The ant genus Aenictus from Laos,with description of a new species (Hymenoptera: Formicidae: Aenictinae)

Ants of the genus Aenictus are recorded from Laos for the first time. Here, we report six species including a new species and a new status: Aenictus binghami Forel, A. cf. dentatus Forel, A. doydeei Jaitrong et Yamane, sp. nov., A. cf. fuchuanensis Zhou, A. hodgsoni Forel, stat. nov., and A. nishimurai Terayama et Kubota. A. doydeei sp. nov. is very probably a nocturnal species.     

Losses of workers and reproductives in colonies of the paper waspPolistes riparius (Hymenoptera: Vespidae) due to the parasitic waspLatibulus sp.

1. Losses in workers and reproductives due to the ichneumonid parasitoid Latibulus sp. were determined in a haplometrotic, temperate paper wasp, Polistes riparius, during a 5-year study.
2. Fifty-four to 79% of pre-emergence nests were parasitized on worker brood, resulting in a worker loss of 25–31% on average. Worker brood reared in outer cells were much more often parasitized than those in central ones, possibly because outer cells are more accessible to the parasitoid. Infestation of worker brood was not random but aggregated among nests.
3. Seventy-eight to 100% of nests were parasitized on reproductive brood, and lost 10–34% of reproductives.
4. The number of emerged workers positively correlated with that of reproductives produced and that of cells made during the season. This suggests that worker loss reduces reproductive output of colonies. Hence, the parasitoid can reduce colonial reproductive output not only by killing reproductive brood but by reducing worker force.

     

Review of the Southeast Asian species of the Aenictus javanus and Aenictus philippinensis species groups (Hymenoptera, Formicidae, Aenictinae)

The Southeast Asian species of the Aenictus javanus and Aenictus philippinensis groups are revised. Six species (four named and two new species) of the Aenictus javanus group occurring in this area are: Aenictus doydeei Jaitrong & Yamane, 2011, Aenictus duengkaei Jaitrong & Yamane, sp. n., Aenictus javanus Emery, 1896, Aenictus longinodus Jaitrong & Yamane, sp. n., Aenictus nishimurai Terayama & Kubota, 1993, and Aenictus piercei Wheeler & Chapman, 1930. Four species (three named and one new species) are recognized in the Aenictus philippinensis group: Aenictus pangantihoni Zettel & Sorger, 2010, Aenictus philippinensis Chapman, 1963, Aenictus punctatus Jaitrong & Yamane, sp. n., and Aenictus rabori Chapman, 1963. Aenictus piercei is removed from the members of the Aenictus piercei group sensu Jaitrong and Yamane (2011) and transferred to the Aenictus javanus group. Lectotypes and paralectotypes are designated for Aenictus piercei and Aenictus rabori. Size variation occurs among individuals from single colonies of the Aenictus javanus group, while the workers in the Aenictus philippinensis group are clearly monomorphic.     

Application of Microsatellite Primers Developed for Polistes in the Independent-Founding Wasp Polists satan Bequaert (Hymenoptera: Vespidae)

Microsatellite primers developed for a given species are sometimes useful for another in the same genus and in other genera within the same family, making possible to search for pre-existing suitable primers in the databanks such as GenBank. We examined whether existing primers developed for Polistes could be used for Polistes satan Bequaert. We tested 50 microsatellite primers from three Polistes species and found that six microsatellite loci show polymorphism in size in P. satan. These six loci were highly polymorphic, having four to 15 alleles in P. satan with an expected heterozygosity of 0.525?C0.832. These loci can be used to study parameters concerning genetic relatedness such as social interactions in colonies and genetic conflicts of interest among nestmate individuals.     

A new species of the Aenictus wroughtonii group (Hymenoptera,Formicidae) from South-East China

A new species of army ant from the Aenictus wroughtonii group is described and illustrated based on the worker caste. Aenictus gutianshanensis Staab, sp. n. is known form a single colony collected in the subtropical mixed evergreen broad-leaved forest of the Gutianshan National Nature Reserve, South-East China. The new species is probably most closely related to A. vieti Jaitrong & Yamane, 2010 known from North Vietnam and Taiwan. It is suggested that the abundant records of A. camposi Wheeler & Chapman, 1925 from East and South-East China should be reevaluated, as they are probably A. gutianshanensis or A. vieti and not A. camposi, which is distributed in Sundaland, the Philippines, and the southernmost part of continental South-East Asia.     

Selection for individual recognition and the evolution of polymorphic identity signals in Polistes paper wasps

Individual recognition (IR) requires individuals to uniquely identify their social partners based on phenotypic variation. Because IR is so specific, distinctive phenotypes that stand out from the crowd facilitate efficient recognition. Over time, the benefits of unique appearances are predicted to produce a correlation between IR and phenotypic variation. Here, we test whether there is an association between elevated phenotypic polymorphism and IR in paper wasps. Previous work has shown that Polistes fuscatus use variable colour patterns for IR. We test whether two less variable wasp species, Polistes dominulus and Polistes metricus, are capable of IR. As predicted, neither species is capable of IR, suggesting that highly variable colour patterns are confined to Polistes species with IR. This association suggests that elevated phenotypic variation in taxa with IR may be the result of selection for identity signals rather than neutral processes. Given that IR is widespread among social taxa, selection for identity signalling may be an underappreciated mechanism for the origin and maintenance of polymorphism.     

Trinucleotide Microsatellite Loci and Increased Heterozygosity in Cross-Species Applications in the Social Wasp, Polistes

Though microsatellite loci are usually found to be most polymorphic in the species in which they are first identified, we have found significant increases in polymorphisms in some cross-species applications. We present eight new trinucleotide microsatellite loci derived from two species of social wasps, Polistes annularis and Polistes bellicosus. We assessed the primers designed from these species and the degree of polymorphism in two additional species, P. dorsalis, which is very closely related to P. bellicosus, and P. dominulus, which is an Old World congener, thought to have diverged from New World Polistes over 80 million years ago. Cross-species applications for these microsatellite loci indicate that the priming sites from P. bellicosus loci are conserved in P. dorsalis and amplified similarly sized products with higher heterozygosities than the original species in two of three cases. A locus that was monomorphic in P. annularis had a heterozygosity of 1.0 in the distantly related P. dominulus. Cross-species applications of these loci indicated that alleles were generally of similar lengths in the new and original species when they retained their heterozygosity.     

The morphology of Van der Vecth's organ as a tool to measure caste dimorphism in Polistes paper wasps: a comparative approach

Given the centrality of chemical communication in social insects, there are many selective pressures acting on morpho‐functional traits that mediate chemical pheromones. On the last gastral sternite of Polistes females, there is an important exocrine surface secreting chemical pheromone, named Van der Vecht's organ. It is involved in chemical defence of the nest, in rank and nestmate recognition, preventing workers from direct reproduction. Allometric differential growth of phenotypic traits between castes of social insects is generally considered as an indication of incipient physical castes. European Polistes present different nesting strategies and reproductive choices. Here, we carry out a comparison of Van der Vecht's organ size between castes of four European Polistes to provide a general measure of dimorphism. We show that Van der Vecht's organ of Polistes dominula and Polistes nimphus foundresses shows an allometric development being enlarged with respect to workers. Otherwise, no allometries have been highlighted in the other two studied species (i.e. Polistes associus and Polistes biglumis). Therefore, our data show that neither rigid monogyny nor specific nesting habits foster the evolution of true morphological castes in primitively eusocial taxa. Thus, at least two other species of Ezuropean Polistes show real evidence of incipient morphological castes.     

Hormonal modulation of reproduction and fertility signaling in polistine wasps

In social insects, it has been suggested that reproduction and the production of particular fertility-linked cuticular hydrocarbons (CHC) may be under shared juvenile hormone (JH) control, and this could have been key in predisposing such cues to later evolve into full-fledged queen pheromone signals. However, to date, only few studies have experimentally tested this “hormonal pleiotropy” hypothesis. Here, we formally test this hypothesis using data from four species of Polistine wasps, Polistes dominula, Polistes satan, Mischocyttarus metathoracicus, and Mischocyttarus cassununga, and experimental treatments with JH using the JH analogue methoprene and the anti-JH precocene. In line with reproduction being under JH control, our results show that across these four species, precocene significantly decreased ovary development when compared with both the acetone solvent-only control and the methoprene treatment. Consistent with the hormonal pleiotropy hypothesis, these effects on reproduction were further matched by subtle shifts in the CHC profiles, with univariate analyses showing that in P. dominula and P. satan the abundance of particular linear alkanes and mono-methylated alkanes were affected by ovary development and our hormonal treatments. The results indicate that in primitively eusocial wasps, and particularly in Polistes, reproduction and the production of some CHC cues are under joint JH control. We suggest that pleiotropic links between reproduction and the production of such hydrocarbon cues have been key enablers for the origin of true fertility and queen signals in more derived, advanced eusocial insects.     

POPULATION STRUCTURE AND KINSHIP IN POLISTES (HYMENOPTERA,VESPIDAE): AN ANALYSIS USING RIBOSOMAL DNA AND PROTEIN ELECTROPHORESIS

Six variable protein loci and one variable ribosomal DNA restriction site were used for an analysis of population structure in five species of Polistes from Texas. A sample-reuse algorithm was developed that estimated F_ST, F_IS, and ø (the coefficient of kinship) from probabilities of identity. Of the four species analyzed in detail only one, Polistes exclamans, had statistically significant values of F_ST. These values may reflect natural constraints on successful nesting for migrants of this species. Three of the four species had significant values of F_IS and three of the four species had significant values of ø. In many cases ø also differed from the expected value under haplodiploidy and random mating. Values of ø did not differ from expectations under haplodiploidy and local inbreeding. These results emphasize that theories of social behavior and evolution based on coefficients of kinship should include some explicit consideration of population structure.     

Polymorphic microsatellite loci for the Japanese paper wasp,Polistes chinensis antennalis (Hymenoptera: Vespidae)

We isolated 13 microsatellite markers for the Japanese paper wasp Polistes chinensis antennalis. Null alleles were ruled out in the loci because we could detect polymerase chain reaction products for 578 haploid males. Our results indicate that these markers are excellent tools for the analysis of pedigrees within colonies for species with a complicated pedigree structure due to worker reproduction (male production by workers in queen‐right colonies).     

Persistence of occluded viruses in the nests of the paper waspPolistes hebraeus (Hym.: Vespidae)

Bioassays were used to study the infectivity of cytoplasmic polyhedrosis viruses (CPVs), nuclear polyhedrosis viruses (NPVs) and entomopoxviruses (EPVs) contained in 16 nests of the paper waspPolistes hebraues F. in Réunion Island. Several virosis were propagated from 6 nest contents in 5 Lepidoptera:Catopsilia thauruma Saalmüller,Catopsilia florella F.,Callixena versicolora Mabille,Polydesma umbricola Boisduval andEagris sabadius Boisduval. Each of the previous species supported the development of one or two virosis and therefore, wasp nests must be considered as accumulating centres where infectivity of occluded entomoviruses is preserved. The accurate origin of the virosis propagated through bioassays was searched for using ecological investigations on similar natural diseases, REN analysis and a cross-transmission test onSpodoptera mauritia Boisduval. The nuclear polyhedrosis and one cytoplasmic polyhedrosis appearing in bioassays can be assigned respectively to viruses produced byC. thauruma andC. versicolora; the other virosis must be considered as developing in alternate or substitution hosts. Wasp nests could therefore be used to detect the presence of specific viruses in an environment and to collect new virus isolates.     

Individual recognition is associated with holistic face processing in Polistes paper wasps in a species-specific way

Most recognition is based on identifying features, but specialization for face recognition in primates relies on a different mechanism, termed ‘holistic processing’ where facial features are bound together into a gestalt which is more than the sum of its parts. Here, we test whether individual face recognition in paper wasps also involved holistic processing using a modification of the classic part-whole test in two related paper wasp species: Polistes fuscatus, which use facial patterns to individually identify conspecifics, and Polistes dominula, which lacks individual recognition. We show that P. fuscatus use holistic processing to discriminate between P. fuscatus face images but not P. dominula face images. By contrast, P. dominula do not rely on holistic processing to discriminate between conspecific or heterospecific face images. Therefore, P. fuscatus wasps have evolved holistic face processing, but this ability is highly specific and shaped by species-specific and stimulus-specific selective pressures. Convergence towards holistic face processing in distant taxa (primates, wasps) as well as divergence among closely related taxa with different recognition behaviour (P. dominula, P. fuscatus) suggests that holistic processing may be a universal adaptive strategy to facilitate expertise in face recognition.     

Trophic relations between vespid wasps (Hymenoptera,Vespidae) and flowering plants in the Crimea

The results of the study of trophic relations between nectar-feeding adult wasps and flowering plants in the Crimea are given. In 2002–2009, a total of 55 wasp species were observed on flowers of 74 plant species belonging to 25 families. The highest number of plant species was visited by Polistes dominulus (36), Ancistrocerus nigricornis (18), Polistes nimphus (16), P. gallicus (13) and Dolichovespula sylvestris (13); these plants belonged to the families Apiaceae (15), Lamiaceae (10), Scrophulariaceae (8), Rosaceae (8) and Asteraceae (7). Plants of the families Scrophulariaceae, Apiaceae, Lamiaceae, Rosaceae, and Fabaceae were visited by 25, 24, 22, 11, and 11 wasp species, respectively. Other plant families were visited by 10 or less wasp species. The highest number of wasp species were observed on flowers of Scrophularia umbrosa (22), Mentha longifolia (12), Bupleurum fruticosum (10), Laserpitium hispidum (10), Koelreuteria paniculata (10), Sambucus ebulus (10), Dorycnium herbaceum (9), Foeniculum vulgare (8), Limonium platyphyllum (7), Torilis arvensis (7), Marrubium peregrinum (7), Butomus umbellatus (7), Eryngium campestre (6), Cynanchum acutum (6) and Scrophularia scopolii (6). One to five wasp species were recorded on flowers of plants belonging to other families. Two wasp species (Polistes nimphus and Vespula germanica) were registered as visitors of the nectarless flowers of the orchid Steveniella satyrioides.     

Species and Colony Components in the Recognition Odor of Young Social Wasps: Their Expression and Learning (Polistes biglumis and P. atrimandibularis; Hymenoptera: Vespidae)

In Polistes, nestmate recognition relies on the learning of recognition cues from the nest. When wasps recognize nestmates, they match the template learned with the odor of the encountered wasp. The social wasp Polistes biglumis use the homogeneous odor of their colony to recognize nestmates. When these colonies become host colonies of the social parasite P. atrimandibularis, colony odor is no longer homogeneous, as the parasite offspring have an odor that differs from that of their hosts. In trying to understand how the mechanism of nestmate recognition works in parasitized colonies and why parasite offspring are accepted by hosts, we tested the responses of resident Polistes biglumis wasps from parasitized and unparasitized colonies to newly emerged parasites and to nestmate and non-nestmate conspecifics. The experiments indicate that immediately upon eclosion both young parasites and young hosts lack a colony odor and that colony odor can be soon acquired from the accepting colony. In addition, while residents of nonparasitized colonies recognize only the odor of their species, resident hosts of parasitized colonies have learned a template that fits the odors of two species.     

The effect of juvenile hormone on Polistes wasp fertility varies with cooperative behavior

Social insects provide good models for studying how and why the mechanisms that underlie reproduction vary, as there is dramatic reproductive plasticity within and between species. Here, we test how the effect of juvenile hormone (JH) on fertility covaries with cooperative behavior in workers and nest-founding queens in the primitively eusocial wasp Polistes metricus. P. metricus foundresses and workers appear morphologically similar and both are capable of reproduction, though there is variation in the extent of social cooperation and the probability of reproduction across castes. Do the endocrine mechanisms that mediate reproduction co-vary with cooperative behavior? We found dramatic differences in the effect of JH on fertility across castes. In non-cooperative nest-founding queens, all individuals responded to JH by increasing their fertility. However, in cooperative workers, the effect of JH on fertility varies with body weight; large workers increase their fertility in response to JH while small workers do not. The variation in JH response may be an adaptation to facilitate resource allocation based on the probability of independent reproduction. This work contrasts with previous studies in closely related Polistes dominulus paper wasps, in which both foundresses and workers form cooperative associations and both castes show similar, condition-dependent JH response. The variation in JH responsiveness within and between species suggests that endocrine responsiveness and the factors influencing caste differentiation are surprisingly evolutionarily labile.     

A ten-year comparative study of the population dynamics and parasitoidism in the native paper wasp Polistes fuscatus and the invasive P. dominulus

The invasive success of Polistes dominulus in North America has been attributed to its greater productivity relative to native Polistes. Liberation from parasites and parasitoids are thought to be major factors contributing to the high productivity of P. dominulus. We analyzed historical records of colony relative abundance and productivity of P. dominulus and the sympatric, native Polistes fuscatus from 1995 to 2010 using historical data from our Michigan Polistes study site. We also analyzed evidence of parasitoids from 294 P. fuscatus and 507 P. dominulus archived combs from 2001 to 2010. Additionally, we examined field and laboratory colonies from outside of our study site for parasites and parasitoids in 2009 and 2010. We documented one parasite and three parasitoids exploiting Polistes in our Michigan study sites. Our historical records document that P. dominulus initially displaced P. fuscatus rapidly, then slowed, and finally the two populations stabilized. Furthermore, the historical pattern of decreasing displacement of P. fuscatus by P. dominulus corresponded temporally with a significant decline in the productivity and a significant increase in Dibrachys cavus infestation of P. dominulus. Our evidence indicates that the parasitoid, D. cavus, is a major factor in stabilizing the populations of the sympatric P. dominulus and P. fuscatus.     

Inhibition of Host Queen Reproductive Capacity by the Obligate Social Parasite Polistes atrimandibularis (Hymenoptera,Vespidae)

Polistes atrimandibularis is the obligate and permanent parasite of the social paper wasp Polistes biglumis bimaculatus. This parasite lives together with the host foundress for a considerable period on the invaded nest. Ovarian development was measured in females of the host species (foundresses and, when present, workers) from 18 parasitized and 14 non-parasitized colonies. The reproductive capacity of foundresses from parasitized nests decreased faster than that of foundresses from non-parasitized nests. These results indicate that the presence of the parasite lowers reproductive capacity of the host queen. Simultaneously, the fertility of the host worker is inhibited.