The evolution of imperfect mimicry in hoverflies

Ideas about the evolution of imperfect mimicry are reviewed. Their relevance to the colours patterns of hoverflies (Diptera, Syrphidae) are discussed in detail. Most if not all of the hoverflies labelled as mimetic actually are mimics. The apparently poor nature of their resemblance does not prevent them from obtaining at least some protection from suitably experienced birds. Mimicry is a dominant theme of this very large family of Diptera, with at least a quarter of all species in Europe being mimetic. Hoverfly mimics fall into three major groups according to their models, involving bumblebees, honeybees and social wasps. There are striking differences in the general levels of mimetic fidelity and relative abundances of the three groups, with accurate mimicry, low abundance and polymorphism characterizing the bumblebee mimics: more than half of all the species of bumblebee mimics are polymorphic. Mimics of social wasps tend to be poor mimics, have high relative abundance, and polymorphism is completely absent. Bumblebee models fall into a small number of Muellerian mimicry rings which are very different between the Palaearctic and Nearctic regions. Social wasps and associated models form one large Muellerian complex. Together with honeybees, these complexes probably form real clusters of forms as perceived by many birds. All three groups of syrphid mimics contain both good and poor mimics; some mimics are remarkably accurate, and have close morphological and behavioural resemblance. At least some apparently 'poor' mimetic resemblances may be much closer in birds' perception than we imagine, and more work needs to be done on this. Bumblebees are the least noxious and wasps the most noxious of the three main model groups. The basis of noxiousness is different, with bumblebees being classified as non-food, whereas honeybees and wasps are nasty-tasting and (rarely) stinging. The distribution of mimicry is exactly what would be expected from this ordering, with polymorphic and accurate forms being a key feature of mimics of the least noxious models, while highly noxious models have poor-quality mimicry. Even if the high abundance of many syrphid mimics relative to their models is a recent artefact of man-made environmental change, this does not preclude these species from being mimics. It seems unlikely that bird predation actually controls the populations of adult syrphids. Being rare relative to a model may have promoted or accelerated the evolution of perfect mimicry: theoretically this might account for the pattern of rare good mimics and abundant poor ones, but the idea is intrinsically unlikely. Many mimics seem to have hour-to-hour abundances related to those of their models, presumably as a result of behavioural convergence. We need to know much more about the psychology of birds as predators. There are at least four processes that need elucidating: (a) learning about the noxiousness of models; (b) the erasing of that learning through contact with mimics (extinction, or learned forgetting); (c) forgetting; (d) deliberate risk-taking and the physiological states that promote it. Johnston's (2002) model of the stabilization of imperfect mimicry by kin selection is unlikely to account for the colour patterns of hoverflies. Sherratt's (2002) model of the influence of multiple models potentially accounts for all the patterns of hoverfly mimicry, and is the most promising avenue for testing.

Ovarian development in a primitively eusocial wasp: Social interactions affect behaviorally dominant and subordinate wasps in opposite directions relative to solitary females

In many primitively eusocial wasp species new nests are founded either by a single female or by a small group of females. In the single foundress nests, the lone female develops her ovaries, lays eggs as well as tends her brood. In multiple foundress nests social interactions, especially dominance-subordinate interactions, result in only one `dominant' female developing her ovaries and laying eggs. Ovaries of the remaining `subordinate' cofoundresses remain suppressed and these individuals function as workers and tend the dominant's brood. Using the tropical, primitively eusocial polistine wasp Ropalidia marginata and by comparing wasps held in isolation and those kept as pairs in the laboratory, we demonstrate that social interactions affect ovarian development of dominant and subordinate wasps among the pairs in opposite directions, suppressing the ovaries of the subordinate member of the pair below that of solitary wasps and boosting the ovaries of dominant member of the pair above that of solitary females. In addition to being of physiological interest, such mirror image effects of aggression on the ovaries of the aggressors and their victims, suggest yet another mechanism by which subordinates can enhance their indirect fitness and facilitate the evolution of worker behavior by kin selection. (C) 2014 Elsevier B.V. All rights reserved.

Caste in the swarming wasps: 'queenless' societies in highly social insects

Morphometric data for 30 species of swarming wasps (Vespidae: Polistinae: Epiponini) are presented, representing all currently recognized genera. Data are coded according to whether females that were shown by dissection to be egglayers are larger, similar, or smaller for each dimension than non-egglayers. These data are analysed in a phylogenetic framework with primitively social Polistes and Mischocyttarus as outgroups. Representative measurements are illustrated to show that most genera of Epiponini appear to have ancestry in a lineage that has no queen caste comparable with either the primitively social outgroups, or the more derived species of the tribe. This analysis indicates that a conspiracy of workers that operates without a queen characterizes the societies of many Epiponini, or their recent ancestors. (c) 2008 The Linnean Society of London.

Checklist of social (polistinae) and solitary (eumeninae) wasps from a fragment of Cerrado campo Sujo in the State of Mato Grosso do sul

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Ants, bees, and wasps : a record of observations on the habits of the social hymenoptera /

Includes bibliographical references (p. [xvii]-xix) and index.

Ants, bees, and wasps : a record of observations on the habits of the social Hymenoptera /

"Lists of the principal books and memoirs referred to": p. [xvii]-xix.

Ants, bees, and wasps : a record of observations on the habits of the social hymenoptera /

Includes index and bibliographical references.

A comparative social network analysis of wasp colonies and classrooms: Linking network structure to functioning

A major question in current network science is how to understand the relationship between structure and functioning of real networks. Here we present a comparative network analysis of 48 wasp and 36 human social networks. We have compared the centralisation and small world character of these interaction networks and have studied how these properties change over time. We compared the interaction networks of (1) two congeneric wasp species (Ropalidia marginata and Ropalidia cyathiformis), (2) the queen-right (with the queen) and queen-less (without the queen) networks of wasps, (3) the four network types obtained by combining (1) and (2) above, and (4) wasp networks with the social networks of children in 36 classrooms. We have found perfect (100%) centralisation in a queen-less wasp colony and nearly perfect centralisation in several other queen-less wasp colonies. Note that the perfectly centralised interaction network is quite unique in the literature of real-world networks. Differences between the interaction networks of the two wasp species are smaller than differences between the networks describing their different colony conditions. Also, the differences between different colony conditions are larger than the differences between wasp and children networks. For example, the structure of queen-right R. marginata colonies is more similar to children social networks than to that of their queen-less colonies. We conclude that network architecture depends more on the functioning of the particular community than on taxonomic differences (either between two wasp species or between wasps and humans).

Assessment of prey overlap between a native (Polistes humilis) and an introduced (Vespula germanica) social wasp using morphology and phylogenetic analyses of 16S rDNA

In newly invaded communities, interspecific competition is thought to play an important role in determining the success of the invader and its impact on the native community. In southern Australia, the native Polistes humilis was the predominant social wasp prior to the arrival of the exotic Vespula germanica (Hymenoptera: Vespidae). Both species forage for similar resources (water, pulp, carbohydrate and protein prey), and concerns have arisen about potential competition between them. The aim of this study was to identify the protein foods that these wasps feed on. As many prey items are masticated by these wasps to the degree that they cannot be identified using conventional means, morphological identification was complemented by sequencing fragments of the mitochondrial 16S rRNA gene. GenBank searches using blast and phylogenetic analyses were used to identify prey items to at least order level. The results were used to construct complete prey inventories for the two species. These indicate that while P. humilis is restricted to feeding on lepidopteran larvae, V. germanica collects a variety of prey of invertebrate and vertebrate origin. Calculated values of prey overlap between the two species are used to discuss the implications of V. germanica impacting on P. humilis. Results obtained are compared to those gained by solely 'conventional' methods, and the advantages of using DNA-based taxonomy in ecological studies are emphasized.

Social evolution - Has nature ever rewound the tape?

Social insects such as ants, bees, wasps and termites exhibit extreme forms of altruism where some individuals remain sterile and assist other individuals in reproduction. Hamilton's inclusive fitness theory provides a powerful framework for investigating the evolution of such altruism. Using the paper wasp Ropalidia marginata, we have quantified and delineated the role of ecological, physiological, genetic and demographic factors in social evolution. An interesting feature of the models we have developed is their symmetry so that either altruism or selfishness can evolve, depending on the numerical values of various parameters. This suggests that selfish/solitary behaviour must occasionally re-emerge even from the eusocial state, It is useful to contemplate expected intermediate states during such potential reversals. We can perhaps envisage three successive steps in such a hypothetical process: i) workers revolt against the hegemony of the queen and challenge her status as the sole reproductive, ii) workers stop producing queens and one or more of them function as egg layers (functional queen/s) capable of producing both haploid as well as diploid offspring and iii) social evolution reverses completely so that a eusocial species becomes solitary, at least facultatively. It appears that the third step, namely transition from eusociality to the solitary state, is rare and has been restricted to transitions from the primitively eusocial state only. The absence of transitions from the highly eusocial state to the solitary state may be attributed to a number of 'preventing mechanisms' such as (a) queen control of workers (b) loss of spermathecae and ability to mate (c) morphological specialization (d) caste polyethism and (e) homeostasis, which must each make the transition difficult and, taken together, perhaps very difficult. However, the discovery of a transition from the highly eusocial to the solitary state can hardly he ruled out, given that little or no effort has gone into its detection. In this paper I discuss social evolution and its possible reversal and cite potential examples of stages in the transition from the social to the solitary.

Behavioural and morphological dimorphism of the sexes: an account of two primitively eusocial wasps

In social Hymenoptera, foraging, nest building, brood care and all other colony maintenance functions are carried out by the females while males function solely as reproductives. This asymmetry in social roles of the two sexes has led social insect researchers to focus almost exclusively on the females whereas males have remained relatively neglected. We studied two sympatric, primitively eusocial wasps, Ropalidia marginata and Ropalidia cyathiformis, and compared the morphological and behavioural profiles of males and females. Males of both species are smaller in size and weigh less compared to females. Males of the two species live in the nest for different durations. Borrowing from the ecological literature we use novel methods to compute and compare behavioural diversity and behavioural richness and show that females of both species are behaviourally richer and more diverse than the males.

Evolution of social behaviour in the primitively eusocial wasp Ropalidia marginata: do we need to look beyond kin selection?

Ropalidia marginata is a primitively eusocial wasp widely distributed in peninsular India. Although solitary females found a small proportion of nests, the vast majority of new nests are founded by small groups of females. In suchmultiple foundress nests, a single dominant female functions as the queen and lays eggs, while the rest function as sterile workers and care for the queen's brood. Previous attempts to understand the evolution of social behaviour and altruism in this species have employed inclusive fitness theory (kin selection) as a guiding framework. Although inclusive fitness theory is quite successful in explaining the high propensity of the wasps to found nests in groups, several features of their social organization suggest that forces other than kin selection may also have played a significant role in the evolution of this species. These features include lowering of genetic relatedness owing to polyandry and serial polygyny, nest foundation by unrelated individuals, acceptance of young non-nest-mates, a combination of well-developed nest-mate recognition and lack of intra-colony kin recognition, a combination of meek and docile queens and a decentralized self-organized work force, long reproductive queues with cryptic heir designates and conflict-free queen succession, all resulting in extreme intra-colony cooperation and inter-colony conflict.

Necrophagy by the social wasp Agelaia pallipes (Hymenoptera : Vespidae, Epiponini): Possible forensic implications

The occurrence of 62 specimens of Agelaia pallipes caught in carrion traps using three types of baits (fish, cow liver and poultry viscera) in three different types of environments (rural, urban and forest area) in seven municipalities in Southeastern Brazil is reported here. This specific necrophagic behavior is discussed, since investigations concerning carrion wasps are scant in literature.

Mass spectrometric characterization of two novel inflammatory peptides from the venom of the social wasp Polybia paulista

The social wasp P. paulista is relatively common in southeast Brazil causing many medically important stinging incidents. The seriousness of these incidents is dependent on the amount of venom inoculated by the wasps into the victims, and the characteristic envenomation symptoms are strongly dependent on the types of peptides present in the venom. In order to identify some of these naturally occurring peptides available in very low amounts, an analytical protocol was developed that uses a combination of reversed-phase and normal-phase high-performance liquid chromatography (HPLC) of wasp venom for peptide purification, with matrix-assisted laser desorption/ionization time-of-flight post-source decay mass spectrometry (MALDI-Tof-PSD-MS) and low-energy collision-induced dissociation (CID) in a quadrupole time-of-flight tandem mass spectrometry (QTof-MS/MS) instrument for peptide sequencing at the sub-picomole level. The distinction between Leu and Ile was achieved both by observing d-type fragment ions obtained under CID conditions and by comparison of retention times of the natural peptides and their synthetic counterparts (with different combinations of I and/or L at N- and C-terminal positions). To distinguish the isobaric residues K and Q, acetylation of peptides was followed by Q-Tof-MS analysis. The primary sequences obtained were INWLKLGKMVIDAL-NH2 (MW 1611.98Da) and IDWLKLGKMVMDVL-NH2 (MW 1658.98Da). Micro-scale bioassay protocols characterized both peptides as presenting potent hemolytic action, mast cell degranulation, and chemotaxis of poly-morphonucleated leukocyte (PMNL) cells. The primary sequences and the bioassay results suggest that these toxins constitute members of a new sub-class of mastoparan toxins, directly involved in the occurrence of inflammatory processes after wasp stinging. Copyright (C) 2004 John Wiley Sons, Ltd.