Sexual size dimorphism and timing of spring migration in birds.

Sexually selected traits are limited by selection against those traits in other fitness components, such as survival. Thus, sexual selection favouring large size in males should be balanced by higher mortality of larger males. However, evidence from red-winged blackbirds (Agelaius phoeniceus) indicates that large males survive better than small males. A survival advantage to large size could result from males migrating north in early spring, when harsh weather favours large size for energetic reasons. From this hypothesis we predicted that, among species, sex differences in body size should be correlated with sex differences in timing of spring migration. The earlier males migrate relative to females, the larger they should be relative to females. We tested this prediction using a comparative analysis of data collected from 30 species of passerine birds captured on migration. After controlling for social mating system, we found that sexual size dimorphism and difference in arrival dates of males and females were significantly positively correlated. This result is consistent with the hypothesis that selection for survival ability promotes sexual size dimorphism (SSD), rather than opposes SSD as is the conventional view. If both natural selection and sexual selection favour large adult males, then limits to male size must be imposed before males become adults.

Caterpillar talk: acoustically mediated territoriality in larval Lepidoptera.

We provide evidence for conspecific acoustic communication in caterpillars. Larvae of the common hook-tip moth, Drepana arcuata (Drepanoidea), defend silk nest sites from conspecifics by using ritualized acoustic displays. Sounds are produced by drumming the mandibles and scraping the mandibles and specialized anal "oars" against the leaf surface. Staged interactions between a resident and intruder resulted in escalated acoustic "duels" that were typically resolved within minutes, but sometimes extended for several hours. Resident caterpillars generally won territorial disputes, regardless of whether they had built the nest, but relatively large intruders occasionally displaced residents from their nests. All evidence is consistent with acoustic signaling serving a territorial function. As with many vertebrates, ritualized signaling appears to allow contestants to resolve contests without physical harm. Comparative evidence indicates that larval acoustic signaling may be widespread throughout the Lepidoptera, meriting consideration as a principal mode of communication for this important group of insects.

Insights into population ecology and sexual selection in snakes through the application of DNA-based genetic markers.

Hypervariable genetic markers have revolutionized studies of kinship, behavioral ecology, and population biology in vertebrate groups such as birds, but their use in snakes remains limited. To illustrate the value of such markers in snakes, we review studies that have used microsatellite DNA loci to analyze local population differentiation and parentage in snakes. Four ecologically distinct species of snakes all show evidence for differentiation at small spatial scales (2-15 km), but with substantial differences among species. This result highlights how genetic analysis can reveal hidden aspects of the natural history of difficult-to-observe taxa, and it raises important questions about the ecological factors that may contribute to restricted gene flow. A 3-year study of genetic parentage in marked populations of the northern water snake showed that (1) participation in mating aggregations was a poor predictor of genetic-based measures of reproductive success; (2) multiple paternity was high, yet there was no detectable fitness advantage to multiple mating by females; and (3) the opportunity for selection was far higher in males than in females due to a larger variance in male reproductive success, and yet this resulted in no detectable selection on morphological variation in males. Thus genetic markers have provided accurate measures of individual reproductive success in this species, an important step toward resolving the adaptive significance of key features including multiple paternity and reversed sexual size dimorphism. Overall these studies illustrate how genetic analyses of snakes provide previously unobtainable information of long-standing interest to behavioral ecologists.

A comparison of RAPD versus microsatellite DNA markers in population studies of the massasauga rattlesnake.

We compared genetic differentiation among populations of the threatened massasauga rattlesnake (Sistrurus c. catenatus) using two types of nuclear molecular markers: randomly amplified polymorphic DNA (RAPD) markers and microsatellites. Analyses of molecular variance (AMOVA) and G(ST) and F(ST) analyses indicated that levels of among-population differentiation between regional populations (>100 km) were comparable for both markers. However, microsatellites were superior in population assignment tests and at discerning fine-scale genetic differentiation between subpopulations separated by tens of kilometers. These results argue that both types of markers are suitable for defining broad-scale genetic structures in snake populations and can provide important inputs into conservation initiatives of focal taxa. However, our analyses suggest that microsatellites 3re better for detecting structure at limited spatial scales.

Behavioral and life history responses of eastern massasauga rattlesnakes (Sistrurus catenatus catenatus) to human disturbance.

Parks and nature reserves protect important natural habitats but also provide public opportunities for outdoor recreational activities that may have unintended negative effects on wildlife. We examined the response of eastern massasauga rattlesnakes (Sistrurus catenatus catenatus) to inadvertent disturbance by humans in Killbear Provincial Park, Ontario, Canada. Radio telemetry of 25 adult snakes over two active seasons revealed that, as disturbance increased, gravid females were less visible to observers, but the visibility of non-gravid females and males did not change. Mean distance moved per day decreased and mean time between moves greater than 10 m increased in gravid females, non-gravid females and males with increasing exposure to human disturbance. However, mark-recapture data revealed no differences in the condition or growth rates of snakes, or in the litter size of gravid females, between individuals captured in disturbed and undisturbed study areas. While it is possible that the behavioral responses we observed are not sufficient to have life history consequences, more detailed information on the exposure of individual snakes to human activity is necessary before the conclusion that disturbance is not detrimental to snakes can be accepted. Similarly, other potential negative effects of human disturbance not investigated here remain to be explored.

Microgeographic population genetic structure in the northern water snake, Nerodia sipedon sipedon detected using microsatellite DNA loci.

We describe the isolation and genetic characterization of eight microsatellite DNA loci from the northern water snake, Nerodia sipedon sipedon and use these loci to analyse levels of genetic differentiation between local (< 2 km apart) populations of these snakes in Ontario. These loci are variable, with expected heterozygosities ranging from 0.28 to 0.91, and can correctly exclude nonsires in parentage analyses with a high probability (0.998). Population analyses reveal significant deviation from expected heterozygosity levels for one population, probably a result of a null allele(s) at a single locus and small but significant levels of genetic differentiation among all three populations. This demonstrates that microgeographic genetic structure exists in this species, possibly due to limited dispersal.

Lekking without a paradox in the buff-breasted sandpiper.

Females in lek-breeding species appear to copulate with a small subset of the available males. Such strong directional selection is predicted to decrease additive genetic variance in the preferred male traits, yet females continue to mate selectively, thus generating the lek paradox. In a study of buff-breasted sandpipers (Tryngites subruficollis), we combine detailed behavioral observations with paternity analyses using single-locus minisatellite DNA probes to provide the first evidence from a lek-breeding species that the variance in male reproductive success is much lower than expected. In 17 and 30 broods sampled in two consecutive years, a minimum of 20 and 39 males, respectively, sired offspring. This low variance in male reproductive success resulted from effective use of alternative reproductive tactics by males, females mating with solitary males off leks, and multiple mating by females. Thus, the results of this study suggests that sexual selection through female choice is weak in buff-breasted sandpipers. The behavior of other lek-breeding birds is sufficiently similar to that of buff-breasted sandpipers that paternity studies of those species should be conducted to determine whether leks generally are less paradoxical than they appear.

Genetic structure of populations of the threatened eastern massasauga rattlesnake, Sistrurus c. catenatus: evidence from microsatellite DNA markers.

Throughout its distribution in North America, the threatened eastern massasauga rattlesnake (Sistrurus c. catenatus) persists in a series of habitat-isolated disjunct populations of varying size. Here, we use six microsatellite DNA loci to generate information on the degree of genetic differentiation between, and the levels of inbreeding within populations to understand how evolutionary processes operate in these populations and aid the development of conservation plans for this species. Samples were collected from 199 individuals from five populations in Ontario, New York and Ohio. Our results show that all sampled populations: (i) differ significantly in allele frequencies even though some populations are < 50 km apart, and may contain genetically distinct subpopulations < 2 km apart; (ii) have an average of 23% of alleles that are population specific; and (iii) have significant FIS values (mean overall FIS = 0.194) probably due to a combination of Wahlund effects resulting from fine-scale genetic differentiation within populations and the presence of null alleles. Our results imply that massasauga populations may be genetically structured on an extremely fine scale even within continuous populations, possibly due to limited dispersal. Additional information is needed to determine if dispersal and mating behaviour within populations can account for this structure and whether the observed differentiation is due to random processes such as drift or to local adaptation. From a conservation perspective, our results imply that these massasauga populations should be managed as demographically independent units and that each has high conservation value in terms of containing unique genetic variation.

Realized reproductive success of polygynous red-winged blackbirds revealed by DNA markers.

Hypervariable genetic markers, including a novel locus-specific marker detected by a mouse major histocompatibility complex probe, reveal that multiple paternity is common in families of polygynous red-winged blackbirds (Agelaius phoeniceus). Almost half of all nests contained at least one chick resulting from an extra-pair fertilization, usually by a neighboring male. Genetically based measures of reproductive success show that individual males realize more than 20% of their overall success from extra-pair fertilizations, on average, and that this form of mating behavior confounds traditional measures of male success. The importance of alternative reproductive tactics in a polygynous bird is quantified, and the results challenge previous explanations for the evolution of avian polygny.