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.

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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.

     

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.     

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.     

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.     

New fossil ants of the subfamily Myrmeciinae (Hymenoptera, Formicidae) from Germany

Two new fossil members of the subfamily Myrmeciinae are described: Archimyrmex wedmannae, sp. nov (Grube Messel, Germany, Middle Eocene) and Prionomyrmex wappleri, sp. nov. (Rott, Germany, Late Oligocene). Members of the genus Archimyrmex were known earlier from Eocene deposits of the United States and Argentina, and members of the genus Prionomyrmex were known from Late Eocene Baltic amber.     

The Dufour gland closing apparatus in Formica sanguinea Latreille (Hymenoptera,Formicidae)

Summary A closing apparatus of the Dufour gland is described in the formicine ant Formica sanguinea Latreille 1798. Four sets of muscles are involved, two of which directly attach to the slit-like duct. The latter shows a considerable cuticular thickening of its intima at this level. Ultrastructural observations reveal that the muscles are attached to a cuticula by means of intracellular microtubules in the duct cells. These microtubules run parallel to the myofilaments. Together with the increased contact area for muscular attachments they are believed to ensure the accurate muscular mechanism assuring a well-controlled spraying activity of this gland. Opening of the duct is probably achieved by active muscular contractions, while its closure may be achieved by a passive return to the rest position of the thickened cuticular intima.Research assistant of the Belgian National Fund for Scientific Research     

Molecular phylogeny of Crematogaster subgenus Decacrema ants (Hymenoptera: Formicidae) and the colonization of Macaranga (Euphorbiaceae) trees

To elucidate the evolution of one of the most species-rich ant-plant symbiotic systems, the association between Crematogaster (Myrmicinae) and Macaranga (Euphorbiaceae) in South-East Asia, we conducted a phylogenetic analysis of the ant partners. For the phylogenetic analysis partial mitochondrial cytochrome oxidase I and II were sequenced and Maximum Parsimony analysis was performed. The analyzed Crematogaster of the subgenus Decacrema fell into three distinct clades which are also characterized by specific morphological and ecological traits (queen morphology, host-plants, and colony structure). Our results supported the validity of our currently used morphospecies concept for Peninsula Malaysia. However, on a wider geographic range (including North and North-East Borneo) some morphospecies turned out to be species complexes with genetically quite distinct taxa. Our phylogenetic analysis and host association studies do not indicate strict cocladogenesis between the subgenus Decacrema and their Macaranga host-plants because multiple ant taxa occur on quite distinct host-plants belonging to different clades within in the genus Macaranga. These results support the view that host-shifting or host-expansion is common in the ants colonizing Macaranga. Additionally, the considerable geographic substructuring found in the phylogenetic trees of the ants suggests that allopatric speciation has also played a role in the diversification and the current distribution of the Decacrema ants.