Benefits of foundress associations in the paper wasp Polistes dominulus: increased productivity and survival, but no assurance of fitness returns

Successful Polistes dominulus nests can be started by one ormore nest founding queens (foundresses). Consequently, thereis much interest in the specific benefits that induce cooperationamong foundresses. Here, we experimentally demonstrate one majorbenefit of cooperation, namely that multiple foundresses increasecolony productivity. This increase is close to the value predictedby subtracting the productivity of undisturbed single-foundresscolonies from the productivity of undisturbed multiple-foundresscolonies. However, we found no evidence that an associatingfoundress' contribution to colony growth is preserved if shedisappears (assured fitness returns). Our correlational datasuggest that cooperation provides survival benefits, multiple-foundresscolonies are more likely to survive to produce offspring thanare single-foundress colonies, and individual foundresses inmultiple-foundress groups are less likely to disappear beforeworker emergence than foundresses nesting alone. Therefore,association provides substantial productivity and survival benefitsfor cooperating foundresses.     

Colony success of the submissive ant Formica fusca within territories of the dominant Formica polyctena

Abstract. 1. Twenty-three nests of the submissive ant Formica fusca L. were sampled in two adjacent territories of the dominant wood ant Formica polyctena Först. The nests were dug up at different distances from the wood-ant mounds. Distance is assumed to be inversely related to the extent of disturbance of F. fusca by F. polyctena.
2. Colony success of F. fusca was assayed by counting the numbers of workers (colony size), and worker and sexual offspring in a nest. For individual size, the head width and dry weight of fifty workers per nest were measured.
3. Colony size correlated positively with the numbers of worker and sexual offspring in one territory, but only with worker offspring in the other.
4. Distance from the wood-ant mound correlated positively with colony size and numbers of worker and female offspring in one territory. In the other territory distance correlated positively with number of females, but negatively with number of worker offspring. No significant correlations between distance and the size measurements were obtained.
5. Because of its nonaggressive behaviour F. fusca may nest fairly close to a wood-ant mound but is likely to show reduced abundances in terms of nest density and forager number, and, ultimately, lowered fitness.     

Adaptations to sexual selection and sexual conflict: insights from experimental evolution and artificial selection

Artificial selection and experimental evolution document natural selection under controlled conditions. Collectively, these techniques are continuing to provide fresh and important insights into the genetic basis of evolutionary change, and are now being employed to investigate mating behaviour. Here, we focus on how selection techniques can reveal the genetic basis of post-mating adaptations to sexual selection and sexual conflict. Alteration of the operational sex ratio of adult Drosophila over just a few tens of generations can lead to altered ejaculate allocation patterns and the evolution of resistance in females to the costly effects of elevated mating rates. We provide new data to show how male responses to the presence of rivals can evolve. For several traits, the way in which males responded to rivals was opposite in lines selected for male-biased, as opposed to female-biased, adult sex ratio. This shows that the manipulation of the relative intensity of intra- and inter-sexual selection can lead to replicable and repeatable effects on mating systems, and reveals the potential for significant contemporary evolutionary change. Such studies, with important safeguards, have potential utility for understanding sexual selection and sexual conflict across many taxa. We discuss how artificial selection studies combined with genomics will continue to deepen our knowledge of the evolutionary principles first laid down by Darwin 150 years ago.     

Two alternate mechanisms contribute to the persistence of interdependent lineages in Pogonomyrmex harvester ants

Some populations of Pogonomyrmex harvester ants comprise pairs of highly differentiated lineages with queens mating at random with several males of their own and of the alternate lineage. These queens produce two types of diploid offspring, those fertilized by males of the queens' lineage which develop into new queens and those fertilized by males of the other lineage which mostly develop into functionally sterile workers. This unusual mode of genetic caste determination has been found in 26 populations and a total of four lineage pairs (F(1)-F(2), G(1)-G(2), H(1)-H(2) and J(1)-J(2)) have been described in these populations. Despite the fact that a few interlineage queens are produced, previous studies revealed that there is a complete lack of genetic introgression between lineages. Here we quantify the proportion of interlineage queens produced in each of the four lineage pairs and determine the fate of these queens. In the F(1)-F(2), G(1)-G(2) and H(1)-H(2) lineage pairs, interlineage queens were produced by a minority of colonies. These colonies exclusively produced interlineage queens and workers, suggesting that interlineage eggs can develop into queens in these three pairs of lineages in the absence of competition with pure-lineage brood. An analysis of three key stages of the colony life cycle revealed that colonies headed by interlineage queens failed to grow sufficiently to produce reproductive individuals. In laboratory comparisons, interlineage queens produced fewer viable eggs, with the effect that they raised fewer workers and lost more weight per worker produced than pure-lineage queens. In the J(1)-J(2) lineage pair, we did not find a single interlineage queen, raising the possibility that interlineage eggs have completely lost the ability to develop into queens in this lineage pair. Hence, two distinct mechanisms seem to account for the complete lack of between-lineage gene flow in the F(1)-F(2), G(1)-G(2), H(1)-H(2) and J(1)-J(2) lineage pairs.     

Will male advertisement be a reliable indicator of paternal care,if offspring survival depends on male care?

Existing theory predicts that male signalling can be an unreliable indicator of paternal care, but assumes that males with high levels of mating success can have high current reproductive success, without providing any parental care. As a result, this theory does not hold for the many species where offspring survival depends on male parental care. We modelled male allocation of resources between advertisement and care for species with male care where males vary in quality, and the effect of care and advertisement on male fitness is multiplicative rather than additive. Our model predicts that males will allocate proportionally more of their resources to whichever trait (advertisement or paternal care) is more fitness limiting. In contrast to previous theory, we find that male advertisement is always a reliable indicator of paternal care and male phenotypic quality (e.g. males with higher levels of advertisement never allocate less to care than males with lower levels of advertisement). Our model shows that the predicted pattern of male allocation and the reliability of male signalling depend very strongly on whether paternal care is assumed to be necessary for offspring survival and how male care affects offspring survival and male fitness.     

Alarm-recruitment behaviour in Pheidole militicida (Hymenoptera: Formicidae)

ABSTRACT.

• 1 Pheidole militicida Wheeler, a seed-harvesting species of the southwestern United States, possesses a major worker caste (soldiers) with unusually large heads. Previous work suggested that these large major workers are specialized defenders against large seed-harvesting species in the genus Pogonomyrmex.
• 2 Experimental introductions of Pogonomyrmex maricopa Wheeler and the omnivorous fire ant, Solenopsis xyloni McCook, demonstrate that P. militicida colonies alarm-recruit major workers against S.xyloni workers but not against the larger Pogonomyrmex workers.
• 3 P.militicida is the second species in which enemy-specific defence against Solenopsis fire ants has been detected. I suggest that the special role of Solenopsis as both competitor and predator in ant communities frequently favours major worker specialization against this genus.
• 4 Major workers of P.militicida and Pheidole dentata Mayr, the other species with enemy-specific defence against Solenopsis, are morphologically very different from one another. It is suggested that interspecific variation in major worker morphology is not necessarily associated with variation in behavioural specialization.

     

Size‐dependent resource allocation and sex allocation in herbaceous perennial plants

Individuals within a population often differ considerably in size or resource status as a result of environmental variation. In these circumstances natural selection would favour organisms not with a single, genetically determined allocation, but with a genetically determined allocation rule specifying allocation in relation to size or environment. Based on a graphical analysis of a simple evolutionarily stable strategy (ESS) model for herbaceous perennial plants, we aim to determine how cosexual plants within a population should simultaneously adjust their reproductive allocation and sex allocation to their size. We find that if female fitness gain is a linear function of resource investment, then a fixed amount of resources should be allocated to male function, and to post‐breeding survival as well, for individuals above a certain size threshold. The ESS resource allocation to male function, female function, and post‐breeding survival positively correlate if both male and female fitness gains are a saturating function of resource investment. Plants smaller than the size threshold are expected to be either nonreproductive or functionally male only.     

Rock selection for nesting inProformica longiseta Collingwood, 1978 (Hymenoptera,Formicidae) in a Mediterranean high mountain

Proformica longiseta, an ant species endemic to certain high mountains, has been studied with regard to its tendency to nest underneath rocks. This behaviour led us to investigate whether a particular type of rock is selected, and, if so, the advantages to this species of the selection. This study was located at 2400 m above sea-level in the Sierra Nevada mountains (Granada, Spain). To determine the range of rock sizes available, in eight control plots the thicknesses were recorded and also the maximum and minimum diameters of all rocks with surface area of more than 6 cm². A total of 1724 rocks were recorded, noting among the rock types available those that were used for nesting. Also as a preliminary estimate of the thermal characteristics of the rocks, we recorded the temperatures of the air, soil surface, soil under a rock covering an ant nest, and soil at 30 cm depth. The results indicate thatP. longiseta selects flat rocks with a surface area/thickness ratio of 2–5, the thickness being not less than 1.5 cm, the surface area being 50–250 cm² and volume less than 1000 cm³. This type of rock offers the nests two beneficial but apparently contradictory effects: (1) protection against high temperatures on hot days, and (2) warmth on colder days by absorbing the diffusing solar heat more effectively than the air or soil around the nest.     

Polyandry in two South American harvester ants

Although monandry (single mating) is the ancestral state in social hymenopteran insects, effective mating frequencies greater than 2 have been confirmed for a fair amount of ant species: Cataglyphis cursor, the leaf-cutters of the genera Atta and Acromyrmex, army ants of the genera Eciton, Dorylus, Aenictus and Neivamyrmex, and some North American seed harvester species of the genus Pogonomyrmex. This last genus spreads throughout open arid habitats from Patagonia to southwestern Canada. Whereas some North American Pogonomyrmex species are thoroughly studied, we know much less about these ants in South America. The objective of this study was to estimate the effective mating frequency of Pogonomyrmex inermis and P. pronotalis, two Pogonomyrmex sensu stricto species from the central Monte desert of Argentina. A total of 477 P. pronotalis workers from 24 colonies and 402 P. inermis workers from 20 colonies were analyzed using six and four highly polymorphic microsatellites, respectively. The multilocus analysis revealed that all colonies were monogynous and all queens multiply-mated. The effective mating frequency was 8.75 and 6.52 for queens of P. pronotalis and P. inermis, respectively; those values increased up to 15.66 and 9.78, respectively, when corrected for sampling errors. This is the first demonstration that queens in at least some members of the South American Pogonomyrmex sensu stricto are strictly polyandrous, with mating numbers per queen at least as high as those previously found for North American species. We suggest that multiple mating probably arose early in the evolution of the genus Pogonomyrmex and may be the basis of its ecological success and wide distribution. Received 11 October 2006; revised 10 August 2007 and 19 November 2007; accepted 21 November 2007.     

Frequency- and density-dependent sexual selection in natural populations of Galician Littorina saxatilis Olivi

Galician exposed shore populations of the direct developing periwinkle Littorina saxatilis are strikingly polymorphic, with an ornamented and banded upper shore form and a smooth and unbanded lower shore form. Intermediates between the two pure forms occur in a narrow mid shore zone together with the parental forms. We have previously shown that the two pure forms share the same gene pool but that mating between them is non-random. This is due to a non-random microdistribution in the zone of overlap, and also to assortative mating. In this study we present data which show that intermediate (hybrid) females mate less often than pure females in micropatches dominated by either of the pure forms, but not in micropatches in which the two pure forms are equally common. Thus, sexual fitness in intermediate females depends on the frequency of both pure morphs. Furthermore, sexual selection against intermediate females also varies with the densities of snails within each micro patch. The biological mechanisms which may explain this particular reduction of female hybrid fitness are discussed. Assortative mating between the pure morphs is sometimes almost complete, while both morphs do not mate the intermediates assortatively. In the light of this, sexual selection against intermediate females may contribute considerably to restrict gene flow between the pure forms.     

The formal Darwinism project: a mid-term report

For 8 years I have been pursuing in print an ambitious and at times highly technical programme of work, the 'Formal Darwinism Project', whose essence is to underpin and formalize the fitness optimization ideas used by behavioural ecologists, using a new kind of argument linking the mathematics of motion and the mathematics of optimization. The value of the project is to give stronger support to current practices, and at the same time sharpening theoretical ideas and suggesting principled resolutions of some untidy areas, for example, how to define fitness. The aim is also to unify existing free-standing theoretical structures, such as inclusive fitness theory, Evolutionary Stable Strategy (ESS) theory and bet-hedging theory. The 40-year-old misunderstanding over the meaning of fitness optimization between mathematicians and biologists is explained. Most of the elements required for a general theory have now been implemented, but not together in the same framework, and 'general time' remains to be developed and integrated with the other elements to produce a final unified theory of neo-Darwinian natural selection.     

Budding dispersal and the sex ratio

There is much interest in understanding how population demography impacts upon social evolution. Here, we consider the impact of rate and pattern of dispersal upon a classic social evolutionary trait--the sex ratio. We recover existing analytical results for individual dispersal, and we extend these to allow for budding dispersal. In particular, while a cancelling of relatedness and kin competition effects means that the sex ratio is unaffected by the rate of individual dispersal, we find that a decoupling of relatedness and kin competition means that budding dispersal favours increasingly female-biased sex ratios. More generally, our analysis illustrates the relative ease with which biological problems involving class structure can be solved using a kin selection approach to social evolution theory.