Person: Giribet, Gonzalo
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Publication A revised dated phylogeny of the arachnid order Opiliones
(Frontiers Media S.A., 2014) Sharma, Prashant P.; Giribet, GonzaloDating the Opiliones tree of life has become an important enterprise for this group of arthropods, due to their ancient origins and important biogeographic implications. To incorporate both methodological innovations in molecular dating as well as new systematic discoveries of harvestman diversity, we conducted total evidence dating on a data set uniting morphological and/or molecular sequence data for 47 Opiliones species, including all four well-known Palaeozoic fossils, to test the placement of both fossils and newly discovered lineages in a single analysis. Furthermore, we investigated node dating with a phylogenomic data set of 24,202 amino acid sites for 14 species of Opiliones, sampling all extant suborders. In this way, we approached molecular dating of basal harvestman phylogeny using different data sets and approaches to assess congruence of divergence time estimates. In spite of the markedly different composition of data sets, our results show congruence across all analyses for age estimates of basal nodes that are well constrained with respect to fossil calibrations (e.g., Opiliones, Palpatores). By contrast, derived nodes that lack fossil calibrations (e.g., the suborders Cyphophthalmi, and Laniatores) have large uncertainty intervals in diversification times, particularly in the total evidence dating analysis, reflecting the dearth of calibration points and undersampling of derived lineages. Total evidence dating consistently produced older median ages than node dating for ingroup nodes, due to the nested placement of multiple Palaeozoic fossils. Our analyses support basal diversification of Opiliones in the Ordovician-Devonian period, corroborating the inferred ancient origins of this arthropod order, and underscore the importance of diversity discovery—both paleontological and neontological—in evolutionary inference.
Publication The Linguistic Problem of Morphology: Structure Versus Homology and the Standardization of Morphological Data
(Wiley-Blackwell, 2010) Vogt, Lars; Bartolomaeus, Thomas; Giribet, GonzaloThe present article discusses the need for standardization in morphology in order to increase comparability and communicability of morphological data. We analyse why only morphological descriptions and not character matrices represent morphological data and why morphological terminology must be free of homology assumptions. We discuss why images only support and substantiate data but are not data themselves. By comparing morphological traits and DNA sequence data we reveal fundamental conceptual shortcomings of the former that result from their high average degree of individuality. We argue that the delimitation of morphological units, of datum units, and of evidence units must be distinguished, each of which involves its own specific problems. We conclude that morphology suffers from the linguistic problem of morphology that results from the lack of (i) a commonly accepted standardized morphological terminology, (ii) a commonly accepted standardized and formalized method of description, and (iii) a rationale for the delimitation of morphological traits. Although this is not problematic for standardizing metadata, it hinders standardizing morphological data. We provide the foundation for a solution to the linguistic problem of morphology, which is based on a morphological structure concept. We argue that this structure concept can be represented with knowledge representation languages such as the resource description framework (RDF) and that it can be applied for morphological descriptions. We conclude with a discussion of how online databases can improve morphological data documentation and how a controlled and formalized morphological vocabulary, i.e. a morphological RDF ontology, if it is based on a structure concept, can provide a possible solution to the linguistic problem of morphology.
Publication Canga Renatae, A New Genus and Species of Cyphophthalmi from Brazilian Amazon Caves (Opiliones: Neogoveidae)
(Magnolia Press, 2010) DaSilva, Marcio Bernardino; Pinto-da-Rocha, Ricardo; Giribet, GonzaloA new genus and species of Cyphophthalmi, Canga renatae gen. nov., sp. nov., is described in the family Neogoveidae from a system of caves in the Serra de Carajás, Pará State, Brazil. Canga can be easily distinguished from other neogoveid genera by the presence of a dentate claw on leg I, a unique character among known cyphophthalmid species, and by the free coxa II, which is fused to coxae III and IV in all the other neogoveid species except for the North American Metasiro. The new genus also differs from other Neotropical neogoveids in the lack of a dorsal crest on the chelicerae and in the lack of opisthosomal glands. The finding of a neogoveid in the Pará State greatly increases the known distribution of South American cyphophtalmids into the Eastern Brazilian Amazon forest.
Publication Including Secondary Structure, Fossils and Molecular Dating in the Centipede Tree of Life
(Elsevier BV, 2010) Murienne, Jerome; Edgecombe, Gregory; Giribet, GonzaloA well-corroborated morphological scheme of interrelationships for centipedes, once broadly accepted, has been in conflict with molecular data with respect to deep branching events. Expanded taxonomic coverage compared to previous analyses adds longer fragments for 28S rRNA and a structural alignment as part of a sample of four genes (two nuclear ribosomal and two mitochondrial) for 111 extant species; these sequence data are combined with morphology under parsimony and maximum likelihood, exploring both traditional multiple sequence alignment and direct optimization approaches. Novel automated procedures to incorporate secondary structure information are also explored. The molecular data in combination yield trees that are highly congruent with morphology as regards the monophyly of all centipede orders as well as the major groups within each of the large orders. Regardless of the optimality criterion or alignment strategy, the Tasmanian/New Zealand Craterostigmomorpha is resolved in a different position by the molecular data than by morphology. Addition of morphology overturns the placement of Craterostigmomorpha in favour of the traditional morphological resolution and eliminates the need to posit major character reversals with respect to developmental mode and maternal care. Calibration of the tree with Palaeozoic and Mesozoic fossils for a relaxed clock analysis corroborates the palaeontological signal that divergences between centipede orders date to the Silurian and earliest Devonian, and familial divergences are likewise almost wholly Palaeozoic.
Publication A molecular palaeobiological exploration of arthropod terrestrialization
(The Royal Society, 2016) Lozano-Fernandez, Jesus; Carton, Robert; Tanner, Alastair R.; Puttick, Mark N.; Blaxter, Mark; Vinther, Jakob; Olesen, Jørgen; Giribet, Gonzalo; Edgecombe, Gregory D.; Pisani, DavideUnderstanding animal terrestrialization, the process through which animals colonized the land, is crucial to clarify extant biodiversity and biological adaptation. Arthropoda (insects, spiders, centipedes and their allies) represent the largest majority of terrestrial biodiversity. Here we implemented a molecular palaeobiological approach, merging molecular and fossil evidence, to elucidate the deepest history of the terrestrial arthropods. We focused on the three independent, Palaeozoic arthropod terrestrialization events (those of Myriapoda, Hexapoda and Arachnida) and showed that a marine route to the colonization of land is the most likely scenario. Molecular clock analyses confirmed an origin for the three terrestrial lineages bracketed between the Cambrian and the Silurian. While molecular divergence times for Arachnida are consistent with the fossil record, Myriapoda are inferred to have colonized land earlier, substantially predating trace or body fossil evidence. An estimated origin of myriapods by the Early Cambrian precedes the appearance of embryophytes and perhaps even terrestrial fungi, raising the possibility that terrestrialization had independent origins in crown-group myriapod lineages, consistent with morphological arguments for convergence in tracheal systems. This article is part of the themed issue ‘Dating species divergences using rocks and clocks’.
Publication Cementing mussels to oysters in the pteriomorphian tree: a phylogenomic approach
(The Royal Society, 2016) Lemer, Sarah; González, Vanessa L.; Bieler, Rüdiger; Giribet, GonzaloMussels (Mytilida) are a group of bivalves with ancient origins and some of the most important commercial shellfish worldwide. Mytilida consists of approximately 400 species found in various littoral and deep-sea environments, and are part of the higher clade Pteriomorphia, but their exact position within the group has been unstable. The multiple adaptive radiations that occurred within Pteriomorphia have rendered phylogenetic classifications difficult and uncertainty remains regarding the relationships among most families. To address this phylogenetic uncertainty, novel transcriptomic data were generated to include all five orders of Pteriomorphia. Our results, derived from complex analyses of large datasets from 41 transcriptomes and evaluating possible pitfalls affecting phylogenetic reconstruction (matrix occupancy, heterogeneity, evolutionary rates, evolutionary models), consistently recover a well-supported phylogeny of Pteriomorphia, with the only exception of the most complete but smallest data matrix (Matrix 3: 51 genes, 90% gene occupancy). Maximum-likelihood and Bayesian mixture model analyses retrieve strong support for: (i) the monophyly of Pteriomorphia, (ii) Mytilida as a sister group to Ostreida, and (iii) Arcida as sister group to all other pteriomorphians. The basal position of Arcida is congruent with its shell microstructure (solely composed of aragonitic crystals), whereas Mytilida and Ostreida display a combination of a calcitic outer layer with an aragonitic inner layer composed of nacre tablets, the latter being secondarily lost in Ostreoidea.
Publication Species limits and phylogeography of Newportia (Scolopendromorpha) and implications for widespread morphospecies
(Pensoft Publishers, 2015) Edgecombe, Gregory D.; Vahtera, Varpu; Giribet, Gonzalo; Kaunisto, PipsaAbstract The genus Newportia Gervais, 1847, includes some 60 nominal species distributed in the Caribbean islands and from Mexico to central South America. Modern keys to species and subspecies are available, greatly facilitating identification, but some species are based on few specimens and have incomplete documentation of taxonomically-informative characters. In order to explore genetic variability and evolutionary relationships within geographically-widespread morphospecies, specimens of Newportia (Newportia) stolli (Pocock, 1896) and Newportia (Newportia) divergens Chamberlin, 1922, two nominal species distinguished principally by differences in suture patterns on T1, were sequenced for mitochondrial 16S rRNA and cytochrome c oxidase subunit I (COI) genes from populations in southern Mexico, Guatemala, Honduras and Brazil. Newportia (Newportia) stolli is paraphyletic with respect to Newportia (Newportia) divergens within a clade from Guatemala, Honduras, and Chiapas (Mexico), most trees being consistent with a single loss of a connection between the anterior transverse suture on T1, whereas specimens of “Newportia (Newportia) stolli” from Brazil are not closely allied to those from the Mesomerican type area. The widespread morphospecies Newportia (Newportia) monticola Pocock, 1890, was sequenced for the same loci from populations in Costa Rica, Colombia and Brazil, finding that specimens from these areas do not unite as a monophyletic group. Samples of Newportia (Newportia) oreina Chamberlin, 1915, from different regions of Mexico form geographic clusters that resolve as each other’s closest relatives. These results suggest that some widespread species of Newportia may be taxa of convenience more so than natural groupings. In several cases geographic proximity fits the phylogeny better than taxonomy, suggesting that non-monophyletic species do not result from use of inappropriate molecular markers. Molecular identification is possible for specimens missing taxonomically informative morphological characters, notably damaged specimens that lack the ultimate leg pair, a protocol that may also apply to other taxonomically difficult genera that are prone to damage (such as Cryptops).
Publication A Phylogeny of Vetigastropoda and Other "Archaeogastropods": Re-Organizing Old Gastropod Clades
(Wiley-Blackwell, 2010) Aktipis, Stephanie W.; Giribet, GonzaloThe phylogenetic relationships among the “archaeogastropod” clades Patellogastropoda, Vetigastropoda, Neritimorpha, and Neomphalina are uncertain; the phylogenetic placement of these clades varies across different analyses, and particularly among those using morphological characteristics and those relying on molecular data. This study explores the relationships among these groups using a combined analysis with seven molecular loci (18S rRNA, 28S rRNA, histone H3, 16S rRNA, cytochrome c oxidase subunit I [COI], myosin heavy-chain type II, and elongation factor-1α [EF-1α]) sequenced for 31 ingroup taxa and eight outgroup taxa. The deep evolutionary splits among these groups have made resolution of stable relationships difficult, and so EF-1α and myosin are used in an attempt to re-examine these ancient radiation events. Three phylogenetic analyses were performed utilizing all seven genes: a single-step direct optimization analysis using parsimony, and two-step approaches using parsimony and maximum likelihood. A single-step direct optimization parsimony analysis was also performed using only five molecular loci (18S rRNA, 28S rRNA, histone H3, 16S rRNA, and COI) in order to determine the utility of EF-1α and myosin in resolving deep relationships. In the likelihood and POY optimal phylogenetic analyses, Gastropoda, Caenogastropoda, Neritimorpha, Neomphalina, and Patellogastropoda were monophyletic. Additionally, Neomphalina and Pleurotomariidae fell outside the remaining vetigastropods, indicating the need for further investigation into the relationship of these groups with other gastropods.
Publication Report of a Cohesive Gelatinous Egg Mass Produced by a Tropical Marine Bivalve
(Wiley-Blackwell, 2010) Collin, Rachel; Giribet, GonzaloGelatinous egg masses are common in a number of animal phyla. However, they are virtually unknown in marine bivalves, with structures that could be thought of as gelatinous egg masses being reported for only five species. We describe the gelatinous egg mass and intracapsular development in the tropical lucinid Phacoides pectinatus. The embryos developed within individual capsules embedded in a large flimsy, spherical mass. Swimming veligers hatch at 198 μm shell length. They did not feed, settled within several days of hatching, and metamorphosis was completed within 2 weeks of hatching. Gelatinous egg masses might be detected in members of more lucinid species if studies of development included field or in vivo observations of reproduction in addition to producing embryos by stripping the gonads.
Publication 'Moa’s Ark' or 'Goodbye Gondwana': Is the Origin of New Zealand’s Terrestrial Invertebrate Fauna Ancient, Recent or Both?
(CSIRO Publishing, 2010) Giribet, Gonzalo; Boyer, Sarah L.Evaluating the hypothesis of New Zealand’s total submersion during the Oligocene requires the strictest tests, including sound phylogenetic data and dating of phylogenies. Although New Zealand has been traditionally considered to host ancient biota that originated by vicariance after it separated from Australia ~80 Mya, the ancient origins of its biota have been recently questioned, with some authors even suggesting that all current land organisms had to arrive to the islands after it re-emerged from the ocean 22 million years ago. Here we examine examples of short-range endemic soil-dwelling invertebrates and find compelling evidence that at least some of them are the result of old lineages that diversified in New Zealand before the hypothesised submersion event 22 million year ago. We conclude that New Zealand indeed has old lineages as well as recently diversified lineages and compare this situation with that of other more stable areas of the Neotropics.