Cytogenetic study in sand spiders (Sicariidae) from the brazilian Caatinga: sex chromosome system diversity in closely related species / Estudio citogenético en arañas de arena (Sicariidae) de la Caatinga brasileña: diversidad del sistema de cromosomas sexuales en especies estrechamente relacionadas

ABSTRACT In this study, we investigated the chromosomes of three species of Sicarius spiders from the Brazilian Caatinga, using classical and molecular cytogenetic techniques. Based on the phylogenetic approach, we also discussed about the variation of diploid number, types of sex chromosome system and changes in the localization of ribosomal genes of Scytodoidea. Sicarius are Synspermiata spiders that together with the genera Loxosceles and Hexophthalma constitute the family Sicariidae. In this group, the available cytogenetic data showed a low diploid number range (2n♂=18 to 2n♂=23) and the presence of only multiple sex chromosome systems (X1X2Y and X1X20). Mitotic metaphase cells exhibited 2n♂=16+X1X2Y for Sicarius cariri and S. ornatus, and 2n♂=18+XY for S. tropicus. In these species, silver impregnation revealed nucleolar organizer region (Ag-NOR) on the terminal region of pair 1. In S. ornatus and S. tropicus, the results obtained with fluorescent in situ hybridization (FISH) using 18S rDNA probe were similar to Ag-NOR, however in S. cariri, the ribosomal sites were localized in the terminal region of the X1 sex chromosome. In this work, we presented the first description of a simple sex chromosome system for Sicariidae, helping to understand how the XY sex chromosome system evolved from the X1X2Y system. Additionally, FISH data incongruous with Ag-NOR indicate that the cytogenetic studies in Sicariidae allow investigating the relation between the karyotype evolution and the distribution and the activity of rDNA genes.

RESUMEN En este estudio, investigamos los cromosomas de tres especies de arañas Sicarius de la Caatinga brasileña, utilizando técnicas de citogenética clásica y molecular. Usando un enfoque filogenético, también discutimos la variación del número diploide, los tipos de sistema cromosómico sexual y los cambios en la localización de los genes ribosómicos en Scytodoidea. Los Sicarius son arañas Synspermiata que, junto con los géneros Loxosceles y Hexophthalma, constituyen a la familia Sicariidae. En este grupo, los datos citogenéticos disponibles mostraron un rango de número diploide bajo (2n♂=18 a 2n♂=23) y únicamente la presencia de sistemas de cromosomas sexuales múltiples (X1X2Y y X1X20). Las células mitóticas en metafase mostraron 2n♂=16+X1X2Y para Sicarius cariri y S. ornatus, y 2n♂=18+XY para S. tropicus. En estas especies, la impregnación de plata reveló la región organizadora nucleolar (Ag-NOR) en la región terminal del par 1. En S. ornatus y S. tropicus, los resultados obtenidos con la hibridación in situ fluorescente (FISH) utilizando la sonda de ADNr 18S fueron similares a los de Ag-NOR, sin embargo, en S. cariri los sitios ribosomales se localizaron en la región terminal del cromosoma sexual X1. En este trabajo, presentamos la primera descripción de un sistema cromosómico sexual simple para Sicariidae, ayudando a entender cómo el sistema cromosómico sexual XY evolucionó a partir del sistema X1X2Y. Además, los datos de FISH incongruentes con Ag-NOR indican que los estudios citogenéticos en Sicariidae permiten investigar la relación entre la evolución del cariotipo y la distribución y la actividad de los genes de ADNr.

Phylogeny of Neotropical Sicarius sand spiders suggests frequent transitions from deserts to dry forests despite antique, broad-scale niche conservatism.

Phylogenetic niche conservatism (PNC) shapes the distribution of organisms by constraining lineages to particular climatic conditions. Conversely, if areas with similar climates are geographically isolated, diversification may also be limited by dispersal. Neotropical xeric habitats provide an ideal system to test the relative roles of climate and geography on diversification, as they occur in disjunct areas with similar biotas. Sicariinae sand spiders are intimately associated with these xeric environments, particularly seasonally dry tropical forests (SDTFs) and subtropical deserts/scrublands in Africa (Hexophthalma) and the Neotropics (Sicarius). We explore the role of PNC, geography and biome shifts in their evolution and timing of diversification. We estimated a time-calibrated, total-evidence phylogeny of Sicariinae, and used published distribution records to estimate climatic niche and biome occupancy. Topologies were used for estimating ancestral niches and biome shifts. We used variation partitioning methods to test the relative importance of climate and spatially autocorrelated factors in explaining the spatial variation in phylogenetic structure of Sicarius across the Neotropics. Neotropical Sicarius are ancient and split from their African sister-group around 90 (57-131) million years ago. Most speciation events took place in the Miocene. Sicariinae records can be separated in two groups corresponding to temperate/dry and tropical/seasonally dry climates. The ancestral climatic niche of Sicariinae are temperate/dry areas, with 2-3 shifts to tropical/seasonally dry areas in Sicarius. Similarly, ancestral biomes occupied by the group are temperate and dry (deserts, Mediterranean scrub, temperate grasslands), with 2-3 shifts to tropical, seasonally dry forests and grasslands. Most of the variation in phylogenetic structure is explained by long-distance dispersal limitation that is independent of the measured climatic conditions. Sicariinae have an ancient association to arid lands, suggesting that PNC prevented them from colonizing mesic habitats. However, niches are labile at a smaller scale, with several shifts from deserts to SDTFs. This suggests that PNC and long-distance dispersal limitation played major roles in confining lineages to isolated areas of SDTF/desert over evolutionary history, although shifts between xeric biomes occurred whenever geographical opportunities were presented.

Phylogeny of Neotropical Sicarius sand spiders suggests frequent transitions from deserts to dry forests despite antique, broad-scale niche conservatism

Phylogenetic niche conservatism (PNC) shapes the distribution of organisms by constraining lineages to parti-cular climatic conditions. Conversely, if areas with similar climates are geographically isolated, diversificationmay also be limited by dispersal. Neotropical xeric habitats provide an ideal system to test the relative roles ofclimate and geography on diversification, as they occur in disjunct areas with similar biotas. Sicariinae sandspiders are intimately associated with these xeric environments, particularly seasonally dry tropical forests(SDTFs) and subtropical deserts/scrublands in Africa (Hexophthalma) and the Neotropics (Sicarius). We explorethe role of PNC, geography and biome shifts in their evolution and timing of diversification. We estimated atime-calibrated, total-evidence phylogeny of Sicariinae, and used published distribution records to estimateclimatic niche and biome occupancy. Topologies were used for estimating ancestral niches and biome shifts. Weused variation partitioning methods to test the relative importance of climate and spatially autocorrelatedfactors in explaining the spatial variation in phylogenetic structure ofSicariusacross the Neotropics. NeotropicalSicariusare ancient and split from their African sister-group around 90 (57–131) million years ago. Most spe-ciation events took place in the Miocene. Sicariinae records can be separated in two groups corresponding totemperate/dry and tropical/seasonally dry climates. The ancestral climatic niche of Sicariinae are temperate/dryareas, with 2–3 shifts to tropical/seasonally dry areas inSicarius. Similarly, ancestral biomes occupied by thegroup are temperate and dry (deserts, Mediterranean scrub, temperate grasslands), with 2–3 shifts to tropical,seasonally dry forests and grasslands. Most of the variation in phylogenetic structure is explained by long-distance dispersal limitation that is independent of the measured climatic conditions. Sicariinae have an ancientassociation to arid lands, suggesting that PNC prevented them from colonizing mesic habitats. However, nichesare labile at a smaller scale, with several shifts from deserts to SDTFs. This suggests that PNC and long-distancedispersal limitation played major roles in confining lineages to isolated areas of SDTF/desert over evolutionaryhistory, although shifts between xeric biomes occurred whenever geographical opportunities were presented

Phylogeny of Neotropical Sicarius sand spiders suggests frequent transitions from deserts to dry forests despite antique, broad-scale niche conservatism

Phylogenetic niche conservatism (PNC) shapes the distribution of organisms by constraining lineages to parti-cular climatic conditions. Conversely, if areas with similar climates are geographically isolated, diversificationmay also be limited by dispersal. Neotropical xeric habitats provide an ideal system to test the relative roles ofclimate and geography on diversification, as they occur in disjunct areas with similar biotas. Sicariinae sandspiders are intimately associated with these xeric environments, particularly seasonally dry tropical forests(SDTFs) and subtropical deserts/scrublands in Africa (Hexophthalma) and the Neotropics (Sicarius). We explorethe role of PNC, geography and biome shifts in their evolution and timing of diversification. We estimated atime-calibrated, total-evidence phylogeny of Sicariinae, and used published distribution records to estimateclimatic niche and biome occupancy. Topologies were used for estimating ancestral niches and biome shifts. Weused variation partitioning methods to test the relative importance of climate and spatially autocorrelatedfactors in explaining the spatial variation in phylogenetic structure ofSicariusacross the Neotropics. NeotropicalSicariusare ancient and split from their African sister-group around 90 (57–131) million years ago. Most spe-ciation events took place in the Miocene. Sicariinae records can be separated in two groups corresponding totemperate/dry and tropical/seasonally dry climates. The ancestral climatic niche of Sicariinae are temperate/dryareas, with 2–3 shifts to tropical/seasonally dry areas inSicarius. Similarly, ancestral biomes occupied by thegroup are temperate and dry (deserts, Mediterranean scrub, temperate grasslands), with 2–3 shifts to tropical,seasonally dry forests and grasslands. Most of the variation in phylogenetic structure is explained by long-distance dispersal limitation that is independent of the measured climatic conditions. Sicariinae have an ancientassociation to arid lands, suggesting that PNC prevented them from colonizing mesic habitats. However, nichesare labile at a smaller scale, with several shifts from deserts to SDTFs. This suggests that PNC and long-distancedispersal limitation played major roles in confining lineages to isolated areas of SDTF/desert over evolutionaryhistory, although shifts between xeric biomes occurred whenever geographical opportunities were presented