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D H Clayton

Publications and source records attributed to D H Clayton.

18 recordsLinked to original sources

Phylogenetic analysis of partial sequences of elongation factor 1alpha identifies major groups of lice (Insecta: Phthiraptera).

As a first attempt to use molecular data to resolve the relationships between the four suborders of lice and within the suborder Ischnocera, we sequenced a 347-bp fragment of the elongation factor 1alpha gene of 127 lice (Insecta: Phthiraptera) as well as outgroup taxa from the order Psocoptera. A number of well-supported monophyletic groups were found but the relationships among many of these groups could not be resolved. While it is probable that multiple substitutions at high divergences and ancient radiation over a short period of time have contributed to the problem, we attribute most of this lack of resolution to the high ratio of taxa to characters. Nevertheless, the sequence data unequivocally support a number of important relationships that are at variance with the conclusions of morphological taxonomy. These include the sister group relationship of Chelopistes and Oxylipeurus, two lice occupying different ecological niches on the same host, which have previously been assigned to different families. These results provide evidence in support of the hypothesis that lice have speciated in situ on the host in response to niche specialization and that this has given rise to convergent morphologies in the lice of different host groups which share similar ecological niches. We discuss our attempts to overcome the limitations of this large data set, including the use of leaf stability analysis, a new method for analyzing the stability of taxa in a phylogenetic tree, and examine a number of hypotheses of relationships based on both traditional taxonomy and host associations.

Animals↗

Molecular systematics of Goniodidae (Insecta: Phthiraptera).

The higher level phylogenetic relationships within the avian feather lice (Insecta: Phthiraptera: Ischnocera) are extremely problematic. Here we investigate the relationships of 1 family (Goniodidae), sometimes recognized as distinct within Ischnocera, using parsimony and likelihood analyses of nuclear and mitochondrial DNA sequences. These data support monophyly for a restricted definition of traditional Goniodidae, but recognition of this family would result in paraphyly of the large heterogeneous family Philopteridae. We show that the New World Chelopistes is not related to other members of Goniodidae, despite similarities in morphology, but rather is the sister taxon to Oxylipeurus. Within Goniodidae, genera are divided into those occurring on Galliformes (the Goniodes complex) and those occurring on Columbiformes (the Coloceras complex). Within the well-sampled Coloceras complex, or Physconelloidinae, several groups are identified. However, traditionally recognized genera such as Coloceras and Phvsconelloides appear to be paraphyletic. Whereas the phylogeny of Goniodidae reflects some aspects of host relationships, biogeography also influences coevolutionary history.

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Critical evaluation of five methods for quantifying chewing lice (Insecta: Phthiraptera).

Five methods for estimating the abundance of chewing lice (Insecta: Phthiraptera) were tested. To evaluate the methods, feral pigeons (Columba livia) and 2 species of ischnoceran lice were used. The fraction of lice removed by each method was compared, and least squares linear regression was used to determine how well each method predicted total abundance. Total abundance was assessed in most cases using KOH dissolution. The 2 methods involving dead birds (body washing and post-mortem-ruffling) provided better results than 3 methods involving live birds (dust-ruffling, fumigation chambers, and visual examination). Body washing removed the largest fraction of lice (>82%) and was an extremely accurate predictor of total abundance (r2 = 0.99). Post-mortem-ruffling was also an accurate predictor of total abundance (r2 > or = 0.88), even though it removed a smaller proportion of lice (<70%) than body washing. Dust-ruffling and fumigation chambers removed even fewer lice, but were still reasonably accurate predictors of total abundance, except in the case of data sets restricted to birds with relatively few lice. Visual examination, the only method not requiring that lice be removed from the host, was an accurate predictor of louse abundance, except in the case of wing lice on lightly parasitized birds.

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Nuclear and mitochondrial genes contain similar phylogenetic signal for pigeons and doves (Aves: Columbiformes).

Molecular systematic studies generally assume that gene trees are reasonable estimates of species trees. We tested the validity of this assumption in the pigeons and doves (Aves: Columbiformes) by comparing phylogenies derived from nuclear (beta-fibrinogen intron 7) and mitochondrial (cytochrome b) genes. Trees derived from the two genes when analyzed separately contained many nodes in common. A partition homogeneity test revealed no significant incongruence between trees derived from the two genes; so, we combined nuclear and mitochondrial data in subsequent phylogenetic analyses. The resulting tree, which was highly resolved and generally well supported, contained a strong biogeographic component. The rate of nucleotide substitution for the nuclear intron was approximately six times slower than that of cytochrome b. This resulted in a much higher consistency index for trees derived from the intron because of the low level of multiple substitution. However, the degree of resolution and support for trees reconstructed from the two genes was similar. We also examined the transition and transversion substitution rates for the genes. Third position transversions for cytochrome b accumulated linearly with intron divergence, suggesting low levels of multiple substitution for third position transversions.

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Taxonomic review of Physconelloides (Phthiraptera: Philopteridae) from the Columbiformes (aves), including descriptions of three new species.

We provide a comprehensive taxonomic review of Physconelloides, a genus of ischnoceran chewing lice found on pigeons and doves (Columbiformes). Thirteen previously known Physconelloides species are redescribed and 16 new synonymies are designated: P. rubripes Carriker, P. rubripes longulus Tendeiro, P. piotrowskii Tendeiro and P. auritae Tendeiro are synonyms of P. zenaidurae (McGregor); P. recurvatus Eichler, P. chocoensis Carriker and P. montana Carriker are synonyms of P. ceratoceps Ewing; P. silvestris Tendeiro is a synonym of P. perijae Carriker; P. keleri Kaddou and P. branderi Kaddou are synonyms of P. spenceri Emerson and Ward; P. wolfdietrichi Kaddou is a synonym of P. anolaimae Carriker; and Goniocotacanthus mattogrossensis Guimaraes, P. passerinae Emerson, P. eurysema pretiosa Carriker, P. talpacoti Carriker and P. picuii Tendeiro are synonyms of P. eurysema (Carriker). Three new species are also described: P. moyeri (type host Geotrygon linearis), P. johnsoni (type host Columbina passerina bahamensis), and P. robbinsi (type host Metriopelia ceciliae). A key is provided for identification of the 16 recognized species.

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A different tempo of mitochondrial DNA evolution in birds and their parasitic lice.

A phylogeny for the lice (Insecta: Phthiraptera: genus Dennyus) parasitic on swiftlets (Aves: Collocalliinae) was constructed based on mitochondrial cytochrome b DNA sequences. This phylogeny is congruent with previous phenetic analyses of morphometric data for the lice. Comparison with a previously obtained phylogeny for the hosts indicates some degree of cospeciation. These cospeciation events are used to compare relative rates of evolution in the birds and their lice for the same segment of the cytochrome b gene. Cytochrome b is evolving two to three times more rapidly in lice than in birds, and louse cytochrome b is highly divergent compared to that of most other insects. Although generation time has been suggested as an explanation for the disparity in evolutionary rates between lice and their hosts, we suggest that the small effective population sizes of lice coupled with founder events occurring during transmission to new host individuals may be an important factor.

Amino Acid Sequence↗

Taxonomic review of Cotingacola (Phthiraptera: Philopteridae) from the cotingas (Passeriformes: Tyrannidae), with descriptions of two new species.

The genus Cotingacola currently contains 14 species and subspecies described by Carriker. A careful examination of these reveals only 6 valid taxa with 8 names placed as junior synonyms. These new synonymies are Cotingacola rupicolae colombiana, C. latigastra, C. temporalis and C. longicrucis, synonyms of C. rupicolae; C. foramina, C. graciligastra and C. tityra, synonyms of C. tergalis; C. acuticeps, a synonym of C. dimorpha. Two new species are described from Peruvian material: C. stotzi (type host: Querula purpurata) and C. fitzpatricki (type host: Lipaugus subalaris). A key is provided for identification of the 8 species.

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Does behavior reflect phylogeny in swiftlets (Aves: Apodidae)? A test using cytochrome b mitochondrial DNA sequences.

Swiftlets are small insectivorous birds, many of which nest in caves and are known to echolocate. Due to a lack of distinguishing morphological characters, the taxonomy of swiftlets is primarily based on the presence or absence of echolocating ability, together with nest characters. To test the reliability of these behavioral characters, we constructed an independent phylogeny using cytochrome b mitochondrial DNA sequences from swiftlets and their relatives. This phylogeny is broadly consistent with the higher classification of swifts but does not support the monophyly of swiftlets. Echolocating swiftlets (Aerodramus) and the nonecholocating "giant swiftlet" (Hydrochous gigas) group together, but the remaining nonecholocating swiftlets belonging to Collocalia are not sister taxa to these swiftlets. While echolocation may be a synapomorphy of Aerodramus (perhaps secondarily lost in Hydrochous), no character of Aerodramus nests showed a statistically significant fit to the molecular phylogeny, indicating that nest characters are not phylogenetically reliable in this group.

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Lice and cospeciation: a response to Barker.

"The student who intends working on the Mallophaga should take warning that he will be tried almost beyond endurance by the paradoxes and complexities which beset his subject but he will also find, in the dual and inter-related aspect of insect and bird, an infinite fascination." (Rothschild & Clay, 1952: pp. 156-157). The study of host louse coevolution will benefit greatly from the phylogenetic perspective offered by recent advantages in molecular systematics. However, in order to make best use of phylogenies we need to appreciate the complexities of the possible relations between host and parasite phylogeny. At the same time, the very complexity of louse-host systems has a potentially useful consequence; the presence of multiple lineages of lice on the same hosts allow for replicated tests of coevolutionary hypotheses. For example, if a number of louse clades infest the same host clade but some lice show more cospeciation than others, we might ask whether there are features of louse biology that correlate with this difference in host tracking fidelity. It may further be possible to ascertain the relative importance of these features in ecological time through controlled transfer experiments. By beginning to appreciate "the paradoxes and complexities" of host-louse evolution, lice may offer us not only "infinite fascination" but also a chance to address important questions in coevolution.

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Sampling effort and parasite species richness.

Comparative studies of parasite species richness among host taxa can be confounded by uneven sampling effort. Sampling ceases to be a confounding factor when extrapolation methods are used to estimate true species richness. Here, Bruno Walther and colleagues review examples of sampling bias and the use of extrapolation methods for circumventing it. They also discuss the confounding effects of phylogenetic association of estimates of species richness.

Journal Article↗

Comparative effects of mites and lice on the reproductive success of rock doves (Columba livia).

We report experimental data comparing the effects of Mesostigmatid mites and Ischnoceran lice on the reproductive performance of a single group of captive rock doves (Columba livia). Several components of host reproductive success were compared for the two groups, including number of eggs laid, hatching success, nestling growth rates, fledging success, post-fledging body mass and survival. Adult body mass and survival were also compared. There was a dramatic difference in the effects of the mites and lice. The former drove host reproductive success to zero, mainly by agitating adults and causing them to incubate eggs less faithfully. Nestling growth rates and post-fledging survival were also significantly reduced by mites. Lice showed no effect on reproductive success whatsoever, even though the feather damage they cause is known to have energetic consequences (Booth, Clayton & Block, 1993). Neither parasite had a significant effect on adult birds. Although Ischnocera are found on most species of birds, our results for lice constitute the first experimental test of the impact of Ischnocera on avian reproductive success (preliminary report by Clayton & Tompkins, 1994). We discuss reasons for the different effects of mites and lice, including the relationship of horizontal (mites) and vertical (lice) transmission to the evolution of virulence.

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Ectoparasite virulence is linked to mode of transmission.

Theory suggests that the evolution of parasite virulence is linked to the dynamics of parasite transmission. All else being equal, parasites transmitted vertically from parents to offspring should be less virulent than parasites capable of horizontal transmission to unrelated hosts. This is because the fitness of vertically transmitted parasites is tightly linked to the reproductive success of the host, whereas the fitness of horizontally transmitted parasites is relatively independent of host reproduction. The virulence-transmission relation has seldom been tested because of difficulties inherent in comparing virulences of different parasite-host systems. We compared the virulence of lice and mites infesting a single group of captive rock doves (Columba livia). Lice, which were vertically transmitted, had no detectable effect on host fitness, whereas horizontally transmitted mites drove host reproductive success to zero. These results, in conjunction with a survey of the literature, support the hypothesis that ectoparasite virulence is linked to the mode of transmission.

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Review of the species of Rallicola (Phthiraptera: Philopteridae) from the antbirds, ovenbirds, and tapaculos (Passeriformes).

Fourteen species of chewing lice in the genus Rallicola are recognized and described from members of the passerine families Thamnophilidae, Furnariidae (subfamily Furnariinae), and Rhinocryptidae. These include 10 previously described species and four new ones: R. mooreae from Premnoplex brunnescens (Sclater), R. leeae from Thripadectes melanorhynchus (Tschudi), R. tompkinsi from Sclerurus caudacutus (Vieillot), and R. hambleri from Syndactyla rufosuperciliata (Lafresnaye). New synonymies include R. acutifrons subsimilis (Carriker) and R. a. chocoana (Carriker), which are both junior synonyms of R. acutifrons (Carriker), and R. mirandae (Carriker), R. punensis (Carriker), R. myrmeciza (Carriker), and R. hylactiphaga canae (Carriker), which are, respectively, junior synonyms of R. laticephala (Carriker), R. titicacae (Carriker), R. heterocephala (Carriker), and R. hylactiphaga (Carriker). A key is provided for identification of the species treated herein.

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Review of the species of Rallicola (Phthiraptera: Philopteridae) from the woodcreepers (Passeriformes: Dendrocolaptinae).

Sixteen species of chewing lice in the genus Rallicola are recognized and described from members of the passerine subfamily Dendrocolaptinae. These include 12 previously described species and 4 new ones: R. keymerae from Dendrocolaptes picumnus, R. lyali from Xiphocolaptes major, R. harveyi from Deconychura longicauda, and R. palmai from Campylorhamphus trochilirostris. New synonymies include R. guttata and R. pyriglena, both of which are junior synonyms of R. chunchotambo, and R. certhia microgenitalia, which is a junior synonym of R. colombiana. A key is provided for identification of the species treated herein.

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The influence of parasites on host sexual selection.

In 1982 Hamilton and Zuk(1) proposed a provocative solution for the unexplained fact that the males of many species exhibit 'showy' traits such as brightly coloured plumage or vigorous courtship displays. They suggested that showy traits are fully expressed only by males who are resistant to parasites and that females examine such traits in order to choose resistant males as mates. Hamilton and Zuk's proposal has been the topic of extensive research and vigorous debate for nearly a decade. This article reviews the research, relevant criticisms and unanswered questions pertaining to the influence of parasites on sexual selection.

Journal Article↗

Host specificity of Strigiphilus owl lice (Ischnocera: Philopteridae), with the description of new species and host associations.

Three new species and 11 new host associations are described for the cursitans species group of the genus Strigiphilus, the only genus of Mallophaga confined to owls (Strigiformes). The new species are S. schemskei ex Bubo coromandus, S. garylarsoni ex Otus leucotis, and S. petersoni ex Ninox punctulata. This paper completes a taxonomic review of the cursitans species group and includes an updated host-parasite list. The host specificity of the group is reviewed, including features of the natural history of lice and their hosts that may have influenced the evolution of host-parasite associations. Although most members of the cursitans group are host specific, several cases of secondary transfer among ecologically related hosts are apparent. The significance of such transfers is discussed in light of attempts to employ data on host-parasite associations for host systematics.

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