Behavioral adaptations for raiding in the slave-making ant,Polyergus breviceps

We studied recruitment behavior of the slavemaking ant Polyergus breviceps,which typically raids colonies of Formica gnava.The first test series demonstrated the importance of social context, by showing that recruitment was high during raiding, but virtually absent during preraid circling and during the return trip after a slave raid. The second test series showed that Formicapupae (alone or together with adults) must be present for workers of Polyegrusto recruit nestmates. The third test series demonstrated that panic alarm by raided Formicais caused by a pheromone, and we suggest that adults of Formicamay be the source of this secretion. Finally, the fourth test series showed that formic acid is lethal to adults of Formicabut has almost no adverse effect on Polyergus.This relative immunity by Polyergusmay enable them to remain organized while entering nests of Formicaduring slave raids.     

Cuticular spectra in the slave-making ant Polyergus rufescens and the slave species Formica rufibarbis

Abstract. The relative proportions of cuticular components having the same retention times were compared between the slave-making ant Polyergus rufescens and the slave ant Formica rufibarbis living in monospecific or mixed colonies. The two species were found to present different spectra. The Formica workers, when enslaved by Polyergus , tend to lose their colony characteristics but they do not seem to adopt the characteristics of Polyergus.     

Wax,sex and the origin of species: Dual roles of insect cuticular hydrocarbons in adaptation and mating

Evolutionary changes in traits that affect both ecological divergence and mating signals could lead to reproductive isolation and the formation of new species. Insect cuticular hydrocarbons (CHCs) are potential examples of such dual traits. They form a waxy layer on the cuticle of the insect to maintain water balance and prevent desiccation, while also acting as signaling molecules in mate recognition and chemical communication. Because the synthesis of these hydrocarbons in insect oenocytes occurs through a common biochemical pathway, natural or sexual selection on one role may affect the other. In this review, we explore how ecological divergence in insect CHCs can lead to divergence in mating signals and reproductive isolation. We suggest that the evolution of insect CHCs may be ripe models for understanding ecological speciation.     

Cuticular spectra and inter-individual recognition in the slave-making ant Polyergus rufescens and the slave species Formica rufibarbis

Abstract. The relationship between behavioural tests and relative proportions of cuticular components were studied in the slave-making species Polyergus rufescens and the slave and Formica rufibarbis living in either monospecific or mixed colonies. A correlation between the relative proportions of the cuticular products and interindividual recognition exists in each of the two species Polyergus and Formica: Polyergus are fiercely aggressive towards individuals which have different cuticular spectra and originate from a geographically isolated nest. This seems to be true also in the case of Formica living in monospecific colonies. A similar correlation also exists between the two species, which have different cuticular spectra: encounters arranged between them show that free-living Formica are always fiercely aggressive towards Polyergus. The reason why no such correlation seems to exist, however, between Polyergus and Formica when the latter are enslaved and the two species coexist peacefully at the same nest is discussed.     

Phylogeny and phylogeography of the Mediterranean species of the parasitic ant genus <Emphasis Type="Italic">Chalepoxenus</Emphasis> and its <Emphasis Type="Italic">Temnothorax</Emphasis> hosts

We analysed the phylogenetic and phylogeographic relationships of four Mediterranean species of the rare slave-making ant genus Chalepoxenus and eleven of its about 20 Temnothorax host species by sequencing the mitochondrial Cytochrome Oxidase I and II genes. Neighbour-Joining, Maximum Parsimony and Bayesian analyses based on 1320 bp indicate that the genus Chalepoxenus constitutes a monophylum. In all three analyses, C. kutteri from Southwest Europe and the workerless, “degenerate slavemaker” C. brunneus from North Africa form a monophyletic group. C. muellerianus and C. tauricus, distributed in Southern Europe and Ukraine, respectively, form a monophylum in the Neighbour-Joining and the Maximum Parsimony analysis. In our limited set of only 11 of several hundred Temnothorax species, T. flavicornis forms the sister group of Chalepoxenus. Our study further indicates paraphyly of the genus Temnothorax with respect to Chalepoxenus. Moreover, the results suggest that speciation in this slave-making genus is possibly caused by the formation of host races as different Chalepoxenus species use different hosts, and some samples seem to cluster by host species rather than by geographical distance. Received 5 September 2006; revised 17 March 2007; accepted 23 March 2007.     

Cuticular hydrocarbon profiles as a taxonomic tool: advantages,limitations and technical aspects

Cuticular hydrocarbons (CHCs) are expressed on an insect's cuticle and are one of the major factors allowing insects to identify members of their own species, colony and gender. As a result of their species‐specificity, CHCs are increasingly used to delimit species in addition to more conventional methods, such as morphology or genetic markers, and so play an important role in chemotaxonomy. Species vary in the type of CHCs that they produce, as well as in the relative quantities of shared compounds. This review summarizes not only how taxonomists may differentiate between species based on CHC profiles, but also the incentive for using CHC composition as taxonomic tool. Benefits regarding the identification of cryptic species and early signs of reproductive isolation are then discussed, giving examples from studies of taxonomy, behaviour and biosynthesis. For taxonomic characters to reliably indicate species boundaries, their limitations need to be known. Potential problems caused by environmental effects, intra‐species variation in profiles and other technical issues are highlighted, and suggestions are made regarding their avoidance. It remains a challenge to determine the variation beyond which two species can be called independent; a problem shared by most methods of delimitation. Recently, there has been a shift towards using a combination of different taxonomic tools, both molecular and non‐molecular, to test observed species differences.     

Incipient sexual isolation in the nasuta-albomicans complex of Drosophila: mating preference in male-, female- and multiple-choice mating experiments

Interracial divergence is an important facet of speciation. Thenasuta-albomicans complex ofDrosophila with sixteen morphologically identical, karyotypically different but cross-fertile races is an excellent system to study a few dimensions of raciation.Drosophila nasuta nasuta, Drosophila nasuta albomicans, Cytorace 1, Cytorace 2, Cytorace 3 and Cytorace 4 of this subgroup have been subjected to male-, female- and multiple-choice mating experiments. Out of 8456 crosses conducted, 7185 had successful matings. The overall impression is that mating is far from random amongst these six closely related races of thenasuta-albomicans complex. The males ofD. n. albomicans, Cytorace 1 and Cytorace 4 in male-choice, the females of Cytorace 1 and Cytorace 2 in female-choice, and the males and females ofD. n. nasuta, D. n. albomicans, Cytorace 1 and Cytorace 4 against the males and females of Cytorace 2 in multiple-choice experiments, had significantly more homogamic matings than expected. Thus in this study of evolutionary experimentation on raciation under laboratory conditions, we have documented the initiation of preference for con-specific matings among closely related and independently evolving members of thenasuta-albomicans complex ofDrosophila.     

Nesting biology of the arboreal fungus-growing ant <Emphasis Type="Italic">Cyphomyrmex cornutus</Emphasis> and behavioral interactions with the social-parasitic ant <Emphasis Type="Italic">Megalomyrmex mondabora</Emphasis>

We describe the extraordinary nesting habits of the fungus-growing ant Cyphomyrmex cornutus (Formicidae, Myrmicinae, Attini) and the natural history of Megalomyrmex mondabora (Formicidae, Myrmicinae, Solenopsidini), a social parasite that inhabits nests of C. cornutus and other small attine ants. The study was carried out at two sites on the Atlantic slope of Costa Rica. The C. cornutus nest is an oblong mass of accreted soil, attached to or suspended from low vegetation in wet forest understory. Less than a fourth of the nest volume has chambers and is inhabited by C. cornutus; the remainder is a semi-solid mass of accreted soil often housing a variety of arthropods, including other unspecialized commensal ant species. Five C. cornutus colonies examined were parasitized by M. mondabora. Colonies of M. mondabora inhabited chambers very near those of the host. In laboratory observations, M. mondabora and C. cornutus workers interacted with little aggression despite the consumption of C. cornutus larvae and fungi by M. mondabora. During most interactions, C. cornutus workers responded submissively, whereas M. mondabora appeared indifferent or nonresponsive. Megalomyrmex mondabora parasitizes several other attine species (Cyphomyrmex costatus, Cyphomyrmex salvini, and Apterostigma goniodes), and it appears therefore a relatively unspecialized social parasite with broad attine hostassociation. The size of M. mondabora workers vary with host species, suggesting M. mondabora sensu lato comprises either cryptic species or the host environment affects worker size. Received 14 September 2006; revised 1 February 2007; accepted 6 February 2007.     

Cuticular hydrocarbons in workers of the slave-making ant Polyergus samurai and its slave, Formica japonica (Hymenoptera: Formicidae)

Comparisons of cuticular hydrocarbons between workers of the dulotic ant Polyergus samurai and its slave, Formica japonica, were carried out. Gas chromatography–mass spectrometry showed that the slave‐maker and its slave shared the major cuticular hydrocarbon compounds, but possessed several minor products unique to each species. No difference in hydrocarbon composition was detected between enslaved and free‐living F. japonica workers, suggesting that association with P. samurai has no qualitative effect on hydrocarbon composition in these ants. Principal component analyses of the cuticular hydrocarbon profiles (CHP) revealed that (i) CHP was species specific in a given mixed colony; and (ii) among mixed colonies, P. samurai workers had species‐colony specific CHP, while the same feature was not always found in enslaved and free‐living F. japonica workers. Therefore, a ‘uniform colony odor’ in terms of CHP is not achieved in naturally mixed colonies of P. samurai nor those of its slaves, F. japonica.     

The effect of B chromosomes on mating success of the grasshopper Eyprepocnemis plorans

The mating ability of E. plorans was tested in laboratory conditions in six experimental units composed of ten males and fifteen females during 31 days. When significant differences were found (three from the six cages, and in totals) they involved a decrease of matings involving males with B chromosomes. The same tendency seems to exist in females, but to a lesser extent, so that a significant effect is only detected when the totals are considered. Accessory chromosomes also delay, in both sexes, the occurrence of the first mating. No mating preferences depending on the number of Bs were detected.     

Cuticular Extracts of Five Common Mantids (Mantodea: Mantidae) of the Eastern United States

We undertook a preliminary investigation of the cuticular extracts of five common mantid species in the eastern United States: Tenodera sinensis (Saussure), T. angustipennis (Saussure) and Mantis religiosa (Linnaeus) introduced from the Old World and Stagmomantis carolina (Johannson) and Bruneria borealis (Scudder), which are New World species. The major components of these mixtures were normal alkanes, predominately hentriacontane, or in the case of the parthenogenic species B. borealis, tritriacontane. Tricontanal was detected in the extracts of all five species, and smaller amounts of other aldehydes and n-tricontanol were detected in some species. Complex mixtures of methyl and dimethylalkanes also were present in these extracts. The composition of the cuticular hydrocarbons of these mantids may be an adaptation for reduction of evaporative water loss in these insects that inhabit open fields.     

三突伊氏蛛雌蛛不同交配史对雄蛛交配行为的影响

近年来越来越多研究表明,雄性产生精子(精液)也需付出代价。在多次交配的动物中,雄性为获得最大生殖潜力,必须依据配偶的质量策略性地调整每次交配的生殖投入。雄性策略性的生殖投入主要体现在两个方面,一是精子竞争(sperm competition),二是柯立芝效应(Coolidge effect)。目前精子竞争研究主要集中于昆虫类群,而柯立芝效应研究集中于高等脊椎动物,同时验证结果也时常与假说不一致。以多次交配的三突伊氏蛛为材料,以雄蛛交配行为为指标,在蜘蛛类群中探讨和验证雄性精子竞争强度假说和柯立芝效应。设定3个交配组合:2只雄蛛依次与1只雌蛛各交配1次(A组)、2只雄蛛依次与2只雌蛛各交配1次(B组)和1只雄蛛与1只雌蛛交配2次(C组),分析比较3个交配组合的三突伊氏蛛第1次交配和第2次交配在交配潜伏期、交配持续时间和交配回合数方面的差异,比较三突伊氏蛛雌蛛不同交配史对雄蛛交配行为的影响,以此验证雄性精子竞争强度假说和柯立芝效应。研究结果表明A和B组的三突伊氏蛛第2次交配的交配潜伏期和交配持续时间显著长于第1次交配。同时,C组的三突伊氏蛛第1次交配的交配潜伏期和交配持续时间与第2次交配都没有显著差异。同时,A、B和C组的三突伊氏蛛第1次交配的交配回合数与第2次交配都没有显著差异。研究结果支持精子竞争强度假说,而不支持柯立芝效应。     

No reversion to single mating in a socially parasitic ant

Social Hymenoptera are ideal biological models for the study of the selective forces affecting the evolution of multiple mating (polyandry), because sister species can evolve different lifestyles and mating strategies. Single mating is predicted in workerless social parasites, because the key benefit of multiple mating in social insects, that is, the increase in genetic diversity among worker offspring, does not hold for workerless species. We compared the queen mating frequency between the ant Plagiolepis pygmaea and its derived social parasite P. xene. Previous studies showed that queens of the host P. pygmaea are obligately polyandrous. Here, pedigree analyses of mother–offspring combinations indicate that queens of the parasite P. xene did not revert to single mating; more than 50% of queens mated multiply, with 2–4 males. This result shows that reversal from multiple to single mating may be not selected in polyandrous social insect workerless parasites. We propose that such reversion does not occur when multiple mating is virtually cost free.     

Interspecific recognition in Chilean parabiotic ant species

Summary In Chile, Camponotus morosus and Solenopsis gayi sometimes co-inhabit a single nest, seemingly in a parabiotic association. To elucidate the nature of this association we conducted behavioural tests that measured aggression between homo- and allospecific ants. These tests revealed that C. morosus was aggressive towards alien conspecific and allospecific ants, but tolerated allospecific individuals from the same parabiotic society as well as allospecific individuals from a different parabiotic colony. In contrast S. gayi was much more tolerant towards alien ants whether homo- or allospecific and irrespective of their colony of origin, parabiotic or non-parabiotic.Chemical analyses showed that each species possess a distinct cuticular hydrocarbons profile. Moreover, each species tended to keep its specific profile even when living in association with the other species, in spite of very little acquired allospecific chemicals in C. morosus, dismissing chemical mimicry as the basis of the peaceful co-existence. We hypothesise that the switch from aggression to tolerance as a consequence of parabiotic association exhibited by C. morosus is due to a familiarisation as well as memorisation of the allospecific colonial odour. The finding that the parabiotic C. morosus was tolerant to S. gayi, even if they originated from a non-parabiotic nest points to an odour generalisation ability in this species.Received 12 July 2002; revised 15 January and 11 April 2003; accepted 23 April 2003.     

Cuticular Hydrocarbons of Reticulitermes (Isoptera: Rhinotermitidae) from Northern California Indicate Undescribed Species

Cuticular hydrocarbon mixtures of Reticulitermes samples from disparate locations in northern California were characterized and correlated with species determinations. Literature records indicate that there are two extant species of Reticulitermes in northern California: Reticulitermes hesperus Banks and R. tibialis Banks. We identified five distinct hydrocarbon phenotypes in Reticulitermes from northern California. Termites were diagnosed morphologically as R. tibialis for all five cuticular hydrocarbon phenotypes based on available keys to soldiers. Commonly accepted distributions of Reticulitermes in northern California indicate that our primary collection sites should only produce R. hesperus. On the basis of cuticular hydrocarbons alone we suggest that there are two or more undescribed taxa of Reticulitermes in northern California, or that described species of Reticulitermes occur in northern California, but have never been noticed before. Therefore, we conclude that the taxonomy of Reticulitermes in North America is in need of revision.     

Body size,male aggression,and male mating success in the cottonwood borer,Plectrodera scalator (Coleoptera: Cerambycidae)

The natural history and mating system ofPlectrodera scalator exhibit several unusual characteristics. Larvae and adults feed on the wood and foliage, respectively, of the same plant,Populus deltoides. The population sex ratio, based on censuses of oviposition areas, is female-biased. Females are significantly larger than males, yet males are intensely aggressive. Larger males tend to win escalated battles, which involve grasping of antennae with mandibles, but smaller males can defeat larger males if they grasp their opponent's antenna first. Most escalated fights involve possession of a female, but prior possession does not play a role in determining the outcome of these fights. The size-dependent fighting advantage does not translate into a mating advantage for larger males. There is no significant difference in elytron length or body mass between mating and single males. Larger females are not preferred as mates. The mating system appears to be a mixture of female-defense and scramble-competition tactics. One advantage to males of aggression may be in its effect on sperm precedence. Males appear to be able to remove previously deposited sperm from a female's reproductive tract.     

Size and mating success in males of the western harvester ant,Pogonomyrmex occidentalis (Hymenoptera: Formicidae)

Mating success in males of the lek mating ant species,Pogonomyrmex occidentalis, increases with increased body size. We estimated the magnitude of the selection coefficients on components of size by collecting males in copula and comparing their morphology to that of males that were collected at the lek but that were not mating. Four characters, body mass, head width, wing length, and leg length, were measured for a sample of 225 mating and 324 nonmating males and 225 females. Significant direct selection favors increased wing length and leg length. Multiple regression of transformed variables (principal components) indicated that the increased mating success of larger males is a function of all four characters. We found no evidence of positive assortative mating on the basis of any individual character or on the multivariate general size variable (the first principal component).