I am an evolutionary biologist with a broad collaborative research program that employs molecular genetic tools to ask basic questions regarding biological evolution. To date my collaborators and I have examined the behavioral underpinnings of genetic mating systems, molecular systematics, population genetics and molecular based phylogeography primarily in avian species.
Abstract: The lowland tailorbirds of Southeast Asia (Orthotomus) offer an excellent opportunity for comparative biogeography because of their diversity in the Greater Sunda and Philippine islands. We reconstructed the phylogeny of all species in the genus using maximum likelihood, Bayesian, and coalescent methods on DNA sequences of three gene segments: an autosomal intron (TGF), a Z-linked intron (MUSK), and a mitochondrial coding gene (ND2). Although resolution is low in parts of the phylogeny, several well defined clades emerge. When considered in light of distribution, these clades indicate that the Greater Sunda and Philippine islands were occupied early in Orthotomus history by the ancestors of O. sericeus in the Greater Sundas and O. frontalis in the Philippines. Subsequently, tailorbirds diversified further in each island group: O. atrogularis, O. ruficeps, and O. sepium arose in the Greater Sundas, and O. castaneiceps castaneiceps, O. c. chloronotus, O. derbianus, O. samarensis, O. nigriceps, and O. cinereiceps in the Philippines. Among the continental taxa (including Sundaic birds), the older lineages (O. sutorius and O. sericeus) are habitat generalists and the recently evolved taxa are more specialized. In the Philippines, several taxa once considered conspecific with O. atrogularis turn out to be highly divergent species (>9% in ND2). Indeed, all Philippine allospecies are well diverged from one another. This finding supports the recent assertion of higher-than-appreciated bird endemicity in the Philippines.
Abstract: Sexing of raptors is important for understanding their ecology and demography. Males and females of monomorphic species such as Red-shouldered Hawks (Buteo lineatus) may be distinguished using molecular and morphometric techniques. We collected blood samples and morphometric measurements from adult and nestling Red-shouldered Hawks in southern Ohio. We determined sex via amplification of the sex-linked chromo-helicase-DNA-binding gene and polymerase chain reaction. We used a suite of morphometric measurements to generate a recursive partitioning classification tree and in a linear discriminant analysis to determine the sex of adults and nestlings. For adults, the recursive partitioning tree utilized only mass to distinguish sexes, with an overall successful classification rate of 94%. For nestling hawks aged approximately 3 wk and older, mass and toepad (footpad) length were used to distinguish the sexes, with an overall successful classification rate of 91%. The ability to sex adults and nestlings in the field is valuable for studies of dispersal, survival, and behavior.
Abstract: Phylogenetic relationships among the 14 manakin genera were inferred from DNA sequence data obtained from both mitochondrial and nuclear DNA loci. Phylogenetic analysis resulted in a well-supported hypothesis that corroborates a sister relationship between tyrant-manakins and the "core" manakins (Antilophia, Chiroxiphia, Corapipo, Dixiphia, Heterocercus, Ilicura, Lepidothrix, Manacus, Masius, Machaeropterus, Pipra, and Xenopipo). Our data strongly support these core manakin genera as a monophyletic group. Consistent with previous work, we find two major clades within the core manakins, although the placement of the genus Xenopipo with regards to these two clades is ambiguous. Generic relationships within these clades are generally well resolved. Although we find some concordance between our study and a previous manakin phylogeny based on syringeal characters, we note several fundamental differences between the phylogenies. Thus, we offer a new phylogenetic hypothesis for Pipridae.
Abstract: The subtropical island of Taiwan is an area of high endemism and a complex topographic environment. Phylogeographic studies indicate that vicariance caused by Taiwan's mountains has subdivided many taxa into genetic phylogroups. We used mitochondrial DNA sequences and nuclear microsatellites to test whether the evolutionary history of an endemic montane bird, Steere's Liocichla (Liocichla steerii), fit the general vicariant paradigm for a montane organism.
Abstract: A growing body of evidence shows that female birds use male plumage coloration as an important criterion in mate choice. In the field, however, males with brighter coloration may both compete better for high-quality territories and be the object of female choice. Positive associations between territory quality, male-male competitive ability and female preferences can make it difficult to determine whether females actively choose the most ornamented males. Male eastern bluebirds, Sialia sialis, display brilliant ultraviolet (UV)-blue plumage coloration on their heads, backs, wings and tails and chestnut coloration on their breasts, which is positively correlated with condition, reproductive effort and reproductive success. We tested the hypothesis that female bluebirds prefer males that display brighter and more chromatic coloration by experimentally widowing males in the field and allowing replacement females to choose partners. We controlled for the influence of territory quality on female choice by widowing dyads of males with adjacent territories. We found no evidence that UV-blue or chestnut plumage coloration, body size or body condition predicted the male with which a female would pair. We found no support for the hypothesis that the coloration of male eastern bluebirds functions as a criterion in female mate choice.
Abstract: This special issue of Genetica brings together empirical researchers and theoreticians to present the latest on the evolutionary ecology of genetic quality in the context of sexual selection. The work comes from different fields of study including behavioral ecology, quantitative genetics and molecular genetics on a diversity of organisms using different approaches from comparative studies, mathematical modeling, field studies and laboratory experiments. The papers presented in this special issue primarily focus on genetic quality in relation to (1) sources of genetic variation, (2) polyandry, (3) new theoretical developments and (4) comprehensive reviews.
Abstract: Data from avian species have played a prominent role in developing and testing theories of female mate choice. One of the most prominent models of sexual selection, the "good genes" model, emphasizes the indirect benefits of female preferences for male ornaments as indicators of a potential sire's additive genetic quality. However, there is growing interest in non-additive sources of genetic quality and mate choice models for self-referential disassortative mating based on optimal levels of genetic dissimilarity. We reviewed the empirical evidence for genetic-complementarity-based female mate choice among birds. We found the evidence for such choice is mixed but in general against the genetic complementarity hypothesis. The lack of evidence for genetic complementarity in many birds may be due to an inability to make the fine distinctions among potential mates based on genes, possibly due to the comparative anosmatic nature of avian sensory system. For some species however there is compelling evidence for genetic complementarity as a criterion used in female mate choice. Understanding the ubiquity of female mate choice based on genetic complementarity and the variation in this source of female preference among and within species remains a challenge.
Abstract: Although sexual differences in birds can be extreme, differences between males and females in body size and plumage color are more subtle in many species. We used a genetic-based approach to determine the sex of male and female Steere's Liocichla (Liocichla steerii) and examine the degree of size dimorphism and plumage dichromatism in this apparently monomorphic species. We found that males were significantly larger than females. In addition, Steere's Liocichla have a prominent yellow plumage patch on the lores that was significantly larger in males than females for both live birds and museum specimens. We also used reflectance spectrometry to quantify the color of the yellow-green breast feathers of Steere's Liocichla and found no significant differences between males and females in brightness, intensity, saturation, or hue. However, females tended to have brighter breast plumage, particularly at long wavelengths. Collectively, these color variables were useful in discriminating birds according to sex when used in a discriminant function analysis. Our study suggests that sexual selection may be more widespread than once assumed, even among birds considered monomorphic, and emphasizes the need for additional data from tropical and subtropical species.
Abstract: For species in which females mate outside of the pair bond, selection should favor male mate-guarding behaviors that minimize investment in genetically unrelated offspring. Mate guarding may impose costs by diverting time and energy from activities such as foraging and seeking extra-pair copulations, so males should adjust their mate-guarding behavior according to the risk of cuckoldry. In this study, we investigated cuckoldry and mate guarding in the blue grosbeak (Passerina caerulea L., 1758), a socially monogamous, territorial songbird. Fifty-three percent of nestlings were extra-pair, and 70% of nests had at least one extra-pair fertilization. Males with dull plumage coloration were more likely to be cuckolded than males with more brightly colored plumage, but duller males did not guard their mates more. Instead, males appeared to adjust mate-guarding behavior according to the attractiveness of neighbors. Males with more brightly colored neighbors guarded their mates more intensely than males with fewer colorful neighbors. These observations suggest that in the blue grosbeak, mate guarding is a context-dependent strategy that is adjusted according to the density and ornamentation of neighboring males.
Abstract: Extra-pair paternity (EPP) is a widespread and highly variable reproductive phenomenon in birds. We tested the effects of habitat, spatial factors, and timing of breeding on the occurrence of EPP in red-winged blackbirds (Agelaius phoeniceus). We used PCR-amplified microsatellites to assess the paternity of 1479 nestlings from 537 broods on 235 territories over four breeding seasons. Over 4 years, 40% of nestlings were extra-pair. At least 27% of actual sires were non-neighbours, suggesting that males or females interacted over longer distances than in other populations of red-winged blackbirds. The level of EPP was significantly clumped within broods and males but not within females across broods. EPP was negatively related to the area of a male's territory. The spatial proximity of a female's nest to the territory boundary had no effect on total EPP, but tended to increase the probability of an EPP by a nearby male. We found no influence on EPP of the type of habitat on the territory or the level of nesting activity nearby. The time in the season when a nest was started and the synchrony of breeding also had no significant effect on the level of EPP. The age of the male, the age of his neighbours, and the interaction between the two had no effect on total EPP. However, older males were less likely to have an offspring sired by a neighbour on their territory. Males with older neighbours were also less likely to have offspring sired by a neighbour, particularly if they were new territory owners. The high variability in who gained and lost paternity, and the limited impact of spatial and temporal factors influencing it, have some interesting implications for theories seeking to explain mating patterns.
Abstract: Female choice for male ornamental traits is widely accepted as a mechanism by which females maximize their reproductive success and/or offspring quality. However, there is an increasing empirical literature that shows a fitness benefit of genetic diversity and a tendency for females to use genetic dissimilarity as a criterion for mate choice. This genetic compatibility hypothesis for female mate choice presents a paradox. How can females use both an absolute criterion, such as male ornamentation, and a relative criterion, such as genetic dissimilarity, to choose their mates? Here, we present potential solutions for this dilemma and the empirical evidence supporting them. The interplay between these two contrasting forms of female mate choice presents an exciting empirical and theoretical challenge for evolutionary ecologists.
Abstract: Studies of the patterns of diversification of birds on islands have contributed a great deal to the development of evolutionary theory. In white-winged fairy-wrens, Malurus leucopterus, mainland males develop a striking blue nuptial plumage whereas those on nearby islands develop black nuptial plumage. We explore the proximate basis for this divergence by combining microstructural feather analysis with an investigation of genetic variation at the melanocortin-1 receptor locus (MC1R). Fourier analysis revealed that the medullary keratin matrix (spongy layer) of the feather barbs of blue males was ordered at the appropriate nanoscale to produce the observed blue colour by coherent light scattering. Surprisingly, the feather barbs of black males also contained a spongy layer that could produce a similar blue colour. However, black males had more melanin in their barbs than blue males, and this melanin may effectively mask any structural colour produced by the spongy layer. Moreover, the presence of this spongy layer suggests that black island males evolved from a blue-plumaged ancestor. We also document concordant patterns of variation at the MC1R locus, as five amino acid substitutions were perfectly associated with the divergent blue and black plumage phenotypes. Thus, with the possible involvement of a melanocortin receptor locus, increased melanin density may mask the blue-producing microstructure in black island males, resulting in the divergence of plumage coloration between mainland and island white-winged fairy-wrens. Such mechanisms may also be responsible for plumage colour diversity across broader geographical and evolutionary scales.
Abstract: A monogamous mating system that includes extrapair fertilization can potentially generate higher variability in male reproductive success than monogamy without extrapair fertilization. That increased variability could provide a correspondingly higher opportunity for sexual selection and, thus, for the origin and persistence of sexual dimorphism in monogamous species. To determine whether extrapair fertilization enhanced the opportunity for sexual selection in a sexually dimorphic, monogamous bird species, we used microsatellite DNA typing to assess the prevalence of extrapair fertilization and its effect on variation in male reproductive success in a population of Chestnut-sided Warblers (Dendroica pensylvanica). We found that the level of extrapair fertilization in our study population was at the upper end of the range reported for bird populations (47% of nestlings had extrapair fathers; 61% of broods contained extrapair offspring). We also discovered that almost all extrapair offspring were sired by paired males resident on nearby territories. In addition, we found that variation in male reproductive success was substantially higher than variation in female reproductive success, and that extrapair fertilizations made a significant contribution to variation in male reproductive success. Together, those finding!; suggest that extrapair fertilization creates an opportunity for sexual selection on male traits in this population.
Abstract: A detailed understanding of the behavioural interactions between males and females is crucial for elucidating the selection pressures shaping mating system evolution; however, these interactions are often difficult to observe, particularly in free-living populations. We simulated extraterritorial intrusions by presenting conspecific models on to territories of a Neotropical migratory passerine, the yellow-breasted chat, Icteria virens. Simulated intrusions elicited different responses based on the sex of the focal individual and the sex and species of the model. Behavioural responses of the focal individuals were largely directed towards conspecific models, whereas simulated heterospecific intrusions, using a Carolina wren, Thryothorus ludovicianus, model, elicited minimal response. Male song was not related to any particular model presentation. Males directed mating behaviours, including courtship displays and copulations, exclusively towards the female conspecific model. Males behaved aggressively towards the conspecific male model. In contrast, females never showed any courtship behaviour towards the male model and showed significant aggression towards the conspecific female model. The results of this study reveal that territorial aggression in yellow-breasted chats is strongly intrasexual. Additionally, simulated female extraterritorial intrusions, but not male intrusions, resulted in extrapair courtship and copulation. This pattern is likely to be typical of many socially monogamous species and points to the behavioural mechanisms underlying both monogamy and extrapair mating systems. (C) 2004 The Association for the Study or Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
Abstract: Sexual dimorphism or dichromatism has long been considered the result of sexual selection. However, for many organisms the degree to which sexual dichromatism occurs has been determined within the confines of human perception. For birds, objective measures of plumage color have revealed previously unappreciated sexual dichromatism for several species. Here we present an unbiased assessment of plumage dichromatism in the yellow-breasted chat Icteria virens. Chats exhibit yellow to orange throat and breast plumage that to the unaided human observer differs only subtly in color. Spectrophotometric analyses revealed that chat throat and breast feathers exhibited reflective curves with two peaks, one in the ultraviolet and one in the yellow end of the spectrum. We found differences in both the shape and magnitude of reflectance curves between males and females. Moreover, for feathers collected from the lower edge and middle of the breast patch, male plumage reflected more light in the ultraviolet and yellow wavelengths compared to females, whereas male throat feathers appeared brighter than those of females only in the ultraviolet. Biochemical analyses indicated that the plumage pigmentation consisted solely of the carotenoid all-trans lutein and we found that males have higher concentrations of plumage carotenoids than females. Feathers that were naturally unpigmented reflected more UV light than yellow feathers, suggesting a potential role of feather microstructure in UV reflectance.
Abstract: Extra-pair paternity is widely documented in birds, but the behaviors that lead to extra-pair copulations (EPCs) have been less well studied, particularly in territorial songbirds. We studied the behaviors associated with extra-territorial forays (ETFs) and male mate guarding in a socially monogamous, but genetically promiscuous, neotropical migrant passerine, the yellow-breasted chat (Icteria virens). Focal observations and radiotelemetry revealed that both males and females engaged in ETFs. 65% of the females in our study engaged in at least one foray onto a neighboring territory. 50% of males also were observed engaging in ETFs, but males were much more conspicuous during these intrusions compared to females. Females preferred to remain close to the ground in areas where vegetation was the densest. Female behavior was difficult to observe during ETFs but females sometimes interacted with neighboring extra-pair males. Males attempted to guard their mates by remaining close to them and following them during extra-territorial intrusions. We observed instances in which a male attacked his mate and appeared to herd her back to his territory. However a male's ability to maintain close proximity to his mate was significantly and negatively correlated with vegetation density. Our results suggest that the behaviors which lead to extra-pair encounters are influenced by the behavior of all participants and are modified by the characteristics of the habitat.
Abstract: Mycoplasma gallisepticum, a major pathogen of chickens and turkeys, has caused significant declines in house finch (Carpodacus mexicanus) populations in the eastern United States since it was first observed in this species in 1994. There is evidence that M. gallisepticum infection is now endemic among eastern house finches, although disease prevalence has declined, suggesting an evolving host-parasite relationship. Studies based on randomly amplified polymorphic DNA (RAPD) have documented the presence of a single, unique RAPD profile in house finch M. gallisepticum isolates, suggesting a single point source of origin, which agrees with the known epidemiologic observations. In the present study, we evaluated the molecular variability of 55 house finch isolates as well as 11 chicken and turkey isolates including reference strains of M. gallisepticum. Molecular variability was evaluated by polymerase chain reaction (PCR)-restriction fragment length polymorphism (RFLP) analysis and nucleotide sequencing of the pvpA gene, which encodes for the putative cytadhesin protein PvpA. Three different RFLP groups and 16 genotypes were evident from the 55 house finch isolates evaluated. Sequence analysis of pvpA gene PCR products showed that although most house finch M. gallisepticum isolates clustered more closely to each other, others clustered more closely to either turkey or chicken field isolates. These findings suggest that house finch isolates are more polymorphic than previously recognized by RAPD studies. This feature may allow us to learn more about the molecular evolution and epidemiology of this emerging disease host-parasite relationship.
