Parasites can be used as unique markers to investigate host evolutionary history, independent of host data. Here we show that modern human head lice, Pediculus humanus, are composed of two ancient lineages, whose origin predates modern Homo sapiens by an order of magnitude (ca. 1.18 million years). One of the two louse lineages has a worldwide distribution and appears to have undergone a population bottleneck ca. 100,000 years ago along with its modern H. sapiens host. Phylogenetic and population genetic data suggest that the other lineage, found only in the New World, has remained isolated from the worldwide lineage for the last 1.18 million years. The ancient divergence between these two lice is contemporaneous with splits among early species of Homo, and cospeciation analyses suggest that the two louse lineages codiverged with a now extinct species of Homo and the lineage leading to modern H. sapiens. If these lice indeed codiverged with their hosts ca. 1.18 million years ago, then a recent host switch from an archaic species of Homo to modern H. sapiens is required to explain the occurrence of both lineages on modern H. sapiens. Such a host switch would require direct physical contact between modern and archaic forms of Homo.

Birds are plagued by an impressive diversity of ectoparasites, ranging from feather-feeding lice, to featherdegrading bacteria. Many of these ectoparasites have severe negative effects on host fitness. It is therefore not surprising that selection on birds has favored a variety of possible adaptations for dealing with ectoparasites. The functional significance of some of these defenses has been well documented. Others have barely been studied, much less tested rigorously. In this article we review the evidence -or lack thereof -for many of the purported mechanisms birds have for dealing with ectoparasites. We concentrate on features of the plumage and its components, as well as anti-parasite behaviors. In some cases, we present original data from our own recent work. We make recommendations for future studies that could improve our understanding of this poorly known aspect of avian biology.

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Introduced parasites threaten native host species that lack effective defenses. Such parasites increase the risk of extinction, particularly in small host populations like those on islands. If some host species are tolerant to introduced parasites, this could amplify the risk of the parasite to vulnerable host species. Recently, the introduced parasitic nest fly Philornis downsi has been implicated in the decline of Darwin's finch populations in the Galápagos Islands. In some years, 100% of finch nests fail due to P. downsi; however, other common host species nesting near Darwin's finches, such as the endemic Galápagos mockingbird (Mimus parvulus), appear to be less affected by P. downsi. We compared effects of P. downsi on mockingbirds and medium ground finches (Geospiza fortis) on Santa Cruz Island in the Galápagos. We experimentally manipulated the abundance of P. downsi in nests of mockingbirds and finches to measure the direct effect of the parasite on the reproductive success of each species of host. We also compared immunological and behavioral responses by each species of host to the fly. Although nests of the two host species had similar parasite densities, flies decreased the fitness of finches but not mockingbirds. Neither host species had a significant antibody-mediated immune response to P. downsi. Moreover, finches showed no significant increase in begging, parental provisioning, or plasma glucose levels in response to the flies. In contrast, parasitized mockingbird nestlings begged more than nonparasitized mockingbird nestlings. Greater begging was correlated with increased parental provisioning behavior, which appeared to compensate for parasite damage. The results of our study suggest that finches are negatively affected by P. downsi because they do not have such behavioral mechanisms for energy compensation. In contrast, mockingbirds are capable of compensation, making them tolerant hosts, and a possible indirect threat to Darwin's finches.

Some species of parasites occur on a wide range of hosts while others are restricted to one or a few host species. The host specificity of a parasite species is determined, in part, by its ability to disperse between host species. Dispersal limitations can be studied by exploring the genetic structure of parasite populations both within a single species of host and across multiple host species. In this study we examined the genetic structure in the mitochondrial cytochrome oxidase I (COI) gene of two genera of lice (Insecta: Phthiraptera) occurring on multiple sympatric species of doves in southern North and Central America. One genus, Columbicola, is generally less host-specific than the other, Physconelloides. For both genera we identified substantial genetic differentiation between populations of conspecific lice on different host species, generally 10-20% sequence divergence. This level of divergence is in the range of that often observed between species of these two genera. We used nested clade analysis to explore fine scale genetic structure within species of these feather lice. We found that species of Physconelloides exhibited more genetic structure, both among hosts and among geographical localities, than did species of Columbicola. In many cases, single haplotypes within species of Columbicola are distributed on multiple host species. Thus, the population genetic structure of species of Physconelloides reveals evidence of geographical differentiation on top of high host species specificity. Underlying differences in dispersal biology probably explain the differences in population genetic structure that we observed between Columbicola and Physconelloides.

Phylogenetic relationships among genera of pigeons and doves (Aves, Columbiformes) have not been fully resolved because of limited sampling of taxa and characters in previous studies. We therefore sequenced multiple nuclear and mitochondrial DNA genes totaling over 9000 bp from 33 of 41 genera plus 8 outgroup taxa, and, together with sequences from 5 other pigeon genera retrieved from GenBank, recovered a strong phylogenetic hypothesis for the Columbiformes. Three major clades were recovered with the combined data set, comprising the basally branching New World pigeons and allies (clade A) that are sister to Neotropical ground doves (clade B), and the Afro-Eurasian and Australasian taxa (clade C). None of these clades supports the monophyly of current families and subfamilies. The extinct, flightless dodo and solitaires (Raphidae) were embedded within pigeons and doves (Columbidae) in clade C, and monophyly of the subfamily Columbinae was refuted because the remaining subfamilies were nested within it. Divergence times estimated using a Bayesian framework suggest that Columbiformes diverged from outgroups such as Apodiformes and Caprimulgiformes in the Cretaceous before the mass extinction that marks the end of this period. Bayesian and maximum likelihood inferences of ancestral areas, accounting for phylogenetic uncertainty and divergence times, respectively, favor an ancient origin of Columbiformes in the Neotropical portion of what was then Gondwana. The radiation of modern genera of Columbiformes started in the Early Eocene to the Middle Miocene, as previously estimated for other avian groups such as ratites, tinamous, galliform birds, penguins, shorebirds, parrots, passerine birds, and toucans. Multiple dispersals of more derived Columbiformes between Australasian and Afro-Eurasian regions are required to explain current distributions.

Synopsis. A recent model by Hamilton and Zuk (1982) suggests that exaggerated sec? ondary sexual traits facilitate mate choice for genetic resistance to parasites. The model predicts that individuals discriminate against parasitized mates by scrutinizing traits indicative of parasite load. In the case of birds and their feather-feeding lice, for example, individuals might avoid parasitized mates by detecting reduced plumage brightness, reduced courtship display, or increased grooming. I conducted a series of mate choice trials in which female Rock Doves (Columba livia) were allowed to choose between "clean" males without lice and "lousy" males with experimentally increased loads. Clean males displayed significantly more often than lousy males and females demonstrated a significant pref? erence for clean males. Lousy males were subject to plumage damage; however, none of the damage was externally visible, and the time spent grooming by clean and lousy males did not differ significantly. Female louse loads, which were also manipulated, were not significantly related to female mating preferences. These results are consistent with the Hamilton-Zuk model. They are also consistent with a model of sexual selection for the avoidance of parasite transmission, which is discussed. The general relevance of lice and other ectoparasites to models of parasite-mediated sexual selection is reviewed.

A parasite's potential effect, or ''pressure'', can influence the life history strategy of its host. In environments with high parasite pressure, hosts invest more in anti-parasite defense, which may limit their investment in other life history components, such as survival. This tradeoff is difficult to study in natural populations because pressure is hard to quantify. Pressure is not necessarily correlated with the abundance of the parasite. A host population can be under high pressure, yet have few parasites, because members of the population have invested heavily in defense. Therefore, the extent to which parasite pressure varies among host populations, and the cause of such variation, remain largely undocumented. In this paper we show that birds in arid regions have fewer ectoparasitic lice than birds in humid regions. We show experimentally that low humidity reduces the number of lice on birds, even when host defense is held constant. Comparisons of ambient humidity to humidity beneath the plumage demonstrate that plumage does not provide a buffer for lice against low humidity. Our results confirm that an abiotic factor can cause substantial variation in parasite pressure among host populations. We suggest that humidity may influence host life history evolution through its impact on ectoparasites.