Ronald A. Van Den Bussche

FASTER LIZARDS SIRE MORE OFFSPRING: SEXUAL SELECTION ON WHOLE-ANIMAL PERFORMANCE

Sexual selection operates by acting on variation in mating success. However, since selection acts on wholeorganism manifestations (i.e., performance) of underlying morphological traits, tests for phenotypic effects of sexual selection should consider whole-animal performance as a substrate for sexual selection. Previous studies have revealed positive relationships between performance and survival, that is, natural selection, but none have explicitly tested whether performance may influence reproductive success (through more matings), that is, sexual selection. Performance predicts dominance in some species, implying the effects of sexual selection, but how it does so has not been established, nor is it certain whether performance might be a by-product of selection for something else, for example, elevated circulating testosterone levels. We investigated the potential for sexual selection on sprint speed performance in collared lizards (Crotaphytus collaris), considering the potential mediating effects of circulating hormone levels. Among territorial, adult male collared lizards, only sprint speed significantly predicted territory area and number of offspring sired as determined by genetic paternity analysis. Body size, head size, and hind limb length had no effect. Neither plasma testosterone levels nor corticosterone levels correlated with sprint speed, territory area, or number of offspring sired. Thus, our results provide a direct link between whole-animal performance and reproductive success, suggesting that intrasexual selection can act directly on sprint speed performance and drive the evolution of underlying morphological traits.

MAJOR-HISTOCOMPATIBILITY-COMPLEX-ASSOCIATED VARIATION IN SECONDARY SEXUAL TRAITS OF WHITE-TAILED DEER (ODOCOILEUS VIRGINIANUS): EVIDENCE FOR GOOD-GENES ADVERTISEMENT

Abstract Good‐genes hypotheses predict that development of secondary sexual characters can be an honest advertisement of heritable male quality. We explored this hypothesis using a cervid model (adult, male white‐tailed deer, Odocoileus virginianus) to determine whether antler development could provide an honest signal of a males genetic quality and condition to adversaries. We compared antler, morphometric, hormonal, and parasitic data collected from hunter‐harvested deer to characteristics of the Mhc‐DRB (Odvi), the most widely studied gene of the major histocom‐patibility complex (MHC) in Artiodactyla. We detected associations between genetic characteristics at Odvi‐DRB and antler development and body mass, suggesting that antler development and body mass may be associated with pathogen resistance in deer and thus may be an honest signal of genetic quality. We also detected associations between Odvi‐DRB characteristics and serum testosterone during the breeding season, suggesting that certain MHC characteristics may help deer cope with stresses related to breeding activity. In addition, we observed a negative relationship between degree of antler development and overall abundance of abomasal helminths. Our observations provide support for the hypothesis that antler development in white‐tailed deer is an honest signal of quality.

Multiple geographic origins of commensalism and complex dispersal history of black rats

The Black Rat (Rattus rattus) spread out of Asia to become one of the worlds worst agricultural and urban pests, and a reservoir or vector of numerous zoonotic diseases, including the devastating plague. Despite the global scale and inestimable cost of their impacts on both human livelihoods and natural ecosystems, little is known of the global genetic diversity of Black Rats, the timing and directions of their historical dispersals, and the risks associated with contemporary movements. We surveyed mitochondrial DNA of Black Rats collected across their global range as a first step towards obtaining an historical genetic perspective on this socioeconomically important group of rodents. We found a strong phylogeographic pattern with well-differentiated lineages of Black Rats native to South Asia, the Himalayan region, southern Indochina, and northern Indochina to East Asia, and a diversification that probably commenced in the early Middle Pleistocene. We also identified two other currently recognised species of Rattus as potential derivatives of a paraphyletic R. rattus. Three of the four phylogenetic lineage units within R. rattus show clear genetic signatures of major population expansion in prehistoric times, and the distribution of particular haplogroups mirrors archaeologically and historically documented patterns of human dispersal and trade. Commensalism clearly arose multiple times in R. rattus and in widely separated geographic regions, and this may account for apparent regionalism in their associated pathogens. Our findings represent an important step towards deeper understanding the complex and influential relationship that has developed between Black Rats and humans, and invite a thorough re-examination of host-pathogen associations among Black Rats.

Molecular phylogeny and biogeography of North American isolates of Anaplasma marginale (Rickettsiaceae: Ehrlichieae)

Anaplasma marginale (A. marginale) is a tick-borne ehrlichial pathogen of cattle that causes the disease anaplasmosis. Six major surface proteins (MSPs) have been identified on A. marginale from cattle and ticks of which three, MSP1a, MSP4 and MSP5, are from single genes and do not vary within isolates. The other three, MSP1b, MSP2 and MSP3, are from multigene families and may vary antigenically in persistently infected cattle. Several geographic isolates have been identified in the United States which differ in morphology, protein sequence and antigenic properties. An identifying characteristic of A. marginale isolates is the molecular weight of MSP1a which varies in size among isolates due to different numbers of tandemly repeated 28-29 amino acid peptides. For these studies, genes coding for A. marginale MSP1a and MSP4, msp1alpha and msp4, respectively, from nine North American isolates were sequenced for phylogenetic analysis. The phylogenetic analysis strongly supports the existence of a south-eastern clade of A. marginale comprised of Virginia and Florida isolates. Analysis of 16S rDNA fragment sequences from the A. marginale tick vector, Dermacentor variabilis, from various areas of the United States was used to evaluate possible vector-parasite co-evolution. Our phylogenetic analysis supports identity between the most parsimonious tree from the A. marginale MSP gene data and the tree that reflected the western and eastern clades of D. variabilis. These phylogenetic analyses provide information that may be important to consider when developing control strategies for anaplasmosis in the United States.

Phylogeography of New World isolates of Anaplasma marginale based on major surface protein sequences

Gene and protein sequences of major surface proteins (MSP) 1a and 4 of Anaplasma marginale (Rickettsiales: Anaplasmataceae) were used to infer phylogenetic relationships between New World isolates from Argentina, Brazil, Mexico and the United States. Seventeen isolates of A. marginale plus two outgroup taxa (A. centrale and A. ovis) were used for maximum-parsimony analysis of MSP4, while 20 isolates were used for phylogenetic analysis of MSP1a. msp4 analysis provided strong bootstrap support for a Latin American clade and, within this clade, support was detected for Mexican and South American clades. Isolates of A. marginale from the United States also grouped into two clades from the southern (isolates from Florida, Mississippi, and Virginia) and west-central (isolates from California, Idaho, Illinois, Oklahoma, and Texas) states. Although little phylogeographic resolution was detected within these higher clades, msp4 sequences appear to be a good genetic marker for inferring phylogeographic patterns of A. marginale isolates. In contrast to the phylogeographic resolution provided by msp4, MSP1a DNA and protein sequence were quite variable and did not provide phylogeographic resolution. Most variation in MSP1a sequences appeared unique to a given isolate and similar DNA sequence variation in msp1alpha was detected within isolates from Idaho and Florida and from Idaho and Argentina. The results of these studies demonstrated that msp4 provided phylogenetic information on the evolution of A. marginale isolates. In contrast MSP1a sequences appeared to be rapidly evolving and these sequences may provide phylogeographic information only when numerous isolate MSP1a sequences are analyzed from a geographic area.

Phylogenetic Relationships among Recent Chiropteran Families and the Importance of Choosing Appropriate Out-Group Taxa

Abstract Results of recent molecular studies cast doubt on the validity of the superorder Archonta, suborders Megachiroptera and Microchiroptera, and infraorder Yinochiroptera and has even led some to consider novel alternatives for the evolution of flight and echolocation in mammals. At present, higher-level relationships within Chiroptera still is without consensus, and much of this controversy is related to how bats are related to other mammals and also to relationships among family-level lineages within Chiroptera. Although this controversy superficially manifests itself as differences in the relative merits of morphologic versus molecular data, both classes of data are themselves conflicting. We contend that much of the discrepancy among these studies is due to improper choice of out-group, limited taxonomic sampling, or both. We examined approximately 3 kb of mitochondrial DNA from 104 bats representing the taxonomic, geographic, and morphologic diversity within all families (except the monotypic Craseonycteridae) and 58 additional taxa representing 12 other orders of mammals. Results of our analyses strongly support other recent work indicating that Archonta is not a natural assemblage and that the sister taxon to Chiroptera may include Cetartiodactyla, Perissodactyla, Carnivora, and possibly Pholidota. Using representatives of these taxa as out-groups to evaluate interfamilial relationships within Chiroptera, we detected strong support for recognition of the suborders Yinpterochiroptera and Yangochiroptera. Within Yangochiroptera, our analyses strongly support expansion of the superfamily Noctilionoidea to include the New World Thyropteridae and Furipteridae.

Phylogeny and evolution of the Piroplasmida as inferred from 18S rRNA sequences

The order Piroplasmida consists of several genera of tick-borne parasites that infect mammals, and to a lesser extent birds, and are therefore of medical and economic importance. Despite their importance, considerable confusion exists concerning the relationship among piroplasmid species, specifically concerning the number of genera and the intergeneric relationships. To examine evolutionary relationships among piroplasmids, we conducted phylogenetic analyses of 192 18S rDNA sequences from the genera Theileria, Babesia and Cytauxzoon. Our analyses revealed eight clades potentially representing distinct genera, and we distinguish the Duncani Group and Microti Group as genetically distinct groups of species requiring detailed analysis of morphology and life-history to allow formal generic description. The piroplasmid phylogeny revealed considerable host diversity and limited host specificity, suggesting piroplasmids have undergone frequent host switches during their evolution. Our analyses provide the first reported evolutionary timescale for piroplasmids independent of the assumption of parasite-host cospeciation, which is invalid for piroplasmids. Evolutionary rate analyses revealed considerable substitution rate heterogeneity, which we attribute to host switching and diversification. Finally, we call for a comprehensive phylogenetic, morphological and life-history analysis for these medically relevant taxa to resolve relationships and understand host specificity.

Anaplasma marginale msp1α Genotypes Evolved under Positive Selection Pressure but Are Not Markers for Geographic Isolates

ABSTRACT Anaplasma marginale (order Rickettsiales, family Anaplasmataceae), a tick-borne pathogen of cattle, is endemic in tropical and subtropical regions of the world. Many geographic isolates of A. marginale occur in the United States and have been identified by major surface protein 1a (MSP1a), which varies in sequence and molecular weight due to different numbers of tandem 28- to 29-amino-acid repeats. The present study was undertaken to examine the genetic variations among isolates of A. marginale obtained during 2001 from infected cattle from east-central Oklahoma, where A. marginale is endemic. The gene and protein sequences of MSP1a and msp4 nucleotide sequences were used to infer the phylogenetic relationships among Oklahoma and New World isolates from Argentina, Brazil, Mexico, and the United States. All 11 A. marginale isolates collected from Oklahoma had different MSP1a sequences but identical MSP4 sequences. The phylogenies of the msp4 sequences of 13 isolates from Oklahoma in comparison with those of 7 Latin American isolates and 12 U.S. isolates by maximum-parsimony (MP) and maximum-likelihood (ML) analyses, with A. centrale and A. ovis sequences used as outgroups, provided strong bootstrap analysis support for a Latin American clade. Isolates of A. marginale from the southern United States (Florida, Mississippi, and Virginia) and the west-central United States (California, Idaho, Illinois, Oregon, Missouri, and Texas) also grouped into two clades. Both clades contained isolates from Oklahoma, suggesting extensive cattle movement. ML analysis of the msp4 sequences of isolates from Oklahoma provided bootstrap analysis support for east-central and north-central clades in Oklahoma, and both clades included isolates from Stillwater, Okla. Analysis of the codon and amino acid changes among the msp4 sequences of isolates with different phylogenies provided evidence that msp4 is not under positive selection pressure. In contrast, the phylogenies of the MSP1a DNA and protein sequences of 13 isolates from Oklahoma in comparison with those of 7 Latin American and 13 isolates from the United States by MP and ML analyses demonstrated no geographic clustering and provided evidence that this gene is under positive selection pressure. The results indicate that msp1α is not a marker for the characterization of A. marginale geographic isolates and suggest that the genetic heterogeneity observed among isolates of A. marginale within Oklahoma could be explained by cattle movement and the maintenance of different genotypes by independent transmission events.

Molecular phylogenetics of Myotis indicate familial-level divergence for the genus Cistugo (Chiroptera)

Abstract The genus Myotis has undergone significant taxonomic revision since the advent of DNA sequencing techniques. Prior morphological examination of Myotis has indicated as many as 4 subgenera correlated with foraging strategies. Recent studies using mitochondrial DNA (mtDNA) sequence data have questioned the validity of these subgenera and have indicated that several taxa may require reevaluation as to their position within Vespertilionidae. Nevertheless, no study has used large-scale nuclear DNA sequencing to examine relationships within Myotis. We generated 4,656 base pairs (bp) of nuclear intron (PRKC1, STAT5A, and THY) and exon (APOB, DMP1, and RAG2) sequence data in addition to 2,866 bp of mtDNA sequence data to test previously hypothesized subgeneric groupings of Myotis. We included 21 species of Myotis from all morphological subgenera previously suggested, representatives of all subfamilies and tribes currently recognized in Vespertilionidae, and multiple representatives of all other families currently included in the superfamily Vespertilionoidea. We also included a representative of the rare African genus Cistugo, because significant doubt exists about its familial position. Our phylogenetic analyses did not support the morphologically defined Myotis subgenera and confirm that morphological similarities among Myotis are the result of convergent evolution. Divergence estimates derived from the total data set were concordant with previous studies, suggesting a middle Miocene trans-Beringian dispersal from Asia colonized North America, with subsequent South American colonization and diversification prior to the formation of the Isthmus of Panama 3–4 million years ago. Myotis latirostris fell outside of Myotis, and the high genetic distance separating it from other Myotis suggested that M. latirostris represented a distinct genus. The genus Cistugo, previously a subgenus within Myotis, fell basal to all vespertilionids, with a high genetic distance separating it from Vespertilionidae. We conclude that Cistugo should constitute a distinct family within Vespertilionoidea.

Present and potential future distribution of common vampire bats in the Americas and the associated risk to cattle.

Success of the cattle industry in Latin America is impeded by the common vampire bat, Desmodus rotundus, through decreases in milk production and mass gain and increased risk of secondary infection and rabies. We used ecological niche modeling to predict the current potential distribution of D. rotundus and the future distribution of the species for the years 2030, 2050, and 2080 based on the A2, A1B, and B1 climate scenarios from the Intergovernmental Panel on Climate Change. We then combined the present day potential distribution with cattle density estimates to identify areas where cattle are at higher risk for the negative impacts due to D. rotundus. We evaluated our risk prediction by plotting 17 documented outbreaks of cattle rabies. Our results indicated highly suitable habitat for D. rotundus occurs throughout most of Mexico and Central America as well as portions of Venezuela, Guyana, the Brazilian highlands, western Ecuador, northern Argentina, and east of the Andes in Peru, Bolivia, and Paraguay. With future climate projections suitable habitat for D. rotundus is predicted in these same areas and additional areas in French Guyana, Suriname, Venezuela and Columbia; however D. rotundus are not likely to expand into the U.S. because of inadequate ‘temperature seasonality.’ Areas with large portions of cattle at risk include Mexico, Central America, Paraguay, and Brazil. Twelve of 17 documented cattle rabies outbreaks were represented in regions predicted at risk. Our present day and future predictions can help authorities focus rabies prevention efforts and inform cattle ranchers which areas are at an increased risk of cattle rabies because it has suitable habitat for D. rotundus.