Dissertations / Theses on the topic 'Species delimitation'

Zoanthidea (Cnidaria, Anthozoa, Hexacorallia) are common on tropical rocky shores and coral reefs. Large numbers of nominate species exist and many are difficult to identify. I used a genetic approach to examine population structure and taxonomy of zoanthids from the Great Barrier Reef and Torres Strait regions of Australia. Genetic population structure was investigated over two different spatial scales, using allozyme electrophoresis at seven polymorphic loci. I collected Palythoa caesia samples from 20 reefs spread over 1800 km. Results show that genetic differentiation among reefs is low but statistically significant, primarily due to variation in two populations. Population structure is characterised by genetic patchiness against a background of high gene flow. In a second study I collected Zoanthus coppingeri samples from three localities. Populations show evidence of asexual reproduction but are not dominated by successful clones. Sites separated by 50 m are genetically differentiated. Gene flow prevents fixed gene differences arising among populations of both these species. Fixed differences in taxonomic surveys (below) are therefore strong indicators of species boundaries. Eight species, from 19 localities throughout the Great Barrier Reef and Torres Strait, were genetically delimited using allozyme analysis of 14 loci. These species are separated by fixed gene differences. Genotype frequencies within species conform to Hardy-Weinberg predicted ratios. There is considerable morphological variation within species. Five species are identified in the family Zoanthidae: Palythoa caesia, Protopalythoa mutuki, Sphenopus marsupialis, Zoanthus coppingeri and Z. vietnamensis. These are probable senior synonyms of eighteen nominate species. Two Protopalythoa species remain unidentified. The eighth species, Parazoanthus dichroicus, is in the Parazoanthidae. A key to species is given. Phylogenetic analysis of zoanthid species used allozyme data and sequence data from 28S rDNA. I discuss results in the light of previous views of zoanthid systematics.

Hintergrund Zu den grundlegendsten Fragestellungen in der Biologie gehört die Frage nach der Natur und Entstehung biologischer Arten. Dieses Problem der Artdefinition (Engl. "Species Problem") war der Ursprung weitläufiger und kontroverser Diskussionen seit der Formulierung der Darwin'schen Evolutionstheorie. Bis heute wurden etwa 30 verschiedene und zum Teil gegensätzliche Konzepte zur Definition und wissenschaftlichen Abgrenzung der Art veröffentlicht. Eine Einigung ist nicht in Sicht. Gleichzeitig ist die Taxonomie mit der Herausforderung konfrontiert, dass ein immenser Teil der weltweiten Artenvielfalt wissenschaftlich noch nicht erfasst und beschrieben ist. Dies erfordert Methoden, die die Beschreibung neuer Arten beschleunigen und gleichzeitig deren Zuverlässigkeit und Nachvollziehbarkeit wahren. DNA-Barcoding, d.h. Artbestimmung an Hand eines kurzen standardisierten Fragments der DNA, soll die Erfassung der Artenvielfalt und das Erkennen unbekannter Arten beschleunigen. Die so genannte "Cybertaxonomie" erlaubt leichteren und schnelleren Zugriff auf vorhandene taxonomische Informationen, indem Daten online und kostenfrei zur Verfügung gestellt werden. Dies trägt zur Steigerung der Effizienz taxonomischer Prozesse bei. Integrative Taxonomie kombiniert verschiedene Beweislinien, wie zum Beispiel morphologische, molekulare und ökologische Daten, um die Zuverlässigkeit und Nachvollziehbarkeit bei der Abgrenzung und Beschreibung von Arten zu erhöhen. In dieser Dissertation untersuche ich zwei verschiedene Studiensysteme, um derzeit als gültig angesehene Modelle der Artbildung und Methoden der Artabgrenzung zu testen. Bei diesen Systemen handelt es sich um die Reptilien der Komoren, einer Gruppe ozeanischer Inseln im westlichen Indischen Ozean, und australische Wasserkäfer. Die Biogeographie dieser beiden Gruppen ist durch höchst unterschiedliche Faktoren geprägt: Die Komoren sind vergleichsweise junge vulkanische Inseln, deren einheimische, landbewohnende und flugunfähige Faunenelemente ausschließlich auf Besiedelung durch Drift über das offene Meer zurückgehen. Dagegen stellt Australien eine alte und isolierte Landmasse dar, deren Lebensgemeinschaften durch Klimaveränderungen in der Erdgeschichte geprägt sind. Ozeanische Inseln wurden schon von frühen Forschern als wichtige Systeme zum Studium der Biogeographie erkannt, und meine Untersuchung dieser beiden so unterschiedlichen Systeme stellt sowohl die Gemeinsamkeiten als auch die Unterschiede der Biogeographie von Inseln und Kontinenten heraus. Methoden und wesentliche Ergebnisse Als Fallbeispiele zur Untersuchung im Rahmen dieser Dissertation wählte ich zwei Teilgruppen der komorischen Reptilien (Geckos der Gattung Paroedura und Schlangen der Gattung Lycodryas) sowie drei Teilgruppen der australischen Wasserkäfer (die Familie Hygrobiidae und die Gattungen Antiporus und Sternopriscus aus der Familie Dytiscidae) aus. In beiden Fällen wurde der Grundstein für weitere Untersuchungen durch DNA-Barcoding gelegt, wie für die Reptilien als Teil dieser Dissertation beschrieben. Als nächsten Schritt führte ich Untersuchungen an mehreren mitochondrialen und nukleären Genmarkern durch, um die Phylogenien der jeweiligen Gruppen zu rekonstruieren und, im Fall der Hygrobiidae, das Alter der Phylogenie durch eine molekulare Uhr abzuschätzen. Ich versuchte, die Phylogenien komorischer Reptilien mit geologischen Daten über die erdgeschichtliche Entstehung der Inseln sowie die Ausbreitungsmöglichkeiten zu und zwischen den Inseln in Verbindung zu bringen. Bei Phylogenien australischer Käfer der Gattungen Antiporus und Sternopriscus suchte ich nach Korrelationen zu Klimaveränderungen in der Erdgeschichte, der Entstehung der australischen Trockengebiete und den Eiszeiten im Pleistozän. Diese Hypothesen konnte ich durch Belege für die ökologische Diversifikation australischer Käfer aus meinen Ökologischen Nischenmodellierungen untermauern. Auf der Grundlage der Ergebnisse von DNA-Barcoding und molekularen Phylogenien unternahm ich taxonomische Revisionen der betreffenden Gruppen nach Methoden der integrativen Taxonomie. Als Beweislinien verwendete ich Daten aus morphologischen Untersuchungen, mitochondrialen und nukleären Genen, sowie kategorische und quantitative ökologische Daten. Dieser Ansatz führte zur Beschreibung einer neuen Art von Käfern (Antiporus occidentalis HAWLITSCHEK, HENDRICH, PORCH, & BALKE, 2011), zweier neuer Arten (Paroedura stellata HAWLITSCHEK & GLAW, 2012 and Lycodryas cococola HAWLITSCHEK, NAGY & GLAW, 2012) und einer Unterart von Reptilien (Lycodryas cococola innocens HAWLITSCHEK, NAGY & GLAW, 2012), sowie zur Bestätigung oder Wiederherstellung der Gültigkeit der zuvor beschriebene Taxa Lycodryas maculatus (GÜNTHER, 1858) und Lycodryas maculatus comorensis (PETERS, 1874). Alle taxonomischen Handlungen wurden gemäß dem Konzept der Cybertaxonomie ausgeführt: es wurden LSID-Nummern vergeben, Einträge in Online-Datenbanken vorgenommen, und nach Möglichkeit Publikationsmodi mit freiem Zugang für Leser gewählt. Zudem verwendete ich die im Rahmen meiner Dissertation gesammelten Daten zur Abschätzung des artenschutzfachlichen Status der Reptilien der Komoren. Außerdem dienten sie als Basis für die Entwicklung von SmartHerper Comoros, einem Naturführer zur Herpetofauna der Komoren als Applikation für Smartphone. Schlussfolgerungen Die Ergebnisse meiner Untersuchungen weisen auf komplexe biogeographische Muster sowohl im insulären als auch im kontinentalen Untersuchungsgebiet hin. Demzufolge haben die Stammformen der dort heimischen Reptilien die Komoren in einem sehr komplizierten Muster besiedelt, das z.B. im Fall der Gecko-Gattung Paroedura mehrere Aussterbe- und Wiederbesiedlungsereignisse beinhaltet und kaum mit der geographischen Lage und dem geologischen Alter der Inseln korreliert. Viele endemische Arten zeigen mögliche morphologische Anpassungen an den Insellebensraum. Molekulare Daten komorischer Reptilien legen nahe, dass Grand Comoro, zuvor als geologisch jüngste Insel angesehen, möglicherweise weit älter ist als bislang angenommen. Über australische Wasserkäfer erhobene Daten zeigten, dass Artbildungsereignisse innerhalb dieser Gruppe von höchst unterschiedlichem erdgeschichtlichem Alter sind und vom Mesozoikum (Hygrobiidae) über das Pleistozän (Antiporus) bis in die jüngste erdgeschichtliche Vergangenheit (Sternopriscus) reichen. Molekulare Unterschiede weisen darauf hin, dass die "Sternopriscus tarsalis radiation" einen der am schnellsten verlaufenen bislang beschriebenen Artbildungsprozesse innerhalb der Insekten darstellt. Der integrativ-taxonomische Ansatz erwies sich in meinen Augen bei der Abgrenzung aller neu beschriebenen Taxa wie auch bei der Bestätigung bestehender Taxa als höchst erfolgreich. Durch diesen Ansatz standen Belege für die Artabgrenzung auch bei unzureichender morphologischer oder genetischer Differenzierung in ausreichendem Maße zur Verfügung. Ökologische Daten, insbesondere solche, die bei Ökologischer Nischenmodellierung gewonnen wurden, haben sich in diesen Fällen als höchst aussagekräftig bei der Artabgrenzung erwiesen. Bei der Anwendung des integrativ-taxonomischen Ansatzes auf Schlangen der Gattung Lycodryas argumentierte ich, den Rang der Unterart auf infraspezifische Einheiten mit einem gewissen Grad der Differenzierung anzuwenden. Schlussendlich liefern die Ergebnisse der Untersuchungen in meiner Dissertation nur einen kleinen, aber meiner Meinung nach dennoch nützlichen Beitrag zu unserem Verständnis darüber, wie biologische Arten entstehen und wie sie wissenschaftlich erfasst werden können. Meine Dissertation präsentiert diese Ergebnisse im Kontext der Debatte über die Artdefinition und stellt auch meine Meinung und Position darin dar. Meiner Ansicht nach ist diese äußerst fruchtbare Debatte von hoher Bedeutung für die zeitgenössische Entwicklung der Evolutionsbiologie und Biodiversitätsforschung.
Background The question of the nature and the origin of biological species is one of the most fundamental issues in biology. This so-called 'species problem' has been intensely debated since the formulation of the theory of evolution by Darwin. To date, about 30 concepts have been published that attempt to define, often conflictingly, what a species is and how it can be recognized by scientists, and a general agreement is not in sight. At the same time, taxonomy faces the challenge of a huge amount of global biodiversity that remains to be scientifically described. Therefore, taxonomic methods are required that make the description of new species faster and at the same time make them more reliable and reproducible. DNA barcoding, i.e., the use of a short standardized fragment of DNA for species identification, means to accelerate biodiversity inventories and the recognition of new species. Cybertaxonomy makes the access to taxonomic information easier and faster and helps increasing the efficiency of the taxonomic workflow by making data available online and free. Integrative taxonomy combines different lines of evidence, such as morphological, molecular, and ecological data, to make species delimitation and species descriptions more reliable and reproducible. In this dissertation I explore two different zoological study systems in order to test current models of speciation and methods of species delimitation. These study systems are the reptiles of the Comoros Archipelago, a group of oceanic islands in the Western Indian Ocean, and aquatic beetles of Australia. The biogeographical backgrounds of these two groups are very different: The Comoros are relatively young volcanic islands whose native terrestrial and non-flying fauna originates exclusively from overseas dispersal. In contrast, Australia is an old isolated landmass whose biota were shaped by past climate change. Oceanic islands have been recognized as prime study systems even by early biogeographers, and my study of these two different systems highlights the common grounds as well as the differences between insular and continental biogeography. Methods and principal findings I selected two groups out of the Comoran reptiles (Paroedura geckos and Lycodryas snakes) and three groups out of the Australian aquatic beetles (family Hygrobiidae and genera Antiporus and Sternopriscus, Dytiscidae) as study groups for this dissertation. In both cases, the data fundament for subsequent studies was laid by DNA barcoding, as included for reptiles in this dissertation. I then conducted analyses of several mitochondrial and nuclear genetic markers to reconstruct the phylogenies of the study groups and, in Hygrobiidae, estimate the divergence times within the phylogeny in a molecular clock approach. In Comoran reptiles, I attempted to correlate phylogenetic hypotheses with the geological history of island emergence and dispersal to and within the archipelago. In Australian Antiporus and Sternopriscus beetles, I attempted to correlate phylogenies with past climate change, the genesis of the Australian arid zone, and the Pleistocene climate oscillations. I used Ecological Niche Modeling to corroborate these hypotheses with evidence for ecological diversification in Australian beetles. Based on the results of DNA barcoding and molecular phylogenies, I used an integrative taxonomic approach to revise the taxonomy of the study groups accordingly. The lines of evidence I used were morphological data, mitochondrial molecular markers, nuclear molecular markers, and categorical and quantitative ecological data. This approach led to the description of one new species of beetle (Antiporus occidentalis HAWLITSCHEK, HENDRICH, PORCH, & BALKE, 2011) and two new species (Paroedura stellata HAWLITSCHEK & GLAW, 2012 and Lycodryas cococola HAWLITSCHEK, NAGY & GLAW, 2012), and one subspecies (Lycodryas cococola innocens HAWLITSCHEK, NAGY & GLAW, 2012), of reptiles, as well as to the confirmation or resurrection of the previously described taxa Lycodryas maculatus (GÜNTHER, 1858) and Lycodryas maculatus comorensis (PETERS, 1874). All taxonomic acts followed a cybertaxonomic concept by using LSID numbers, online databases, and, as far as possible, open access publication. Additionally, I used data collected in the course of this dissertation for estimating the conservation status of Comoran reptiles and for the development of SmartHerper Comoros, a field guide to the herpetofauna of the Comoros as a mobile application for smartphone. Conclusions The results of my studies show complex biogeographic patterns in both the insular and the continental study system. According to these results, the ancestors of native reptiles have colonized the Comoros Archipelago in a very complex pattern, including several events of extinction and re-colonization, e.g., in the case of the gecko genus Paroedura, with little correlation to the geographic positions or geological ages of the islands. Many endemic species show possible morphological adaptations to the island environment. Molecular data of reptiles suggest that Grand Comoro, the presumably geologically youngest island, may be considerably older than previously estimated. In Australian aquatic beetles, speciation events were shown to be of very different ages from Mesozoic (Hygrobiidae) to Pleistocene (Antiporus) and very recent (Sternopriscus). Molecular divergences indicate that speciation in the Sternopriscus tarsalis radiation was one of the fastest speciation events so far described among insects. I applied an integrative taxonomical approach in the delimitation of all newly described taxa and in the confirmation of previously described taxa. This approach provided sufficient evidence for species delimitation even in the absence of morphological differentiation (Antiporus), or when genetic data did not provide any clear evidence (Sternopriscus tarsalis radiation). In these cases, ecological data, particularly such data from Ecological Niche Modeling, was shown to be highly useful in integrative species delimitation. In the same approach applied to Lycodryas snakes, I argued for the usefulness of the subspecies rank for infraspecific entities with some degree of differentiation. I conclude that my research in the study systems I investigated in this dissertation are but small pieces that nevertheless advance our understanding of speciation and species delimitation by contributing to the ongoing debate on the species problem. My dissertation presents these results and represents my position in the debate. I see this debate as a very fruitful process that is highly important for the current development of evolutionary biology and biodiversity research.

The plant family Gesneriaceae is represented in Sri Lanka by six genera: Aeschynanthus, Epithema, Championia, Henckelia, Rhynchoglossum and Rhynchotechum, with 13 species (plus one subspecies/variety) of which ten are endemic including the monotypic genus Championia, according to the last revision in 1981. They are exclusively distributed in undisturbed habitats, and some have high ornamental value. The species are morphologically diverse, but face a problem of taxonomic delineation, which is further complicated by the presence of putative hybrids. Sri Lanka and Indian Peninsula, represent the Deccan plate of the ancient Gondwanan supercontinent. The presence of a relict flora may indicate the significance of the geological history of the Deccan plate for the evolution of angiosperms. The high degree of endemism here, along with their affinities to the global angiosperm flora paints a complex picture, but its biogeographic history is still unclear. The pantropical family Gesneriaceae distributed in Sri Lanka and South India is therefore an appropriate study group in this context. Besides, the family itself has a complex but largely unresolved biogeographical history especially concerning the origin and diversification of Old World Gesneriaceae. Modern approaches for the taxonomic studies were applied, integrating morphological and molecular data. Multiple samples were collected for each species across their geographical distribution. Nuclear ITS and chloroplast trnL-F sequences for the taxa from Sri Lanka were used to generate regional genus phylogenies of all six genera, using maximum parsimony. The rate of evolution of the nuclear ITS region versus chloroplast trnL-F was varied greatly across the six genera studied. Molecular delimitations were mostly congruent with the classical taxonomy. Over 65 taxonomic characters were studied in detail to recognize synapomorphies for clades and taxa. A complete taxonomic revision of Gesneriaceae in Sri Lanka, including lectotypification, was conducted based on both, the molecular and morphological data. This resulted in the recognition of 14 species in the six genera, including one newly described species H. wijesundarae Ranasinghe and Mich. Möller. Henckelia communis and H. angusta were not supported molecularly as two separate entities but are recognized as two species because of consistent morphological differences between them. Henckelia humboldtiana is proposed to represent a species complex due to its highly variable and inconsistent molecular and morphological diversity and overlap with H. incana and H. floccosa; more research is needed here. National conservation assessments were conducted, and all 14 species were recognized as threatened. Biogeographic affinities of Sri Lankan Gesneriaceae were elucidated, generating a dated phylogeny using an existing matrix of four plastid gene regions; trnL-F, matK, rps16 and ndhF, amended by sequences generated in this study. The final combined matrix included 175 taxa including newly generated sequences for the 13 Sri Lankan taxa. Phylogenetic trees were generated using parsimony, maximum likelihood and Bayesian inference. Molecular dating was carried out using BEAST and ancestral area reconstruction using BioGeoBears. These analyses indicated that the six genera of Gesneriaceae arrived in Sri Lanka separately and sometimes different time periods. One lineage dated back to the early diversification of the subfamily Didymocarpoideae (generally regarded as the Old World Gesneriaceae), which occurred around the KT boundary, before the Deccan plate was connected to Asia.

Heteromys pictus-spectabilis is a species complex within the subfamily Heteromyinae (Family: Heteromyidae) that is distributed along the western and southern Mexican coast and surrounding environments. Currently, the species complex is accepted as being 2 separate species (H. pictus and H. spectabilis), but this also renders H. pictus paraphyletic. Therefore, the species complex requires re-evaluation in order to resolve the paraphyly. Mitochondrial DNA sequences from a previously existing ~720 specimen database compiled by Victoria Vance were used in conjunction with new nuclear DNA sequences sequenced for the purpose of this study to generate multiple phylogenetic trees via the software programs RAxML, BEAST, and MrBayes to evaluate how different haplotype networks were related to each other. Using these molecular datasets in consideration with Kimura two-parameter values, time calibrations via BEAST, and the relative geographic locations of the haplotype networks, the results strongly indicate this species complex is composed of multiple cryptic species and potentially multiple genera. This was a preliminary exploration into this species complex however, and future research will be required to verify these findings.

BACKGROUND:Traditional morphological and biological species concepts are difficult to apply to closely related, asexual taxa because of the lack of an active sexual phase and paucity of morphological characters. Phylogenetic species concepts such as genealogical concordance phylogenetic species recognition (GCPSR) have been extensively used
however, methods that incorporate gene tree uncertainty into species recognition may more accurately and objectively delineate species. Using a worldwide sample of Alternaria alternata sensu lato, causal agent of citrus brown spot, the evolutionary histories of four nuclear loci including an endo-polygalacturonase gene, two anonymous loci, and one microsatellite flanking region were estimated using the coalescent. Species boundaries were estimated using several approaches including those that incorporate uncertainty in gene genealogies when lineage sorting and non-reciprocal monophyly of gene trees is common.RESULTS:Coalescent analyses revealed three phylogenetic lineages strongly influenced by incomplete lineage sorting and recombination. Divergence of the citrus 2 lineage from the citrus 1 and citrus 3 lineages was supported at most loci. A consensus of species tree estimation methods supported two species of Alternaria causing citrus brown spot worldwide. Based on substitution rates at the endo-polygalacturonase locus, divergence of the citrus 2 and the 1 and 3 lineages was estimated to have occurred at least 5, 400 years before present, predating the human-mediated movement of citrus and associated pathogens out of SE Asia.CONCLUSIONS:The number of Alternaria species identified as causing brown spot of citrus worldwide using morphological criteria has been overestimated. Little support was found for most of these morphospecies using quantitative species recognition approaches. Correct species delimitation of plant-pathogenic fungi is critical for understanding the evolution of pathogenicity, introductions of pathogens to new areas, and for regulating the movement of pathogens to enforce quarantines. This research shows that multilocus phylogenetic methods that allow for recombination and incomplete lineage sorting can be useful for the quantitative delimitation of asexual species that are morphologically indistinguishable. Two phylogenetic species of Alternaria were identified as causing citrus brown spot worldwide. Further research is needed to determine how these species were introduced worldwide, how they differ phenotypically and how these species are maintained.

Includes bibliographical references.
Ecklonia maxima and Ecklonia radiata are both kelp bed forming macroalgae along the South African coast, and the latter is also found in considerable abundance in Australia and New Zealand. Genetically they exist as two distinct species and can usually be differentiated morphologically, especially when occurring as geographically separate entities. However, they do appear to intergrade when growing together, where plants of apparent intermediate and indeterminate morphology have been observed. This study tested the reliability of morphology in separating these two Ecklonia species across their intraspecific range of morphological variation, from locations where the species co-occur, as well as where they appear in isolation in South Africa (both species) and Australia (E. radiata only). No individual characters reliably separated between species, yet overall size distinctions as well as the morphometric separation of hollow and solid-stiped sporophytes provide good evidence for morphological differentiation of E. radiata and E. maxima. While E. radiata clusters morphometrically, a distinction between Australian and South Africa specimens is observed. In localities where the two species are reported to co-exist morphological distinction is less clear, particularly in deep water at Buffels Bay. The blade morphology of these deep water sporophytes is distinct from both E. radiata and E. maxima across all locations, while overall size distinctions contribute most prominently to the morphological separation of E. radiata and E. maxima at De Hoop. Environmental data in combination with more detailed genetic analyses, especially those aimed at hybrid identification, are necessary to resolve the nature of these subtidal plants as well as to investigate the relationship between genetic differentiation and overlapping morphology in plants at De Hoop.

Brachyglottis (Asteraceae) is a genus of approximately 30 species in the Brachyglottidinae, a recently recognised sub-tribe of tribe Senecioneae. Within Brachyglottis is a clade of five species of rosette-forming herbs: B. bellidioides, B. haastii, B. lagopus, B. southlandica and B. traversii. A sixth species, B. saxifragoides, has recently been synonymised with B. lagopus. The rosette-Brachyglottis have historically been recognised as a taxonomically problematic group because species overlap in both morphology and geographical distribution. A recent molecular study of rosette-Brachyglottis using AFLP data indicated that genetic distances among populations of rosette-Brachyglottis in the South Island appear to be correlated with geographical distance between populations rather than taxonomic identification. This is problematic as the currently described rosette-Brachyglottis species have overlapping ranges which implicitly hypothesises reproductive barriers other than geographic distance. We conducted an investigation into the species delimitation of rosette-Brachyglottis with the aim of answering two related questions: Does the current delimitation of rosette-Brachyglottis accurately reflect patterns of genetic similarity? Do the patterns of genetic structure in rosette-Brachyglottis support the presence of multiple biological species? A total of 46 populations of rosette-Brachyglottis were represented in this study. Herbarium specimens collected from these populations were identified following the taxonomic treatment of Allan (1961). Twenty one discrete and numerical morphological characters were measured from herbarium specimens including those collected for this study and previously collected herbarium specimens. Morphological dissimilarity of 354 herbarium specimens was investigated by performing a PCoA on Gower’s pairwise morphological distances among individuals. The pattern of genetic similarity was explored using DNA fragment length variation in nine markers for 273 individuals and this resulted in a total 177 unique alleles. Bayesian clustering analysis was performed on this data set using STRUCTURE, in addition, pairwise genetic distances were calculated among individuals and populations using Jaccard and Nei’s dissimilarity coefficient’s respectively. Jaccard genetic distances among individuals were analysed using PCoA and Nei’s genetic distances among populations were analysed using a Neighbour-Net analysis. The relationship between pairwise genetic and geographic distances among populations was analysed using a combination of linear regression and a Mantel Test. The pattern of morphological similarities among specimens was generally congruent with the currently delimited species in rosette-Brachyglottis. However, many morphologically intermediate specimens confound the recognition of distinct morphological entities. Comparison of patterns of genetic similarity and the current morphologically-based species delimitation showed that the delimitation does not accurately reflect the genetic structure of rosette-Brachyglottis. Furthermore, patterns of genetic dissimilarity did not indicate discrete genetic groups at the individual or population levels. The finding of incongruence between patterns of genetic and morphological similarity and absence of morphologically or genetically discrete groups suggests that rosette-Brachyglottis are best considered a single, yet morphologically diverse, biological species. In addition genetic structure within this species appears to be primarily driven by geographical isolation.

Phylogeography and molecular phylogenetics have proven remarkably useful for understanding the patterns and processes influencing historical diversification of biotic lineages at and below the species level, as well as delimiting morphologically cryptic species. In this dissertation, I used an integrative approach coupling comparative phylogeography and coalescent-based species delimitation to improve our understanding of the biogeography and species limits of Central American freshwater fishes. In Chapter 1, I conducted a literature review of the contributions of phylogeography to understanding the origins and maintenance of lower Central American biodiversity, in light of the geological and ecological setting. I highlighted emerging phylogeographic patterns, along with the need for improving regional historical biogeographical inference and conservation efforts through statistical and comparative phylogeographic studies. In Chapter 2, I compared mitochondrial phylogeographic patterns among three species of livebearing fishes (Poeciliidae) codistributed in the lower Nicaraguan depression and proximate uplands. I found evidence for mixed spatial and temporal divergences, indicating phylogeographic “pseudocongruence” suggesting that multiple evolutionary responses to historical processes have shaped population structuring of regional freshwater biota, possibly linked to recent community assembly and/or the effects of ecological differences among species on their responses to late Cenozoic environmental events. In Chapter 3, I used coalescent-based species tree and species delimitation analyses of a multilocus dataset to delimit species and infer their evolutionary relationships in the Poecilia sphenops species complex (Poeciliidae), a widespread but morphologically conserved group of fishes. Results indicated that diversity is underestimated and overestimated in different clades by c. ±15% (including candidate species); that lineages diversified since the Miocene; and that some evidence exists for a more probable role of hybridization, rather than incomplete lineage sorting, in shaping observed gene tree discordances. Last, in Chapter 4, I used a comparative phylogeographical analysis of eight codistributed species/genera of freshwater fishes to test for shared evolutionary responses predicted by four drainage-based hypotheses of Neotropical fish diversification. Integrating phylogeographic analyses with paleodistribution modeling revealed incongruent genetic structuring among lineages despite overlapping ancestral Pleistocene distributions, suggesting multiple routes to community assembly. Hypotheses tests using the latest approximate Bayesian computation model averaging methods also supported one pulse of diversification in two lineages diverged in the San Carlos River, but multiple divergences of three lineages across the Sixaola River basin, Costa Rica, correlated to Neogene sea level events and continental shelf width. Results supported complex biogeographical patterns illustrating how species responses to historical drainage-controlling processes have influenced Neotropical fish diversification.

The inference of species boundaries and phylogenetic relationships are fundamental for evolutionary, ecological, and conservation studies. The resolution of species boundaries and the inference of phylogenetic relationships among species are required to define the units of analysis and to find the most closely related units for evaluating alternative models of speciation. I highlight lizards as model organisms for ecological and evolutionary studies, emphasizing their contributions to advances in understanding linkages between phylogeography and speciation. In this dissertation, I focus on the phylogenetic relationships of the lizards in the Liolaemus darwinii group, and the species boundaries of a nested clade within the group, the L. darwinii complex, because of several advantages that make these taxa ideal for phylogeographic studies of speciation. I infer a phylogeny for the L. darwinii group based on DNA sequences of 20 loci (19 nuclear and 1 mitochondrial) using species trees methods that take into account the incongruence among gene trees. I found the minimum number of loci, number of sequences per species, and number of base pairs per locus that should be included in an analysis for an accurate and precise estimate of the species tree. The species tree based on all available data support a clade of closely related species (L. darwinii, L. grosseorum, and L. laurenti) known as the L. darwinii complex. A new method for species delimitation using Approximate Bayesian Computation is introduced and is shown to accurately delimit species given that limited or no gene flow has occurred after divergence and despite biased estimates of demographic parameters. ABC analyses supported the distinctness of two lineages within L. darwinii under a model of speciation with gene flow. Based on the species tree and the species limits obtained in this dissertation, phylogenetic comparative methods can be carried out to address the morphological and ecological evolution in the L. darwinii group and several sister species can be used for testing the alternative speciation models via correlation analyses of genetic, morphological, and ecological datasets. Future studies should assess the role speciation due to adaptive processes and its association the species' ecological niches and life histories.

L'introgression est un processus fréquent et qui a d'importantes conséquences évolutives. L'objectif de ce travail était de tester un modèle neutre d'introgression chez des épicéas du Plateau tibétain et des régions voisines. Le travail a permis de mettre en évidence que la direction de l'introgression pouvait être prédite par la dynamique passée des populations d'arbres, et que l'importance de cette introgression était inversement proportionnelle à l'intensité des échanges génétiques au sein de l'espèce invasive, grâce à la comparaison de la structure génétique basée sur des marqueurs chloroplastiques (à hérédité paternelle) et mitochondriaux (à hérédité maternelle)
Introgression is a widespread phenomenon with potentially profound evolutionaryconsequences. Recently, significant progress in our understanding of introgression hasbeen made with the development of a neutral model. This model predicts that, whenone species invades an area already occupied by a related species, introgression ofneutral genes takes place mainly from the local species towards the invading ones. Inaddition, following a contact between two hybridizing species, the model predicts thatintrogression should be particularly frequent for genome components experiencinglittle gene flow. However, to date, there was no empirical example available, in whichone species expanded into the range of a closely related one and two markers withcontrasted rates of gene flow had been studied for both species. Only in such a casecould the two predictions outlined above be tested simultaneously. In addition, basedon these two predictions, species delimitation should be more efficient when usingmolecular markers experiencing high rates of gene flow. The present thesis was designed to test the hypotheses of this model. The biologicalmodel used was conifers, a group in which introgression and hybridization arecommon because of incomplete reproductive isolation. The species investigatedbelong to the genus Picea (spruce). We focused on two species complexes,represented by monographic clades in a phylogenetic study using the chloroplast genematK. All species studied occur in the Qinghai-Tibetan Plateau (QTP) and adjacenthighlands. The phylogeography of these species complexes was reconstructed usingorganelle markers (mitochondrial DNA, mtDNA and chloroplast DNA, cpDNA). Inconifers, mtDNA and cpDNA have contrasted modes of inheritance. The former ismaternally inherited, transmitted by seeds experiencing little gene flow while thelatter is paternally inherited, transmitted by both pollen and seeds experiencing highlevels of gene flow. Therefore, uniparentally inherited mtDNA and cpDNA markersexperience different rates of gene flow in such a group, providing an ideal model to test the relationship between rates of gene flow, introgression and species delimitation.Two mtDNA fragments (nad1intron b/c; nad5 intron1) and three cpDNA fragments(ndhK-C；trnL-trnF；trnS-trnG) were sequenced for nine species belonging to thePicea asperata and P. likiangensis species complex.(1) Nine mtDNA and nine cpDNA haplotypes were detected in 459 individualsfrom 46 natural populations in five species of P. asperata complex. As found in mostconifer species studied so far, low variation is present in the two mtDNA intronsalong with a high level of differentiation among populations (GST = 0.90). In contrast,higher variation and lower differentiation among populations was found at cpDNAmarkers (GST = 0.56). The cpDNA, although far from being fully diagnostic, is morespecies-specific than mtDNA: four groups of populations were identified usingcpDNA markers, all of them related to species or groups of species, whereas formtDNA, geographical variation prevails over species differentiation. A literaturereview shows that mtDNA variants are often shared among related conifer species,whereas cpDNA variants are more species-specific. Hence, increased intraspecificgene flow appears to decrease differentiation within species but not among species.[...]

In this thesis I have connected different evolutionary studies of Lioalemus lizards. In Chapter 1, I followed an integrative approach to delimit species in the Liolaemus walkeri complex. Using mitochondrial markers, morphological data, bioclimatic information and methods appropriate for each data type, we found that the name L. walkeri was covering three new lineages. Three new species were described and one of them (L. chavin) is now categorized as Near Threatened in the IUCN red list. In Chapter 2, I change the subject from species boundaries to the study of viviparity and placentation. In this paper we employed scanning electron and confocal microscopy to compare the placental ultra-structure and pattern of blood vessels in two Liolaemus species. One of the most remarkable traits found is the complete reduction of the eggshell in both placentae, a possible adaptation to improve gas exchange in the hypoxic environments of the high Andes. In chapter 3, I returned to the issue of species delimitation and employed two integrative approaches: a hypothetical deductive framework and a model-based procedure. I applied both approaches in lowland and highland Liolaemus species of the montanus group. I found that in only one case (of four) an unnamed lowland lineage ("Nazca") was delimited concordantly by both procedures. In Chapter 4, I focus on a study of convergent evolution of desert phenotype in Liolaemus species and Ctenoblepharys adspersa. I performed a Bayesian time calibrated and maximum likelihood tree based on 55 taxa and seven molecular markers. We employed quantitative and categorical traits based on 400 specimens and non-metric multidimensional scaling to obtain new quantitative variables. I used three phylogenetic comparative methods to identify and measure the strength of convergence. My results found a strong case of convergent traits in C. adspersa, L. lentus, L. manueli, L. poconchilensis and L. stolzmanni that are probably related to predator avoidance in the Peruvian-Atacama and Monte deserts. In addition, my time calibrated tree resolves the origin of these traits first in C. adspersa at about 80 million years (My) and later independently in Liolaemus species at about 25 My suggesting the present of evolutionary constraints.

The remarkable richness of the Cape Floristic Region (CFR) and the high in situ diversification inferred for the region prompt interest in two key areas: first, to what extent has the true species richness of the Cape been discovered and described, and second, what are the key drivers of speciation? Steady efforts in taxonomy dating back to the early 17th century have led some to estimate that over 99% of species in the Cape flora have already been described. However, taxonomic research in the Cape has, as elsewhere, relied on morphology for delimiting species, implying that undiscovered species diversity among cryptic taxa may be substantial. Early ideas regarding the drivers of diversification in the Cape flora emphasised climatically-induced vicariant speciation. Since that time, both vicariance and ecological speciation have been invoked as drivers of diversification. However, the relative contributions of either of these modes to the richness of the flora remains unclear. The present work focuses on Seriphium plumosum, a species complex in the daisy tribe Gnaphalieae with a recent evolutionary origin and a core distribution in the Cape Floristic Region. The species’ problematic taxonomic history, its substantial morphological and ecological variability, as well as its large geographic distribution in southern Africa suggest that the current concept of the species houses multiple independent evolutionary species. Species limits within the complex are reevaluated using the Bayes Factor Delimitation method of Leaché et al. (2014) within an integrative taxonomic approach, incorporating evidence from next-generation DNA sequence data, previous taxonomic treatments, morphology, ecology and geographic distribution. The drivers of lineage divergence are then investigated at the population level within selected species in the group, with a focus on neutral and adaptive processes, and the spatial contexts within which each of these processes is thought to operate. The evidence presented here provides support for ten or eleven independent evolutionary species housed within the current concept of S. plumosum. The species currently considered sibling to the complex, Seriphium cinereum, is strongly supported as embedded within the complex, which I thus term the S. plumosum clade. In order to better facilitate identification of the species, several of the evolutionary species uncovered are consolidated. As such, a total of nine species are described in the taxonomic treatment of the clade, each of which is morphologically and ecologically cohesive, and five of which qualify as biological species sensu Mayr (1969). These include S. plumosum1 and S. cinereum under new circumscriptions, four new species; Seriphium alto-argillaceum, S. burrowsii, S. crypticum and S. dunensis, and three species housed within the current concept of S. plumosum and formerly treated within Stoebe; Seriphium burchellii Levyns, S. vulgaris Levyns, and S. virgatum Thunb. At the population level within selected species in the S. plumosum clade, neutral processes and weak divergent selection within a spatially discontinuous context are supported as the key driver of differentiation. These findings suggest that phylogenetic niche conservatism and/or the limited dispersal abilities characteristic of many Cape clades have been important in diversification by neutral and adaptive processes in the region. If extrapolatable to other Cape lineages, this work implies that undiscovered species diversity in the Cape may considerably exceed recent estimates, and that the role of non-adaptive processes in generating species diversity may be considerably underestimated.

Accurate species delimitation has critical implications for ecological and conservation studies; and for understanding factors driving diversification. However, a growing body of evidence indicates that morphology-based species circumspection in lichenized ascomycetes often fails to accurately represent the number of fungal species. The use of molecular data in lichen systematics provides an important alternative to traditional morphological characters for identifying natural groups and assessing evolutionary histories in challenging lichen taxa. In this work, I examined two common lichen-forming genera in western North America, Rhizoplaca and Xanthoparmelia, as models for investigating character evolution, species delimitation in morphologically and chemically diverse species, and identification of lineages in the early stages of divergence. Phylogenetic hypotheses were reconstructed to assess character evolution using sequence data from four nuclear ribosomal markers and fragments from two nuclear loci. I applied a multifaceted approach to delimit species in Rhizoplaca and Xanthoparmelia by assembling multiple lines of evidence using DNA sequence data, and genealogical and population genetic analyses. I have found that traditionally circumscribed species are not supported by molecular data. For example, in Rhizoplaca previously unrecognized lineages were identified within what has thus far been considered a single species. In contrast, morphologically and chemically distinct species within Xanthoparmelia were not supported by molecular data. Distinct medullary chemistries, growth forms, and the production of vegetative diaspores appear to have evolved independently multiple times in Xanthoparmelia. This work clearly indicates that morphological and chemical characters do not always accurately reflect lichen species diversity within even the best known and studied genera. My study of the Rhizoplaca melanophthalma species complex demonstrates that the genus Rhizoplaca, as presently circumscribed, is more diverse in western North American than previously thought. I present these analyses as a working example of species delimitation in morphologically cryptic lichenized fungi. In Xanthoparmelia diagnostic morphological and chemical characters have evolved in a highly homoplasious manner. In contrast to other studies documenting previously undiscovered fungal lineages masked within lichen species circumscribed by traditional morphological and chemical characters, my work suggests that species diversity has been overestimated in the lichen genus Xanthoparmelia.

Changes to traditional taxonomic methods to incorporate new technologies and methods have already improved the quality of species hypotheses, but more work can be done to improve the speed of new species documentation. The mitochondrial COI DNA barcode has been successfully used to identify species with high accuracy since the early 2000s, and has been used in conjunction with morphological examinations and other DNA markers to discover and delimit new species. This thesis explores the application of DNA barcodes as the primary data for delimitation and diagnosis of new species of ichneumonoids. The genera Zelomorpha and Hemichoma are revised and 18 new species from the Área de Conservación Guanacaste in Costa Rica are diagnosed based on COI barcodes. Two additional species are described based on morphology. An illustrated morphological key and morphological diagnoses for each species are also included.

Despite their documented importance in maintaining ecosystem functioning, many groups of soildwelling invertebrates have been inadequately studied with regards to many facets of their biology. In this thesis, I focussed my attention on two poorly understood aspects of the diversity of soildwelling invertebrates: (i) local values of species richness and (ii) species boundaries. As a study system, I used one of the most neglected groups of soil invertebrates, i.e. the centipedes (Myriapoda: Chilopoda), and I carried out my research in the area of the Southeastern Prealps, which are known to harbour a large amount of biodiversity. The present work is in the form of a paper collection and contains a general introduction, two chapters and a conclusive paragraph. An original research article, dealing with the aforementioned topics, is presented at the end of each chapter. In chapter I, I addressed the dearth of knowledge on the local species richness of centipede communities. I applied statistical models to estimate the actual species richness of 10 centipede communities in the Southeastern Prealps and to compare richness values between these communities. I demonstrated that up to 27-28 species of centipedes can coexist in syntopy in temperate forests of the Southeastern Prealps, and that richness values can vary significantly among communities. In chapter II, I addressed the issue of species delimitation in poorly-vagile endogeic invertebrates, for which traditional morphology-based taxonomy has proven ineffective in identifying species boundaries. In particular, I applied a rigorous integrative approach, in order to test for the existence of species boundaries in a set of populations traditionally referred to a single species of endogeic centipede, i.e., Clinopodes carinthiacus (Chilopoda: Geophilomorpha). Taking advantage of the integration between different lines of evidence (sequences from three DNA loci and morphological characters) and different methods for species discovery (ABGD, GMYC, PTP and Expectation Maximisation cluster analysis with v-fold cross-validation), I found evidence for the existence of at least two candidate species within the investigated population system.

Taxonomic delimitation of rare species is vital for accurate assessments of diversity and for their conservation. Cycladenia humilis, the sole species of Cycladenia, is an enigmatic perennial widely dispersed across the western United States. Within this species there are three currently recognized varieties: C. humilis var. humilis in Northern California, C. humilis var. venusta in Southern California, and C. humilis var. jonesii in Utah and Northern Arizona. Some populations occur geographically in areas between the typical distribution of each variety and the presently accepted taxonomy inadequately addresses these populations. Using five nDNA regions, we seek to clarify relationships between current varieties and assess the pattern of variation throughout the species. Analyses including K-means clustering, principle component analysis, fields for recombination, AMOVA, and ecological niche modeling were applied. Results indicate significant genetic structure between varieties and supports recognition of C. jonesii at the species level as distinct from C. humilis. Well defined intraspecific groupings are evident in the data, with evidence supporting the recognition of an additional variety in C. humilis, and two varieties in C. jonesii. Haplotype diversity and relationships between metapopulation clusters inform conservation efforts regarding diversity within Cycladenia and offer insights into the historical demography of this genus.

I constructed a best estimate phylogeny based on congruence of multiple data sources. In recent years molecular data has been used both to construct phylogenies of taxonomic groups and to aid in the delimitation of new species. I generated and analyzed sequence data for forest spiny pocket mice (Genus Heteromys) for the mitochondrial gene cyt b (1143 bp) and two nuclear gene segments MYH2 (252 bp) and EN2 (189 bp). I used maximum parsimony and Bayesian analyses to infer relationships among species and to provide a framework for using a species delimitation method to investigate the possibility of multiple species within the widespread Heteromys desmarestianus. I found several well-supported lineages within the H.desmarestianus complex, including H. goldmani and H. oresterus. Incorporating karyotype and allozyme data from earlier studies, I found sufficient supporting evidence to justify maintaining H. goldmani and H. oresterus as species as well as identifying four lineages as candidate species. I present a revised taxonomic arrangement within the genus; the subgenus Heteromys should be divided into three species groups: anomalus (H. anomalus and H. australis), gaumeri (H. gaumeri), and desmarestianus (H. desmarestianus, H. goldmani, H. oresterus, and the four candidate species).

I highlight two squamate lineages endemic to Southeast Asia, the Asian water monitors (Varanus salvator Complex) and the Angle-headed lizards (Agamidae: Gonocephalus), and elucidate their systematic affinities and historical biogeography. My results represent novel phylogenetic inferences, with biogeographic histories and diversification events corresponding to major climatic fluctuations over the past 30 million years. Additionally, I solidify the taxonomy and systematics of the Varanus salvator Complex, and enumerate more than a dozen Gonocephalus lineages as candidate species in need of taxonomic scrutiny. Lastly, I investigate contemporary and historical patterns of dispersal throughout Sundaland, and between Sundaland and the Philippines.

Orientador: Antonia Cecilia Zacagnini Amaral
Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia
Made available in DSpace on 2018-08-22T01:09:52Z (GMT). No. of bitstreams: 1 Quast_MonicaPaiva_D.pdf: 2160555 bytes, checksum: a8f9e004b6d58d12222f57e63c865d89 (MD5) Previous issue date: 2013
Resumo: Espécies são unidades fundamentais da biologia e sua identificação é essencial para a pesquisa nos mais diversos campos. Esta tarefa, no entanto, é dificultada por limites interespecíficos naturalmente mal definidos, especialmente em ambientes marinhos, onde complexos de espécies crípticas são comuns. Assim, a delimitação de espécies tem recebido grande atenção nos últimos anos e técnicas moleculares têm se mostrado de grande importância para a questão. Corbula (Bivalvia: Corbulidae) é um gênero frequente e ecologicamente importante em comunidades bentônicas marinhas de sublitoral. A taxonomia do grupo é bastante confusa, em parte devido à plasticidade fenotípica das conchas que dificulta o estabelecimento de limites morfológicos entre as espécies. O presente estudo teve como objetivo estudar, com base em sequências de dois marcadores moleculares (COI e 16S), os limites entre seis espécies de Corbula morfologicamente identificadas da costa sudeste e sul do Brasil, de forma a testar a delimitação morfológica. Como se trata de espécies predominantemente de sublitoral, o material analisado encontrava-se preservado em álcool, havendo sido fixado em formol. Dessa forma, fez-se necessário o desenvolvimento de protocolos específicos de extração e amplificação. Uma combinação de extração orgânica com adsorção em sílica mostrou-se o melhor método de extração de DNA total. Para as reações de amplificação, a utilização de nested PCR produziu resultados superiores à PCR direta. As análises de delimitação utilizaram quatro métodos diferentes, dois baseados em árvores (GMYC e Brownie) e dois não (regra das 4x e ABGD). Os resultados divergiram entre métodos e marcadores, mas a combinação das diferentes linhas de evidência permitiu corroborar a delimitação morfológica de três espécies (Corbula caribaea, Corbula tryoni e Corbula lyoni). Os indivíduos identificados como Corbula patagonica dividiram-se em duas espécies distintas. O único indivíduo de Corbula aequivalvis foi considerado distinto das outras espécies e um indivíduo atribuído a Corbula sp1 não pôde ser distinguido de C. caribaea
Abstract: Species are fundamental unities in many biological studies and, being so, their identification is essential for researches in many different fields. This task, however, is complicated by badly defined interspecies boundaries, especially in the sea, where cryptic species are quite common. Species delimitation has been receiving much attention, and molecular techniques have been proved of great value to the matter. Corbula (Bivalvia, Corbulidae) is frequent and ecologically important genus in benthic marine communities. Nevertheless, its taxonomy is confusing, in part due to a plastic shell, which makes it difficult to establish species boundaries. This study aimed to analyze the COI and 16S sequences of six morphologically identified Corbula species occurring off the South-Southeastern Brazilian coast. Being a mainly sublittoral genus, most of the analyzed material had been previously sampled, fixed in formalin and preserved in alcohol. Hence, initially specific protocols for DNA extraction and PCR were developed. Better results were obtained with an extraction protocol combining organic extraction and silica adsorption. The nested PCR yielded more product than the direct PCR. Delimitation analyses were conducted with four different methods: two tree based (GMYC and Brownie) and two non-tree based (4x rule and ABGD). Different methods and markers produced different delimitations, but the combined evidence supports the morphological delimitation of three species: Corbula caribaea, Corbula tryoni and Corbula lyoni. Individuals assigned to Corbula patagonica were separated into two molecular species. Only one individual of Corbula aequivalvis was analyzed and it was distinguished from other species. One individual assigned to Corbula sp1 could not be distinguish from C. caribaea
Doutorado
Ecologia
Doutora em Ecologia

This thesis reports the results of a systematic investigation of the genus Bitis, Africa's most taxonomically diverse and geographically wid espread genus of viperid snakes, in addition to phylogeographic investigations of its most geographically widespread representative, Bitis orietans. The phylogeny of Bitis is inferred using mitochondrial sequences plus those from two phylogenetically informative nuclear markers, and analysed using recently developed multispecies coalescent methods. These analyses successfully resolve the phylogenetic position of several rare and previously unstudied species and confirm the majority of intrageneric relationships. Several instances where current taxonomy may not adequately portray evol utionary relationships among these snakes are also revealed. The development of five anonymous nuclear markers for the genus is also reported that will be valuable for future studies, and are utilised here in more focussed investigations of the puff adder, B. arietans. A pan-African phylogeographic invest igation of B. arietans revea ls the presence of multiple parapatric mitochondrial clades. The historical processes responsible for generating phylogeographic st ructure in southern Africa are invest igated using species distribution modelling and genetic approaches, which support isolation in multiple southern refugia during Pleistocene cold phases. Nuclear markers indicate varying degrees of admixture between these southern refugial populations upon secondary contact. The occurrence of divergent mitochondrial lineages elsewhere in the range of B. arietans indicates the possibility of cryptic speciation. This hypothesis is tested using recently developed coalescent species delimitation approaches applied to data from anonymous nuclear markers. This supports the existence of six candidate species, although corroborative evidence and/or increased geographic sampling will be requ ired before taxonomic changes are justified. Overall, this thesis provides a significant advance in our understanding of the evolutionary history of Bitis. The results also highlight several priorities for future research on these snakes.

China harbors 10% of the world's salamander species. Studying their evolutionary history provides critical insights into the evolution of the fauna of the Far East. The stout newts (Pachytriton, also known as paddle-tailed newts) are a genus of aquatic montane salamanders that are widely distributed in southeastern China. Despite their longstanding popularity among the global pet trade, little is known of their biology beyond external morphology. My thesis presents the first systematic study to elucidate phylogenetic relationships, character evolution, biogeographic patterns, species delimitation, and speciation mechanisms in this genus.

Les écosystèmes des eaux souterraines sont de plus en plus reconnus pour leur faune endémique, phylogénétiquement ancienne et écologiquement spécialisée. Avec plus de 425 espèces décrites, les amphipodes niphargidés constituent la famille des eaux souterraines la plus riche en espèces au monde et un système modèle intéressant pour la biologie de l'évolution. Cependant, les scientifiques doivent faire face à des données incomplètes et biaisées en raison de trois déficits majeurs: le déficit Linnéen pour la taxonomie, le déficit Darwinien pour la phylogénie, et le déficit Wallacien pour la biogéographie. La présente thèse vise à évaluer l'importance de ces déficits chez les niphargidés, ouvrant ainsi la voie pour y remédier. Le premier chapitre est une évaluation des effets de la découverte d'espèces cryptiques (une des causes du déficit Linnéen) sur notre compréhension des modèles de distribution à grande échelle de la diversité des niphargidés. Contrairement à ce que l'on attendait, les espèces cryptiques putatives sont réparties de manière homogène le long des gradients environnementaux, et leur découverte ne modifie donc pas notre compréhension des modèles de distribution. Le deuxième chapitre analyse l'importance de l'application des techniques moléculaires à la taxonomie des niphargidés. En étudiant le genre Microniphargus, la morphologie seule s'est avérée peu informative en raison de la pédomorphose et de l'homoplasie. L'utilisation de marqueurs ADN a permis d'attribuer le genre à une famille différente (Pseudoniphargidae), venant éclaircir les relations phylogénétiques au sein des Niphargidae (et contribuant ainsi à remédier au déficit Darwinien). Le troisième chapitre traite du rôle des régressions et transgressions marines sur la distribution des niphargidés en utilisant la biogéographie moléculaire et une modélisation biogéographique innovante (afin de remédier au déficit Wallacien). Les résultats soutiennent l'idée que la dispersion a joué un rôle essentiel dans la biogéographie historique des niphargidés, en montrant que leurs voies de dispersion sont corrélées à des événements paléogéographiques anciens. Enfin, le quatrième chapitre traite de la taxonomie, de la phylogénie et de la biogéographie d'un clade de niphargidés distribué dans la région des Alpes et des Carpates, et illustre un cas de discordance mitonucléaire dans la délimitation d'espèces vivant dans des zones affectées par les glaciations quaternaires. Une histoire complexe de divergence de lignées évolutives et de contacts secondaires pendant les fluctuations climatiques du Pléistocène explique la plus grande variabilité de l'ADN mitochondrial par rapport aux marqueurs nucléaires. Dans une telle situation, la description formelle d'espèces cryptiques basée sur le seul barcodage de l'ADN mitochondrial, comme dans certains articles récents sur les niphargidés, n'est pas recommandée. Cette thèse ouvre plusieurs perspectives pour des recherches futures basées sur la taxonomie intégrative et la modélisation biogéographique, permettant aux niphargidés très diversifiés de jouer un rôle majeur dans la surveillance des écosystèmes des eaux souterraines.
Doctorat en Sciences
info:eu-repo/semantics/nonPublished

Understanding the processes that generate diversity is key to interpreting the patterns we see in the present. New developments in modelling these processes have promised unprecedented prospects for unravelling the evolutionary past. However, the empirical behaviour of these models in many of their practical applications is not well understood. This thesis investigates the influence of diversification models in a variety of contexts. First, I consider the Generalised Mixed Yule-Coalescent (GMYC) method for molecular species delimitation. This method identifies transition points between species- and population-level diversification processes on a time-resolved evolutionary tree. I show that this method is sensitive to the choice of mitochondrial marker used, and that the best marker can vary widely across study groups. Next, I investigate the influence of diversification models used to place prior distributions on time-resolved trees in molecular dating. Specifically, I look at the influence of the tree prior in analysing data sets with multiple individuals per species. These data sets can arise by accident where species boundaries are not well understood, and violate the assumptions of both population- and species-level tree priors. I use simulation to show that molecular date estimates can be seriously affected by the choice of tree prior in some circumstances, but are remarkably robust in general. Finally, I extend the analysis of tree prior sensitivity to new methods for dating the origins of human language families. I show that these methods are also robust to the choice of tree prior, and that speciation priors are preferred for language data sets regardless of taxonomic scale. My work will contribute to an improved understanding of the role of diversification models in empirical studies and will increase confidence in these methods across multiple realms of enquiry.

Abstract Species are the fundamental units of biological diversity, but their identification and delimitation is often difficult. The difficulties are pronounced in diverse taxa such as insects. DNA barcodes, short standardized segments of the genome, have recently become a popular tool for identifying specimens to species, and are increasingly used as one of the sources of information for species delimitation. In this thesis, I studied the utility of DNA barcodes in species identification and delimitation in beetles (Coleoptera). Beetles are one of the most diverse animal groups, with nearly 400 000 known species. The Nordic beetle fauna is among the most thoroughly studied on the planet, providing excellent conditions for these studies. I also approached barcode sequences from a new angle, exploring amino acid variation and its connections to life history in a sample of the entire animal kingdom. I also studied variation and evolution at the amino acid level in large-scale samples of beetles and moths & butterflies (Lepidoptera). DNA barcodes proved to be a feasible tool for identifying species of Nordic beetles: depending on the criteria for successful identification, 95-98% of specimens could be identified to the species level based on DNA barcodes. Regardless of the delimitation method used, approximately 90% of the currently accepted species were perfectly recovered based on barcode data, and simple rules for forming consensus between delimitations improved the fit between species and barcode clusters even further. Several species that were split into two or more sequence clusters apparently include species new to science that have been previously overlooked. This conclusion is supported by preliminary morphological analysis. The study on amino acid variation revealed both a general pattern of structural conservation throughout the animal kingdom, and some interesting amino acid substitutions with potential to affect enzymatic function. Amino acid variation was more extensive in Coleoptera than in Lepidoptera, potentially due to differences in selection pressure and patterns of molecular evolution in the barcode region between the two orders
Tiivistelmä Laji on luonnon monimuotoisuuden perusyksikkö, mutta lajien tunnistaminen ja rajaaminen on usein vaikeaa. Vaikeudet korostuvat erityisesti hyvin monimuotoisissa eliöryhmissä kuten hyönteisissä. DNA-viivakoodit ovat lyhyitä standardoituja DNA-sekvenssejä, joiden käyttö lajien tunnistamisessa sekä yhtenä tiedon lähteenä lajien rajaamisessa on viime aikoina yleistynyt nopeasti. Tutkin väitöskirjatyössäni DNA-viivakoodien soveltuvuutta lajinmääritykseen ja lajien rajaamiseen kovakuoriaisilla. Kovakuoriaiset ovat yksi maailman lajirikkaimmista eliöryhmistä: lajeja on kuvattu lähes 400000. Pohjois-Euroopan lajisto tunnetaan koko maailman mittakaavassa poikkeuksellisen hyvin, mikä tarjoaa erinomaiset edellytykset tutkia DNA-viivakoodeihin liittyviä kysymyksiä kuoriaisilla. Tutkin DNA-viivakoodeja myös kokonaan uudesta näkökulmasta, selvittäen aminohappotason muuntelua koko eläinkunnan kattavassa otoksessa, sekä laajalla perhos- ja kuoriaisaineistolla. DNA-viivakoodit osoittautuivat erinomaiseksi työkaluksi lajinmääritykseen: riippuen onnistuneen määrityksen kriteereistä 95–98 % kuoriaislajeista voitiin tunnistaa luotettavasti viivakoodien perusteella. Käytetystä menetelmästä riippumatta noin 90 % nykykäsityksen mukaisista lajeista voitiin rajata viivakoodien perusteella oikein, ja soveltamalla yksinkertaisia konsensus-sääntöjä yhteensopivuus lajien ja viivakoodiklustereiden välillä kasvoi entisestään. Useat kuoriaislajit, jotka jakautuivat kahteen tai useampaan viivakoodiklusteriin, sisältävät alustavien morfologisten tutkimusten perusteella aiemmin huomaamatta jääneitä uusia lajeja. Aminohappo- ja proteiinitason tutkimus osoitti, että viivakoodijakson koodaaman proteiinin rakenne on yleisesti ottaen konservoitunut kautta eläinkunnan. Havaitsin kuitenkin myös useita kiinnostavia aminohappo-muutoksia, jotka saattavat vaikuttaa entsyymitoimintaan. Aminohapposekvenssi muuntelee kuoriaisilla paljon enemmän kuin perhosilla, mahdollisesti johtuen taksonien välisistä eroista molekyylievoluutiossa ja viivakoodisekvenssiin kohdistuvassa valintapaineessa

Molecular taxonomy is the science that classifies and identifies organism basing on molecular information. Its application to species-rich groups of organisms, as insects, could improve and accelerate taxa identification and delimitation. The efficiency and the limits of the methods for molecular taxonomy have to be properly evaluated for achieving unbiased results with this taxonomic approach. The main aims of this thesis are: i) to develop COI barcode libraries for Euro-Mediterranean Chrysomelidae identification and test DNA-barcoding efficiency in the identification of the species of the family; ii) to estimate group-specific nucleotide distance thresholds for the molecular identification of Chrysomelidae taxa and compare the error related to their use with the error related to the use of a general threshold value estimated for the whole family; iii) to apply molecular species delimitation methods to other insects taxa with the aim of resolving their taxonomic status in an integrative taxonomy framework; vi) to test the influence of factors intrinsic to the analysed data on molecular species delimitation efficiency. The results obtained confirmed the high efficiency of DNA-Barcoding as tool for Chrysomelidae molecular identification (94-99% efficiency) and the usefulness of molecular delimitation methods in integrative taxonomy for resolving taxonomic debated issues. The use of groups specific thresholds resulted to be related to a significantly lower identification error than the use of a general threshold for the whole Chrysomelidae, suggesting how the use of fixed thresholds for taxa identification is unwise. Finally, some factors, i.e. the species delimitation methods used, the geographic intraspecific distance among individuals collection localities and the difficulties in morphological identification of species, resulted to affect the efficiency of molecular species delimitation.

Chez les insectes phytophages, l'adaptation à la plante hôte peut correspondre à l'une des premières étapes de la spéciation. Dans ce contexte, ce manuscrit s'intéresse à Spodoptera frugiperda (Lepidoptera : Noctuidae), un ravageur responsable de dommages importants sur de nombreuses cultures en Amérique et dans les Caraïbes. Spodoptera frugiperda présente deux variants, l'un adapté au riz et l'autre au maïs, et constitue un modèle biologique pertinent pour étudier ce mécanisme. En effet, les deux variants sont morphologiquement identiques, mais ils sont génétiquement différenciés et présentent des différences écologiques et comportementales. L'ensemble de ces caractéristiques suggèrent la présence d'un isolement reproducteur entre les deux variants, qui pourraient alors correspondre à deux espèces différentes. Cependant la présence d'hybrides dans la nature ainsi que des résultats controversés sur le succès d'accouplement entre les deux variants, rendent ambiguë le statut d'espèce de S. frugiperda. L'objectif de ce travail est donc de mieux estimer, à travers une étude intégrative, le niveau de différenciation génétique présent entre les deux variants de S. frugiperda. Grâce à un premier niveau d'analyse, menées à partir de population naturelles, il a été possible de mettre en évidence un niveau de différenciation génétique élevé entre les deux variants, qui est compatible à celui attendu entre deux espèces. Le deuxième niveau d'étude réalisé à partir de population de laboratoire nous a permis de mettre en évidence la présence d'un isolement reproducteur entre les deux variants, se traduisant notamment par une distorsion de ségrégation méiotique des marqueurs moléculaires chez les hybrides. L'ensemble de ces résultats supportent l'hypothèse selon laquelle le variant riz et maïs de Spodoptera frugiperda seraient des entités situées à une étape avancée dans le continuum de la spéciation
In phytophagous insects, adaptation to host plant could be the first step leading to speciation. In this context, this document focus on moth Spodoptera frugiperda (Lepidoptera: Noctuidae), a pest responsible for serious damages in several crops in the Western hemisphere. Spodoptera frugiperda consists of two host-plant strains, one adapted to rice and the other adapted to maize, which made it a relevant model to study this mechanism. Though the two variants are morphologically identical, they are nonetheless genetically distinguishable and present some ecological and behavioral differences. The species status of S. frugiperda is also highly controversial because hybrids naturally occur in the wild, not to mention the fact of the discrepancies among published results concerning mating success between the two strains. The aim of this thesis is thus to better estimate, through an integrative approach, the level of genetic differentiation between the two variants of S. frugiperda. Starting with natural populations, various phylogenetic methods allowed us to highlight a high level of genetic differentiation between the two variants, compatible with what is expected between distinct species. Furthermore, studies on laboratory populations, including crossing experiments, showed a significant unidirectional bias in inter-strain mating success and the presence of meiotic segregation distortion of molecular markers in hybrid progenies. These results as a whole support the assumption that the two strains of S. frugiperda are well-advanced in the continuum of speciation

Thesis (Ph.D.)--Indiana University, Dept. of Biology, 2009.
Title from PDF t.p. (viewed on Feb. 5, 2010). Source: Dissertation Abstracts International, Volume: 70-04, Section: B, page: 2009. Adviser: Loren H. Rieseberg.

Until relatively recently, many groups of terrestrial reptiles were considered to contain only few species vastly distributed across the Arabian Peninsula. This was mostly the result of the apparent morphological uniformity of these groups in combination with their geographic ranges, often overlapping with politically unstable regions in which scientific research has been hampered for centuries. However, our understanding of the diversity inhabiting these unexplored areas changed with the onset of molecular phylogenetic methods that became extensively available and routinely applied to tackle a broad set of evolutionary questions. This thesis focuses on resolving the systematics, biogeography and evolution of four selected groups of reptiles from Arabia, using an unprecedented sampling of over 1,000 tissues for genetic analyses and more than 900 voucher specimens. For some taxa, we inferred time-calibrated phylogenies, and we reconstructed their biogeographical history and their morphological evolution. For many others, we unexpectedly discovered high levels of genetic diversity and several lineages potentially representing unknown species. In the latter case, testing different species delimitation methods led to more detailed examination of the genetic, morphological and ecological data in order to resolve the taxonomy of these candidate species. As a result of that, four new species of geckos and two lacertid lizards have been formally described as part of this thesis. Essentially, most of the new species were found in the mountain regions of Arabia, stressing the importance of these poorly studied areas as recognized hotspots in terms of reptile diversity and endemicity.
Fins fa relativament poc, es considerava que la majoria de grups de rèptils terrestres comprenien només unes poques espècies àmpliament distribuïdes per tota la Península Aràbiga. Aquesta percepció es devia, en gran part, a l’aparent uniformitat morfològica d’aquests grups i les seves àrees de distribució geogràfica, sovint coincidents amb regions políticament inestables on la investigació científica s’ha vist limitada durant segles. No obstant, el coneixement que teníem sobre aquestes àrees poc explorades va canviar amb l’aparició de les filogènies moleculars, aviat àmpliament disponibles i aplicades de manera rutinària per abordar un nou ventall de qüestions evolutives. Aquesta tesi doctoral es centra en resoldre la sistemàtica, la biogeografia i l’evolució de quatre grups concrets de rèptils, utilitzant un mostreig sense precedents que inclou més de 1.000 teixits analitzats genèticament i més de 900 espècimens. Per alguns tàxons, hem inferit filogènies calibrades tot reconstruint la seva història biogeogràfica i la seva evolució morfològica. Per a d’altres, hem descobert inesperadament alts nivells de diversitat genètica i diversos llinatges que podrien representar espècies desconegudes. En el darrer cas, l’ús de diferents mètodes de delimitació d’espècies va conduir a examinar més detalladament les dades genètiques, morfològiques i ecològiques per tal de resoldre la taxonomia d’aquestes espècies candidates. Com a resultat, quatre espècies de dragons i dues sargantanes de la família dels lacèrtids han estat formalment descrites en el decurs d’aquesta tesi. Donat que la majoria d’aquestes espècies habiten a diferents zones muntanyoses d’Aràbia, aquesta tesi destaca la importància d’aquestes regions com a punts calents de diversitat biològica i endemicitat pel que fa a la seva fauna reptiliana.

Collembola (=springtails) is one of the most abundant, widespread and ancient lineages of basal hexapods. During their long evolutionary history, springtails have adapted to most damp environments on Earth, including those of South Pole. Antarctic springtails are endemic to the frozen Continent and among the few invertebrate taxa adapted to its strictly terrestrial ecosystem. These species have evolved when Antarctica was still linked to the Gondwanaland at lower latitudes and have adapted and survived to the cooling, isolation and southwards migration of the landmass. Antarctic springtails’ habitats are restricted to the few coastal areas, seasonally ice-free and accounting for less than the 0.5% of the entire continental area and off-shore islands. The niche fragmentation, together with springtails poor dispersal capability (due to the primary absence of wings), entail a severe degree of isolation among populations, with very low levels of gene flow. The Antarctic springtail species composition is limited without overlap among the two main Antarctic bioregions (i.e., the maritime and the continental Antarctica), with Friesea antarctica being the only species found both in the Antarctic Peninsula and Victoria Land (continental Antarctica). The high levels of endemism and fragmentation among populations, as well as the low invertebrate biodiversity and the complex and delicate array of physiological adaptation these species evolved, make Antarctic taxa particularly susceptible to anthropogenic climate changes, that we are all experiencing since the second industrial revolution in the XIXth century. In this respect, studying the molecular mechanisms underlying springtail adaptation to such a harsh environment, as well as the genetic structure of the populations and the way in which specimens may have been and can be influenced by the Antarctic terrestrial environment, may greatly assist the development of adequate and biogeographically-specific (thus, effective) conservational plans. In order to address these issues, different studies have been carried out during the current PhD project. A genetics of population study was performed to investigate the genetic structure of the Antarctic springtail species Cryptopygus terranovus. As previously observed in other Antarctic species (i.e, F. antarctica), high levels of genetic divergence were detected, with very few haplotypes shared among populations nearly suggesting the absence of gene flow, as well as the presence of cryptic species. One way to address this issue is the integration of morphological, molecular and biogeographic data to assess whether the detection of cryptic species is due to an ongoing phylogenetic niche conservatism process or to overlooked morphological differences. In this respect, an integrative taxonomic analysis has been carried out on the Antarctic springtail F. antarctica. Nuclear and mitochondrial markers were used in bioinformatic analyses of species delimitation. Although applied tools rely on different algorithms and biological assumptions, and the chosen molecular markers are generally subject to different evolutionary pressures, the results obtained in this study would suggest that at least two more species may be hidden within the F. antarctica complex. Specimens from the same localities were also morphologically re-described so that new species could be possibly established. These analyses would suggest an even higher species richness of the Antarctic terrestrial ecosystem, that should be taken into account when developing conservational plans. Our ability to safeguard Antarctica relies also on our in-depth understanding of the terrestrial ecosystem functioning and dynamics. In this perspective, an initial descriptive analysis of the microbial communities associated to four Antarctic springtail species was performed (specifically on: Cryptopygus antarcticus antarcticus and F. antarctica from the Antarctic Peninsula and C. terranovus and F. antarctica collected along Victoria Land). The results obtained are in line with previous studies on Collembola microbiomes. In addition, the occurrence of particularly interesting OTUs, such as those of the genera Streptomyces and Bacillus, was, to my knowledge, firstly detected among springtails and may break new grounds for biotechnology development, especially starting from such an unspoiled ecosystem, like the Antarctic one. Finally, the mitochondrial genomes from 13 springtail species, both living at low and high latitudes, have been applied to maximum likelihood analyses of positive selection in order to investigate whether or not the organelle chromosome may have been involved in Antarctic springtails adaptation to such an extreme environment. The results pointed out that some mitochondrial genes involved in the oxidative phosphorylation process may have been under positive selection, thus suggesting the development of additional thermoregulatory mechanism within the mitochondrion, complementary to the well-known cold hardiness strategies.

Orientadores: Karina Lucas da Silva Brandão, André Victor Lucci Freitas
Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia
Made available in DSpace on 2018-08-19T17:32:03Z (GMT). No. of bitstreams: 1 Seraphim_Noemy_M.pdf: 28481674 bytes, checksum: 287c682142f5d5cd09f6635fbb8a291f (MD5) Previous issue date: 2011
Resumo: O gênero Hermeuptychia Forster (Nymphalidae, Satyrinae, Euptychiina) está amplamente distribuído no continente Americano, desde a Argentina até o sul dos Estados Unidos. O gênero foi anteriormente considerado um complexo de espécies, e atualmente são reconhecidas oito espécies . Todas as espécies possuem um padrão alar muito parecido, o que compromete a identificação taxonômica correta. Em adição, a posição filogenética do gênero dentro da subtribo Euptychiina permanece incerta. Para o presente estudo foram obtidos espécimens de 45 localidades de cinco países, com maior ênfase em uma amostragem no Brasil. Três marcadores moleculares, dois do DNA mitocondrial (cox1 5' e nad6) e um do DNA nuclear (RpS5) foram utilizados para gerar hipóteses filogenéticas (Máxima Parcimônia e Inferência Bayesiana), para delimitar espécies, e para gerar estimativas de tempo de divergência e distribuição ancestral. Adicionalmente, o desempenho da região anterior da cox1 como 'barcode - código de barras' para delimitar as espécies de Hermeuptychia foi testado. Além disso, análise morfológica da genitália masculina foi empregada para a delimitação e identificação de espécies. Os indivíduos amostrados agruparam-se em dez clados nas análises moleculares, correspondendo a sete espécies reconhecidas mais H. gisella, que havia sido anteriormente sinonimizada com H. cucullina. A análise morfológica dos indivíduos possibilitou o estabelecimento de caracteres diagnósticos para todas as espécies de Hermeuptychia - incluindo H. cucullina, que não está presente nas análises moleculares - e concordou com os agrupamentos obtidos através das análises moleculares. As relações filogenéticas entre as espécies de Hermeuptychia permanecem incertas, possivelmente devido a um padrão de evolução rápida, descrito anteriormente para outros Satyrini. Entretanto, dois grupos de espécies-irmãs podem ser identificados, H. pimpla + H. harmonia, e H. gisella + H. fallax, ambos sustentados por ocorrência simpátrica. Em adição, H. gisella e H. fallax parecem apresentar um isolamento reprodutivo incompleto, com formação de híbridos. Algumas espécies de Hermeuptychia estão distribuídas amplamente, como H. atalanta, H. hermes e H. gisella, sendo que H. atalanta é a espécie mais comum encontrada no Brasil. H. fallax é uma espécie restrita a Mata Atlântica; H. pimpla e H. harmonia são espécies restritas à região andina, encontradas em altitudes moderadas; H. maimoune pode ser encontrada na região andina e no sul da Amazônia brasileira, correspondendo a duas espécies crípticas; H. cucullina foi encontrada no centro-oeste brasileiro e na região andina, e é a espécie mais rara de Hermeuptychia; e H. sosybius pode ser encontrada do sul dos Estados Unidos até a região norte da Colômbia. O gênero diversificou-se de seu grupo-irmão a cerca de 8,2 milhões de anos (mya), a diversificação das espécies ocorreu entre 3,5 e 1,4 mya, e a distribuição ancestral estimada é a cordilheira dos Andes. Apenas com a região 'barcode' e análise de distância usando Neighbor-Joining, foi possível separar as espécies de Hermeuptychia com uma taxa de 2% de erro. O limite entre as distâncias intra e interespecíficas estimado fica em torno de 2% de divergência genética
Abstract: The Hermeuptychia genus Forster (Nymphalidae: Satyrinae: Euptychiina) is widely distributed in the American continent, from Argentina to South United States. Previously considered a species complex, the genus presents eight valid species taxa at the moment. Wing pattern is very similar in all Hermeuptychia species resulting in difficult and prone to error taxonomic identification. Additionally its position within the subtribe Euptichiina remains uncertain. Samples from 45 locations in five countries, with major emphasis in Brazilian territory sampling, were obtained for the present study. Three molecular markers, two from mitochondrial DNA (cox1 5' and nad6) and one from nuclear DNA (RpS5), were used to generate phylogenetic hypothesis (Maximum Parsimony and Bayesian Inference), to delimit species, and to estimate divergence time and ancestral distributions. The 'barcode' region performance (cox1 5') was tested for Hermeuptychia species. Male genitalia morphology was also used to identify and delimitate species. Sampled individuals are grouped in ten molecular clusters, corresponding to seven valid species and H. gisella, previously synonymized to H. cucullina. Morphological analysis of individuals revealed morphological diagnose traits to identify all Hermeuptychia species, including H. cucullina, which is not present in the molecular analysis, and was congruent with molecular analysis. Phylogenetic relationships among Hermeuptychia species remain unresolved due to a possible rapid evolutionary pattern common to Satyrini. However, two pairs of sister species could be identified: H. pimpla + H. harmonia and H. gisella + H. fallax, both sympatric. Additionally, H. gisella + H. fallax present incomplete reproductive isolation, with hybrids. Some Hermeuptychia are widely distributed as H. atalanta, H. hermes, and H. gisella, and H. atalanta is the most common species found in Brazil. H. fallax is restricted to Atlantic Forest; H. pimpla and H. harmonia are restricted to Andes region, at moderately high altitudes; H. maimoune is found in the Andine and in the Amazonian regions, corresponding to two cryptic species; H. cucullina is the rarest Hermeuptychia and was found in Central Brazil and in Andes; and H. sosybius is the only Hermeuptychia found in North America, been present from South United States to north Colombia. The genus diverged from its sister group around 8.2 my, species diversification occurred between 3.5 and 1.4 my and the ancestral estimate distribution is Andine region. Only the 'barcode' region was able to identify each Hermeuptychia species, with an 2% error rate and the molecular threshold for intra and interspecific distance was around 2% of genetic divergence
Mestrado
Ecologia
Mestre em Ecologia

Genomes provide windows into the evolutionary histories of species. The recent accessibility of genome-scale data in non-model organisms and the proliferation of powerful statistical models are now providing unprecedented opportunities to uncover evolutionary relationships and to test hypotheses about the processes that generate and maintain biodiversity. This dissertation work reveals shallow-scale species boundaries and population genetic structure in two imperiled groups of salamanders and demonstrates that the number and information content of genomic regions used in species delimitation exert strong effects on the resulting inferences. Genome scans are employed to test hypotheses about the mechanisms of genetic sex determination in cryptobranchid salamanders, suggesting a conserved system of female heterogamety in this group. At much deeper scales, phylogenetic analyses of hundreds of protein-coding genes across all major amphibian lineages are employed to reveal the backbone topology and evolutionary timescales of the amphibian tree of life, suggesting a new set of hypotheses for relationships among extant amphibians. Yet, genomic data on their own are no panacea for the thorniest questions in evolutionary biology, and this work also demonstrates the power of a model testing framework to dissect support for different phylogenetic and population genetic hypotheses across different regions of the genome.

Emplectonema gracile (Johnston 1837) is a hoplonemertean of marine intertidal hard-bottom communities and is distributed throughout the Northern Hemisphere. Although possessing a planktonic larval stage in its life history, the range of such cosmopolitan marine invertebrate species is often explained by cryptic speciation and anthropogenic transport. The purpose of this study is to test for possible cryptic species using mtDNA markers (COI and 16S rDNA) and to investigate population structure in E. gracile over a portion of its geographic range using mtDNA markers and ddRADseq nuclear SNP data. The results of both phylogenetic- and tree-based species delimitation revealed that E. gracileis a morphotype containing cryptic species. Three North Atlantic and one Pacific coast population are inferred as one species (E. gracile sensu stricto) and two Pacific coast populations (Akkeshi, Japan and Charleston, Oregon) are inferred as another species (Emplectonemasp 1), strongly confirming an earlier study and extending the range of the latter species to the Pacific coast of Japan. Anthropogenic transport is suggested as the likely mode of transport for E. gracile.Both Fst, PCA and haplotype network analyses suggest a lack of differentiation between E. gracile populations separated by large geographic distances.In contrast corresponding analyses forEmplectonemasp. 1 indicate differentiation between the two populations sampled. Further research will be necessary to reveal if rare anthropogenic transport or natural dispersal (larval transport, rafting) between geographically adjacent yet to be delimitedE. gracile morphotype populations is responsible for its seemingly disjunct distribution.