The control of Myrmica rubra workers by queens of their own and other Myrmica species and the interaction between queenless groups of workers

ABSTRACT. Queens of two species of the ant genus Myrmica bonded to workers of the species M. rubra L. as the latter emerge from the pupal skin can use these workers nearly 6 months later to arrest gyne formation in sex-competent larvae of the same species. Queens of M. ruginodis Nylander var. microgyria (Brian & Brian, 1949) are as good at this as the natural M. rubra , but those of M. sabuleti Meinert (of a race close to M. scabrinodis) are not. Though the M. sabuleti queens induce normal aggression against sexualizing larvae, they are unable to prevent some or all of the workers feeding larvae as though they were queenless. However, queens from different colonies of M. rubra adopted by queenless populations of workers in spring, control their brood-rearing behaviour perfectly. M. rubra workers from different colonies bring gynes to maturity from female sexual larvae at different average sizes. When workers from two such sources are mixed in equal proportions, the size of gyne larva produced after a week's culture corresponds with that of one of the worker populations; it is not intermediate in size. Also, large workers can rear larger gyne-larvae than small workers of the same age. This is only true if the workers have been living with queens all the time from emergence as an imago to the moment the experiment was set. Size mixtures only achieve the same size larvae as a pure culture of small workers would. A possible reason for this is that small workers exclude the larger ones from the nursery areas of the nest.     

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.     

Molecular cloning and sequencing of 18S rDNA gene fragments from six different ant species

Summary The evolutionary relationship between socially parasitic ants and their hosts is still an unsolved problem. We have compared a 1.2 kb sequence of the 18 S ribosomal RNA genes of the parasitic antsDoronomyrmex kutteri, Harpagoxenus sublaevis andChalepoxenus muellerianus to the sequence of the host speciesLeptothorax acervorum andL. recedens (all subfamily Myrmicinae, tribe Leptothoracini) and to an out-group antCamponotus ligniperda (Formicinae). We found that parasitic species and the host species and alsoCamponotus ligniperda differ at less than 1% of the base positions of the 1.2 kb segment of the 18S rRNA gene. The sequences showed 80.3% identity to the 18 S ribosomal RNA genes of the beetleTenebrio molitor and only 66.5% to that of the dipteranDrosophila melanogaster.     

Colony size predicts division of labour in attine ants

Division of labour is central to the ecological success of eusocial insects, yet the evolutionary factors driving increases in complexity in division of labour are little known. The size–complexity hypothesis proposes that, as larger colonies evolve, both non-reproductive and reproductive division of labour become more complex as workers and queens act to maximize inclusive fitness. Using a statistically robust phylogenetic comparative analysis of social and environmental traits of species within the ant tribe Attini, we show that colony size is positively related to both non-reproductive (worker size variation) and reproductive (queen–worker dimorphism) division of labour. The results also suggested that colony size acts on non-reproductive and reproductive division of labour in different ways. Environmental factors, including measures of variation in temperature and precipitation, had no significant effects on any division of labour measure or colony size. Overall, these results support the size–complexity hypothesis for the evolution of social complexity and division of labour in eusocial insects. Determining the evolutionary drivers of colony size may help contribute to our understanding of the evolution of social complexity.     

Sequence comparisons of the internal transcribed spacer region of ribosomal genes support close relationships between parasitic ants and their respective host species (Hymenoptera: Formicidae)

Summary A fragment of the internal transcribed spacer (ITS-1) adjacent to the 5.8S rRNA gene of 20 myrmicine ant species was sequenced. Sequence comparisons were carried out between 11 species of the tribe Leptothoracini, five species of the tribe Tetramoriini, three species of the tribe Solenopsidini and one species of the tribe Myrmicini. Additionally, the formicine antCamponotus ligniperda (tribe Camponotini) was analyzed as an outgroup species. Among all investigated species, the fragment had a variable length of 230–380 bp with only a few conserved sequence elements. The sequences of this fragment were perfectly identical within four palearctic populations ofLeptothorax acervorum indicating that intraspecific variation is rather low. Within the species of Tetramoriini (includingAnergates atratulus) 94.1% of sequence positions were identical, 95.6% within the species of theLeptothorax s.str.-group and 64.6% within the species of theMyrafant-group. Phylogenetic analysis reveals that the social parasitesHarpagoxenus sublaevis, Doronomyrmex goesswaldi, D. kutteri andD. pacis, Chalepoxenus muellerianus as well asStrongylognathus alpinus andTeleutomyrmex schneideri are most closely related to the groups of their respective host species, which generally confirms the taxonomical classifications of the subfamily Myrmicinae based on morphological criteria. The taxonomical positions of the speciesA. atratulus has as yet been uncertain, however, sequence comparison of the ITS-1 fragment leads to the conclusion thatA. atratulus rather belongs to the tribe Tetramoriini than to the Solenopsidini.Author who provided the sequence data from his doctoral thesis.     

How can a character be developmentally constrained despite variation in developmental pathways?

A fundamental riddle of evolutionary developmental biology is the conservation of adult morphological patterns (Hall, 1992). Conservative patterns are either called body plans if they concern overall body design, or homologues if they concern parts of the body (Riedl, 1978; Roth, 1982; Sattler, 1984; Van Valen, 1982; Wagner, 1989a, 1989b). An adult pattern is considered conservative if it remains unchanged in spite of changes in function, as indicated by the original definition of homology by Owen, as a similarity of organs regardless of form and function (Owen, 1848). Conservation of anatomical features despite different adaptive pressures is naturally explained by developmental constraints (Wagner, 1986). However, this approach to explain the biological basis of homology is plagued by the fact that developmental pathways are often more variable than the characters that they produce (see Tab. 1) (Hall, 1992; Roth, 1988, 1991; Spemann, 1915; Wagner, 1989b). This is also true for any other application of the concept of developmental constraints. The widely held opinion that early stages of development are conservative because any early perturbation is likely to interfere with later development, is far from absolute, since a vast amount of data in comparative developmental biology speaks to developmental variation (see e.g. the examples in Tab. 1). The question then is, how can developmental constraints on adult variation be reconciled with the fact of developmental variation?     

Comparison of acoustical signals in Maculinea butterfly caterpillars and their obligate host Myrmica ants

An acoustical comparison between calls of parasitic butterfly caterpillars and their host ants is presented for the first time. Overall, caterpillar calls were found to be similar to ant calls, even though these organisms produce them by different means. However, a comparison of Maculinea caterpillars with those of Myrmica ants produced no evidence suggesting fine level convergence of caterpillar calls upon those of their species specific host ants. Factors mediating the species specific nature of the Maculinea-Myrmica system are discussed, and it is suggested that phylogenetic analysis is needed for future work.     

Yearly variation in polygyny in the termite <Emphasis Type="Italic">Macrotermes michaelseni</Emphasis> (Sjöstedt)

Summary Macrotermes michaelseni, an abundant termite species in semi-arid ecosystems of eastern Africa, exhibits considerable fluctuations in the proportion of polygynous colonies across years (between 5 and 50%). We found a negative correlation between rainfall and the proportion of polygynous colonies with a time lag of two years. A decrease in the percentage of polygynous colonies was particularly obvious in 1999 and 2000, two years after an El Niño-event. Rainfall and subsequent primary production seem to influence the degree of polygyny. Variation in the proportion of polygynous colonies could be due to several mutually non-exclusive factors, including better survival of polygynous colonies during years with low resource availability and increased aggression between nestmate queens with decreasing resource availability. Evaluating several lines of circumstantial evidence we suggest that the degree of polygyny depends on ecological factors.Received 11 July 2003; revised 14 October and 3 February 2004; accepted 9 March 2004.     

Multiple convergent evolution of arboreal life in oribatid mites indicates the primacy of ecology

Frequent convergent evolution in phylogenetically unrelated taxa points to the importance of ecological factors during evolution, whereas convergent evolution in closely related taxa indicates the importance of favourable pre-existing characters (pre-adaptations). We investigated the transitions to arboreal life in oribatid mites (Oribatida, Acari), a group of mostly soil-living arthropods. We evaluated which general force—ecological factors, historical constraints or chance—was dominant in the evolution of arboreal life in oribatid mites. A phylogenetic study of 51 oribatid mite species and four outgroup taxa, using the ribosomal 18S rDNA region, indicates that arboreal life evolved at least 15 times independently. Arboreal oribatid mite species are not randomly distributed in the phylogenetic tree, but are concentrated among strongly sclerotized, sexual and evolutionary younger taxa. They convergently evolved a capitate sensillus, an anemoreceptor that either precludes overstimulation in the exposed bark habitat or functions as a gravity receptor. Sexual reproduction and strong sclerotization were important pre-adaptations for colonizing the bark of trees that facilitated the exploitation of living resources (e.g. lichens) and served as predator defence, respectively. Overall, our results indicate that ecological factors are most important for the observed pattern of convergent evolution of arboreal life in oribatid mites, supporting an adaptationist view of evolution.     

Feeding Competition and Group Size in Alouatta pigra

Researchers consider group size in primates to be determined by complex relationships among numerous ecological forces. Antipredator benefits and better resource defense are the primary pressures for large groups. Conversely, intragroup limited food availability, can result in greater intragroup feeding competition and individual energy expenditure in larger groups, creating energetic advantages for individuals in small groups and placing an upper limit group size. However, the extent to which food availability constrains group size remains unclear for many species, including black howlers (Alouatta pigra), which ubiquitously live in small social groups (≤10 individuals). We studied the relationship between group size and 2 key indices of feeding competition—day journey length and activity budgets—in 3 groups of wild Alouatta pigra at a hurricane-damaged site in Belize, Central America. We controlled for differences in food availability between home ranges (food tree density) and compared both indicators of feeding competition directly with temporal variation in food availability for each group. Our results show no consistent association between resource availability, group size, and either index of competition, indicating that feeding competition does not limit group size at the site—i.e., that larger groups can form without increased costs of feeding competition. The results support the search for other explanations, possibly social ones, for small group size in the primates, and we conclude with suggestions and evidence for such alternative explanations.