The concept of areas of endemism (AoEs) has rarely been discussed in the literature, even though the use of methods to ascertain them has recently increased. We introduce a grid-based protocol for delimiting AoEs using alternative criteria for the recognition of AoEs that are empirically tested with harvestmen species distributions in the Atlantic Rain Forest. Our data, comprising 778 records of 123 species, were analysed using parsimony analysis of endemicity and endemicity analysis on four different grids (two cell sizes and two cell placements). Additionally, we employed six qualitative combined criteria for the delimitation of AoEs and applied them to the results of the numerical analyses in a new protocol to objectively delimit AoEs. Twelve AoEs (the most detailed delimitation of the Atlantic Rain Forest so far) were delimited, partially corroborating the main divisions previously established in the literature. The results obtained with the grid-based methods were contradictory and were plagued by artefacts, probably due to the existence of different endemism patterns in one cell or to a biogeographical barrier set obliquely to latitudinal and longitudinal axes, for example. Consequently, the congruence patterns found by them should not be considered alone; qualitative characteristics of species and clade distributions and abiotic factors should be evaluated together. Mountain slopes are the main regions of endemism, and large river valleys are the main divisions. Refuges, marine transgressions and tectonic activity seem to have played an important role in the evolution of the Atlantic Rain Forest.
Goniosomatine harvestmen have strongly armed pedipalps, generally large bodies and, commonly, very long legs (sometimes more than 20 cm), and are distributed in the Brazilian Atlantic forest, from southern Bahia to Santa Catarina. Since they are conspicuous animals and individuals of some species tend to concentrate in caves (and also under rock boulders), they have been (and still are) the target of several studies, especially those focusing on reproductive and defensive behavior, population ecology, physiology, chromosomes, etc. In spite of their importance for biological studies (some species constitute important and frequently used models for these studies), the taxonomy of Goniosomatinae has faced some problems, including misidentification, a large number of undescribed species and the lack of a phylogenetic hypothesis for the relationships among its species (which would allow evolutionary studies to be made). The last taxonomic changes in the subfamily were made 60 years ago. Considering a taxonomic revision and cladistic analysis of the subfamily to be of paramount importance, the main scope of the present paper is to provide a cladistic analysis and taxonomic revision of the species of Goniosomatinae and a new arrangement of genera (and species). The main taxonomic changes are given as follows. Six genera are recognised within the subfamily: Goniosoma; the newly described genus Pyatan; the reestablished genera Serracutisoma, Heteromitobates and Mitogoniella; and Acutisoma. New generic synonyms include: Glyptogoniosoma = Goniosomella = Lyogoniosoma = Metalyogoniosoma = Xulapona = Goniosoma, Acutisomelloides = Pygosomoides = Spelaeosoma = Serracutisoma; and Acutisomella = Heteromitobates. Newly described species include: Goniosoma capixaba; G. apoain; Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini; Serracutisoma pseudovarium; S. fritzmuelleri; S. guaricana; Heteromitobates anarchus; H. harlequin; H. alienus; Mitogoniella taquara; M. unicornis; and Acutisoma coriaceum. New combinations include: Goniosoma macracanthum (Mello-Leitão, 1922); G. unicolor (Mello-Leitão, 1932); G. carum (Mello-Leitão, 1936); Serracutisoma proximum (Mello-Leitão, 1922); S. banhadoae (Soares & Soares, 1947); S. molle (Mello-Leitão, 1933); S. thalassinum (Simon, 1879); S. catarina (Machado, Pinto-da-Rocha & Ramires, 2002); S. inerme (Mello-Leitão, 1927); S. spelaeum (Mello-Leitão, 1933); Heteromitobates inscriptus (Mello-Leitão, 1922); H. albiscriptus (Mello-Leitão, 1932); Mitogoniella modesta (Perty, 1833); and M. badia (Koch, 1839). Reestablished combinations include: Mitogoniella indistincta Mello-Leitão, 1936 and Acutisoma longipes Roewer, 1913. New specific synonyms include: Acutisomella cryptoleuca = Acutisomella intermedia = Goniosoma junceum = Goniosoma patruele = Goniosoma xanthophthalmum = Metalyogoniosoma unum = Goniosoma varium, Goniosoma geniculatum = Goniosoma venustum; Goniosomella perlata = Progoniosoma minense = Goniosoma vatrax, Glyptogoniosoma perditum = Progoniosoma cruciferum = Progoniosoma tijuca = Goniosoma dentipes; Leitaoius iguapensis = Leitaoius viridifrons = Serracutisoma proximum; Acutisoma marumbicola = Acutisoma patens = Serracutisoma thalassinum; Progoniosoma tetrasetae = Serracutisoma inerme; and Acutisoma monticola = Leitaoius nitidissimus = Leitaoius xanthomus = Mitogoniella mutila = Acutisoma longipes. The following species are considered species inquirenda: Goniosoma lepidum Gervais, 1844; G. monacanthum Gervais, 1844; G. obscurum Perty, 1833; G. versicolor Perty, 1833; and Mitogoniella badia (Koch, 1839). The monotypic genus Goniosomoides Mello-Leitão, 1932 (and its species, G. viridans Mello-Leitão, 1932) is removed from Goniosomatinae and considered incertae sedis.
Based on a cladistic biogeographic analysis of 6 species-level phylogenies of harvestman taxa, we searched for congruence in the historical relationships of 12 areas of endemism of the Brazilian Atlantic Rain Forest. We constructed general area cladograms using Primary Brooks Parsimony Analysis (BPA), BPA of nodes, and paralogy-free subtree analysis. These analyses resulted in 6 general area cladograms, that allow to infer a general pattern of the relationships among areas of endemism from the Brazilian Atlantic Rain Forest. Northern areas resulted related basally showing main disjunctions at the Doce River Valley and Todos os Santos Bay/São Francisco River Valley. The remaining areas of endemism were included in a southern and a southeastern block, separated by the Ribeira do Iguape Valley. Incongruence Length Differences tests showed no significant incongruence among the resulting cladograms and other matrix partitions. We concluded that tectonism and ancient marine transgressions were the probable processes responsible for the main disjunctions, whereas Neogene refugia seem to have caused the more recent disjunctions. The general pattern and redundancy in area relationships suggest a model of main reiterative barriers in diversification at multiple times for the evolution of the Atlantic Rain Forest. The renewal of cladistic biogeography and the search for common biogeographic patterns are discussed.
The harvestmen subfamily Hernandariinae is reviewed and a new classification is proposed based on cladistic analysis using 67 morphological characters. The subfamily is composed of six genera and 23 species and occurs in south-southeastern Brazil, Paraguay, and northeastern Argentina. Fourteen new combinations are proposed: Hernandaria armatifrons (Roewer, 1917); H. una (Mello-Leitão, 1927); Acrogonyleptes granulatus (H. Soares, 1966); A. pectinifemur (Soares & Soares, 1947); Acanthogonyleptes alticola (Mello-Leitão, 1922); A. editus (Roewer, 1943); A. fallax (Mello-Leitão, 1932); A. fulvigranulatus (Mello-Leitão, 1922); A. marmoratus (Mello-Leitão, 1940); A. pictus (Piza, 1942); A. singularis (Mello-Leitão, 1935); A. soaresi (Mello-Leitão, 1944); A. variolosus (Mello-Leitão, 1944). Seven synonymies are proposed: Proweyhia Mello-Leitão, 1927 and Metaxundarava Mello-Leitão, 1927 = Hernandaria Sørensen, 1884; Apembolephaenus calcaratus Soares & Soares, 1945 = H. armatifrons (Roewer, 1917); Sphaerobunus Rower, 1917 and Paraproweyhia Soares & Soares, 1947 = Acrogonyleptes Roewer, 1917; Paraproweyhia curitibae Soares & Soares, 1947 = Acrogonyleptes exochus (Mello-Leitão, 1931); and Melloleitaniana curitibae B. Soares, 1943 = Acrogonyleptes spinifrons Roewer, 1917. Three species are revalidated: Acrogonyleptes granulatus (H. Soares, 1966), A. pectinifemur (Soares & Soares, 1947), and A. spinifrons Roewer, 1917. Seven new species are described: Hernandaria sundermannorum sp. nov. (São Paulo State, Brazil), Hernandaria anitagaribaldiae sp. nov. (Santa Catarina State, Brazil), Hernandaria zumbii sp. nov. (Santa Catarina State, Brazil), Hernandaria chicomendesi sp. nov. (Santa Catarina State, Brazil), Acrogonyleptes cheguevarai sp. nov. (Rio Grande do Sul State, Brazil), Pseudotrogulus pagu sp. nov. (São Paulo State, Brazil), Pseudotrogulus trotskyi sp. nov. (Paraná State, Brazil)
The integration of ecology and historical biogeography is fostering the investigation of diversity patterns. We studied alpha and beta diversity patterns of Brazilian Atlantic Forest harvestman (Arachnida: Opiliones) communities and related them to environmental and historical factors. Our data bank contains 508 species from 63 sites, encompassing almost the entire latitudinal range of Atlantic Forest. Alpha diversity was higher in coastal sites in the south and southeast regions and decreased in sites inland, as well as in sites in the coastal northeast region, especially in northern Bahia state. Alpha diversity was positively influenced by precipitation and altitudinal range, but the low number of species in northeastern coastal sites seems to be more related to the historical distribution of Neotropical harvestman lineages than to recent environmental factors. Geographic distance was the most influential factor for beta diversity. Compositional changes were also remarkably congruent with areas of endemism delimited for Atlantic Forest harvestmen. The percentage of protected areas for each area of endemism was very unbalanced, and Espírito Santo and Pernambuco states were the least protected areas. The turnover process observed in the compositional changes indicates that conservation strategies should include as many reserves as possible because every community presents a unique set of species.
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