H. Lisle Gibbs

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.

Genetic evidence for female host-specific races of the common cuckoo

The common cuckoo Cuculus canorus is divided into host-specific races (gentes). Females of each race lay a distinctive egg type that tends to match the hosts eggs, for instance, brown and spotted for meadow pipit hosts or plain blue for redstart hosts. The puzzle is how these gentes remain distinct. Here, we provide genetic evidence that gentes are restricted to female lineages, with cross mating by males maintaining the common cuckoo genetically as one species. We show that there is differentiation between gentes in maternally inherited mitochondrial DNA, but not in microsatellite loci of nuclear DNA. This supports recent behavioural evidence that female, but not male, common cuckoos specialize on a particular host, and is consistent with the possibility that genes affecting cuckoo egg type are located on the female-specific W sex chromosome. Our results also support the ideas that common cuckoos often switched hosts during evolution, and that some gentes may have multiple, independent origins, due to colonization by separate ancestral lineages.

Phylogeography and genetic structure of northern populations of the yellow warbler ( Dendroica petechia )

Phylogeographic patterns of intraspecific variation can provide insights into the population‐level processes responsible for speciation and yield information useful for conservation purposes. To examine phylogeography and population structure in a migratory passerine bird at both continental and regional geographical scales, we analysed 344 bp of mitochondrial DNA (mtDNA) control region sequence from 155 yellow warblers (Dendroica petechia) collected from seven locations across Canada and from Alaska. There is a major subdivision between eastern (Manitoba to Newfoundland) and western (Alaska and British Columbia) populations which appears to have developed during the recent Pleistocene. Some localities within these two regions also differ significantly in their genetic composition, suggesting further subdivision on a regional geographical scale. Eastern and western birds form distinct phylogeographic entities and the clustering of all western haplotypes with two eastern haplotypes suggests that the western haplotypes may be derived from an eastern lineage. Analyses based on coalescent models support this explanation for the origin of western haplotypes. These results are consistent with important features of Mengel’s model of warbler diversification. From a conservation perspective they also suggest that individual populations of migrant birds may form demographically isolated management units on a smaller scale than previously appreciated.

Relatedness and helping in fish: examining the theoretical predictions

Many studies have attempted to explain the evolution of cooperation, yet little attention has been paid to what factors control the amount or kind of cooperation performed. Kin selection theory suggests that more cooperation, or help, should be given by relatives. However, recent theory suggests that under specific ecological and demographic conditions, unrelated individuals must ‘pay to stay’ in the group and therefore may help more. We tested these contrasting predictions using the cooperatively breeding fish, Neolamprologus pulcher, and found that the degree of work effort by helpers depended on which helping behaviours were considered and on their level of relatedness to the breeding male or female. In the field, helpers unrelated to the breeding male performed more territory defence, while helpers unrelated to the breeding female contributed less to territory defence. In the laboratory, unrelated group members helped more. Our work demonstrates that a number of factors in addition to kinship shape cooperative investment patterns.

Inferring Species-Level Phylogenies and Taxonomic Distinctiveness Using Multilocus Data in Sistrurus Rattlesnakes

Phylogenetic relationships and taxonomic distinctiveness of closely related species and subspecies are most accurately inferred from data derived from multiple independent loci. Here, we apply several approaches for understanding species-level relationships using data from 18 nuclear DNA loci and 1 mitochondrial DNA locus within currently described species and subspecies of Sistrurus rattlesnakes. Collectively, these methods provide evidence that a currently described species, the massasauga rattlesnake (Sistrurus catenatus), consists of two well-supported clades, one composed of the two western subspecies (S. c. tergeminus and S. c. edwardsii) and the other the eastern subspecies (S. c. catenatus). Within pigmy rattlesnakes (S. miliarius), however, there is not strong support across methods for any particular grouping at the subspecific level. Monophyly based tests for taxonomic distinctiveness show evidence for distinctiveness of all subspecies but this support is strongest by far for the S. c. catenatus clade. Because support for the distinctiveness of S. c. catenatus is both strong and consistent across methods, and due to its morphological distinctiveness and allopatric distribution, we suggest that this subspecies be elevated to full species status, which has significant conservation implications. Finally, most divergence time estimates based upon a fossil-calibrated species tree are > 50% younger than those from a concatenated gene tree analysis and suggest that an active period of speciation within Sistrurus occurred within the late Pliocene/Pleistocene eras.

Functional basis of a molecular adaptation: prey-specific toxic effects of venom from Sistrurus rattlesnakes.

Understanding the molecular bases of adaptations requires assessing the functional significance of phenotypic variation at the molecular level. Here we conduct such an assessment for an adaptive trait (snake venom proteins) which shows high levels of interspecific variation at the molecular level. We tested the toxicity of venom from four taxa of Sistrurus rattlesnakes with different diets towards 3 representative prey (mice, lizards and frogs). There were significant differences among prey in their overall susceptibility to Sistrurus venom, with frogs being an order of magnitude more resistant than mice or lizards. However, only in mice was there substantial variation in the toxicity of venom from different Sistrurus taxa, with the variation being roughly correlated with the incidence of mammals in the snakes diet. A comparative analysis using published data of the toxicity of rattlesnake and outgroup (Agkistrodon) venoms to mice confirms that both the gain and loss of toxicity to mammals were major modes of venom evolution in Sistrurus catenatus and Sistrurus miliarius. Our findings identify toxicity to mammals as a major axis along which venom evolution has occurred among Sistrurus rattlesnakes, with little evidence for evolutionary changes in toxicity towards the other prey tested. They also emphasize the need to consider ecological and evolutionary factors other than diet alone as causes of variation in venom toxicity.

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.

The Imprint of History on Communities of North American and Asian Warblers

The ecology of the component species of an adaptive radiation is likely to be influenced by the form of the founding ancestor to the radiation, its timing, and rates of speciation and extinction. These historical features complement environmental selection pressures. They imply that, if the history of the species’ radiations are very different, ecological communities are unlikely to be completely convergent even when placed in identical environments. We compare the adaptive radiation of the Dendroica warblers of North America with that of the Phylloscopus warblers of Asia. We consider the ecology of the species in two localities where species’ diversity is very high (New Hampshire, U.S.A., and Kashmir, India, respectively) and contrast the history of the two radiations on the basis of a molecular (mitochondrial cytochrome b) phylogeny. By comparison with the Phylloscopus, the Dendroica are on average larger and morphologically more similar to one another. Although there is some similarity between the Dendroica and Phylloscopus communities, they differ in foraging behavior and in associations of morphology with ecological variables. The Dendroica likely reflect an early Pliocene radiation and are two to four times younger than the Phylloscopus. They probably had a colorful sexually dichromatic ancestor, implicating sexual selection in the production of the many ecologically similar species. The Phylloscopus are much older and probably had a drab, monomorphic ancestor. Given the difference in ages of the two radiations, it is plausible that the close species’ packing of the Dendroica warblers is a transient phenomenon. If this is the case, community structure evolves on the timescales of millions of years. Differences in ancestry and timing of the species’ radiations can be related to the different biogeography of the two regions. This implies that the historical imprint on adaptive radiations could be predicted on the basis of the attributes of ancestors and biogeographical context.

Ancestral polymorphisms in genetic markers obscure detection of evolutionarily distinct populations in the endangered Florida grasshopper sparrow ( Ammodramus savannarum floridanus )

Genetic analyses of bird subspecies designated as conservation units can address whether they represent units with independent evolutionary histories and provide insights into the evolutionary processes that determine the degree to which they are genetically distinct. Here we use mitochondrial DNA control region sequence and six microsatellite DNA loci to examine phylogeographical structure and genetic differentiation among five North American grasshopper sparrow (Ammodramus savannarum) populations representing three subspecies, including a population of the endangered Florida subspecies (A. s. floridanus). This federally listed taxon is of particular interest because it differs phenotypically from other subspecies in plumage and behaviour and has also undergone a drastic decline in population size over the past century. Despite this designation, we observed no phylogeographical structure among populations in either marker: mtDNA haplotypes and microsatellite genotypes from floridanus samples did not form clades that were phylogenetically distinct from variants found in other subspecies. However, there was low but significant differentiation between Florida and all other populations combined in both mtDNA (FST = 0.069) and in one measure of microsatellite differentiation (θ = 0.016), while the non‐Florida populations were not different from each other. Based on analyses of mtDNA variation using a coalescent‐based model, the effective sizes of these populations are large (∼80 000 females) and they have only recently diverged from each other (< 26 000 ybp). These populations are probably far from genetic equilibrium and therefore the lack of phylogenetic distinctiveness of the floridanus subspecies and minimal genetic differentiation is due most probably to retained ancestral polymorphism. Finally, levels of variation in Florida were similar to other populations supporting the idea that the drastic reduction in population size which has occurred within the last 100 years has not yet had an impact on levels of variation in floridanus. We argue that despite the lack of phylogenetic distinctiveness of floridanus genotypes the observed genetic differentiation and previously documented phenotypic differences justify continued designation of this subspecies as a protected population segment.

Limited differentiation in microsatellite DNA variation among northern populations of the yellow warbler: evidence for male‐biased gene flow?

Comparisons of the patterns of differentiation among genetic markers with different modes of inheritance can provide insights into patterns of sex‐biased dispersal and gene flow. Here, we compare the patterns of differentiation in six microsatellite loci among eight northern breeding populations of the yellow warbler (Dendroica petechia) with results obtained with mitochondrial DNA. Significant but low levels of differentiation (overall FST = 0.014; overall RST = 0.015) were present across all populations. The level of differentiation is substantially less than that observed in the same samples based on mitochondrial DNA control region variation. The presence of low population imbalance index values and significant isolation‐by‐distance relationships for both FST and RST suggests that these populations are at evolutionary equilibrium and that the high degree of similarity between populations may be due to high levels of male‐biased gene flow. This suggests that there may be significant but previously unappreciated differences in the long‐distance and/or episodic dispersal behaviour of males and females in these birds.