Filogenética E a Evolução Da Morfológica Da Família Cosmetidae Koch, 1839 (Arachnida: Opiliones): O Curioso Caso De Cosmetidae

1 Capa Brittany Nicole Damron Filogenética e a Evolução da Morfológica da família Cosmetidae Koch, 1839 (Arachnida: Opiliones): O Curioso Caso de Cosmetidae. Phylogenetics and morphology evolution of the family Cosmetidae Koch, 1839 (Arachnida: Opiliones): The Curious Case of Cosmetidae. São Paulo 2020 2 X. Resumo A família Cosmetidae (Arachnida: Opiliones) é a família de Opiliones mais diversificada da América Central, e depois da família Gonyleptidae nos Neotrópicos. Continuam sendo um grupo pouco estudado e se beneficiam de qualquer trabalho sistemático. Esta pesquisa de PhD é uma tentativa de entender as relações dentro da família. Foram utilizados múltiplos critérios de otimalidade, evidência total usando otimização direta em POY e máxima verossimilhança em RAxML, e molecular apenas usando Bayesiano em BEAST, datação por divergência foram calculadas em BEAST. Foram utilizados cinco marcadores (28S rRNA, 12S rRNA, 16S rRNA, COI e H3) e 95 caracteres morfológicos (i.e. caracteres genitais, contorno do escuto dorsal, armadura do escuto dorsal) para determinar os clados. As análises demonstram que as subfamílias atuais (Discosomaticinae e Cosmetinae) não são válidas e a subfamília Cosmetinae é um grupo parafilético. Enquanto parte da monofilia de Taito Metalibitia Ferkeria gêneros está implícita ( , , ), outros gêneros Cynorta Paecileama diagnosticados na literatura recente não são ( , , Flirtea ). Foi demonstrado aqui que o alcance geográfico e, portanto, o histórico compartilhado são uma característica importante a ser considerada na definição de clados. Uma nova subfamília é proposta aqui, apoiada por hipóteses filogenéticas e morfologia. Análises preliminares de datação por divergência mostram que a família tem mais de 10 my a mais do que o inicialmente estimado, com uma idade de 47±5 my. Novos diagnósticos de espécies, gêneros e subfamílias da família devem se concentrar nos caracteres da genitália e na região biogeográfica das espécies nos grupos. 3 XI. Abstract The family Cosmetidae (Arachnida: Opiliones) is the most diverse harvestmen family in Central America, second to the family Gonyleptidae in the Neotropics. They remain a poorly studied group, and benefit from any systematic work. This PhD research is an attempt at understanding the relationships within the family Cosmetidae. Multiple optimality criteria were used, total evidence using direct optimization in POY and Maximum Likelihood in RAxML, and molecular only using Bayesian in BEAST, divergence dating were also calculated in BEAST. Five molecular markers were used (28S rRNA, 12S rRNA, 16S rRNA, COI, and H3), and 95 morphological characters (i.e. genitalia characters, dorsal scutum outline, dorsal scutum armature) to determine clades. Analyses demonstrate that the current subfamilies of Cosmetidae (Discosomaticinae and Cosmetinae) are not valid, and the subfamily Cosmetinae is a paraphyletic group. While some monophyly of genera is Taito Metalibitia Ferkeria implied in this analysis (see , , and ), other Cynorta Paecileama genera rediagnosed in recent literature are not ( , , and Flirtea ). It was demonstrated in these analyses that geographic range and therefore shared history is an important characteristic to consider when defining clades. A new subfamily is proposed here, supported by phylogenetic hypotheses and morphology. Preliminary divergence dating analyses shows that the family Cosmetidae is more than 10 million years older than originally estimated, with an age of approximately 47±5 myr. New diagnoses of species, genera, and subfamilies within the family Cosmetidae should focus on genitalia characters, and biogeographic region of species in the groups. 4 Introdução A. Phylogenetic and Systematic History of the family Cosmetidae To-date no work published has done any substantial investigation of the relationships within Cosmetidae. The few cosmetid species that have had molecular data collected were simply used as an outgroup in other analyses (Pinto-da-Rocha et al., 2014; Bragagnolo, Hara & Pinto-da-Rocha, 2015), or as exemplars in a larger analysis at order level (Sharma & Giribet, 2011; 2014). This is unlike the family Gonyleptidae, which has had multiple phylogenetic analyses, one for the entire family (Pinto-da-Rocha et al., 2014), and many examining the relationships within subfamilies (e.g, DaSilva & Pinto-da-Rocha, 2010; DaSilva & Gnaspini, 2010; Mendes, 2011; Bragagnolo & Pinto-da-Rocha, 2012). The relationships in Gonyleptidae have continued to evolve as more molecular data is collected and further analyses are completed (e.g. Pinto-da-Rocha, et al., 2014, Peres et al., 2019), and while many advances have been made in understanding the systematic relationships in Gonyleptidae, there are still groups within the family in need of revision (see Introduction in Pinto-da-Rocha et al., 2014). Cosmetidae has not received the same level of interest beyond some cladistic analyses completed with only morphological characters, but these have focused mostly on the subfamily Discosomaticinae (Medrano & Kury, 2018) or a single genus with a limited number of species from across the family’s richness (Coronato-Ribeiro & Pinto-da-Rocha, 2017). Given that many characters historically used to define clades have been shown to be highly homoplastic and therefore problematic (Ringuelet, 1959) and these analyses being limited in scope, no conclusions about 5 relationships within the Cosmetidae in its entirety can be made with confidence. Homoplasy is especially evident in characters such as number of tarsal segments on legs, the armature of the DS, and pectination on claws III and IV. As mentioned, this is especially evident for the subfamily Discosomaticinae, with a single synapomorphy of pectinate tarsal claws on legs III and IV. To date it has been stated that this character had arisen in various forms, but this assertion has not been tested across the diversity of the family, only within a small proportion of species (Medrano & Kury, 2018). The morphology of the Cosmetidae has in the past been considered conserved (Kury and Pinto-da-Rocha, 2007a) raising concerns regarding conducting strictly morphological analyses, especially when morphological characters may have a high level of homoplasy. This is only problematic due to the conserved patterns in external morphology in a single lineage, genus or species group, though perhaps not the case when the entirety of the family is taken into consideration. A relatively small number of characters, and the continued use of historic (and problematic) characters have led to hypotheses of relatedness that may not be supported by more robust and modern morphological characters and molecular data. Analyses to date have only addressed a very small proportion of the family’s total species richness (~3%; Medrano & Kury, 2018), which has not allowed characters that have been proposed as diagnostic features to be tested as valid synapomorphies or diagnostic features across the richness of the family. Given a well sampled phylogeny the evolutionary history of Cosmetidae can be explored for the first time, and morphology in the context of a phylogeny can be examined to attempt a subfamilial classification. 6 B. Use of Morphology in Phylogenetic Analyses and the Evolution of Morphology in the Family Cosmetidae. Methods for collecting molecular data has advanced significantly, with difficulty of methods and costs falling (Giribet, 2015). With this, there is a fear that some phylogeneticists will ignore morphological data, when genetic data can easily be extracted and sequenced (Mooi & Gill, 2010). It has been proposed that this can be due to morphological data and how collecting these data are not using more modern methodology, such as that of molecular methods (Wiens, 2001). The most pressing problems being a lack of appropriate experts and taxonomists, and issues with “character analysis” such as translating anatomical data into codified characters, standardization of that coding, ordering, and weighting characters (Wiens, 2001). Some authors proposed that phylogenetic signal of morphological data could be lost when accompanied by molecular data, especially when consensus methods were employed and the data was not congruent with that of molecular data (Miyamoto, 1985). This was disputed by others on the principle that congruence between morphological characters and covariation of this data would still play a role in relationships that would be hypothesized (Donoghue, 1989; Eernisse and Kluge, 1993). Additionally, it is likely that the perceived benefits of using just DNA sequences, such as designating an associated sequence in a database with a holotype specimen (Tautz et al., 2003), come with their own host of similar issues, such as sequences being a subjective source of data. This is countered with the argument that alignment methods, differentiation between paralogs and orthologs, and 7 selection of genes to sequence are all issues that cannot easily be solved or standardized (Lipscomb et al., 2003). Morphological data should not necessarily be excluded from being used in large phylogenetic analyses. They are still informative and important in understanding the evolution of lineages of taxa on Earth, both extant and extinct, the latter containing the vast majority of diversity of life on Earth (Edgecombe 2010). It has been shown in various studies in the last decade that there can be some phylogenetic signal for morphological characters, especially for genitalia characters of animals with

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