Fishing directly selects on growth rate via behaviour: implications of growth-selection that is independent of size

Size-selective harvest of fish and crustacean populations has reduced stock numbers, and led to reduced growth rates and earlier maturation. In contrast to the focus on size-selective effects of harvest, here, we test the hypothesis that fishing may select on life-history traits (here, growth rate) via behaviour, even in the absence of size selection. If true, then traditional size-limits used to protect segments of a population cannot fully protect fast growers, because at any given size, fast-growers will be more vulnerable owing to bolder behaviour. We repeatedly measured individual behaviour and growth of 86 crayfish and found that fast-growing individuals were consistently bold and voracious over time, and were subsequently more likely to be harvested in single- and group-trapping trials. In addition, there was some indication that sex had independent effects on behaviour and trappability, whereby females tended to be less active, shyer, slower-growing and less likely to be harvested, but not all these effects were significant. This study represents, to our knowledge, the first across-individual support for this hypothesis, and suggests that behaviour is an important mechanism for fishing selectivity that could potentially lead to evolution of reduced intrinsic growth rates.

Consequences of intraspecific seed-size variation in Sparganium emersum for dispersal by fish

1. The potential for seed dispersal by fish (ichthyochory) is likely to vary within aquatic plant species, depending on intraspecific variation in phenotypic seed traits.

2. We studied the effect of seed size variation within the unbranched burreed (Sparganium emersum) on the potential for internal dispersal by the common carp (Cyprinus carpio), by feeding them light (< 10 mg), medium (10–20 mg) and heavy ( > 20 mg) seeds, seed mass being positively related to seed size.

3. We hypothesized: (i) that ingestion, retention time, survival during gut passage and viability after gut passage of S. emersum seeds would be affected by seed size; and (ii) that this would translate into intraspecific variation in dispersal probability and dispersal distance among seed size categories.

4. Ingestion was negatively related to seed size, while survival during gut passage was positively related to seed size. Seed viability after gut passage was not affected by seed size. Since the negative effect of ingestion was counterbalanced by an equally strong but positive effect on seed survival, the probability of dispersal did not differ between the tested seed-size categories.

5. The time that seeds remained in the digestive tract of carp did not differ between seed sizes, suggesting equal potential dispersal distances for all seeds. Based on optimum swimming speeds of carp, ranging from 0·9 to 1·8 km h−1, maximum dispersal distances will most likely range from 13·5 to 27 km.

6. This study highlights the importance of studying all stages of the endozoochorous dispersal process in order to estimate the effect of a phenotypic seed trait on plant dispersal.

Rapid depletion of genotypes with fast growth and bold personality traits from harvested fish populations

The possibility for fishery-induced evolution of life history traits is an important but unresolved issue for exploited fish populations. Because fisheries tend to select and remove the largest individuals, there is the evolutionary potential for lasting effects on fish production and productivity. Size selection represents an indirect mechanism of selection against rapid growth rate, because individual fish may be large because of rapid growth or because of slow growth but old age. The possibility for direct selection on growth rate, whereby fast-growing genotypes are more vulnerable to fishing irrespective of their size, is unexplored. In this scenario, faster-growing genotypes may be more vulnerable to fishing because of greater appetite and correspondingly greater feeding-related activity rates and boldness that could increase encounter with fishing gear and vulnerability to it. In a realistic whole-lake experiment, we show that fast-growing fish genotypes are harvested at three times the rate of the slow-growing genotypes within two replicate lake populations. Overall, 50% of fast-growing individuals were harvested compared with 30% of slow-growing individuals, independent of body size. Greater harvest of fast-growing genotypes was attributable to their greater behavioral vulnerability, being more active and bold. Given that growth is heritable in fishes, we speculate that evolution of slower growth rates attributable to behavioral vulnerability may be widespread in harvested fish populations. Our results indicate that commonly used minimum size-limits will not prevent overexploitation of fast-growing genotypes and individuals because of size-independent growth-rate selection by fishing.

Flocking and feeding in the fiddler crab (UCA tangeri): prey availability as risk-taking behaviour

For a full understanding of prey availability, it is necessary to study risk-taking behaviour of the prey. Fiddler crabs are ideally suited for such a study, as they have to leave their safe burrow to feed on the surface of the intertidal flats during low tide, thereby exposing themselves to avian predators. A study in an intertidal area along the coast of Mauritania showed that small crabs always stayed in the vicinity of their burrow, but large crabs wandered in large flocks (also referred to as droves) to feed on sea-grass beds downshore. Transplanting downshore feeding substrate to the burrowing zone of the small crabs proved that they too preferred to feed on it. Since small crabs can be preyed upon by more species of birds, this suggests that the decision not to leave the burrowing zone might be related to the risk of being fed upon by birds. We calculated predation risk from measurements on the density and feeding activity of the crabs, as well as the feeding density, the intake rate and the size selection of the avian predators. Per hour on the surface, crabs in a flock were more at risk than crabs feeding near their burrow. Thus, though flocking crabs may have benefited from ‘swamping the predator’ by emerging in maximum numbers during some tides only, this did not reduce their risk of predation below that of non-flocking crabs. Furthermore we found that irrespective of activity, large crabs suffered a higher mortality per tide from avian predators than small crabs. This suggests that large crabs could not sufficiently reduce their foraging time to compensate for the increased risk while foraging in a flock, even though they probably experienced better feeding conditions than small crabs staying near their burrow. The greater energy demands of large crabs were reflected in a greater surface area grazed. Thus, with increasing size a fiddler crab has to feed further away from its burrow and so may derive less protection from staying near to it. It seems that growing big does not reduce the risk of predation for fiddler crabs, as it does in many other species with indeterminate growth. As in such species, the most probable advantage of growing big is increased mating success. Ultimately, therefore, prey availability must be understood from the life-history decisions of the prey species.

Prey selection, size, and breakage differences in Turbo undulatus opercula found within Pacific Gull (Larus pacificus) middens compared to Aboriginal middens and natural beach deposits, southeast Australia

Qualitative discrimination criteria are employed commonly to distinguish cultural shell middens from natural shell deposits. Quantitative discrimination criteria remain less developed beyond an assumption that natural shell beds tend to contain a wider range of shell sizes compared to cultural shell middens. This study further tests this assumption and provides the first comparative quantitative analysis of shell sizes from cultural middens, bird middens, and beach shell beds. Size distributions of opercula of the marine gastropod Turbo undulatus within two modern Pacific Gull (Larus pacificus) middens are compared with two Aboriginal middens (early and late Holocene) and two modern beach deposits from southeast Australia. Results reveal statistically significant differences between bird middens and other types of shell deposits, and that opercula size distributions are useful to distinguish Aboriginal middens from bird middens but not from beach deposits. Supplementary qualitative analysis of taphonomic alteration of opercula reveal similar opercula breakage patterns in human and bird middens, and further support previously recognised criteria to distinguished beach deposits (water rolling and bioerosion) and human middens (burning). Although Pacific Gulls are geographically restricted to southern Australia, the known capacity of gulls (Larus spp.) in other coastal contexts around the world to accumulate shell deposits indicates the broader methodological relevance of our study.

Nest-tree selection by the threatened brush-tailed phascogale (phascogale tapoatafa) (marsupialia: dasyuridae) in a highly fragmented agricultural landscape

The conservation of roosting and nesting resources is of critical concern for many hollow-dependent species around the world. We investigated the nest-tree requirements of the threatened brush-tailed phascogale (Phascogale tapoatafa) in a highly cleared agricultural landscape in south-eastern Australia. We documented the physical characteristics of selected nest trees and describe the spatial and temporal patterns of nest-tree use as revealed by radio-tracking. Nine phascogales (seven females, two males) were radio-tracked between March and July 1999 in an area where most woodland habitat is confined to linear strips along roads and streams or small patches and scattered trees in cleared farmland. Female phascogales were monitored for 13–35 days over periods of 5–15 weeks and two males were monitored for 2 and 9 days respectively. A total of 185 nest-tree fixes was collected and all nests occupied by phascogales were in standing trees. Eighty-three nest trees were identified, ranging in diameter at breast height (dbh) from 25 to 171 cm, with a mean dbh for the trees used by each individual phascogale of >80 cm. Phascogales did not discriminate between canopy tree species in selecting nest trees, but showed highly significant selection for trees in the largest size class. All individuals used multiple nest trees, with the seven females occupying an average of 11.4 nest trees from a mean of 25 diurnal locations. The number of nest trees continued to increase throughout the study, suggesting that more would be identified during a longer or more intensive study. Occupied nest trees were located throughout each individual’s home range, highlighting the importance of a continuous spatial distribution of suitable nest trees across the landscape. Nest trees were also located in adjacent farmland up to 225 m from roadside vegetation, demonstrating the value that scattered clumps and even single trees in farmland can have for wildlife conservation.

Song sharing and repertoire size in the sedge warbler, Acrocephalus schoenobaenus: changes within and between years

The complex song of the male sedge warbler functions mainly in sexual attraction and the evolution of repertoire size is driven primarily by female choice. As male song ceases upon pairing, male–male singing interactions are relatively brief and have not been studied to our knowledge. This study shows that young males in their first breeding season shared significantly more syllables with their nearest neighbour than with their fathers or more distant males. Moreover, daily recordings revealed that rapid learning and modification of syllable repertoires occurred, resulting in a progressive increase in sharing within just a few days. This does not lead to a gradual increase in repertoire size as some syllables are dropped and new ones are acquired. This turnover process allows males to share syllables with their neighbours, whilst repertoire size, known to be important in female choice, remains relatively constant in any one year. Individual males were followed for several years and also showed considerable syllable turnover between years. However, in this case, repertoire size was found to increase between years, the largest increase occurring between the first and second years. We obtained a significant positive correlation between repertoire size and age, suggesting that females choosing males with larger repertoires may gain indirect (genetic) benefits for their offspring, such as good genes for viability. Whilst these results reveal a more flexible picture of repertoire turnover than previously suspected, the relative stability of repertoire size within a season and the increase with age suggests that repertoire size remains a likely target for sexual selection by female choice.

Sexual selection and individual genetic diversity in a songbird

Here, we report for the first time, to our knowledge, a strong correlation between a measure of individual genetic diversity and song complexity, a sexually selected male trait in sedge warblers, Acrocephalus schoenobaenus. We also find that females prefer to mate with males who will maximize this diversity in individual progeny. The genetic diversity of each offspring is further increased by means of nonrandom fertilization, as we also show that the fertilizing sperm contains a haplotype more genetically distant to that of the egg than expected by chance. These findings suggest that species' mating preferences may be subject to fine tuning aimed at increasing offspring viability through increased genetic diversity. This includes external and internal mechanisms of selection, even within the ejaculate of a single male.

Pheromone production, male abundance, body size, and the evolution of elaborate antennae in moths

The males of some species of moths possess elaborate feathery antennae. It is widely assumed that these striking morphological features have evolved through selection for males with greater sensitivity to the female sex pheromone, which is typically released in minute quantities. Accordingly, females of species in which males have elaborate (i.e., pectinate, bipectinate, or quadripectinate) antennae should produce the smallest quantities of pheromone. Alternatively, antennal morphology may be associated with the chemical properties of the pheromone components, with elaborate antennae being associated with pheromones that diffuse more quickly (i.e., have lower molecular weights). Finally, antennal morphology may reflect population structure, with low population abundance selecting for higher sensitivity and hence more elaborate antennae. We conducted a phylogenetic comparative analysis to test these explanations using pheromone chemical data and trapping data for 152 moth species. Elaborate antennae are associated with larger body size (longer forewing length), which suggests a biological cost that smaller moth species cannot bear. Body size is also positively correlated with pheromone titre and negatively correlated with population abundance (estimated by male abundance). Removing the effects of body size revealed no association between the shape of antennae and either pheromone titre, male abundance, or mean molecular weight of the pheromone components. However, among species with elaborate antennae, longer antennae were typically associated with lower male abundances and pheromone compounds with lower molecular weight, suggesting that male distribution and a more rapidly diffusing female sex pheromone may influence the size but not the general shape of male antennae.

The importance of neighbourhood size in self organising systems

In recent times, the analysis of SOM (self-organising map) performance has concentrated on optimising the gain decay, rather than the size, form and decay of the neighbourhood function. We propose that the size, form and decay of region size plays a much more significant role in the learning, and especially in the development, of topographic feature maps. In this paper, a biologically-derived SOM model is presented. This model is able to select a single winning neuron and to form Gaussian outputs about this winner, without the need for a meta-level decision-making structure to artificially select a winner and fit a Gaussian output to that winner. Using this model, some fundamental characteristics of the relationship between neighbourhood size and SOM output states are demonstrated.

Modelling the combined effect of grain size and grain shape on plastic anisotropy of metals

Within each columnar grain of a metallic film, the resistance to dislocation glide varies in function of the orientation of the slip plane with regard to the grain long axis. Plastic slip is impeded across grain boundaries and this contributes to the anisotropy of the overall mechanical response. A simplified (Taylor-type) crystal plasticity model is proposed that accounts for such effect of grain shape on the slip system selection. Assuming that dislocation density gradients are normal to the grain boundaries, backstresses developed at the onset of plasticity are estimated based on two definitions of the effective grain boundary spacing ‘‘seen’’ by individual slip systems. The first one reduces to the mean area-to-perimeter ratio of cross-sections of the grain cut parallel to the slip plane. Closed-form expressions of the average backstresses developed inside grains with spheroidal shapes are introduced in the crystal hardening law. The model reproduces the very high plastic anisotropy of electro-deposited pure iron with a strong c-fiber and a refined columnar grain structure [Yoshinaga, N., Sugiura, N., Hiwatashi, S., Ushioda, K., Kada, O., 2008. Deep drawability of electro-deposited pure iron having an extremely sharp h111i//ND texture. ISIJ Int. 48, 667–670]. It also provides valid estimates of the texture development and the influence of grain size on the yield strength.

Kin selection, not group augmentation, predicts helping in an obligate cooperatively breeding bird

Kin selection theory has been the central model for understanding the evolution of cooperative breeding, where non-breeders help bear the cost of rearing young. Recently, the dominance of this idea has been questioned; particularly in obligate cooperative breeders where breeding without help is uncommon and seldom successful. In such systems, the direct benefits gained through augmenting current group size have been hypothesized to provide a tractable alternative (or addition) to kin selection. However, clear empirical tests of the opposing predictions are lacking. Here, we provide convincing evidence to suggest that kin selection and not group augmentation accounts for decisions of whether, where and how often to help in an obligate cooperative breeder, the chestnut-crowned babbler (Pomatostomus ruficeps). We found no evidence that group members base helping decisions on the size of breeding units available in their social group, despite both correlational and experimental data showing substantial variation in the degree to which helpers affect productivity in units of different size. By contrast, 98 per cent of group members with kin present helped, 100 per cent directed their care towards the most related brood in the social group, and those rearing half/full-sibs helped approximately three times harder than those rearing less/non-related broods. We conclude that kin selection plays a central role in the maintenance of cooperative breeding in this species, despite the apparent importance of living in large groups.

Sexual selection and the evolution of complex color patterns in dragon lizards

Many species have elaborate and complex coloration and patterning, which often differ between the sexes. Sexual selection may increase the size or intensity of color patches (elaboration) in one sex or drive the evolution of novel signal elements (innovation). The latter potentially increases color pattern complexity. Color pattern complexity may also be influenced by ecological factors related to predation and environment; however, very few studies have investigated the effects of both sexual and natural selection on color pattern complexity across species. We used a phylogenetic comparative approach to examine these effects in 85 species and subspecies of Australian dragon lizards (family Agamidae). We quantified color pattern complexity by adapting the Shannon–Wiener diversity index. There were clear sex differences in color pattern complexity, which were positively correlated with both sexual dichromatism and sexual size dimorphism, consistent with the idea that sexual selection plays a significant role in the evolution of color pattern complexity. By contrast, we found little evidence of a link between environmental factors and color pattern complexity on body regions exposed to predators. Our results suggest that sexual selection rather than natural selection has led to increased color pattern complexity in males.

The role of testosterone in bib size determination in the male house sparrow Passer domesticus, is age dependent

Secondary sexual signals are thought to indicate individual quality. In order to understand the evolutionary pressures that give rise to such traits it is important to understand the physiological mechanisms underlying their production. The black bib of the house sparrow Passer domesticus is known to function as a badge of social status in males. Past studies have found that the size of the bib in older males is determined, at least partly, by the androgen testosterone. The immunocompetence handicap hypothesis suggests that testosterone has a key role in maintaining honest signalling – it is both involved in the development or expression of sexual signals and is immunosuppressive. In this paper we test experimentally two hypo theses relating to bib size development, whether 1) testosterone is only immunosuppressive in conditions where the natural feedback loop from the testes has been removed, and 2) testosterone is, in addition to influencing the bib size of older males, responsible for the size of the bib in juvenile sparrows. In the first experiment we found that exogenous testosterone administered to intact males during the winter (when LH and FSH levels are very low and were not artificially increased by castration) caused significant immunosuppression, albeit in interaction with the stress hormone corticosterone. Second, we found that exogenous testosterone administration in castrated fledgling male house sparrows had no effect on subsequent post-juvenile moult bib size relative to controls. Our results suggest that in some circumstances testosterone can be immunosuppressive, but that its role in bib size determination is age-dependent.

Sex-dependent selection differentially shapes genetic variation on and off the guppy Y chromosome

Because selection is often sex-dependent, alleles can have positive effects on fitness in one sex and negative effects in the other, resulting in intralocus sexual conflict. Evolutionary theory predicts that intralocus sexual conflict can drive the evolution of sex limitation, sex-linkage, and sex chromosome differentiation. However, evidence that sex-dependent selection results in sex-linkage is limited. Here, we formally partition the contribution of Y-linked and non-Y-linked quantitative genetic variation in coloration, tail, and body size of male guppies (Poecilia reticulata)—traits previously implicated as sexually antagonistic. We show that these traits are strongly genetically correlated, both on and off the Y chromosome, but that these correlations differ in sign and magnitude between both parts of the genome. As predicted, variation in attractiveness was found to be associated with the Y-linked, rather than with the non-Y-linked component of genetic variation in male ornamentation. These findings show how the evolution of Y-linkage may be able to resolve sexual conflict. More generally, they provide unique insight into how sex-specific selection has the potential to differentially shape the genetic architecture of fitness traits across different parts of the genome.