Site specificity in advertisement calls and responses to playbacks of local and foreign call variants in satin bowerbirds, <Emphasis Type="Italic">Ptilonorhynchus violaceus</Emphasis>

Avian vocalisations often show patterns of geographic variation. Previous work on the satin bowerbird has shown that although spatial variation in this species’ advertisement calls is strongly associated with habitat structure, some variation is apparent within habitat types. Seventeen populations located throughout the species’ distribution were used to examine whether spatial call variation could be influenced by other processes such as random drift or the presence of fine-scale vocal traditions; if this were the case, differing call variants would be expected at geographically discrete sampling sites both within and among habitat types. There were population-specific call variants at each of the sites sampled, with different variants apparent even within habitat types. At most sites, individuals gave only a single variant of advertisement call, and the call variant at one site, sampled after a 5-year interval, appears to have been relatively stable. Playback experiments were conducted at three populations to examine whether local call variants invoked a greater response than several non-local variants differing in their degree of similarity to the local variant. Birds responded strongly to local call variants but not to either of two foreign variants, one of which was similar to their local variant and one of which was very different. A pattern of geographic variation across populations, the fact that local and non-local variants evoke different responses and circumstantial evidence indicating that individuals can learn new calls all suggest that factors affecting song learning and the ability of males to establish and defend a bower site may have contributed to the establishment of geographically variable vocal cultures in this species.     

Variation in sperm morphometry and sperm competition among barn swallow (Hirundo rustica) populations

Spermatozoa vary greatly in size and shape among species across the animal kingdom. Postcopulatory sexual selection is thought to be the major evolutionary force driving this diversity. In contrast, less is known about how sperm size varies among populations of the same species. Here, we investigate geographic variation in sperm size in barn swallows Hirundo rustica, a socially monogamous passerine with a wide Holarctic breeding distribution. We included samples from seven populations and three subspecies: five populations of ssp. rustica in Europe (Czech, Italy, Norway, Spain, and Ukraine), one population of ssp. transitiva in Israel, and one population of ssp. erythrogaster in Canada. All sperm traits (head length, midpiece length, tail length, and total length) varied significantly among populations. The variation among the European rustica populations was much lower than the differences among subspecies, indicating that sperm traits reflect phylogenetic distance. We also performed a test of the relationship between the coefficient of between-male variation in total sperm length and extrapair paternity levels across different populations within a species. Recent studies have found a strong negative relationship between sperm size variation and extrapair paternity among species. Here, we show a similar negative relationship among six barn swallow populations, which suggests that the variance in male sperm length in a population is shaped by the strength of stabilizing postcopulatory sexual selection.     

Female preferences for spectral call properties in the western genetic lineage of Cope's gray treefrog (Hyla chrysoscelis)

Female frogs discriminate among potential mates based on individual variation in male advertisement calls. While considerable data have accumulated allowing comparisons of female preference functions among species, we still lack fundamental knowledge about how and why the shapes of preference functions for particular call properties vary among populations within all but a few species. Here, we report results from a study aimed at describing female preference functions for spectral call properties in Cope's gray treefrog (Hyla chrysoscelis). Widespread throughout the eastern half of North America, Cope's gray treefrog is the diploid member of the cryptic diploid–tetraploid Hyla versicolor species complex, and its populations are divided into two distinct genetic lineages (eastern and western). In this study of a western lineage population, we recorded and analyzed the spectral properties of 1,000 advertisement calls from 50 males and conducted two-stimulus phonotaxis experiments to estimate a population-level preference function. Females preferred calls with average frequencies over calls with frequencies that were 2 or 3 semitones (1.4 or 2.1 standard deviations, respectively) lower than the population mean. We observed no behavioral discrimination between calls with average and higher-than-average frequencies. Preferences discriminating against low-frequency calls were weak and were abolished by attenuating the preferred average call by 3?dB. We discuss these results in light of previous studies of eastern lineage populations, geographic variation in female preference functions, and the potential adaptive value of discriminating against calls with low frequencies.     

Congenital predispositions and early social experience determine the courtship patterns of males of the Amarillo fish

Mounting evidence indicates that learning shapes fish foraging and social behavior, which suggests that it may also underlie the frequently reported geographic variation in courtship. In this study, we analyzed how early social experience influences courtship patterns in males of the Amarillo fish (Girardinichthys multiradiatus). In a first experiment, we quantified the courtship of males from two allopatric populations grown in isolation and estimated the heritability of a key attribute of their courtship patterns. Then, we raised males from two allopatric populations with either sympatric or allopatric females and at two different densities, and registered their adult courtship patterns towards females from a third population. We found that the courtship pattern of males has a significant heritable component, yet in the second experiment, it was influenced by what type of females males were raised with, by their social rank (dominant/subordinate/singleton), and by the behavior of the female that they were courting. We conclude that courtship behavior in this fish has a congenital predisposition and is affected during development by different aspects of early experience and seemingly remains flexible throughout life; a complex ontogenetic trajectory that under natural conditions may have led to the locally stable dialect types observed in the courtship patterns of males from allopatric populations.     

Range-wide effects of breeding- and nonbreeding-season climate on the abundance of a Neotropical migrant songbird

Geographic variation in the population dynamics of a species can result from regional variability in climate and how it affects reproduction and survival. Identifying such effects for migratory birds requires the integration of population models with knowledge of migratory connectivity between breeding and nonbreeding areas. We used Bayesian hierarchical models with 26 years of Breeding Bird Survey data (1982-2007) to investigate the impacts of breeding- and nonbreeding-season climate on abundance of American Redstarts (Setophaga ruticilla) across the species range. We focused on 15 populations defined by Bird Conservation Regions, and we included variation across routes and observers as well as temporal trends and climate effects. American Redstart populations that breed in eastern North America showed increased abundance following winters with higher plant productivity in the Caribbean where they are expected to overwinter. In contrast, western breeding populations showed little response to conditions in their expected wintering areas in west Mexico, perhaps reflecting lower migratory connectivity or differential effects of winter rainfall on individuals across the species range. Unlike the case with winter climate, we found few effects of temperature prior to arrival in spring (March-April) or during the nesting period (May-June) on abundance the following year. Eight populations showed significant changes in abundance, with the steepest declines in the Atlantic Northern Forest (-3.4%/yr) and the greatest increases in the Prairie Hardwood Transition (4%/yr). This study emphasizes how the effects of climate on populations of migratory birds are context dependent and can vary depending on geographic location and the period of the annual cycle. Such knowledge is essential for predicting regional variation in how populations of a species might vary in their response to climate change.     

High male reproductive success in a low-density Antarctic fur seal (Arctocephalus gazella) breeding colony

Understanding how population density influences mating systems may lead to important insights into the plasticity of breeding behavior, but few natural systems allow for such studies. Antarctic fur seals (Arctocephalus gazella) provide an interesting model system because they breed in colonies of varying densities. Previous studies have largely focused on a high-density site at Bird Island, South Georgia. Here, 13 highly polymorphic microsatellite loci were used to conduct a genetic analysis of a low-density breeding colony of this species at Livingston Island, approximately 1,600 km south of South Georgia. The majority of adults seen ashore (n?=?54) were sampled together with every pup born (n?=?97) over four consecutive years. Paternities were confidently assigned for 34 out of the 97 pups. Two out of 23 sampled males accounted for the paternity of 28 % of all pups sampled during the study and 82 % of the pups with an assigned father. Moreover, a full likelihood pedigree inference method assigned a further eight paternities to a single unsampled male seal that is inferred to have held a territory during the season before the study began. The most successful males in our study easily surpassed the previous record for the total number of pups sired per male seal for the species. Furthermore, we identified two triads of full siblings implying that their parents remated in three consecutive years. These findings suggest that territorial male fur seals may achieve greater success in monopolizing access to breeding females when population density is relatively low.     

Microsatellite Analysis of Population Structure in the Endangered Hawaiian Monk Seal

Abstract: The endangered Hawaiian monk seal breeds at six locations in the northwestern Hawaiian Islands. To determine whether significant genetic differentiation exists among these sites, we used microsatellite loci to examine the monk seal population structure at the five largest breeding colonies. Of 27 loci isolated from other seal species, only 3 were polymorphic in an initial screening of one individual from each breeding site. Only two alleles were found at each of these 3 loci in samples of 46–108 individuals. This extremely low variation is consistent with other measures of genetic variability in this species and is probably the result of a recent severe population bottleneck, combined with a long-term history of small population sizes. Although the smallest monk seal subpopulation in this study ( Kure Atoll) showed some evidence of heterozygote deficit, possibly indicative of inbreeding, the next smallest ( Pearl and Hermes Reef) had an apparent excess of heterozygous individuals. Genetic differentiation was detected between the two subpopulations at extreme ends of the range ( Kure and French Frigate Shoals). This trend was significant only at the microsatellite locus for which we had the largest sample size ( Hg6.3: R _ST = 0.206, p = 0.002; allelic goodness of fit G _h = 15.412, p < 0.005). French Frigate Shoals is the source population for translocated animals that have been released primarily at Kure Atoll. Differentiation between these sites consisted of allele frequency differences (with the same allele predominant in each location at all three loci), rather than the preservation of alternative alleles. Although the translocations have had positive demographic effects, we recommend continued genetic monitoring of both the source and recipient populations because translocated individuals are now entering the breeding population.     

Limited geographic variation in the acoustic structure of and responses to adult male alarm barks of African green monkeys

The global diversity of human languages is a remarkable feature of our species, which requires a capacity for rapid vocal learning. Given that primate alarm calling systems have played an important role in the language origin debate, identifying geographic variation in primate alarm calls and understanding the underlying causal mechanisms are important steps to help uncover evolutionary precursors to language. This study investigates geographic variation in the alarm bark of the widely distributed African green monkey (Chlorocebus). To quantify geographic variation in spectral and temporal call structure, acoustic analysis was used to compare the adult male barks of green monkeys (Chlorocebus sabaeus) and two subspecies of vervet (Chlorocebus pygerythrus pygerythrus and Chlorocebus pygerythrus hilgerti). Playback experiments were also carried out to test whether adult male vervets would distinguish between the barks of own-group males, unknown conspecific males and green monkey males. Acoustic analysis showed that, whilst similar in overall structure, the barks of green monkeys could be distinguished from vervet barks with a high degree of accuracy; the barks of vervet subspecies could also be discriminated, although to a lesser degree. Males responded most strongly to unknown conspecific males’ barks, and exhibited responses typical of leopard-avoidance and territorial defence. Taken together, these findings indicate that variation in alarm calls can be best explained by phylogenetic distance, and that intra- and inter-species differences are relevant during social interactions. Moreover, barks may function as an alarm and display call, which could explain the observed sexual dimorphism in barks in this genus.     

Temporary alteration of local social structure in a threatened population of Cuban iguanas (Cyclura nubila)

In small, insular populations, behavioral patterns that lead to increased variance in individual reproductive success can accelerate loss of genetic variation. Over a 1-year period, we documented behavior and hormone levels in a breeding group of adult Cuban iguanas (Cyclura nubila) at Guantánamo Bay. Male dominance was associated with body and head size, display behavior, testosterone levels, home-range size, and proximity to females. Based on their success in agonistic encounters, we ranked males in a linear dominance hierarchy. During the subsequent breeding season, we conducted a removal experiment in which the five highest-ranking males were temporarily relocated from the study site. Although we were unable to assess reproductive success directly, previously lower-ranking males assumed control of vacated territories, won more fights, and increased their proximity to females in the absence of the dominant males. When it results in greater mating opportunities for otherwise socially suppressed individuals, temporary alteration of local social structure may help limit erosion of genetic variation in small, insular populations. Electronic Publication     

Effects of early auditory experience on the development of local song preference in female swamp sparrows

A growing number of studies show that learning about male mating signals can shape the way females discriminate among males and may influence the evolution of both female preferences and the male traits under selection. Female songbirds commonly prefer local songs over foreign songs from a different population. In some species, however, the extent of variation among songs sung by different males within a population is as great as the variation observed between populations, raising the question of how females are able to discriminate local from foreign songs. Here, we report that laboratory-reared female swamp sparrows (Melospiza georgiana) not only show a preference for the particular song types with which they were tutored as compared with both foreign songs and unfamiliar local songs but also show preference for unfamiliar local songs over unfamiliar foreign songs. An acoustic analysis comparing tutor songs and those presented as unfamiliar local and foreign songs suggests that female swamp sparrows might be attuned to the specifics of local note phonology when assessing the attractiveness of unfamiliar songs. Our results demonstrate that early auditory experience influences response to geographic song variation in female swamp sparrows, and suggest the possibility that female songbirds may generalize what they learn from songs early in life to novel songs heard in adulthood. Additional work is needed to evaluate the contribution of unlearned predispositions for local song.     

Nest distribution affects behaviour and mating success in a marine fish

The distribution of breeding resources, such as nest sites, can have a pronounced impact on a population by affecting the proportion of individuals that succeed to breed and hence, the variation in reproductive success. Aggregation of important resources can lead to resource monopolisation by a limited number of individuals and thus affect the intensity of sexual selection. In this study, we tested, by contrasting two experimental treatments (dispersed vs. aggregated), how nest distribution affects: (1) mating behaviour, (2) male nest occupation and mating success, and (3) reproductive success and the opportunity for selection. We used the two-spotted goby (Gobiusculus flavescens), a small marine fish with a resource-based mating system, as our model species. When nests were aggregated, a larger proportion of the males behaved aggressively, fewer males succeeded in occupying a nest, fewer males became mated, and those males that mated received fewer eggs from spawning females. These effects resulted in a higher variance in reproductive success and hence, a higher opportunity for selection (I _rs ), in the aggregated treatment. We suggest that the results are a direct consequence of males defending a territory around their nest, preventing competitively inferior males from breeding. However, we found no significant selection differentials for body length or condition of males in either treatment. Our results support the hypothesis that aggregation of essential resources like nests promotes resource monopolisation. In species facing highly clumped nesting resources in the wild, monopolisation may negatively impact population productivity but could lead to strong selection on traits that promote male competitive ability.     

Are penguins and seals in competition for Antarctic krill at South Georgia?

The Antarctic fur seal (Arctocephalus gazella) and macaroni penguin (Eudyptes chrysolophus) are sympatric top predators that occur in the Southern Ocean around South Georgia where they are, respectively, the main mammal and bird consumers of Antarctic krill (Euphausia superba). In recent years the population of fur seals has increased, whereas that of macaroni penguins has declined. Both species feed on krill of similar size ranges, dive to similar depths and are restricted in their foraging range at least while provisioning their offspring. In this study we test the hypothesis that the increased fur seal population at South Georgia may have resulted in greater competition for the prey of macaroni penguins, leading to the decline in their population. We used: (1) satellite-tracking data to investigate the spatial separation of the Bird Island populations of these two species whilst at sea during the breeding seasons of 1999 and 2000 and (2) diet data to assess potential changes in their trophic niches between 1989 and 2000. Foraging ranges of the two species showed considerable overlap in both years, but the concentrations of foraging activity were significantly segregated spatially. The size of krill taken by both species was very similar, but over the last 12 years the prevalence of krill in their diets has diverged, with nowadays less krill in the diet of macaroni penguins than in that of Antarctic fur seals. Despite a significant degree of segregation in spatial resource use by the study populations, it is likely that the South Georgia populations of Antarctic fur seal and macaroni penguin exploit the same krill population during their breeding season. For explaining the opposing population trends of the two species, the relative contributions of independent differential response to interannual variation in krill availability and of interspecies competition cannot be resolved with available evidence. The likely competitive advantage of Antarctic fur seals will be enhanced as their population continues to increase, particularly in years of krill scarcity.     

Identification of genetic populations of the Pacific blue shrimp Penaeus stylirostris of the Gulf of California, Mexico

Species are often composed of discrete breeding units (i.e. populations or stocks) which, while not reproductively isolated from other such groups, may have limited opportunities to exchange genetic material because of geographic distance, barriers to migration, or spawning asynchrony. Low levels of gene flow between stocks may result, over time, in their genetic divergence, and species that are subdivided into morphologic or genetically distinct stocks are said to be structured. The aim of our investigation was to test whether or not Penaeus stylirostris from the Gulf of California (Mexico) was structured into genetically distinct populations. Shrimp samples were collected in 1996 from six regions of the Gulf where specimens with distinct morphologic characteristics had previously been identified. Statistical analysis of 324 RAPD loci (randomly amplified polymorphic DNA), resolved through polyacrylamide gels and scored for each of 78 specimens, permitted the quantification and comparison of between-stock genetic differences. The finding that genetically discrete stocks of P. stylirostris can be found in a small portion of the geographic distribution range of the species, disagrees with the long-held perception that this resource is panmictic in nature. This new evidence is not only of interest for selective breeding programs in the shrimp aquaculture industry, but is also relevant to the management of the Mexican shrimp fishery which, at present, is perceived and managed as a single stock. Received: 10 May 1999 / Accepted: 27 July 2000     

Male's return rate, rather than territory fidelity and breeding dispersal, explains geographic variation in song sharing in two populations of an oscine passerine (Oreothlypis celata)

Males of some oscine passerines learn and share songs of neighboring males. This process can lead to the formation of song pattern neighborhoods or microhabitat song dialects. The degree to which song sharing occurs between populations and the spatial scale over which neighboring males share songs, however, can vary widely, and interpopulation comparisons have suggested that song sharing is more common in residents than in migrants. Here, we examine two populations of the orange-crowned warbler (Oreothlypis celata) to quantify patterns of song sharing at the northern (long-distance migrant) and southern (short-distance migrant) edges of the breeding distribution and to test if return rate, territory fidelity, and breeding dispersal explain the patterns found in the two populations. The southern population (O. celata sordida breeding on Santa Catalina Island, California; 33°N) had a higher annual return rate to their territories and exhibited higher song sharing in neighborhoods than their counterparts (O. celata celata breeding in Fairbanks, Alaska; 64°N). Year-to-year patterns of territory fidelity and breeding dispersal distances were similar between populations. Our results suggest that if migratory distance generally covaries with the proportion of returning males, this could explain different levels of song sharing between the short- and long-distance migrants.     

Patch use and vigilance by sympatric lemmings in predator and competitor-driven landscapes of fear

Prey living in risky environments can adopt a variety of behavioral tactics to reduce predation risk. In systems where predators regulate prey abundance, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, patterns of habitat use also reflect interspecific competition over habitat. Collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings represent such a system and possess distinct upland tundra versus mesic meadow habitat preferences consistent with interspecific competition. Yet, we do not know whether this habitat preference might also reflect differences in predation risk or whether the two species differ in their behavioral tactics used to avoid predation. We performed experiments where we manipulated putative predation risk perceived by lemmings by increasing protective cover in upland and meadow habitats while we recorded lemming activity and behavior. Both lemming species preferentially used cover more than open patches, but Dicrostonyx was more vigilant than Lemmus. Both species also constrained their activity to protective patches in upland and meadow habitats, but during different periods of the day. Use of cover and vigilance were independent of habitat, suggesting that both species live in a fearsome but flattened landscape of fear at Walker Bay (Nunavut, Canada), and that their habitat preference is a consequence of competition rather than predation risk. Future studies aiming to map the contours of fear in multi-prey–predator systems should consider how predation and competition interact to modify prey species’ habitat preference, patch use, and vigilance.     

Effective Population Size in Red-Cockaded Woodpeckers: Population and Model Differences

Loss of genetic variability in isolated populations is an important issue for conservation biology. Most studies involve only a single population of a given species and a single method of estimating rate of loss. Here we present analyses for three different Red-cockaded Woodpecker ( Picoides borealis ) populations from different geographic regions. We compare two different models for estimating the expected rate of loss of genetic variability, and test their sensitivity to model parameters. We found that the simpler model (Reed et al. 1988) consistently estimated a greater rate of loss of genetic variability from a population than did the Emigh and Pollak (1979) model. The ratio of effective population size (which describes the expected rate of loss of genetic variability) to breeder population size varied widely among Red-cockaded Woodpecker populations due to geographic variation in demography. For this species, estimates of effective size were extremely sensitive to survival parameters, but not to the probability of breeding or reproductive success. Sensitivity was sufficient that error in estimating survival rates in the field could easily mask true population differences in effective size. Our results indicate that accurate and precise demographic data are prerequisites to determining effective population size for this species using genetic models, and that a single estimate of rate of loss of genetic variability is not valid across populations.     

Connectivity of wood thrush breeding,wintering, and migration sites based on range‐wide tracking

Many migratory animals are experiencing rapid population declines, but migration data with the geographic scope and resolution to quantify the complex network of movements between breeding and nonbreeding regions are often lacking. Determining the most frequently used migration routes and nonbreeding regions for a species is critical for understanding population dynamics and making effective conservation decisions. We tracked the migration of individual Wood Thrushes (Hylocichla mustelina) (n = 102) from across their range with light‐level geolocators and, for the first time, quantified migration routes and wintering regions for distinct breeding populations. We identified regional and species‐level migratory connectivity networks for this declining songbird by combining our tracking results with range‐wide breeding abundance estimates and forest cover data. More than 50% of the species occupied the eastern wintering range (Honduras to Costa Rica), a region that includes only one‐third of all wintering habitat and that is undergoing intensive deforestation. We estimated that half of all Wood Thrushes in North America migrate south through Florida in fall, whereas in spring approximately 73% funnel northward through a narrow span along the central U.S. Gulf Coast (88–93°W). Identifying migratory networks is a critical step for conservation of songbirds and we demonstrated with Wood Thrushes how it can highlight conservation hotspots for regional populations and species as a whole. Conectividad de Sitios de Reproducción, Invierno y Migración del Zorzal con Base en Rastreo de Cobertura Amplia     

Sperm storage reflects within- and extra-pair mating opportunities in a cooperatively breeding bird

In passerine birds, storage and maturation of sperm takes place in the cloacal protuberance (CP), an external swelling of the reproductive organ. The considerable variation in CP size among species is presumed to be a consequence of varying levels of sperm-competition, but whether individual variation in CP size within a species also reflects sperm competition is not well established. Here, we study temporal variation in male CP size in relation to within-pair and extra-pair mating opportunities and cuckoldry risk in purple-crowned fairy-wrens Malurus coronatus. This is a socially monogamous cooperatively breeding passerine that can breed year-round and has low levels of extra-pair paternity (in 6?% of broods). We show that male CP size sharply increased a few weeks before, and rapidly regressed after his partner laid eggs, consistent with a cost of its maintenance and/or sperm production. Surprisingly, despite low levels of extra-pair paternity, CP size of non-breeding and pre-breeding males was positively correlated with the number of breeding females in the population, suggesting that CP size is sensitive to extra-pair mating opportunities. However, CP sizes do not seem to reflect cuckoldry risk: CP size of dominant males was unaffected by the presence of a subordinate that was unrelated to the dominant female, although those subordinates occasionally sire offspring, and had a larger CP than subordinates living with their mother. Our results suggest that, even in a species with low levels of extra-pair paternity, individual investment in sperm storage reflects both within-pair and, albeit to lesser extent, extra-pair mating opportunities.     

Species divergence and seasonal succession in rates of mate desertion in closely related Neotropical cichlid fishes

Across animal taxa, exclusive female offspring care has evolved repeatedly from biparental care, suggesting that the latter becomes evolutionarily unstable under certain conditions. Both the attributes of a species and the environment it experiences can help to predict shifts from one particular care mode to another. Nevertheless, factors inducing differences in care strategies among closely related species, or seasonal variation within species, have been subject to surprisingly little empirical testing. Here, we report the results of a field-based study that examined both among and within species variation in mate desertion in five species of closely related Nicaraguan cichlid fish in the genera Amphilophus and Amatitlania. The results show a link between female body size and male involvement in offspring care. Specifically, the larger the species the less often males were found to provide extended care. Furthermore, we found that solitary females became more common towards the end of the breeding season. We discuss the implications of this finding in the context of previous theoretical and empirical contributions regarding the frequency of offspring desertion by males.     

Variation in male sailfin molly, preference for female size: does sympatry with sexual parasites, drive preference for smaller conspecifics?

Male sailfin mollies (Poecilia latipinna) are sexually parasitized by gynogenetic Amazon mollies (Poecilia formosa). In areas of sympatry, Amazon mollies are frequently larger than female sailfin mollies. In sympatry, selection may favor males that prefer smaller conspecific mates (avoid mismating with Amazon mollies), or selection may favor males that prefer larger conspecific mates (higher fecundity). To explore this potential species and mate-quality recognition conflict, we examined male preference variation across populations. Males from one sympatric population showed stabilizing preference functions, whereas in another sympatric population, males showed directional preference functions. Variation across sympatric populations may be related to the length of time of co-evolution with Amazon mollies. In the allopatric populations, we found flat preference functions. Variation in male preferences could have important ramifications for the maintenance of Amazon mollies, as well as for the evolution of female size.