Incest avoidance and bonding between siblings in gerbils

Summary Social and sexual preferences for siblings versus non-siblings have been investigated in Mongolian gerbils (Meriones unguiculatus) in a series of laboratory and outdoor enclosure (100 m²) studies. When first social contact took place at pairing in adulthood full siblings and nonsiblings did not differ with respect to onset of reproduction. Correspondingly, both siblings and non-siblings paired at weaning age had a delayed onset of reproduction. Thus, no effect could be established in either case, related to the individuals' relatedness, but the importance of social exposure was clear. Non-siblings paired when 2 months old had a delayed onset of reproduction, which did not differ statistically from the onset in siblings, raised together, and housed pairwise since the same age. When sexually mature animals were paired at 3 months of age, siblings were found to start reproducing later than non-siblings (Ågren 1981). Following an initial delay in both siblings and nonsiblings paired when 5 months old, all non-sibling pairs reproduced before any sibling pair. At pair formation, at least the females in both groups were known to be sexually immature. These results indicate that for sibling-bonding to occur social exposure may be delayed until the animals are old enough, or somatically developed, to mature sexually. Thus, social exposure before weaning age during the period for primary socialization seem not to be crucial, nor the actual maturational state, sexually, in older females. To study the stability of the social bonds between siblings and non-siblings, animals kept pairwise in the laboratory were released in large outdoor enclosures. Pairs brought together before the age of sexual maturity did not remain stable, whether or not the former cagemates were siblings. Non-sibling pairs formed after this age, which had also reproduced successfully in the laboratory, did remain stable. Social and sexual interactions were studied within and between two mixed-sex sibling groups of four. The sibling groups had the possibility to establish group territories before being exposed to each other. Although the sibling groups remained stable, the females still preferred non-sibling males as sexual partners. However, socially dominant brothers copulated more frequently than subordinant brothers. This rank difference was not evident in non-siblings.     

Effects of kinship or familiarity? Small thrips larvae experience lower predation risk only in groups of mixed-size siblings

In many species of insects, larvae are distributed in an aggregated fashion. As they may differ in size and size matters to predation risk, small larvae may be less likely to fall prey to predators when near large and therefore better-defended larvae. We hypothesize that the small larvae may profit even more when these large larvae are siblings. We tested this hypothesis on kinship-dependent survival in groups of larvae of the Western flower thrips (Frankliniella occidentalis) exposed to a predatory mite (Iphiseius degenerans). Our experiments showed that small larvae in sibling groups survive significantly better than in non-sibling groups, but only when such groups consisted of a mixture of small and large larvae. To test whether the survival effect we found is due to familiarity of thrips larvae growing up together (i.e. on one leaf), we also measured survival in sibling groups of larvae grown up on different leaves and in non-sibling groups of larvae grown up on the same leaf. These experiments showed an increased survival of small thrips larvae only in groups of sibling larvae from the same leaf. Non-sibling larvae did not show an increased survival when they come from the same leaf. Our results indicated that the increased survival in sibling groups was only partly due to the familiarity effect we tested. Growing up together did not return the same survival effect for non-siblings as it did for siblings. We conclude that growing up together is a necessary but not sufficient condition for discrimination in thrips larvae.     

Kin recognition by juvenile predatory mites: prior association or phenotype matching?

Kin recognition, the biased treatment of conspecific individuals based on genetic relatedness, is a widespread phenomenon in animals. The most common mechanisms mediating kin recognition are prior association (familiarity) and phenotype matching. Recognition based on prior association allows identifying familiar individuals. Recognition based on phenotype matching is an extension of prior-association-based recognition and allows identifying familiar and unfamiliar individuals due to a shared phenotypic trait. I investigated which of the two mechanisms is used by cannibalistic juvenile predatory mites, Phytoseiulus persimilis. Protonymphs that were associated with either siblings or non-kin discriminated familiar and unfamiliar larvae and preferentially cannibalized the latter irrespective of genetic relatedness. In contrast, despite previous association with either siblings or non-kin, protonymphs did not discriminate unfamiliar sibling and unfamiliar non-kin larvae. Association in the larval stage therefore mediated kin recognition based on familiarity, but not phenotype matching in cannibalistic P. persimilis protonymphs. Furthermore, in the presence of a familiar prey individual, sibling cannibalism occurred significantly sooner than non-kin cannibalism. This quick sibling cannibalism may have been the consequence of preferential association of siblings and/or may indicate the occurrence of an alternative cannibal phenotype. I discuss the adaptive significance of prior-association-based recognition for P. persimilis juveniles and emphasize the ability of P. persimilis to use multiple recognition mechanisms in dependence of the ontogeny and the ecological context.     

Young-boy networks without kin clusters in a lek-mating manakin

I use 10 years of data from a long-term study of lek-mating long-tailed manakins to relate the social network among males to their spatial and genetic structure. Previously, I showed that the network connectivity of young males predicts their future success. Here, I ask whether kinship might shape the organization of this “young-boy network”. Not surprisingly, males that were more socially distant (linked by longer network paths) were affiliated with perch zones (lek arenas) that were further apart. Relatedness (r) among males within the network decreased as social distance increased, as might be expected under kin selection. Nevertheless, any role for indirect inclusive fitness benefits is refuted by the slightly negative mean relatedness among males at all social distances within the network (overall mean r = −0.06). That is, relatedness ranged from slightly negative (−0.04) to more negative (−0.2). In contrast, relatedness in dyads for which at least one of the males was outside the social network (involving at least one blood-sampled male not documented to have interacted with other banded males) was slightly above the random expectation (mean r = 0.05). The slight variations around r = 0 among males of different categories likely reflect dispersal dynamics, rather than any influence of kinship on social organization. Relatedness did not covary with the age difference between males. These results, together with previous results for lack of relatedness between alpha and beta male partners, refute any role for kin selection in the evolution of cooperative display in this lek-mating system. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau, and R. James).     

Seasonal decline in intracolony genetic relatedness in eastern tent caterpillars: implications for social evolution

Summary Genetic relatedness in social insect colonies may vary spatially or temporally as a result of changes in colony membership due to immigration or to variation in patterns of maternity and paternity. We estimated relatedness for eastern tent caterpillars (Malacosoma americanum) in laboratory colonies derived from egg masses using multilocus genotypic data derived from electrophoresis. This estimate is compared with estimates obtained from colony samples taken in the field at four intervals spanning the larval developmental season. We found that average intracolony relatedness is close to 0.5 initially but declines through the developmental season due to colony merging, showing that caterpillars do not discriminate between siblings and nonsiblings in order to preserve colony family structure. Using the intracolony values together with relatedness values for higher levels of population structure, we estimated the effective mean number of simple families represented in single colonies through the season. The overall effective number of families per tent increased from one at the time of eclosion to 1.3 by the end of the season. Average intracolony relatedness remained relatively high despite the occurrence of colony merging, apparently as a result of the low density of tents on most trees, combined with high relatedness within the original colonies. Thus, high intracolony relatedness is maintained in M. americanum populations through the effects of adult dispersal, mating, and oviposition patterns, rather than through behavioral discrimination mechanisms of the larvae. These findings underscore the importance of considering the causes of temporal variation in genetic relatedness as well as the consequences for the indirect component of inclusive fitness. Correspondence to: J.T. Costa     

How does salinity tolerance influence the distributions of <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> sibling species?

Environmental salinity is important in defining Brachionus plicatilis sibling species distributions. However, while salinity influences distributions, sibling species often co-exist. Three different mechanisms potentially account for the partial co-occurrence of sibling species: (1) siblings have differing salinity tolerances that partially overlap; (2) siblings physiological tolerances may be commonly broad, but relatively small differences in tolerances differentiate distributions via interactions e.g. competition; or (3) siblings distributions may be influenced by physical factors other than salinity. Here, we assess the extent of salinity tolerance in three B. plicatilis sibling species (B. plicatilis 6TUR, B. plicatilis IOM and B. rotundiformis 6TOS) by measuring population growth rate (μ, day⁻¹) and egg development time in response to salinity (5–60‰) and salinity fluctuations (≤ Δ40‰). Sibling species were identified by analysis of the mitochondrial COI gene, and salinity responses were compared by regression analysis. Responses differed significantly between siblings, although the broad trends were similar. Positive growth occurred at all salinities, and highest growth rates ranged between 0.93 and 1.08 day⁻¹ at 16–18‰. Rapid changes in salinity reduced growth rates, but net mortality occurred only in one treatment (100% mortality on transfer from 10 to 40‰). Egg development time was largely invariant with salinity except for B. plicatilis IOM and where rotifers were transferred from 30 to 60‰. We indicate that several siblings are similarly euryhaline and tolerate salinity fluctuations. Undoubtedly, wide tolerances in B. plicatilis are adaptations to ephemeral and seasonally variable habitats. Given common broad salinity tolerances, it is unlikely that the differential distributions of sibling species are a direct result of physiological constraints. Instead, we illustrate using a simple model that subtle differences in physiological tolerances may have important impacts on interactions between sibling species, which may in turn influence distributions.     

Local mate competition in the solitary parasitoid wasp <Emphasis Type="Italic">Ooencyrtus kuvanae</Emphasis>

Local mate competition (LMC) occurs when brothers compete with each other for mating opportunities, resulting in selection for a female-biased sex ratio within local groups. If multiple females oviposit in the same patch, their sons compete for mating opportunities with non-brothers. Females, in the presence of other females, should thus produce relatively more sons. Sex ratio theory also predicts a more female-biased sex ratio when ovipositing females are genetically related, and sex-ratio responses to foundress size if it differentially affects fitness gains from sons versus daughters. The mating system of the parasitoid wasp Ooencyrtus kuvanae meets assumptions of LMC. Females insert a single egg into each accessible egg of gypsy moth, Lymantria dispar, host egg masses. Wasps complete development inside host eggs and emerge en masse, as sexually mature adults, resulting in intense competition among brothers. We tested the hypothesis that O. kuvanae exhibits LMC by manipulating the number of wasp foundresses on egg masses with identical numbers of eggs. As predicted by LMC theory, with increasing numbers of wasp foundresses on an egg mass, the proportions of emerging sons increased. In contrast, the presence of a sibling compared to a non-sibling female during oviposition, or the size of a female, did not affect the number or sex ratio of offspring produced. The O. kuvanae system differs from others in that larvae do not compete for local resources and thus do not distort the sex ratio in favor of sons. With no resource competition among O. kuvanae larvae, the sex ratio of emergent son and daughter wasps is due entirely to the sex allocation by ovipositing wasp foundresses on host egg masses.     

Kin discrimination in the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>): nepotism among siblings

We examined patterns of affiliation, association, and aggression to inquire whether spotted hyenas (Crocuta crocuta) can distinguish among various groups of maternal and paternal siblings. If so, and if these animals conform to predictions of kin selection theory, then behavioral interactions among hyenas should vary with relatedness. We also considered familiarity-based recognition and phenotype matching as mechanisms hyenas might use to recognize kin. Patterns of affiliative behavior indicated that hyenas favored full-sibling littermates over half-sibling littermates or any other group of half-siblings. Rates of dyadic aggression generally did not vary with kinship. Hyenas associated more closely with half-sibling littermates than with non-littermate half-siblings, and hyenas affiliated more with maternal half-siblings than with paternal half-siblings, suggesting that familiarity-based cues might mediate discrimination among these sibling classes. In addition, operation of a phenotype-matching mechanism was suggested by the preference hyenas demonstrated during affiliative interactions for full- over half-sibling littermates, and by their lack of preference in these interactions for half-sibling littermates over non-littermate half-siblings. Phenotype matching was also suggested by our observation that paternal half-siblings cooperated more, and fought less, than did non-kin. Our data indicate that hyenas can discriminate among various types of siblings, that their social behavior conforms to predictions of kin selection theory, and that they recognize kin using mechanisms of both familiarity and phenotype matching.Communicated by S. Alberts     

Kin discriminators in the eusocial sweat bee <Emphasis Type="Italic">Lasioglossum malachurum</Emphasis>: the reliability of cuticular and Dufour’s gland odours

The ability to discriminate degrees of relatedness may be expected to evolve if it allows unreciprocated altruism to be preferentially directed towards kin (Hamilton in J Theor Biol 7:1–16, 1964). We explored the possibility of kin recognition in the primitively eusocial halictid bee Lasioglossum malachurum by investigating the reliability of worker odour cues that can be perceived by workers to act as indicators of either nest membership or kinship. Cuticular and Dufour’s gland compounds varied significantly among colonies of L. malachurum, providing the potential for nestmate discrimination. A significant, though weak, negative correlation between chemical distance and genetic relatedness (r = −0.055, p < 0.001) suggests a genetic component to variation in cuticular bouquet, but odour cues were not informative enough to discriminate between different degrees of relatedness within nests. This pattern of variation was similar for Dufour’s gland bouquets. The presence of unrelated individuals within nests that are not chemically different from their nestmates suggests that the discrimination system of L. malachurum is prone to acceptance errors. Compounds produced by colony members are likely combined to generate a gestalt colony chemical signature such that all nestmates have a similar smell. The correlation between odour cues and nest membership was greater for perceived compounds than for non-perceived compounds, suggesting that variability in perceived compounds is a result of positive selection for nestmate recognition despite potentially stabilising selection to reduce variability in odour differences and thereby to reduce costs derived from excessive intracolony nepotistic behaviour.     

Association patterns and kinship in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) of southeastern Australia

Kinship has been shown to be an important correlate of group membership and associations among many female mammals. In this study, we investigate association patterns in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) inhabiting an embayment in southeastern Australia. We combine the behavioral data with microsatellite DNA and mitochondrial DNA data to test the hypotheses that genetic relatedness and maternal kinship correlate with associations and social clusters. Mean association between females was not significantly different from a random mean, but the standard deviation was significantly higher than a random standard deviation, indicating the presence of nonrandom associates in the dataset. A neighbor-joining tree, based on the distance of associations between females, identified four main social clusters in the area. Mean genetic relatedness between pairs of frequent female associates was significantly higher than that between pairs of infrequent associates. There was also a significant correlation between mtDNA haplotype sharing and the degree of female association. However, the mean genetic relatedness of female pairs within and between social clusters and the proportion of female pairs with the same and different mtDNA haplotypes within and between clusters were not significantly different. This study demonstrates that kinship correlates with associations among female bottlenose dolphins, but that kinship relations are not necessarily a prerequisite for membership in social clusters. We hypothesize that different forces acting on female bottlenose dolphin sociality appear to promote the formation of flexible groups which include both kin and nonkin.     

Shoaling decisions in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)—familiarity,kinship and inbreeding

Shoaling with familiar kin is a well-known phenomenon. It has been described both for adult and for fry of three-spined sticklebacks (Gasterosteus aculeatus). However, evidence of preference for kin independent of familiarity is scarce. Furthermore, inbreeding effects have not been studied for shoaling preferences and there is a lack of studies about changes in individuals’ ability to recognise kin during different phases of life history. We gave inbred and outbred, nonreproductive sticklebacks of different age the choice to shoal with a group of familiar siblings vs unfamiliar non-siblings and with unfamiliar siblings vs unfamiliar non-siblings. Subadult sticklebacks preferred to shoal with familiar kin over unfamiliar non-kin, but inbred and outbred individuals had similar preferences. When given the choice between unfamiliar siblings and unfamiliar non-sibs, adult outbred fish preferred the shoal of kin, while inbred fish behaved indifferently. Body characteristics of the group such as body mass, standard length and condition of its members did not significantly explain the shoaling preferences for kin. Thus, sticklebacks were capable of recognising familiar and unfamiliar kin. The latter capability was lost by inbreeding.     

Testosterone increases siblicidal aggression in black-legged kittiwake chicks (Rissa tridactyla)

To compete for parental food deliveries nestling birds have evolved diverse behaviors such as begging displays and sibling aggression. Testosterone has been suggested to be an important mechanism orchestrating such competitive behaviors, but evidence is scarce and often indirect. Siblicidal species provide an interesting case in which a clear dominance hierarchy is established and the dominant chicks lethally attack siblings. We experimentally elevated testosterone in chicks of a facultatively siblicidal species, the black-legged kittiwake, Rissa tridactyla, and showed that testosterone-treated chicks were more aggressive toward their sibling than were control chicks. In such facultatively siblicidal species, chicks normally exhibit intense aggression only when threatened by starvation. Indeed, we found that chicks in relatively poorer condition were more aggressive than were chicks in better condition, even among testosterone-treated chicks, suggesting the action of an additional signal modulating aggression. Relatively larger siblings were also more aggressive than were relatively smaller siblings, confirming the importance of size advantage in determining dominance hierarchies within the brood. In addition, testosterone increased aggression toward a simulated predator, indicating that in kittiwakes testosterone can increase aggression in contexts other than siblicide. Testosterone promoted aggression-mediated dominance, which increased begging although testosterone treatment did not have a significant separate effect on begging. Therefore, testosterone production in the kittiwake and most likely other siblicidal species seems an important fitness mediator already early in life, outside the sexual context and not only manifesting itself in aggressive behavior but also in dominance-mediated effects on food solicitation displays toward parents.     

Differential aggressiveness between fire salamander (<Emphasis Type="Italic">Salamandra infraimmaculata</Emphasis>) larvae covaries with their genetic similarity

Responding differentially to kin and non-kin is known to be adaptive in many species. One example is the inclusive fitness benefits of reducing aggression toward closer relatives. Little is known, however, about the ability of animals to assess differential degrees of genetic relatedness and to respond accordingly with differential levels of aggression. In the present study, we tested whether aggressiveness between body mass-matched pairs of fire salamander (Salamandra infraimmaculata) larvae covaried with the genetic similarity between them. We quantified aggressiveness at three levels of genetic similarity by selecting pairs within and across pools from recently genotyped populations. We also assessed aggression between pairs of siblings. Aggression and associated injuries decreased as genetic similarity increased across the groups. These findings suggest that cannibalistic salamanders can assess their degree of genetic relatedness to conspecifics and vary their behavioral responses depending on the degree of similarity between them along a genetic relatedness continuum.     

Population genetics of the family ostreidae. I. Intraspecific studies of Crassostrea gigas and Saccostrea commercialis

The levels of genetic variation and estimates of genetic similarity and distance were determined for populations of Crassostrea gigas (Thunberg) and Saccostrea commercialis (Iredale and Roughley). The proportion of polymorphic loci in Japanese populations of C. gigas ranged from 0.58 to 0.63, while the individual heterozygosities ranged from 0.20 to 0.22. The proportion of polymorphic loci in Australian populations of S. commercialis ranged from 0.43 to 0.46, the individual heterozygosities from 0.17 to 0.19. The genetic similarity between different geographical populations of both species was approximately 99%. The genetic distance and similarity data derived from this investigation has resulted in some provisional reclassifications. The Portuguese oyster C. angulata (Lamarck) appears to be more appropriately regarded as a recent colonized isolate of C. gigas. The New Zealand oyster S. glomerata (Gould) is considered a subspecies of the eastern Australian rock oyster S. commercialis. The Japanese Kumamoto population of C. gigas warrants reclassification as the non-sibling species C. sikamea.     

No apparent genetic basis to demographic differences in scarid fishes across continental shelf of the Great Barrier Reef

 Two species of parrot fish, Scarus frenatus and Chlorurus sordidus, are known to exhibit demographic and life-history differences across the continental shelf of the northern Great Barrier Reef (GBR). DNA sequences from the mitochondrial (mt) control region were analysed to determine whether there were genetic differences between the populations from the mid- and outer-shelf reefs. Analysis of molecular variance (AMOVA) indicated high levels of gene exchange for both species at a local scale between reefs on mid- and outer continental shelf positions (20 km apart) and at a broader scale along the length of the GBR province (>1000 km apart). There was no evidence to suggest that local differences in scarid life-history characteristics on the northern GBR have a genetic basis. Rather it appears more likely that phenotypically plastic responses to prevailing social and environmental conditions explain differences in the life-history characteristics of both taxa. However, analysis of genetic variation and historical demography revealed striking differences between the two species. S. frenatus haplotypes differed from one another at relatively few nucleotide sites (mean = 3.30), and the pairwise mismatch distribution suggested this species has undergone a population expansion within the limit of the resolution of the marker. C. sordidus haplotypes, however, differed from one another at a number of sites (mean = 7.67). Mismatch distribution analysis suggested that the population size of this species has remained at equilibrium over time. These patterns could also reflect differences in the metapopulation sizes or generation times between taxa. Some of the implications for fisheries management are discussed. Received: 28 January 2000 / Accepted: 9 July 2000     

Kelp (<Emphasis Type="Italic">Laminaria digitata</Emphasis>) increases germination and affects rooting and plant vigour in crops and native plants from an arable grassland in the Outer Hebrides,Scotland

Kelp and other seaweeds are traditionally used in many parts of the world as a soil amendment on arable fields. Seaweeds contain biochemical compounds that can act as plant growth regulators in terrestrial plants. In a low-intensity arable grassland in northwest Scotland an organic fertilizer, kelp (Laminaria digitata) has been used for hundreds of years, due to its anticipated positive effect as a soil conditioner and provider of plant nutrients. In this study the effects of kelp on germination and rooting of crops and native plants from this area were investigated in soil-free media. Germination was studied by incubation in the presence of kelp solutions. Rooting of plant cuttings was assessed after a pulse treatment with kelp solutions, and indole-3 acetic acid (IAA) as a reference plant growth regulator. Germination percentage of Plantago lanceolata, Trifolium repens and Avena strigosa seeds increased significantly when incubated with 0.05% kelp solutions. Total root weight and the individual weight of roots produced in cuttings of Vigna radiata and P. lanceolata were significantly increased when exposed to a 0.5% solution of kelp. Plant vigour, assessed visually, decreased significantly for P. lanceolata exposed to kelp at concentrations of 0.5 and 5.0% indicating the presence of a threshold level for an inhibitory effect of kelp at these concentrations, which may be due to high salinity. The results confirmed the presence of plant growth regulators in kelp, and indicates that amendment with kelp may potentially affect plant community composition. The threshold levels where some plants responded negatively to kelp amendment were close to or lower than the theoretical concentrations of kelp in soil water at field conditions with the current doses used on the machair, indicating that care should be taken in either administering kelp at the appropriate dose or leaching out salt before application.     

Mate choice in the face of both inbreeding and outbreeding depression in the intertidal copepod Tigriopus californicus

 In species vulnerable to both inbreeding and outbreeding depression, individuals might be expected to choose mates at intermediate levels of genetic relatedness. Previous work on the intertidal copepod Tigriopus californicus has repeatedly shown that crosses between populations result in either no effect or hybrid vigor in the first generation, and hybrid breakdown in the second generation. Previous work also shows that mating between full siblings results in inbreeding depression. The present study again found inbreeding depression, with full sibling mating causing significant fitness declines in two of the three populations assayed. In the mate choice assays, a single female was combined with two males. Despite the costs of both inbreeding and outbreeding, mate choice showed clear inbreeding avoidance but no clear pattern of outbreeding avoidance. This lack of outbreeding avoidance may be attributed either to the temporary increase in fitness in the F₁ generation or to the absence of selection for premating isolation in wholly allopatric populations with infrequent migration. If this inability to avoid unwise matings is common to other taxa, it may contribute to the problem of outbreeding depression when allopatric populations are mixed together. Received: 18 May 1999 / Accepted: 25 January 2000     

Food sharing by sibling vs nonsibling spiny mice (Acomys cahirinus)

Summary In comparison to paired unfamiliar unrelated age-mates, pairs of familiar littermate (sibling) spiny mice (Acomys cahirinus) were more active in exploring a novel environment containing limited food resources and engaged in a greater degree of cooperative feeding or food sharing. When animals born in successive litters were tested, however, familiar siblings did not differ from unfamiliar nonsiblings on measures of food sharing even though subsequent data indicate that such siblings could recognize one another. Young animals were found to feed for longer periods of time than their older cagemate regardless of whether the latter animal was a familiar sibling or an unfamiliar nonsibling.     

Familiarity with adults,but not relatedness,affects the growth of juvenile red-backed salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

The relative roles of kinship and familiarity in affecting an individual’s growth and fitness are not easy to disentangle. Not only is an individual more likely to have prior behavioral interactions with conspecifics in close proximity, it may also be related (in terms of kinship) to those nearby conspecifics. While some studies have inferred that kin discrimination affects fitness correlates, other studies found that familiarity alone can reduce aggressive interactions, thus increasing fitness. These studies have all focused on intra-age class pairs or groups. However, many animals interact with conspecifics from different cohorts. In many populations, adults of Plethodon cinereus territorially defend rocks and logs that retain moisture and food resources. We investigated whether juveniles of P. cinereus grew more in the presence of adults that were relatives or familiar. We collected pairs of juveniles and adults found under the same cover objects in the forest (familiar) and pairs of juveniles and adults found under different cover objects, approximately 10 m apart (unfamiliar). We determined parentage and relatedness of the adult–juvenile pairs and then placed these pairs in semi-natural mesocosms for 17 days. We found that juveniles housed with familiar adults had significantly greater increases in mass and snout–vent length than juveniles housed with unfamiliar adults in 2006 but not in 2007. Relatedness had no effect on growth. In addition, juveniles cohabitating with adults were not more likely to be their offspring. At least under certain environmental conditions, familiarity with adults, independent of relatedness or parentage, increased the growth of juvenile salamanders.     

Biochemical genetic variation and evidence of speciation in Chthamalus barnacles of the tropical eastern Pacific Ocean

Gene-enzyme variation was studied electrophoretically within and between barnacle populations of the genus Chthamalus from 8 intertidal stations from central California to the Pacific coast of Panamá. Horizontal starch-gel electrophoresis separated and resolved 18 enzymes from 512 individual barnacles. A maximum of 25 gene-enzyme systems was interpretable from the resulting zymograms. Electrophoretic phenotypes and patterns of phenotypic variation generally conformed with those observed in other organisms. The amount of genetic variation within barnacle populations varied; average heterozygosity, for example, ranged from a low of 3.4% in a Mexican population of the C. fissus group to a high of 10.4% in C. anisopoma from the Gulf of California. Observed and expected average heterozygosities agreed in all population samples, indicating that these species are outbreeding. In contrast to the prediction stemming from a hypothesis that trophic stability regulates the amount of genetic variation in marine species, average heterozygosity tends to be positively correlated with latitude. Data from these and other barnacle species may support a hybrid environmental heterogeneity-trophic diversity model recently proposed to explain genetic variation in decapod crustaceans. Juxtaposition of individuals from different localities revealed numerous genetic differences among populations of the C. fissus group. At least three partially sympatric sibling species are separated by genetic distances as large as those observed between the C. fissus group and the distinct species C. dalli and C. anisopoma. A cladistic analysis places C. anisopoma close to the Mexican and Panamanian sibling species, with C. fissus from San Diego and C. dalli successively farther removed.