ants (Formica rufa group)

Red wood ants (Formica rufa group) constitute a group of species that are considered to be among the most promising bioindicators in forest ecosystems. However, because of their morphological similarity and intraspecific variability, morphological species identification can be difficult. Considerable expertise is necessary to discriminate between the sibling species F. lugubris and F. paralugubris, two species that often live in sympatry in the same Alpine forests. New taxonomic tools providing rapid and reliable species identification are needed. We present a simple and reliable molecular technique based on mtDNA (COI gene) and a restriction enzyme for discriminating between F. lugubris and F. paralugubris. We confirm the validity of this method with a Bayesian analysis based on microsatellites. This new molecular tool represents a clear breakthrough for discriminating between F. lugubris and F. paralugubris and is likely to be helpful in large‐scale biomonitoring.     

Molecular phylogeny and biogeography of the spider ants, genus Leptomyrmex Mayr (Hymenoptera: Formicidae)

This study provides the first phylogenetic reconstruction of the ant genus Leptomyrmex Mayr, a prominent endemic component of rain forest and wet sclerophyll forest in Australia, New Guinea and New Caledonia. Five genes are used to reconstruct phylogeny and estimate of ages of diversification in order to test congruence of the history of nuclear and mitochondrial genes: three protein-coding nuclear genes: arginine kinase (argK, 897 bp), long wavelength rhodopsin (LW Rh, 546 bp) and wingless (Wg, 409 bp), as well as the large subunit ribosomal gene 28S (482 bp) and the mitochondrial gene cytochrome oxidase I (COI, 658 bp). Four different partitioning schemes were tested for optimal resolving power; results show that partitioning by gene, translational pattern and codon position were uniformly favoured over less complex partitions. Nuclear markers showed relatively minor sequence divergence and provided strongly supported topology; phylogeny based solely on mtDNA produced somewhat conflicting topology but offered little power to resolve species complexes. Monophyly of the genus Leptomyrmex was recovered, as was the sister-group relationship of 'micro-' and 'macro-'Leptomyrmex species. Divergence dating analyses estimate that Leptomyrmex arose in the Eocene (stem age ～ 44 million years ago (ma)), and that the 'macro-' species diverged from the 'micro-' species in the early Oligocene (～ 31 ma). Diversification of the crown group 'macro-' and 'micro-'Leptomyrmex occurred in the Miocene (～ 15 ma and 7.9 ma, respectively). New Guinean and New Caledonian lineages appear to have diverged from Australian lineages only recently (～ 4.7 ma and 10.3 ma, respectively), and the latter clade is inferred to have reached New Caledonia from Australia via long distance dispersal. These results challenge previous hypotheses of Leptomyrmex classification and assumptions about their historical dispersal, but are in agreement with the current knowledge of the geological history of Melanesia.     

Response of the foraging behaviour of red wood ants (Formica rufa group) to exclusion from trees

Abstract 1 Because of the large numbers within a colony and their aggressive nature, red wood ants (Formica rufa group) have a potential to greatly influence the cold‐temperate forest ecosystem. Wood ants are omnivorous and hunt in trees as well as on the forest floor. 2 A field experiment in a mixed forest in central Sweden was carried out to examine (i) the foraging behaviour of wood ants on the forest floor and (ii) the impact of increased numbers of wood ants on the soil fauna. The foraging behaviour of wood ants was manipulated by excluding the ants from their food resources in the tree canopy, with the intention to increase ant activity on the forest floor. To estimate this activity, the number of trees with foraging ants, the numbers of ants going to and from their nests and the prey carried by home‐running wood ants were determined during the summer period. Pitfall traps were placed in the soil to determine effects on mobile soil invertebrates. 3 When excluded from local trees, wood ants searched other trees further away from the nests rather than searching more intensively for prey on the forest floor. By contrast to the initial hypothesis, more soil‐living prey were caught by ants in the control plots than in the plots where the local trees were not accessible to the wood ants. The proportion of soil‐living to tree‐living prey tended to be greater in the control plots. 4 In the treated plots (no access to the trees), wood ants had a negative effect on the activity of Linyphiidae spiders. There was little effect of wood ants on other soil invertebrates. 5 This study suggests that the role of wood ants as top predators in the forest soil food‐web in central Sweden is limited.     

Further records of ants (Hymenoptera: Formicidae) from Iran

Wide ranging surveys of the ant fauna of Iran have enabled us to add a further 30 named species to the country list. A review of almost all the published literature and of photographs of unidentified specimens within the public domain gives a grand total of 248 species, from seven subfamilies and 37 genera. In the majority of instances, our own specimens were compared with type images available from antweb.com. This has led us to propose new or revised status for Cataglyphis turcomanica Crawley 1920, Lepisiota integrisquama (Kuznetsov-Ugamsky, 1929), Lepisiota surchanica (Kuznetsov-Ugamsky, 1929) and Messor obscurior Crawley 1920. We note that the total includes a number of what may be misidentifications and a small number of named species that seem unlikely to occur in Iran.     

Cold-hardiness of the phoretic mites Anoetus myrmicarum (Acariformes,Anoetidae) from the ants Formica lemani (Hymenoptera,Formicidae) in the Kolyma Highland

In the sparse larch forests of the upper Kolyma River, hypopi of the mite Anoetus myrmicarum (Scheucher, 1957) were found in several nests of the ant Formica lemani Bondroit, 1917. These mites were not found in hundreds of nests of the other 10 ant species examined in northeastern Asia. A possible ecological and physiological conditionality of the restricted distribution of phoretic mites was analyzed. For this purpose, coldhardiness of mites and their ant hosts, the biotopic distribution and the structure of nests, and the temperature conditions of overwintering were examined. At the stage of hypopus, the mites overwintered on ants in the overcooled stage; their mean supercooling temperatures (SCP) varied from ?25.8 ± 0.3°C to ?27.7 ± 0.4°C (min ?32.2°C, n = 157). These values were by 0.1 to 7.0°C lower than the mean SCP of the ants from 8 tested nests of F. lemani (?20.7 ± 0.5°C to ?25.7 ± 0.8°C). The soil temperatures at the level of winter chambers varied from ?12°C to ?15°C. Scarcity of findings of Anoetus myrmicarum in the Kolyma Highland is not associated with the limited cold-hardiness of the examined stages, but is most probably determined by interrelations between mites and ants.     

Decyl acetate synthesis in the ant,Formica schaufussi (Hymenoptera: Formicidae)

Decyl acetate occurs in high amounts in the Dufour's gland of Formica schaufussi. When worker ants were fed [¹⁴C]-acetate the acetate moiety of decyl acetate was labelled preferentially indicating that decyl acetate is synthesized via an esterification reaction.     

Atypical vitellins in ponerine ants (Formicidae: Hymenoptera)

Higher hymenopteran vitellogenin/vitellins have been characterized as containing one large apoprotein. We show that in the ant subfamily Ponerinae, species in the tribes Odontomachini, Platythyrini, and Amblyoponini, also have a vitellin with this simple structure, containing a single apoprotein of 180-190kDa. Species in tribes Ponerini and Ectatommini, however, have vitellins containing multiple subunits. The size and number of the subunits varies, with differences even among species within the same genus. This is the first report of diversity in vitellogenin structure in the higher Hymenoptera. Vitellin and vitellogenin in Harpegnathos saltator (Ponerini) contain two large subunits of about 165kDa and two small subunits of about 45 and 43kDa. N-terminal sequence analysis suggests that provitellogenin is cleaved at two different sites, producing two large and two small subunits differing slightly in size. Diversity of vitellin types in Ponerini and Ectatommini is similar to that found in the more primitive tenthredinoid sawflies (Hymenoptera, Symphyta), and may indicate polyphyly in the Ponerinae. Insect vitellogenins and yolk proteins show considerably more diversity than originally believed, and the possibility of the functional significance of these differences should be considered.     

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.     

Karyological studies of Japanese ants (Hymenoptera,Formicidae)

The karyotypes of nine Japanese ants in three subfamilies (Ponerinae, Formicinae, and Myrmicinae) were successfully analysed by the improved squash technique. Three ponerine species had 2n=7 and n=4 (Ponera scabra), 2n=22 (Brachyponera sinensis), and 2n=28 and n=14 (Cryptopone sauteri). Four formicine species had 2n=18 and n=9 (Camponotus sp. and C. tokioensis), 2n=26 and n=13 (Camponotus japonicus), and 2n=30 and n=15 (Lasius niger). Two myrmicine species had 2n=18 and n=9 (Leptothorax congruus), and 2n=37, 38, 39, and n=17, 18, 19, 20 (Pheidole nodus). It was found that the variation of chromosome number observed in P. nodus was caused by Robertsonian type polymorphism.     

Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria

The present catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria is made on a base of critical reconsideration of literature (covering the period from 1892 till 2009 and part of 2010) as well as on examination of the authors' and several museum's collections. A lot of data were omitted in the previous Bulgarian monograph on ants, lots of new data were recently added and many important additions and alterations were made due to taxonomic revisions of Eurasian Formicidae during the last three decades. Two new species are reported for the country [Temnothorax graecus (Forel, 1911) and Temnothorax cf. korbi (Emery, 1924)].This catalogue contains a list of 163 ant species belonging to 40 genera of 6 subfamilies now known from Bulgaria. Synonyms and information on the previously reported names in relevant publications are given. Known localities of the species are grouped by geographic regions. Maps with concrete localities or regions for each species were prepared. The conservation status of 13 ant species is given as they are included in IUCN Red List of Threatened Species and Bulgarian Biodiversity Act. In comparison with adjacent Balkan regions the ant fauna of Bulgaria is quite rich and its core is composed of South European elements.     

Polygyny and Monoandry in the Ant Formica japonica (Hymenoptera: Formicidae)

Queen number, mating frequency and nest kin-structure of the ant Formica japonica were studied in the field and the laboratory. Nest excavation in the study site, the east slope of Mt. Fuji, Gotenba, Japan, revealed that F. japonica is weakly polygynous all year round and the queen number increases after the nuptial flight season, suggesting the adoption of newly mated queens by established nests. Dissection and laboratory rearing demonstrated that nearly all queens in polygynous nests had mated and were fertile with mature oocytes in their ovaries. Multilocus DNA fingerprinting was used to examine kin relationships among ants found in the same nests. The fingerprint band patterns were apparently governed by a simple genetic rule and suggested monoandry (single mating per queen). The mean band sharing score of DNA fingerprints among full sisters was 0.90, and the mean value between queens and their daughters was 0.75. Comparison of DNA fingerprints of adult and pupal workers with pupal gynes suggested that multiple queens in a nest may contribute unequally to gyne (new queen) production.     

Common names for Australian ants (Hymenoptera: Formicidae)

Abstract Most insects do not have common names, and this is a significant barrier to public interest in them, and to their study by non-specialists. This holds for even highly familiar insect groups such as ants. Here, I propose common names for all major native Australian ant genera and species-groups, as well as for many of the most abundant and distinctive species. Sixty-two genera, 142 species-groups and 50 species are given names. The naming system closely follows taxonomic structure; typically a genus is given a general common name, under which species-group and species names are nested.     

Testing the reproductive groundplan hypothesis in ants (Hymenoptera: Formicidae)

The evolution of complex societies with obligate reproductive division of labor represents one of the major transitions in evolution. In such societies, functionally sterile individuals (workers) perform many of fitness‐relevant behaviors including allomaternal ones, without getting any direct fitness benefits. The question of how such worker division of labor has evolved remains controversial. The reproductive groundplan hypothesis (RGPH) offers a powerful proximate explanation for this evolutionary leap. The RGPH argues that the conserved genetic and endocrinological networks regulating fitness‐relevant behavior (e g. foraging and brood care) in their solitary ancestors have become decoupled from actual reproduction in the worker caste and now generate worker behavioral phenotypes. However, the empirical support for this hypothesis remains limited to a handful of species making its general validity uncertain. In this study, we combine data from the literature with targeted sampling of key species and apply phylogenetically controlled comparative analysis to investigate if the key prediction of the RGPH, namely an association between allomaternal behavior and an allomaternal physiological state holds in the largest and most species‐rich clade of social insects, the ants. Our findings clearly support the RPGH as a general framework to understand the evolution of the worker caste and shed light on one of the major transition in evolutionary history.