Occlusal relief changes with molar wear in Pan troglodytes troglodytes and Gorilla gorilla gorilla

Most research on primate tooth form-function relationships has focused on unworn teeth. This study presents a morphological comparison of variably worn lower second molars (M(2)s) of lowland gorillas (Gorilla gorilla gorilla; n=47) and common chimpanzees (Pan troglodytes troglodytes; n=54) using dental topographic analysis. High-resolution replicas of occlusal surfaces were prepared and scanned in 3D by laser scanning. The resulting elevation data were used to create a geographic information system (GIS) for each tooth. Occlusal relief, defined as the ratio of 3D surface area to 2D planometric area of the occlusal table, was calculated and compared between wear stages, taxa, and sexes. The results failed to show a difference in occlusal relief between males and females of a given taxon, but did evince differences between wear stages and between taxa. A lack of significant interaction between wear stage and taxon factors suggests that differences in occlusal relief between chimpanzees and gorillas are maintained throughout the wear sequence. These results add to a growing body of information on how molar teeth change with wear, and how differences between primate species are maintained at comparable points throughout the wear sequence. Such studies provide new insights into form-function relationships, which will allow us to infer certain aspects of diet in fossils with worn teeth.     

Measurements of tooth wear among Australian Aborigines: I. Serial loss of the enamel crown

The dental casts made from Aboriginal children during the course of a longitudinal growth study in Central Australia provided material for analyzing tooth wear under known environmental conditions. The wear facets produced on the occlusal surfaces were clearly preserved on the dental stone casts and recorded the progress of enamel attrition from ages 6 to 18. These casts were photographed and traced by electronic planimetric methods that automatically recorded the location and size of wear facets on the first and second permanent molars. These areas of worn tooth surface were compared to the total tooth surface. The worn surface was regressed on age to calculate wear rates of each tooth. Discriminant analyses were also performed to determine the significance of dental attrition differences between the sexes at each age group. The total wear on each tooth was highly correlated with age as expected but females wore their teeth at a significantly higher rate than males. The mandibular molars wore more rapidly than maxillary teeth in both sexes. The discriminant analysis successfully grouped 91% of the cases according to age and sex. Pattern of wear, the location, and size of wear facets also differed between age groups and sex. The questions of why there is a difference between male and female wear or why there is greater wear on one arch or arch region have no ready answers. The differing rates and pattern of dental wear do suggest that arch shape and growth rates may be the answer though it has yet to be tested. However, the occlusal surface wear is useful for age estimation in a population and provides a record of shifting masticatory forces during growth.     

Wear Fast,Die Young: More Worn Teeth and Shorter Lives in Iberian Compared to Scottish Red Deer

Teeth in Cervidae are permanent structures that are not replaceable or repairable; consequently their rate of wear, due to the grinding effect of food and dental attrition, affects their duration and can determine an animal''s lifespan. Tooth wear is also a useful indicator of accumulative life energy investment in intake and mastication and their interactions with diet. Little is known regarding how natural and sexual selection operate on dental structures within a species in contrasting environments and how these relate to life history traits to explain differences in population rates of tooth wear and longevity. We hypothesised that populations under harsh environmental conditions should be selected for more hypsodont teeth while sexual selection may maintain similar sex differences within different populations. We investigated the patterns of tooth wear in males and females of Iberian red deer (Cervus elaphus hispanicus) in Southern Spain and Scottish red deer (C. e. scoticus) across Scotland, that occur in very different environments, using 10343 samples from legal hunting activities. We found higher rates of both incisor and molar wear in the Spanish compared to Scottish populations. However, Scottish red deer had larger incisors at emergence than Iberian red deer, whilst molars emerged at a similar size in both populations and sexes. Iberian and Scottish males had earlier tooth depletion than females, in support of a similar sexual selection process in both populations. However, whilst average lifespan for Iberian males was 4 years shorter than that for Iberian females and Scottish males, Scottish males only showed a reduction of 1 year in average lifespan with respect to Scottish females. More worn molars were associated with larger mandibles in both populations, suggesting that higher intake and/or greater investment in food comminution may have favoured increased body growth, before later loss of tooth efficiency due to severe wear. These results illustrate how independent selection in both subspecies, that diverged 11,700 years BP, has resulted in the evolution of different longevity, although sexual selection has maintained a similar pattern of relative sex differences in tooth depletion. This study opens interesting questions on optimal allocation in life history trade-offs and the independent evolution of allopatric populations.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Variation in social rank acquisition influences lifetime reproductive success in black-capped chickadees

Dominance relationships structure many animal societies, yet the process of rank attainment is poorly understood. We investigated acquisition of social dominance in winter flocks and its fitness consequences in male black-capped chickadees (Poecile atricapillus) over a 10-year period. Age was the best predictor of rank, and paired comparisons showed high-ranked males to be older than their low-ranked flock-mates. When controlling for age, morphological variables did not predict male social rank, but high-ranked males were heavier, had lower fat scores and were in leaner condition than low-ranked males. Males that survived between years tended to increase in rank over time; however, the rate of rank advancement varied individually. Rank reversals between familiar contestants were rare, and changes in male social rank were associated with changes in flock membership. Average lifetime reproductive success (LRS) of males and females was variable and best predicted by lifespan. Male rank history also influenced realized reproductive success. Birds with higher average rank over their lifespan were more likely to reproduce successfully. However, among successful birds, average rank did not significantly predict LRS. Thus, birds that lived longer and attained high social rank earlier had higher fitness, but this effect was not manifested as fine-scale differences among successful individuals. Taken together, these findings demonstrate the importance of social factors influencing individual fitness.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 85–95.     

Genetic detection of sperm from forced copulations between sympatric populations of Poecilia reticulata and Poecilia picta

The reproductive success of the sneaky mating tactic is difficult to determine in fish with internal fertilization. We approached this problem by developing an assay that allows the recovery of sperm DNA from the reproductive tract of females. This assay was used to test whether sperm transfer occurs between sympatric populations of the guppy, Poecilia reticulata , and its putative sister species, Poecilia picta . Six hundred females of both species from two sympatric sites in Trinidad were collected, and then the contents of the females' reproductive tracts were analysed using reciprocally diagnostic microsatellite markers. Conspecific sperm occurred in approximately 86% of females, whereas heterospecific sperm were found in only 4% of females. Because females of either species do not mate willingly with heterospecific males, the results indicate that sneaky mating results in the transfer of sperm. The data are consistent with the idea that sexual conflict can result in differences in the strength of behavioural isolation between the sexes, and they suggest that behavioural isolation is unlikely to have driven speciation between the guppy and P. picta .  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 397–402.     

Sex,androgens, and whole‐organism performance in an Australian lizard

Understanding underlying physiological differences between the sexes in circulating androgens and how hormonal variation affects morphology–performance relationships may help clarify the evolution of sexual dimorphism in diverse taxa. Using a widely distributed Australian lizard (Eulamprus quoyii) with weak sexual dimorphism and no dichromatism, we tested whether circulating androgens differed between the sexes and whether they covaried with morphological and performance traits (bite force, sprint speed, endurance). Males had larger head dimensions, stronger bite force, faster sprint speed, and longer endurance compared to females. We found that the sexes did not differ in androgen concentrations and that androgens were weakly associated with both morphological and performance traits. Interestingly, high circulating androgens showed a nonlinear relationship with bite force in males and not females, with this relationship possibly being related to alternative male reproductive tactics. Our results suggest that androgens are not strongly correlated with most performance and morphological traits, although they may play an important organizational role during the development of morphological traits, which could explain the differences in morphology and thus performance between the sexes. Differences in performance between the sexes suggest differential selection on these functional traits between males and females. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 834–849.     

Quantification of sexual dimorphism in Asellus aquaticus (Crustacea: Isopoda) using outline approaches

A marked sexual dimorphism is often observed in arthropods species in which males perform precopulatory mate guarding. It is generally thought to reflect the influence of sexual selection. Until now, sexual dimorphisms associated with mate guarding have mainly been qualitatively described. However, assessing the effects of sexual selection on sexual dimorphims requires a preliminary quantitative assessment of differences in morphology between sexes. Using Fourier analyses, we tested if morphological dimorphisms could be quantitatively assessed in the isopod Asellus aquaticus . In addition, we checked whether sexual dimorphism in shape was exclusively related to mate guarding through considering characters that are not, a priori , implicated in mating behaviour. To assess the potential role of sexual selection in shaping morphology, we then examined how dimorphic characters could influence males' pairing success. Three characters (pleotelson, paraeopods 4 and 5) differed significantly in shape between males and females. In addition, two characters (pleotelson and paraeopods 4) differed in shape between guarding males and non-guarding males, with the latter being closer in shape to females. This suggests that sexual selection may be partly responsible for the observed morphological divergence between sexes in A. aquaticus .  © 2002 The Linnean Society of London, Biological Society of the Linnean Society , 2002, 77 , 523–533.     

Bite-force performance and head shape in a sexually dimorphic crevice-dwelling lizard, the common chuckwalla [Sauromalus ater (= obesus)]

The Common Chuckwalla [ Sauromalus ater (=  obesus )] is a large, sexually dimorphic lizard with a flattened head that takes refuge from predators in rock crevices. Males use their relatively large heads to bite competing males during territorial fights and to restrain females during copulation. Flattened heads with an antipredator function (i.e. seeking refuge in crevices) and enlarged heads with intrasexual competition and reproductive functions suggest possible antagonism between selective pressures on head morphology in males. To examine this hypothesis, we performed a morphometric analysis and measured the bite-force performance of 49 adult chuckwallas. Males had disproportionately wider heads than females, but did not have deeper heads. Males bit with nearly four times the force of females, consistent with the notion of sexual selection for high bite force in males. Although constrained by crevice-wedging behaviour, head depth was a good predictor of bite force in both sexes. In males, however, osteological head width also was a good predictor of bite force. These results are consistent with the hypothesis that head shape in males is under antagonistic selective pressures, which may partly explain the pattern of head shape dimorphism. The disproportionately wide head of males may reflect anatomical modifications to enhance bite force in response to sexual selection in spite of presumed constraints on head shape for crevice-wedging behaviour  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 215–222.     

Nutrition‐ and sex‐dependent utilization of body resources in relation to reproduction in a scorpionfly

Reproduction often comes at a cost of a reduction in body functions. In order to enhance their reproductive output, some insect species degenerate their thoracic muscles, typically resulting in reduced flight ability. From a life‐history trade‐off perspective, we expect the importance of body resource utilization to be amplified both with increased reproductive expenditure and with increased resource limitation. In this study, we measured age‐related changes in thorax weight, as a measure of flight muscle size, during a major part of the adult lifespan in males and females of the scorpionfly Panorpa vulgaris. The aim of the study was twofold: first to investigate whether scorpionflies have the potential to degenerate their flight muscles; second, and more importantly, to determine whether the magnitude of flight muscle degeneration is a plastic response in relation to resource availability, and if it differs between the sexes. The results clearly demonstrate that food availability does influence investment in flight muscle development. The build‐up of the thoracic muscles was strongly influenced by nutrient availability. Furthermore, the age‐related decrease in thorax weight was significantly different for males and females. Only females showed a strong age‐dependent decrease in thorax weight, indicative of muscle degeneration, yet no difference between food treatments was detected. For males, there was no significant directional change in thorax weight. Nevertheless, with increasing age, the difference in thorax weight between food treatments increased significantly. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 199–207.     

Locomotor capacity and dominance in male lizards Lacerta monticola: a trade‐off between survival and reproductive success?

Trade-offs between reproduction and survival are important determinants of life-history characteristics of lizards. Organisms cannot increase the allocation of limited resources to reproduction without diverting a proportional amount of energy from another trait. Locomotor performance is an ecologically relevant trait that potentially influences survival by affecting the ability to escape from predators. Most studies have used female lizards as subjects because pregnancy is known to reduce their locomotor abilities, whereas little is known on costs of reproduction in males. In this study we suggest that in males of the lizard Lacerta monticola reproductive investment in morphological traits that confer dominance (i.e. head size) might lead to a low probability of survival by decreasing investment in other traits that affect locomotor performance (i.e. limb symmetry). We staged laboratory agonistic encounters between males and measured their morphology and burst speed on a race track to examine possible relationships between morphology, social dominance and locomotor capacity. Our results indicate that social dominance was positively related to relative head height, and that escape speed was negatively related to levels of fluctuating asymmetry in femur length, but also negatively related to relative head height. Males with greater relative head height also had more asymmetrical femurs, thus dominant males suffered a decrease in locomotor performance. Males with higher heads tend to dominate male–male interactions and hence may gain access to reproductive females, thus increasing their current reproduction success. However, this might occur at the expense of future survivorship mediated by a decrease in escape speed. Therefore, in male L. monticola there might be a trade-off between current reproductive success and survival.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002; 77 , 201–209.     

On the sex-specific mechanisms of butterfly flight: flight performance relative to flight morphology, wing kinematics, and sex in Pararge aegeria

Many evolutionary ecological studies have documented sexual dimorphism in morphology or behaviour. However, to what extent a sex-specific morphology is used differently to realize a certain level of behavioural performance is only rarely tested. We experimentally quantified flight performance and wing kinematics (wing beat frequency and wing stroke amplitude) and flight morphology (thorax mass, body mass, forewing aspect ratio, and distance to centre of forewing area) in the butterfly Pararge aegeria (L.) using a tethered tarsal reflex induced flight set-up under laboratory conditions. On average, females showed higher flight performance than males, but frequency and amplitude did not differ. In both sexes, higher flight performance was partly determined by wing beat frequency but not by wing stroke amplitude. Dry body mass, thorax mass, and distance to centre of forewing area were negatively related to wing beat frequency. The relationship between aspect ratio and wing stroke amplitude was sex-specific: females with narrower wings produced higher amplitude whereas males show the opposite pattern. The results are discussed in relation to sexual differences in flight behaviour.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 675–687.     

Reproductive trait divergence and hybrid fertility patterns between chromosomal races of the house mouse in Tunisia: analysis of wild and laboratory-bred males and females

The divergence in reproductive features and hybrid fertility patterns between two chromosomal races (2 n  = 40, 40St, and 2 n  = 22, 22Rb) of the house mouse in Tunisia were re-assessed on a larger sample of wild and laboratory-bred individuals than studied hitherto. Results showed that litter sizes were significantly smaller in 40St than in 22Rb mice, contrary to previous analyses. This suggests that variation in litter size between the two chromosomal races is more likely related to selective and/or environmental factors acting locally than to interracial reproductive trait divergence. However, the significantly reduced litter size of F₁ hybrids compared with parental individuals was confirmed, and further highlighted a sex difference in hybrid infertility, as F₁ females produced fewer litters and of smaller size than males. Histological analyses of F₁ and backcrosses showed a breakdown of spermatogenesis in males and a significantly reduced primordial follicle pool in females. The degree of gametogenic dysfunction was not related to the level of chromosomal heterozygosity per se , but a significant effect of two Rb fusions on follicle number was observed in hybrid females. These results suggest that genetic incompatibilities contribute to primary gametogenic dysfunction in hybrids between the chromosomal races in Tunisia.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 407–416.     

The relationship between tooth wear, habitat quality and late-life reproduction in a wild red deer population

1. Molar tooth wear is considered an important proximate mechanism driving patterns of senescence in ungulates but few studies have investigated the causes of variation in molar wear or their consequences for reproductive success. 2. In this study, we assessed molar tooth wear at death among red deer Cervus elaphus of known age on the Isle of Rum, Scotland. 3. First molar height showed a decelerating decline with age. In females, the rates of molar wear with age varied with location of home range and individuals experiencing low resource competition showed reduced molar wear. We suggest that this spatial variation in molar wear is related to differences in the availability of high-quality grazing habitat and levels of resource competition. 4. There was no evidence that females with more heavily worn molars had reduced reproductive performance late in life or that first molar height was associated with reproductive senescence.     

Intrapopulation variation in macro tooth wear patterns—a case study from Igloolik,Canada

The pattern of human tooth wear—the way it varies between teeth in the mouth—is crucial to our understanding of important questions in archeology and paleoanthropology, such as the contrasts in diet and behavior between Neanderthals and early modern humans in Europe and Asia, or with the adoption of agriculture in the Americas. Little is known, however, about the way in which wear patterns develop with increasing age or the way in which they differ between males and females. One explanation is that few living people show the high rates of tooth wear seen worldwide throughout the preindustrial archaeological record. The study described here investigates the macroscopic pattern of tooth wear in a unique group of known age and sex dental casts from living Canadian Inuit from Igloolik. The results show that the Igloolik people possessed a pattern of extremely heavy anterior tooth wear, relative to the first molar and the other posterior teeth, which is attributed to the use of the anterior teeth in cultural practices as well as the extreme and marginal environments in which they lived. Heavy anterior tooth wear was established at an early age and maintained throughout life; statistically significant differences were found between the wear patterns of males and females and are explained in terms of sexual division of labor within the community. This study highlights the need to understand both intra‐ and interpopulation variation in tooth wear patterns when interpreting patterns in past human groups. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Head shape allometry and proximate causes of head sexual dimorphism in Podarcis lizards: joining linear and geometric morphometrics

Podarcis bocagei and Podarcis carbonelli are two lacertid species endemic to the western Iberian Peninsula, and both show head size and shape sexual dimorphism. We studied immature and adult head sexual dimorphism and analysed ontogenetic trajectories of head traits with body and head size, aiming to shed light on the proximate mechanisms involved. Immatures were much less dimorphic than adults, but geometric morphometric techniques revealed that head shape sexual differences are already present at this stage. Males and females differed in allometry of all head characters with body size, with males showing a disproportionate increase of head size and dimensions. On the other hand, head dimensions and head shape changed with increasing head size following similar trends in both sexes, possibly indicating developmental restrictions. Consequently, adult sexual dimorphism for head characters in these species is the result of both shape differences in the immature stage and hypermetric growth of the head in relation to body size in males.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 111–124.     

Hybridization patterns and the evolution of reproductive isolation in ducks

Much of our knowledge of the evolution of reproductive isolation comes from studies of Drosophila . This body of work has revealed the following patterns: (1) reproductive isolation increases with phylogenetic distance between hybridizing species; (2) reproductive isolation is greater between sympatric than allopatric species with the same level of divergence; and (3) hybrid crosses conform to Haldane's rule. We tested for the existence of these patterns in ducks (subfamily Anatinae, sensu Livezey, 1997b ) based on 1037 hybrids of known parentage. Our analyses of the number of interspecific crosses in relation to phylogenetic distance found a significant deviation between the observed and expected distribution of crosses controlling for the topology of the Anatinae phylogeny. In particular, we found both an excess of hybrid crosses among closely related species and a scarcity among distantly related species. The number of hybrid males also decreased with increasing phylogenetic distance between parental species, although the number of hybrid females remained low and constant. Sympatric species produced higher numbers of hybrid males than allopatric ones, despite no difference in phylogenetic distance among parental species in compared groups. The number of hybrid males exceeded the number of hybrid females, consistent with Haldane's rule. This was evident even though the analysis was restricted to a reduced set of phylogenetically independent crosses. However, the pattern was no longer significant after correction for the number of hybrid males by the male-biased sex ratio of adult ducks.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 193−200.     

Connecting morphology, function and tooth wear in microchiropterans

A key objective in understanding the dentition of mammals is the ability to predict the function of teeth from their shape. Very few studies have used dental measurements that allow the prediction of comparative tooth effectiveness, particularly when modification in shape due to tooth wear is considered. Here, dental parameters are used in which a change in the parameter is readily interpretable in terms of change in factors such as increased force or energy required for cusps or crests to break down food. The functional parameters were measured for 3-D digital tooth reconstructions of the upper molars of the microchiropteran Chalinolobus gouldii at various stages of tooth wear. The changes in the majority of the parameters, such as decreased tip, edge and cusp sharpnesses, cusp occlusion relief, rake angle and fragment clearance, predict a deterioration in efficacy with increased wear. This conclusion has not been possible with alternative approaches; for instance, there was no significant change in crest length with wear, and so no change in function would be predicted from that measure. Some of the parameters did not change significantly with heavy wear, such as capture area of a crest, pointing to geometrical and design characteristics for the maintenance of shape with wear in the dilambdodont tooth form. Attrition and abrasion can be considered as wear on the relief and rake surfaces of tribosphenic-like crests, respectively. The differences in function of these two surfaces account for the differences in wear patterns.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 81–96.     

Ontogeny, anatomy and attachment of the dentition in mosasaurs (Mosasauridae: Squamata)

Aspects of mosasaur dental ontogeny are well preserved in many fossils of these giant marine squamates. Replacement teeth on the tooth-bearing elements (TBEs) first appear as small enamel crowns positioned posterolingual to the attached tooth (posterolabial for the pterygoid). Several developing crowns, of progressively larger size, are aligned in rows relating to a specific tooth position. The crowns rest in a dental groove that varies in width and depth depending on the TBE. The crown closest to the attached tooth is always the largest and is found in a small resorption pit. As resorption proceeds, the pit expands in volume (cementum and alveolar bone), and the crown increases in size and settles into the pit. Once mature crown size is achieved, the dentine root and cementum portion of the root develop rapidly, the attached tooth is lost and the replacement tooth erupts out of the alveolus. Mosasaurid teeth develop along a 'zig-zag'-shaped movement path: horizontally along the dental groove, down into the alveolus, and up and out of the alveolus prior to attachment to the alveolar wall. At no point in mosasaurid tooth development are the crowns observed in a horizontal position. The mosasaurid dental lamina appears to have been a continuous strip of dental epithelium as it is in other squamates. Mosasaurid tooth attachment is thecodont (histologically and geometrically) not subpleurodont. Most aspects of mosasaurid tooth attachment and ontogeny are autapomorphic for the group.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 687–700.     

Influence of sociality on allometric growth and morphological differentiation in sponge-dwelling alpheid shrimp

Eusocial societies are defined by a reproductive division of labour between breeders and nonbreeders that is often accompanied by morphological differentiation. Some eusocial taxa are further characterized by a subdivision of tasks among nonbreeders, often resulting in morphological differentiation among different groups (subcastes) that specialize on different sets of tasks. We investigated the possibility of morphological castes in eusocial shrimp colonies ( Zuzalpheus , formerly part of Synalpheus ) by comparing growth allometry and body proportions of three eusocial shrimp species with three pair-forming species (species where reproductive females and males occur in equal sex ratios). Allometry of eusocial species differed in several respects from that of pair-forming species in both lineages. First, allometry of fighting claw size among individuals other than female breeders was steeper in eusocial than in pair-forming species. Second, breeding females in eusocial colonies had proportionally smaller weapons (fighting claws) than females in pair-forming species. Finally, claw allometry changed with increasing colony size in eusocial species; large colonies showed a diphasic allometry of fighting claw and finger size, indicating a distinctive group of large individuals possessing relatively larger weapons than other colony members. Shrimp are thus similar to other eusocial animals in the morphological differentiation between breeders and nonbreeders, and in the indication that some larger nonbreeders might contribute more to defence than others.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 527–540.