Interplay between Developmental Flexibility and Determinism in the Evolution of Mimetic Heliconius Wing Patterns

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Old mice, young islands and competing biogeographical hypotheses

Naturally occurring variation within a small rodent species native to the southeastern USA, Peromyscus polionotus, has interested biologists for nearly a century. This species has contributed significantly to our understanding of geographical variation and has often been presented as an example of adaptive evolution. Much of the interest in this organism has been predicated on assumptions that the species is relatively young (<300 000 bp) and that coastal populations have a very recent history (<10 000 bp). To test these assumptions and the prevailing biogeographical hypothesis (Recurrent Invasion), we examined nucleotide sequence data from the cytochrome b and D-loop mitochondrial regions (2449 bp) for 79 samples of P. polionotus collected across the Gulf Coast region of Florida and Alabama. Samples representing Peromyscus maniculatus bairdii, P. m. sonoriensis, P. m. pallescens, and P. keeni were used as outgroups. The degree of cytochrome b divergence (approximately 4.4%) between P. maniculatus and P. polionotus was higher than expected. Analyses consistently indicated that three distinct groups are represented within P. polionotus from the Gulf Coast region. Among these, coastal populations (beach mice) form a monophyletic group and apparently represent a substantially older group (approximately 200 000 year. separation) than previously recognized. Our results were counter to the core assumptions of the existing biogeographical model but were consistent with an alternative hypothesis (Shore-line Tracking) which provides a more parsimonious explanation for the observed patterns. This research provides new insight into the evolutionary history of P. polionotus and highlights the importance of considering biogeographical history when evaluating extant patterns of natural variation.     

Biogeography of a southern hemisphere freshwater fish: how important is marine dispersal?

Galaxias maculatus is one of the world's most widely distributed freshwater fish. This species has a marine-tolerant juvenile phase, and a geographical range extending through much of the southern hemisphere. We conducted phylogeographic analyses of 163 control region haplotypes of G. maculatus, including samples from New Zealand (five locations), Tasmania (one location) and Chile (one location). A lack of genetic structure among New Zealand samples suggests that marine dispersal facilitates considerable gene flow on an intra-continental scale. The discovery of a Tasmanian-like haplotype in one of 144 New Zealand samples indicates that inter-continental marine dispersal occurs but is insufficient to prevent mitochondrial DNA differentiation among continents. The sister relationship of Tasmanian and New Zealand clades implies that marine dispersal is an important biogeographical mechanism for this species. However, a vicariant role in the divergence of eastern and western Pacific G. maculatus cannot be rejected.     

Delimiting species boundaries and the conservation genetics of the endangered maritime ringlet butterfly (Coenonympha nipisiquit McDunnough)

Species delimitation is a difficult problem that has implications across organismal biology, yet no single method has proved wholly satisfactory. We tested the utility of combining species-delimitation methods based on phylogeny and gene flow statistics using two parapatric members of the Coenonympha tullia group as an example: the endangered maritime ringlet butterfly (Coenonympha nipisiquit McDunnough) and the common inornate ringlet butterfly (Coenonympha inornata Edwards). We reconstructed the phylogeny of the nearctic C. tullia-group taxa from mitochondrial DNA (mtDNA) sequences (cytochrome oxidase I and mitochondrial control region) to explore the ancestry of the C. nipisiquit lineage within the group. We investigated the extent of gene flow between the two taxa with F-statistics using 587 nuclear amplified fragment length polymorphism markers, accounting for the effect of potential scoring 'collisions' where a marker may represent more than one DNA fragment. Combining species-delimitation methods was especially effective because it uncovered both historical and recent evolutionary patterns. Phylogenetic analysis of mtDNA revealed the early divergence of C. nipisiquit from other C. tullia-group taxa, including the morphologically similar C. inornata. F-statistics and gene-by-gene introgression profiles demonstrated clear isolation between the two taxa and revealed strong population structure within C. nipisiquit. C. nipisiquit is the first taxon in the nearctic C. tullia group showing strong evidence of genetic isolation. The methods we used are relatively inexpensive and can be widely used to delimit taxonomic boundaries near the species level, both generally and in particular for taxa that may be targets of conservation efforts.     

Phylogenetic relationships of Dalyat mirabilis Mateu, 2002, with a revised molecular phylogeny of ground beetles (Coleoptera, Carabidae)

Dalyat mirabilis Mateu 2002 (Coleoptera: Carabidae) is a cave species recently described from SE Spain, which, based on morphological analyses, has been related to the Promecognathinae (with one genus in western North America and four genera in South Africa). In this paper, we investigated the phylogenetic relationships of the main lineages of family Carabidae, and the placement of Dalyat among them, with the 18S rRNA full sequence and a fragment of wingless with the use of parsimony, a fast maximum likelihood algorithm (implemented in the program phyml ), and Bayesian posterior probabilities. Although with wingless alone the relationships of Dalyat were not robustly supported, both with 18S rRNA and in a combined analysis there was a strong support for a sister relationship between Promecognathus and Dalyat with the three methods used. Using a molecular-clock approach the two lineages were estimated to have diverged at a similar (or slightly earlier) age than the origin of Harpalinae, known to have radiated in the Cretaceous. This is compatible with a vicariant origin of the lineage leading to Dalyat because of the isolation of the Iberian plate from Pangea in the late Jurassic to early Cretaceous. Other robust relationships within the Carabidae are the monophyly of Harpalinae (including Morionini, Peleciini and Pseudomorphini), its sister relationship with Brachininae, and the inclusion of these two subfamilies together with Scaritini and the austral Psydrinae in a strongly supported clade (the 'higher' Carabidae).     

An empirical test of freshwater vicariance via river capture

River capture is a geomorphological process through which stream sections are displaced from one catchment to another, and it may represent a dominant facilitator of interdrainage transfer and cladogenesis in freshwater-limited taxa. However, few studies have been conducted in a manner to explicitly test the biological significance of river capture. Here we present a multispecies phylogeographical analysis to test whether the nonmigratory fish fauna of the Von River (South Island, New Zealand) is the product of a well-documented, Late Quaternary capture of a section of the Oreti River (Southland drainage). Specifically, we predict that nonmigratory fishes of the Von River will exhibit closer genetic affinities with those of Southland, rather than those of the Clutha system, into which the Von River presently drains. Mitochondrial DNA phylogeography (control region and cytochrome b sequence data) and analysis of nuclear orthologues of mtDNA sequences indicate that 'flathead'Galaxias of the Von River (n = 31, three sites) have greatest genetic affinities with those of Southland (Galaxias 'southern', n = 216, 38 sites), rather than with those of the Clutha River (Galaxias sp. 'D', n = 73, 32 sites). Likewise, Von River 'roundhead'Galaxias (n = 52, four sites) have greatest genetic affinities with those of Southland drainages (Galaxias gollumoides, n = 223, 58 sites), rather than with those of the Clutha River (Galaxias pullus, Galaxias anomalus, Galaxias gollumoides of the Nevis tributary; n = 68, 32 sites). These findings are consistent with our predictions that genetic affinities of the nonmigratory fish fauna in the Von River would reflect past, rather than present, drainage connections. Consequently, river capture is responsible for the nonmigratory fish fauna of the Von River. In a broader context, river capture has frequently influenced the distribution of genetic lineages among catchments in New Zealand freshwater-limited fish, and its biogeographical significance may have been underestimated in other regions.     

Coming to America: multiple origins of New World geckos

Geckos in the Western Hemisphere provide an excellent model to study faunal assembly at a continental scale. We generated a time-calibrated phylogeny, including exemplars of all New World gecko genera, to produce a biogeographical scenario for the New World geckos. Patterns of New World gecko origins are consistent with almost every biogeographical scenario utilized by a terrestrial vertebrate with different New World lineages showing evidence of vicariance, dispersal via temporary land bridge, overseas dispersal or anthropogenic introductions. We also recovered a strong relationship between clade age and species diversity, with older New World lineages having more species than more recently arrived lineages. Our data provide the first phylogenetic hypothesis for all New World geckos and highlight the intricate origins and ongoing organization of continental faunas. The phylogenetic and biogeographical hypotheses presented here provide an historical framework to further pursue research on the diversification and assembly of the New World herpetofauna.     

The historical ecology of yellow perch (Perca flavescens[Mitchill]) and their parasites

Aim To determine the origins of the host–parasite association between among yellow perch (Perca flavescens[Mitchill]) and the parasites Crepidostomum cooperi Hopkins, Proteocephalus pearsei La Rue and Urocleidus adspectus Beverly Burton. Of secondary interest are the parasites Bunodera luciopercae (Muller) and Proteocephalus percae (Muller) predictably associated with the Eurasian perch. Location The areas considered are the Holarctic, since the upper‐Cretaceous, and contemporary North America. Methods Published and new information from host and parasite phylogenies, palaeontology, palaeogeography and plate tectonics and host biology is incorporated to assess the origins of yellow perch and several of its parasites. This information is used to determine the origins for these host–parasite associations. Results Cladistic analysis suggests a Laurasian origin for Percidae and Perca, and that Perca is sister to the other genera in the family. Parasite phylogenies support a North American origin for the three species associated with yellow perch and a Laurasian origin for B. luciopercae. Proteocephalus pearsei and P. percae are not sister taxa. The fossil record for Perca dates to the Miocene in Europe and the Pleistocene in North America. North America and Europe were connected across the North Atlantic since at least the upper Cretaceous with separation complete by the Miocene. Europe was separated from Asia by the Obik Sea from the late Cretaceous until the Oligocene. Western cordillera orogeny and its accompanying high rates of water flow and Pleistocene glaciation represent barriers to Perca dispersal. Main conclusions The origin of Perca in North America dates at least to the late Oligocene when North America and Europe were connected across the North Atlantic and Europe and Asia were separate landmasses, and does not result from Pleistocene dispersal across Beringia from Asia. The present disjunction of Perca species in North America and Europe is due to the vicariant separation of North America and Europe. Based on the available information, yellow perch and its parasites have a North America origin. The association between yellow perch and the parasites in all cases is a consequence of host switching from other sympatric host species in North America and is not explained by co‐speciation. Even the association between the host‐specific Urocleidus adspectus and yellow perch originated with a host switch and is not due to co‐speciation. The basis for this host switching is geographical and ecological sympatry, especially shared feeding habits, with other North American fish hosts.     

Systematics of the extinct reed warblers Acrocephalus of the Society Islands of eastern Polynesia

In the Society archipelago (French Polynesia), Acrocephalus reed warblers are known only from four islands: Tahiti, Mo'orea, Huahine and Raiatea. All populations are now extinct except on Tahiti. Our knowledge of these birds is based on a small number of specimens preserved in museums, collected mostly during the 19th century. We present here a review of the past and present distribution, habitat and threats to the Society Islands reed warblers, including details on the specimens in museum collections. We compare the external morphology of the different populations, and use samples from museum specimens to propose a molecular phylogeny of all taxa based on partial cytochrome b gene sequences. The genetic data do not support the monophyly of the Society Islands reed warblers, which probably derived from three different lineages, found in Tahiti, Mo'orea and in the cluster Raiatea–Huahine. We outline the taxonomic consequences of this phylogeny. Our results support the hypothesis that evolutionary pattern, not distance between islands, shaped the long-distance colonization of oceanic islands by reed warblers.