Jason D. Weckstein

Biogeography Explains Cophylogenetic Patterns in Toucan Chewing Lice

Historically, comparisons of host and parasite phylogenies have concentrated on cospeciation. However, many of these comparisons have demonstrated that the phylogenies of hosts and parasites are seldom completely congruent, suggesting that phenomena other than cospeciation play an important role in the evolution of host-parasite assemblages. Other coevolutionary phenomena, such as host switching, parasite duplication (speciation on the host), sorting (extinction), and failure to speciate can also influence host-parasite assemblages. Using mitochondrial and nuclear protein-coding DNA sequences, I reconstructed the phylogeny of ectoparasitic toucan chewing lice in the Austrophilopterus cancellosus subspecies complex and compared this phylogeny with the phylogeny of the hosts, the Ramphastos toucans, to reconstruct the history of coevolutionary events in this host-parasite assemblage. Three salient findings emerged. First, reconstructions of host and louse phylogenies indicate that they do not branch in parallel, and their cophylogenetic history shows little or no significant cospeciation. Second, members of monophyletic Austrophilopterus toucan louse lineages are not necessarily restricted to monophyletic host lineages. Often, closely related lice are found on more distantly related but sympatric toucan hosts. Third, the geographic distribution of the hosts apparently plays a role in the speciation of these lice. These results suggest that for some louse lineages biogeography may be more important than host associations in structuring louse populations and species, particularly when host life history (e.g., hole nesting) or parasite life history (e.g., phoresis) might promote frequent host switching events between syntopic host species. These findings highlight the importance of integrating biogeographic information into cophylogenetic studies.

Temporal and spatial diversification of Pteroglossus araçaris (AVES: Ramphastidae) in the neotropics: Constant rate of diversification does not support an increase in radiation during the Pleistocene

We use the small-bodied toucan genus Pteroglossus to test hypotheses about diversification in the lowland Neotropics. We sequenced three mitochondrial genes and one nuclear intron from all Pteroglossus species and used these data to reconstruct phylogenetic trees based on maximum parsimony, maximum likelihood, and Bayesian analyses. These phylogenetic trees were used to make inferences regarding both the pattern and timing of diversification for the group. We used the uplift of the Talamanca highlands of Costa Rica and western Panama as a geologic calibration for estimating divergence times on the Pteroglossus tree and compared these results with a standard molecular clock calibration. Then, we used likelihood methods to model the rate of diversification. Based on our analyses, the onset of the Pteroglossus radiation predates the Pleistocene, which has been predicted to have played a pivotal role in diversification in the Amazon rainforest biota. We found a constant rate of diversification in Pteroglossus evolutionary history, and thus no support that events during the Pleistocene caused an increase in diversification. We compare our data to other avian phylogenies to better understand major biogeographic events in the Neotropics. These comparisons support recurring forest connections between the Amazonian and Atlantic forests, and the splitting of cis/trans Andean species after the final uplift of the Andes. At the subspecies level, there is evidence for reciprocal monophyly and groups are often separated by major rivers, demonstrating the important role of rivers in causing or maintaining divergence. Because some of the results presented here conflict with current taxonomy of Pteroglossus, new taxonomic arrangements are suggested.

The perils of using host relationships in parasite taxonomy: phylogeny of the Degeeriella complex

The taxonomy of lice (Insecta: Phthiraptera) is often heavily influenced by host taxonomy. The use of host information to define genera of avian lice in the widespread Degeeriella complex has been prevalent but has created problems. Several workers have suggested that genera defined on the basis of host association are not monophyletic. We used sequences of nuclear (elongation factor-1alpha) and mitochondrial (cytochrome oxidase I) genes to test the monophyly of several genera in the Degeeriella complex. Parsimony and likelihood analyses of these data indicated that many genera in the Degeeriella complex are not monophyletic, such that species occurring on the same host groups do not form monophyletic groups. Biological features of hosts (including predaceous habits, brood parasitism, and hole nesting) for species in the Degeeriella complex likely provide opportunities for switching of lice between host groups. In addition, dispersal of lice via phoresy on hippoboscid flies also likely provides opportunities for host switching in the Degeeriella complex. This study indicates that the overuse of host taxonomy in louse taxonomy can result in classifications that do not reflect phylogenetic history.

MOLECULAR PHYLOGENETICS OF THE RAMPHASTOS TOUCANS: IMPLICATIONS FOR THE EVOLUTION OF MORPHOLOGY, VOCALIZATIONS, AND COLORATION

Abstract I reconstructed the phylogeny of 12 Ramphastos toucan taxa using mitochondrial DNA (mtDNA) sequences. This analysis identified two major groups, including a monophyletic smooth-billed yelping clade and a clade including most, but not all, the channel-keel-billed croakers. Within the R. tucanus and R. vitellinus groups, uncorrected mtDNA divergences are relatively low and mtDNA sequences from several subspecies are paraphyletic. One exception to low divergences within the R. vitellinus group is R. v. ariel from southeastern Brazil, which on average differs from all other R. vitellinus sampled by 2.9%. Character reconstructions on the phylogeny indicate that the ancestral Ramphastos was most likely a large-bodied channel- keel-billed croaker. Furthermore, an assessment of the patterns of bill shape, voice, and both plumage and bare-part coloration characters suggests that bill shape and voice have significant phylogenetic signal but that color characters do not. Sympatric Ramphastos taxa are not closely related in the phylogeny; therefore, character reconstructions indicate that the extreme similarity in coloration patterns between many sympatric Ramphastos pairs is most likely attributable to a combination of convergence or parallelism (homoplasy) and shared ancestral character states (symplesiomorphy). Filogenética Molecular de los Tucanes del Género Ramphastos: Implicaciones para la Evolución de la Morfología, las Vocalizaciones y la Coloración

Comparative phylogenetic histories of two louse genera found on Catharus thrushes and other birds.

Abstract The louse genera Brueelia (Ischnocera) and Myrsidea (Amblycera) are broadly codistributed on songbirds (Passeriformes), but differ in a variety of life history characteristics. We used mitochondrial and nuclear DNA sequences to assess levels of genetic divergence and reconstruct phylogenies of these 2 genera, focusing especially on Catharus thrushes in North America. We then qualitatively compared the phylogenies and levels of divergence within these 2 genera of codistributed parasites. Neither Brueelia nor Myrsidea appears to cospeciate with Catharus thrushes or passerine birds in general. The Myrsidea phylogeny exhibits significant levels of biogeographic structure, whereas the Brueelia phylogeny does not. Myrsidea and Brueelia also differ in their levels of intra-generic genetic divergence, with Myrsidea showing higher levels of genetic divergence and host specificity than Brueelia. Our genetic data support traditional morphology-based taxonomy in several instances in which the same species of Brueelia has been reported on multiple host taxa, e.g., all migrant Catharus spp. carry B. antiqua, with little haplotype divergence. Myrsidea found on each Catharus sp. are in general genetically distinct, except for M. incerta, which parasitizes both Catharus ustulatus and Catharus minimus. The strong biogeographic signal in the Myrsidea phylogeny and higher relative levels of host specificity of Myrsidea spp. suggest that infrequent host-switching, followed by speciation, is shaping the evolutionary history of this group. In contrast, the relatively lower host specificity of Brueelia spp. suggests that host-switching, combined with more frequent ongoing dispersal, has been more important in the evolutionary history of Brueelia.

RECENT EVOLUTIONARY HISTORY OF THE FOX SPARROWS (GENUS: PASSERELLA)

Abstract On the basis of plumage coloration and mitochondrial DNA variation, four main groups are recognized within the Fox Sparrow (Passerella iliaca): the red group (iliaca, RE), sooty group (unalaschcensis, SO), thick-billed (megarhyncha, TB), and slate-colored (schistacea, SC). To establish phylogenetic relationships among those four groups, we analyzed 2119 base pairs of sequence from four mitochondrial regions: ND2, ND3, cytochrome b, and control region. The control region is less variable than the coding genes surveyed. Both maximum parsimony and maximum likelihood resolved the same ingroup relationships (RE(SC(TB,SO))). However, placement of the root could not be established, even with four outgroups. Lack of resolution of the root is due to the nearest living relative of the Fox Sparrow being over 11% divergent. Despite lacking a clear root, the data suggest that the two taxa connected by a hybrid zone (TB, SC) are not sister species, which has implications for species limits because under the biological species concept they should be lumped. We recommend that all four main groups be recognized as species.

Phylogenetic relationships in the louse genus Penenirmus based on nuclear (EF-1α) and mitochondrial (COI) DNA sequences

Abstract. Ischnoceran lice in genus Penenirmus are parasites of birds in orders Piciformes and Passeriformes. No comprehensive revision of this genus has been published, but a few host‐based revisions have been done. Here we present a phylogenetic analysis of Penenirmus based on nuclear (elongation factor‐1 alpha) and mitochondrial (cytochrome oxidase I) gene sequences. Sequences from portions of both these genes provide a well resolved tree that is relatively stable across methods of analysis and to bootstrap resampling. Some aspects of the Penenirmus phylogeny reflect the phylogeny of their avian hosts. We identified monophyly of a group of Penenirmus species occurring on Passeriformes as well as monophyly of a group containing species sometimes placed in a genus Picophilopterus. Species of Penenirmus occurring on Old World barbets fall in several positions at the base of the tree, suggesting that other lineages of Penenirmus may be derived from those occurring on Old World barbets.

Biogeography and spatio-temporal diversification of Selenidera and Andigena Toucans (Aves: Ramphastidae)

Andean uplift, Plio-Pleistocene climatic fluctuation, and river dynamics in the Amazon basin have all been implicated in the diversification of the South American avifauna. We reconstructed phylogenetic relationships in the genus Selenidera, which has served as a classic case of putative refugial speciation, and the closely related genus Andigena, to better understand the processes driving their diversification. Using mitochondrial and nuclear DNA sequences, we constructed a phylogeny to estimate the pattern and timing of divergence within and between seven lowland Selenidera toucanets and the five species of Andigena mountain-toucans, which together form a single clade. All phylogenetic analyses supported the monophyly of the montane genus Andigena, but indicated that the genus Selenidera is likely paraphyletic with respect to Andigena. Our time tree analysis is consistent with the orogenic uplift of the northern Andean range having initiated the divergence between Selenidera and Andigena, and that the movement and fragmentation of montane habitats in response to Pleistocene climatic oscillations likely influenced diversification within Andigena. Estimated divergence times for lowland Amazonian Selenidera did not support the Last Glacial Maximum (LGM) refuge hypothesis as an important biogeographic factor for the diversification of lineages studied here. The timing of divergence within Selenidera is consistent with the hypothesis that geographic isolation of areas of endemism generated by Amazonian river dynamics during the Plio-Pleistocene contributed to Selenidera speciation and current species distributions.

HYBRIDIZATION AND POPULATION SUBDIVISION WITHIN AND BETWEEN ROSS'S GEESE AND LESSER SNOW GEESE: A MOLECULAR PERSPECTIVE

Abstract We reanalyzed Quinns (1992) mtDNA control region data set including new sequences from nine Lesser Snow Geese (Chen caerulescens caerulescens) and 10 Rosss Geese (Chen rossi) and found the same divergent lineages that Quinn (1992) attributed to vicariant separation of Lesser Snow Goose populations during the Pleistocene. However, peculiar patterns of mtDNA control region sequence variation, including a multimodal mismatch distribution of mtDNA sequences with two levels of population structuring and the sharing of two divergent haplotype lineages, are consistent with two hybridization episodes in Chen geese. Comparisons of mtDNA variation with historical and allozyme data sets compiled by Cooke et al. (1988) are consistent with the hypothesis that sharing of two mtDNA haplotype lineages between Rosss Goose and Lesser Snow Goose resulted from hybridization (Avise et al. 1992). Furthermore, population structure found within one haplotype cluster is consistent with Cooke et al.‘s (1988) hypothesis of past allopatry between blue and white Lesser Snow Geese. Hibridización y Subdivisión dentro y entre Poblaciones de Chen rossi y Chen caerulescens caerulescens: Una Perspectiva Molecular Resumen. Reanalizamos los datos de la región de control del ADN mitocondrial (ADNmt) de Quinn (1992), junto con nuevas secuencias de nueve individuos de la especie Chen caerulescens caerulescens y 10 de Chen rossi. Encontramos los mismos linajes divergentes que Quinn (1992) atribuyó a la separación vicariante de las poblaciones de C. c. caerulescens durante el Pleistoceno. Sin embargo, encontramos que las dos especies comparten dos linajes de haplotipos divergentes, y la distribución de “mismatch” en secuencias del ADNmt mostró multimodalidad con dos niveles de estructuración de la población. Estos patrones peculiares están de acuerdo con la hipótesis de que hubo dos episodios de hibridización en gansos del género Chen. Los datos históricos y de aloenzimas compilados por Cooke et al. (1988) también apoyan esta hipótesis (Avise et al. 1992). Además, la estructura de la población dentro de un grupo de haplotipos es consistente con la hipótesis de Cooke et al. (1988) acerca de la pasada alopatría entre los morfos azul y blanco de C. c. caerulescens.

Unlocking the black box of feather louse diversity: A molecular phylogeny of the hyper-diverse genus Brueelia

Songbirds host one of the largest, and most poorly understood, groups of lice: the Brueelia-complex. The Brueelia-complex contains nearly one-tenth of all known louse species (Phthiraptera), and the genus Brueelia has over 300 species. To date, revisions have been confounded by extreme morphological variation, convergent evolution, and periodic movement of lice between unrelated hosts. Here we use Bayesian inference based on mitochondrial (COI) and nuclear (EF-1α) gene fragments to analyze the phylogenetic relationships among 333 individuals within the Brueelia-complex. We show that the genus Brueelia, as it is currently recognized, is paraphyletic. Many well-supported and morphologically unified clades within our phylogenetic reconstruction of Brueelia were previously described as genera. These genera should be recognized, and the erection of several new genera should be explored. We show that four distinct ecomorphs have evolved repeatedly within the Brueelia-complex, mirroring the evolutionary history of feather-lice across the entire order. We show that lice in the Brueelia-complex, with some notable exceptions, are extremely host specific and that the host family associations and geographic distributions of these lice are significantly correlated with our understanding of their phylogenetic history. Several ecological phenomena, including phoresis, may be responsible for the macroevolutionary patterns in this diverse group.