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Paragoniosomatinae, a new subfamily of Gonyleptidae (Arachnida: Opiliones), based on a new species from the Chapada Diamantina relict cloud forests, Brazil
Abstract
In this paper, we describe a new subfamily of Gonyleptidae, Paragoniosomatinae subfam. nov., based on a new genus and species, Paragoniosoma cachaceiro gen. nov., sp. nov., found in an upper montane cloud forest of Chapada Diamantina mountains, one of the small interior patches of Atlantic Forest. This discovery suggests that the narrow-restricted cloud forests are old relicts of an Atlantic Forest hotspot. Paragoniosoma cachaceiro gen. nov., sp. nov. is characterized by the presence of four areas on the dorsal scutum, serose dry mark in areas III–IV, seven basal setae in penis (macrosetae A and B) in two rows, and very large tarsal process. We performed a phylogenetic analysis based on morphological characters that revealed a close relationship of the new species with Goniosomatinae, a subfamily that includes six genera and 38 valid species distributed throughout the coastal Atlantic Forest. The new species also has the same habitat and other behavioural and ecological traits as Goniosomatinae, furnishing great insights on the evolution of their characteristic biology, e.g., subsocial behavior. Field surveys of harvestmen from the type locality and nearby lowland seasonal forests indicate a unique fauna, including additional new, undescribed species. In addition to phylogenetics, this dissimilarity with other regions and the presence of endemic species of other animal and plant taxa provide support for the consideration of the cloud montane patches of Chapada Diamantina as relicts. We discuss this rule in the historical biogeographic context of Atlantic Forest and suggest that the new species represents evidence of an Atlantic Forest that was more widespread in the past.
... After the original proposal, the AE11 pattern has been routinely applied by different research groups to species descriptions in the families Cosmetidae (e.g., Damron et al. 2018;Friedrich & Lehmann 2020), Epedanidae (e.g., Zhang & Martens 2018), Gonyleptidae (e.g., Hara et al. 2018;Acosta 2020;Araujo-da-Silva et al. 2020) and Stygnopsidae (e.g., Cruz-López & Francke 2016, 2019, 2020. Kury & Villarreal (2015) suggested that the AE11 pattern may also occur in other superfamilies of Laniatores, such as Assamioidea and Epedanoidea (sensu Ferna´ndez et al. 2017), but the authors did not further pursue this matter. ...
A description of the pattern of structure and organization of the penial macrosetae was recently put forward for Gonyleptoidea Sundevall, 1833 and demonstrated to occur also in other families of Grassatores Kury, 2002, such as Assamiidae Srensen, 1884, Epedanidae Srensen, 1886 and Pyramidopidae Sharma, Prieto & Giribet, 2011. This set of homology hypotheses is herein called AE11 pattern. In this work, the monotypic Sri Lankan genus Eurytromma Roewer, 1949 of the family Podoctidae Roewer, 1912 (which is currently assigned to the Epedanoidea in the Grassatores) is studied. Male genitalia of Eurytromma are described for the first time and the first attempt is made to reconcile the chaetotaxy of ventral plate of podoctid species with the AE11 pattern. The podoctid genera Hoplodino Roewer, 1915, Strandibalonius Roewer, 1912 and Santobius Roewer, 1949 are also exemplified here; their set of macrosetae is described and the AE11 pattern is demonstrated to occur in all of them. Therefore, AE11 is more phylogenetically widespread in the Grassatores than previously thought. Eurytromma pictulum (Pocock, 1903) from Sri Lanka is redescribed. It is characterized by the presence of a meso-frontal stridulatory saw of denticles on the cheliceral hand and extreme reduction of mesotergal sutures. A further refinement is made on the current terminology system of cheliceral dentition of Podoctidae.
Background
K92 is a clade of Gonyleptidae harvestmen of which the greatest diversity lies within the large non-monophyletic subfamily Gonyleptinae. K92 formally includes five subfamilies: Caelopyginae Sørensen, 1884, Gonyleptinae Sundevall, 1833, Hernandariinae Sørensen, 1884, Progonyleptoidellinae Soares & Soares, 1985 and Sodreaninae Soares & Soares, 1985. The basic taxonomy of K92 presently finds itself in an unsatisfactory state, with many species poorly known and supraspecific taxa defined by a combination of few trivial and/or misconstrued features.
Questions
Three lines of investigation regarding K92 are pursued in this project: (1) some formal gonyleptine genera, such as Liogonyleptoides Mello-Leitão and de, 1925 and Parapachyloides Roewer, 1913a were suspected to be especially related to each other as members of an undiscovered non-gonyleptine taxon. Do those species, distributed along the semi-arid diagonal in South America plus the Atlantic Forest in eastern Brazil really form a clade? (2) Has the acquisition of the caelopygine facies by species of Progonyleptoidellinae and Sodreaninae been gradual and independent, associated with diurnal and arboreal habits? (3) Does the attenuation of sexual dimorphism present in K92 have a phylogenetic signal?
Methods
A cladistic analysis of some critical taxa in K92 is performed by means of maximum parsimony and Bayesian inference. A matrix with 112 morphological characters and 61 terminals has been analyzed for this project, focusing on (a) potential relatives of Liogonyleptoides, and (b) species of K92 with varied degrees of sexual dimorphism.
Main results
For question (1), a phylogenetic hypothesis is proposed herein for the larger clades of K92 as follows: (a) a branch in the phylogeny of K92 is here recognized and formally described as the new subfamily Cearininae. The Cearininae subfam. nov. includes the following genera: Cearinus Roewer, 1929 (type genus), Inhuma Piza, 1938, Liogonyleptoides Mello-Leitão and de, 1925 (incl. Pachyleptes Mello-Leitão and de, 1932), Pajeuia gen. nov., Parapachyloides Roewer, 1913b and Ypsilonurus Mello-Leitão and de, 1933a. (b) Gonyleptinae did not resolve here as a clade. (c) A monophylum never reported before is recovered here (clade I, here nicknamed CAPS), consisting of the nominal subfamilies Caelopyginae, Progonyleptoidellinae and Sodreaninae. Progonyleptoidellinae is polyphyletic. All of these subfamilies have their internal composition altered. For question (2), we found that the caelopygine facies (diurnal, arboreal, triangular bodied, brightly colored animals with long legs, high tarsal counts and pectinated claws) probably arose independently in (a) the classic Caelopyginae (including pectinated tarsal claws, the full caelopygine facies), (b) partial caelopygine facies of extended Sodreaninae, as represented by Heliella, and (c) partial caelopygine facies of extended Caelopyginae, as represented by Progonyleptoidellus. For question (3), we found that attenuated sexual dimorphism may evolve via two different processes: females acquiring armature and males losing armature. The three different forms of sexual dimorphism provided insight to the phylogeny of K92: (a) strong sexual dimorphism, where females are plump and unarmed at leg IV in contrast with lean and strongly armed males is typical for most members of K92 and other Gonyleptidae and is the primitive condition; (b) a process of masculinization, where the lean, polygonal females resemble males, only with the armature of leg IV somewhat attenuated, occurs in some Caelopyginae and Progonyleptoidellinae in the traditional sense and reaches the most extreme degree in the members of the genus Moreiranula Roewer, 1930; (c) a process of feminization, where males have weakly developed coxa IV, no differential armature on leg IV, and are almost indistinguishable from the females. This occurs in some Hernandariinae such as Piassagera Roewer, 1929, one species of Moreiranula and some Progonyleptoidellinae, such as Iporangaia Mello-Leitão and de, 1935a and Mitopernoides B. Soares, 1945, and one Caelopyginae strictu sensu, Thereza Roewer, 1923.
Other taxonomic results
As byproducts of this project, not direct consequences of the phylogenetic analysis, taxonomic changes are proposed for some genera and species in K92. For conciseness, the equality sign (x = y) is here used instead of the expression “x is here newly considered a junior subjective synonym of y”: (1) Cearinus was hitherto monotypic. A second species is described here – Cearinus torulosus sp. nov. – from Mato Grosso State, Central Brazil. (2) Liogonyleptoides is reviewed. There are currently 5 species included, of which one (Liogonyleptoides heliae Kury 2003) is a species inquirenda, another is a synonym (Liogonyleptoides capichaba = Liogonyleptoides tetracanthus) and a third (Liogonyleptoides minensis) is here transferred to the new monotypic genus Pajeuia. A new diagnosis of the genus Liogonyleptoides is proposed and emended diagnoses are given to Liogonyleptoides inermis and L. tetracanthus. Two new species are described in Liogonyleptoides – Liogonyleptoides mirificus sp. nov. and Liogonyleptoides venatus sp. nov., from northern Espírito Santo. (3) Inhuma Piza, 1938, hitherto monotypic, is expanded to include another three species, each of which originally constituted a monotypic genus. Accordingly, the genera Schubartesia B. Soares, 1944a, Hugoesia H. Soares 1968 and Stefanesia Soares and Soares, 1988 all = Inhuma. (4) Metagonyleptes Roewer, 1913a is dismantled, the genus is newly considered a junior subjective synonym of Mischonyx Bertkau, 1880 (because the synonymy of the type species), while its species are transferred to varied genera, notably to Guascaleptes (herein revalidated from the synonymy with Geraeocormobius Holmberg, 1887) and to the undescribed genera SOD-01 and SOD-02. (5) Gonyleptes carinatus Sørensen, 1884 (type species of Metagonyleptes) = Gonyleptes scaber Kirby 1819 (currently Mischonyx scaber (Kirby 1819)). (6) Geogonys Mello-Leitão 1937, currently a synonym of Metagonyleptes, is newly transferred to the synonymy of Guascaleptes Mello-Leitão and de, 1933b to include some species formerly in Metagonyleptes. (7) Geogonys pallidipalpis Mello-Leitão and de, 1937, type species of Geogonys and currently a valid species of Metagonyleptes is newly synonymized with Metagonyleptes grandis Roewer, 1913b, both newly combined under Guascaleptes. (8) Gonyleptes saprophilus Mello-Leitão and de, 1922 and Sodreana curupira Pinto-da-Rocha and Bragagnolo, 2011 are newly transferred to Moreiranula Roewer, 1930, forming the new combinations Moreiranula saprophila and Moreiranula curupira. (9) Gonyleptes espiritosantensis B. Soares, 1944a = Proctobunoides tuberosus Mello-Leitão and de, 1944.
During a study of Discocyrtus Holmberg, 1878 and related genera, several species of alleged Pachylinae, mostly Discocyrtus, from southern Brazil were suspected of belonging in Roeweriinae. Herein, a maximum parsimony phylogenetic analysis of morphological characters was performed to test this hypothesis. Accordingly, the following taxonomic changes are herein proposed: the genus Bunopachylus Roewer, 1943 (currently in Pachylinae) is herein revalidated from synonymy with Discocyrtus, transferred to the Roeweriinae and now includes three species: B. armatissimus comb. nov. (for Discocyrtus armatissimus Roewer, 1913), B. orientalis comb. nov. (for Pachyloides orientalis Roewer, 1913) and B. occultus sp. nov. (which had been misidentified in the literature). A total of five new synonymies are proposed here: Discocyrtus milloti Roewer, 1943 = Discocyrtus coronatus Mello-Leitão, 1935 = Paradiscocyrtus trochanteralis Roewer, 1929 = Discocyrtus calcarifer Roewer, 1917 = Pachyloides orientalis Roewer, 1913, and Bunopachylus magnicalcar Roewer, 1943 (the type-species of Bunopachylus) = Discocyrtus armatissimus Roewer, 1913.
We herein propose the transfer of Metaphareus punctatus Roewer, 1913, to the genus Eutimesius Roewer, 1913 (Heterostygninae) based on the genital morphology of the species and morphology of tarsi III and IV, resulting in the new combination Eutimesius punctatus (Roewer, 1913), comb. nov. New geographical records for E. albicinctus (Roewer, 1915) are offered, recording sympatry between the congeneric species E. albicinctus and E. punctatus and data on the reproductive behavior of E. punctatus are presented, suggesting the existence of parental care in this species.
The generic name Paecilaema C.L. Koch, 1839 is anchored to the nomen dubium Cosmetus u-flavum Perty, 1833, and the fixation of the latter is proposed by designating a neotype. The choice was made on a potential topotype, being a common species in metropolitan Rio de Janeiro city, SE Brazil, which has been described as new five times and is currently best known as Metavononoides bellus (Mello-Leitão, 1932). Cosmetus u-flavum has been originally included in Paecilaema along with Paecilaema limbatum Kollar in C.L. Koch, 1839, the latter here synonymized with the former. Therefore, our choice of this species is adequate in conveying Koch’s original concept of Paecilaema. The most striking diagnostic feature of the species of Paecilaema is the presence of a lyra, a unique shape of blots in layers of shades of white to pale yellow on the dorsal scutum. Other patterns of scutal white blots are hereby defined and named. Paecilaema is here considered most closely related to Metavononoides Roewer, 1928. Comments are made on the diagnostic characters and the terminology of parts of the lyra is expanded with the introduction of new terms. Comments are made on the formation of the generic name Paecilaema. Taxonomic adjustments are made in the closest relatives of Paecilaema u-flavum, and the following nomenclatural acts are proposed: (1) MNRJ 3433 is designated as male neotype of Cosmetus u-flavum, type species of Paecilaema. (2) Proposed new synonymies: Poecilaema soerenseni Henriksen, 1932 = Paecilaemula bella Mello-Leitão, 1932 = Eucynorta brasiliensis Mello-Leitão, 1923 = Paecilaema limbatum Kollar in C.L. Koch, 1839 = Cosmetus u-flavum Perty, 1833. (3) Nine species originally described elsewhere but currently placed in Metavononoides are transferred to Paecilaema, either as new or as restored combinations: Paecilaemula albisecta Mello-Leitão, 1942, Poecilaemula albosigillata Mello-Leitão, 1941, Paecilaema lyra Sørensen, 1932, Metavononoides melanacanthus Ferreira, Pedroso & Kury, 2005, Paecilaema muticum Sørensen, 1932, Paecilaema ornatissimum Mello-Leitão 1942, Poecilaemula peculiare Roewer, 1917, Paecilaemula preciosa Roewer 1928 and Metavononoides renneri Ferreira, Pedroso & Kury, 2007. (4) The following genera, currently in the synonymy of Paecilaema, are all revalidated and given emended diagnoses: Meterginoides Roewer, 1912, Meterginulus Roewer, 1912, Paecilaemella Roewer, 1925, Pararhauculus Roewer, 1933, Soaresella Goodnight & Goodnight 1947. (5) Their respective type species, currently combined under Paecilaema, are transferred back to their revalidated genera: Paecilaema rastelliferum Pickard-Cambridge 1905 to Meterginulus; Meterginus inermis Banks, 1909 to Meterginoides; Pararhauculus lineatus Roewer, 1933 to Pararhauculus; Paecilaemella festae Roewer, 1925 to Paecilaemella and Soaresella gracilis Goodnight & Goodnight, 1947 to Soaresella.
A group of Amazonian harvestmen is recognized and described as Amazochroma gen. nov. This taxon includes Discocyrtus carvalhoi Mello-Leitão, 1941 (type species), the only species of Discocyrtus previously thought to occur in Amazonia, and Amazochroma pedroi gen. et sp. nov., described here from the Brazilian states of Acre and Rondônia. New records are added for Amazochroma carvalhoi gen. et comb. nov, expanding its distribution from the Brazilian state of Mato Grosso northwards also to Pará and Amazonas in Brazil and additionally French Guiana and Suriname. Diagnostic features of Amazochroma gen. nov. include: trichromatic pattern of legs, dry marks on the dorsal scutum and base of legs and diastema in the row of macrosetae C of the penis ventral plate. A morphological maximum parsimony analysis (1022 scorings; 16 taxa; 64 characters) is performed to test whether Amazochroma gen. nov. is a member of Discocyrtus and if the traditional allocation of Discocyrtus in Pachylinae is defendable. A clade is retrieved containing three groups: (1) Mitobatinae, (2) Discocyrtus and (3) a clade containing Amazochroma gen. nov., Discocyrtanus and Roeweria, here described as a new subfamily of Gonyleptidae – Roeweriinae subfam. nov. Discocyrtanus Roewer, 1929 and Roeweria Mello-Leitão, 1923 are accordingly here transferred from Pachylinae to Roeweriinae subfam. nov.
A new troglobitic harvestman, Relictopiolus galadriel gen. nov et sp. nov., is described from Olhos d’Água cave, Itacarambi, Minas Gerais State, Brazil. Morphological characters, including male genitalia and exomorphology, suggest that this species belongs to the family Kimulidae, and it appears to share the greatest similarities with Tegipiolus pachypus. Bayesian inference analyses of a molecular dataset strongly support the inclusion of this species in Kimulidae and confirm the hypothesized sister-group relationship between R. galadriel and T. pachypus. A time calibrated phylogeny indicates that these sister-taxa diverged from a common ancestor approximately 40 Mya, during the Paleogene. The current range of Kimulidae illustrates a remarkable disjunct distribution, and leads us to hypothesize that the ancestral distribution of Kimulidae was once much more widespread across eastern Brazil. This may be attributed to the Eocene radiation associated with the warming (and humidifying) events in the Cenozoic when the best conditions for evergreen tropical vegetation in South America were established and followed by the extinction of kimulid epigean populations together with the retraction of rain forests during the Oligocene to Miocene cooling. The discovery of this relictual troglobite indicates that the Olhos d’Água cave was a stable refugium for this ancient lineage of kimulids and acted as a "museum" of biodiversity. Our findings, considered collectively with the diverse troglofauna of the Olhos d’Água cave, highlight it as one of the most important hotspots of troglobite diversity and endemism in the Neotropics. Given the ecological stresses on this habitat, the cavernicolous fauna are at risk of extinction and we emphasize the urgent need for appropriate conservation actions. Finally, we propose the transfer of Acanthominua, Euminua, Euminuoides and Pseudominua from Kimulidae to Zalmoxidae, resulting in two new synonymies and 13 new combinations.
Based on a cladistic biogeographic analyses of six 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 (PF). These analyses resulted in six 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 (ILD) 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.
INTRODUCTION As indicated in the introductory chapter to this book, knowledge of tropical montane cloud forest (TMCF) occurrence, biodiversity, hydrology, and ecological functioning has increased considerably since the ground-breaking publications of Zadroga (1981), Stadtmüller (1987), and the proceedings of the first international symposium on TMCF held in 1993 in Puerto Rico (Hamilton et al., 1995a). Cloud forests continue to be threatened in several ways, notably by their conversion to pasture and various forms of agriculture, as well as by climatic drying – the numerous hydrological and ecological consequences of which are only poorly understood as yet (Bubb et al., 2004; Mulligan and Burke, 2005a; Pounds et al., 2006; Zotz and Bader, 2009). The collection of chapters in the present volume further advances our knowledge in the three broad and interrelated areas that were defined in the introduction, viz. (i) cloud forest biogeography and biodiversity, (ii) biophysical and ecological processes, and (iii) management and conservation strategies. As the recognition of the value of TMCFs as treasure houses of biodiversity and as providers of high-quality water continues to increase, an array of initiatives aimed at their conservation has emerged in recent years, often within a Payment for Ecosystem Services (PES) context (Asquith and Wunder, 2008; Muñoz-Piña et al., 2008; Porras et al., 2008; Garriguata and Balvanera, 2009; Tognetti et al., this volume). Such PES schemes, but also land and forest managers and policy-makers in general, need to determine (amongst others) which forests under their jurisdiction are the most diverse and valuable biologically, which ones provide the best water supplies, which forests are the most vulnerable to climate change or most threatened by encroachment, and which degraded TMCFs have the best chances for rehabilitation.
We used DNA sequence data to test the morphology-based taxonomy and examine the biogeography of the Brazilian Atlantic Rain Forest genus Promitobates. Most species are well differentiated morphologically, and a previous morphological phylogeny recovered the genus as monophyletic. However, some of these species have overlapping geographical distributions and considerable intraspecific variation, perhaps representing a species complex. Mitochondrial (12S rRNA and COI) and nuclear (ITS2 and 28S) genes were sequenced from 132 specimens collected from 27 localities. The results are consistent with significant mitochondrial introgression among the species P. ornatus, P. hatschbachi, P. lager, P. bellus and P. intermedius (the 'P. ornatus species complex'), with one specimen identified as a hybrid between P. nigripes and this complex. A phylogeographic study of the complex was conducted using mitochondrial haplotypes. This revealed remarkably poor dispersal among populations, with only one case of a shared haplotype, and very low genetic diversity. The phylogeny showed a clear break between populations on either side of a narrow region of forest, suggesting an important historical event separated these lineages in the genus. The analyses also pointed to population breaks that date back several millions of years or extremely small effective population sizes, depending on the mutation rate.
During Paleogene the Neotropical region was mainly covered by rainforests and ancient Amazon (AM) and Atlantic Forest (AF) were continuous and interconnected. The Andean uplift and drastic climate changes along the Eocene/Miocene resulted in the formation of a drier area separating AM from AF. However, multiple evidences have indicated recurrent connections between Neotropical rainforests during Quaternary. In this study we predict biogeographical connections between Neotropical Forests during the last glacial maximum (LGM—21 ka) and establish the climatic conditions that favored such connections. We tested if the general climatic conditions suitable to both Amazon (AM) and AF were uniform across current Cerrado and Caatinga areas or did subsets of these climatic spaces have independent spatial displacements connecting specific regions of AM and AF. For these, 50 occurrence points equidistant in the climatic space were sampled along the western and eastern AM and northern and southern AF and used to built ecological niche models (ENM) for each region. Potential distributions were predicted for the current and the LGM climatic scenarios using an ensemble approach. The ENMs detected the disjunct distribution of the two Neotropical rainforests in the present and showed three
Serracutisoma gnaspinii sp. nov. is described from the coastal Ombrophylous Dense Atlantic Rain Forest of Parana and Sao Paulo states. It differs from other Serracutisoma by having a retrolateral row of spines on femur IV, a rounded white-mark on the carapace and a low dorsal apophysis on trochanter IV. It is likely the sister-species of S. proximum, and with it and other five species forms the S. proximum group, herein formally proposed. Given that the biologic, phylogenetic and biogeographic aspects of Serracutisoma have been studied, the implications of the discovery of a new species are discussed, for instance the evolution of male fight, aggregation and maternal care, traits that are shared with other species. The geographic range of Serracutisoma gnaspinii sp. nov. is compatible with the "Parana" area of endemism, but with an unique distribution pattern compared to other endemic harvestmen species, which could indicate a new area of endemism also occupied by the primate Leontopithecus caissara Lorini & Persson, 1990.
For the first time a hypothesis of homology is proposed for the macrosetae which compose the armature of the distal truncus penis of the Gonyleptoidea. Previous attempts to name them in the literature have not been continued because they started by referring to macrosetae based on their position, which may change widely, and all of them were very narrow in scope. The present project instead names groups of macrosetae that do not mandatorily refer to their position, thus compensating for their hypothesized secondary position shifts. Using criteria of topology, shape and exclusion, six groups of setae are recognized, A–E, and their topological and phylogenetic distribution is studied and described in all families of Gonyleptoidea and two other related families of Grassatores. A cladistic analysis is performed, providing the following results: (1) the Microsetata are recovered including Metasarcidae/ Cosmetidae sister to an expanded Gonyleptidae; (2) the genera Quindina (Cranaidae) and Zygopachylus (Manaosbiidae) are transferred to Nomoclastidae rank nov., hitherto regarded as a subfamily of Stygnidae, now a sister group of the Microsetata; (3) Zamorinae, currently placed in Cranaidae, is the sister group to Nomoclastidae, and it is therefore transferred to this family; (4) an expanded Gonyleptidae is recovered, including Manaosbiidae and Cranaidae, but the independence of these families is also recovered because the Gonyleptidae stricto sensu form a clade; (5) the Ampycinae, in spite of many particularities, are recovered inside Gonyleptidae; (6) the zamorine-less Cranaidae are recovered as a monophyletic sister group to Gonyleptidae, but not nested inside it. Jabbastygnus gen. nov. is described in Stygnidae along with its type species J. huttorum, from Colombia.
Much of the debate around speciation and historical biogeography has focused on the role of stabilizing selection on the physiological (abiotic) niche, emphasizing how isolation and vicariance, when associated with niche conservatism, may drive tropical speciation. Yet, recent re-emphasis on the ecological dimensions of speciation points to a more prominent role of divergent selection in driving genetic, phenotypic, and niche divergence. The vanishing refuge model (VRM), first described by Vanzolini and Williams (1981), describes a process of diversification through climate-driven habitat fragmentation and exposure to new environments, integrating both vicariance and divergent selection. This model suggests that dynamic climates and peripheral isolates can lead to genetic and functional (i.e., ecological and phenotypic) diversity, resulting in sister taxa that occupy contrasting habitats with abutting distributions. Here, we provide predictions for populations undergoing divergence according to the VRM that encompass habitat dynamics, phylogeography, and phenotypic differentiation across populations. Such integrative analyses can, in principle, differentiate the operation of the VRM from other speciation models. We applied these principles to a lizard species, Coleodactylus meridionalis, which was used to illustrate the model in the original paper. We incorporate data on inferred historic habitat dynamics, phylogeography and thermal physiology to test for divergence between coastal and inland populations in the Atlantic Forest of Brazil. Environmental and genetic analyses are concordant with divergence through the VRM, yet physiological data are not. We emphasize the importance of multidisciplinary approaches to test this and alternative speciation models while seeking to explain the extraordinarily high genetic and phenotypic diversity of tropical biomes.
Bryophytes constitute an important component of tropical rain forests, which provide microhabitats favorable for their establishment. Bryophytes are also quite responsive to changes in microclimate, which makes them good bioindicators. This study aimed to determine the diversity and distribution of bryophytes in upper and lower montane forests of the Chapada Diamantina region of the state of Bahia, Brazil. To that end, we studied community aspects such as richness, diversity, substrates colonized, life forms and floristic similarity between areas and regions. In 2007 and 2008, we collected specimens from six forest sites, located from the north to the south of the Chapada Diamantina region. We identified a total of 205 infrageneric taxa. In comparison with the lower montane forests, the upper montane forests presented higher diversity and species richness, as well as greater numbers of substrates colonized, life form types, species of restricted geographic distribution and species typical of shaded areas. We also found low similarity in the species composition, the populations of the upper and lower montane forests forming two large and distinct groups. Although presenting relatively high floristic homogeneity among themselves, the Chapada Diamantina areas presented little similarity with those of the Atlantic Forest. This can be explained by the differences between the two regions in terms of environmental conditions, precipitation, seasonality, elevation and continentality.
Gonyleptidae is the second most diverse harvestmen family and the most studied in terms of morphology, behaviour, and ecology. Despite this, few phylogenetic studies have focused on gonyleptids, and those are based on a very limited number of taxa. We addressed this gap by constructing a phylogenetic hypothesis of the family using 101 taxa from all 16 gonyleptid sub-families and four mitochondrial and nuclear loci (COI, 28S rRNA, 12S rRNA, and 16S rRNA). These were analysed under par-simony and likelihood optimality criteria (and using direct optimization for the former). Relationships among Gonyleptoidea and within each subfamily of Gonyleptidae were largely congruent between parsimony and maximum-likelihood approaches. Taxonomic actions from our phylogeny include the following: Tricommatidae, new status, is restored as a family; Metasarcidae, new status, is recognized as a family and considered sister to the Cosmetidae; and Cranainae and Manaosbiinae are suggested as members of Gonyleptidae, restoring Roewer's concept of the family. Within Gonyleptidae, the "K92" group—composed of Sod-reaninae, Caelopyginae, Hernandariinae, Progonyleptoidellinae, and Gonyleptinae—forms a clade, although the latter two subfamilies are not monophyletic. The genus Parampheres is here transferred to Caelopyginae, and "Multumbo" dimorphicus to Gonyleptinae. Gonyleptidae is characterized by the presence of a ventral process on the penis glans and a bifid apophysis on the male coxa IV. The long-legged Mitobatinae can be considered monophyletic only if some short-legged pachylines are included, or if we assume that elongate legs arose twice independently (in the true mitobatine genera and in Longiperna). Pachylinae, the most diverse gonyleptid subfamily, represents several distinct lineages. We further conclude that the traditional use of a small set of morphological characters in the systematics of Gonyleptidae is unable to explain the complex evolution of the family.
There is much evidence to support past contact between the Atlantic and the Amazon forests through the South American dry vegetation diagonal, but the spatiotemporal dynamics of this contact still need to be investigated to allow a better understanding of its biogeographic implications for birds. Here, we combined phylogenetic data with distributional data using a supermatrix approach in order to depict the historical connection dynamics between these biomes for New World suboscines. We examined the variation in divergence time and then compared the spatial distributions of taxon pairs representing old and recent divergences. Our results pointed to two distinct spatiotemporal pathways connecting the Atlantic and the Amazonian forests in the past: (1) old connections (middle to late Miocene) through the current southern Cerrado and Mato Grosso and the transition towards the Chaco and palm savannas of Bolivia and Paraguay; (2) young connections (Pliocene to Pleistocene) that possibly occurred through the Cerrado and Caatinga in northeastern Brazil. We suggest that the main events that played important roles in these connections were geotectonic events during the late Tertiary associated with the uplift of the Andes (old connections) and Quaternary climate changes that promoted the expansion of gallery forest through the Cerrado and Caatinga in northeastern Brazil (young connections). Our results provide the first general temporal and spatial model of how the Atlantic and Amazonian forests were connected in the past, which was derived using bird data.
All the systematic literature (774 references) of the suborder Laniatores of the Americas up
to year 2002 is tabulated to generate a thorough annotated classification. 2372 species in
746 genera of Laniatores of the New World are listed. 26 families of Laniatores are
recognized as valid, of which 21 occur in the New World. The most diverse family is
Gonyleptidae (823 species), followed by Cosmetidae (710 species), both endemic to the
New World. Synonymies, revalidations, replacement names and emended spellings are
done when necessary. The complete list of nomenclatural acts herein proposed is given. The
new family Escadabiidae Kury & Pérez is proposed, the new subfamily Ampycinae Kury is
proposed in Gonyleptidae. Countries and ultramarine departments included are 1) South
America: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, French Guyana, Guyana,
Paraguay, Peru, Suriname, Trinidad and Tobago, Uruguay and Venezuela; 2) Central
America: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama; 3)
Antilles: Bahamas, Bermuda Island, Caicos Islands, Cayman Islands, Cuba, Dominican
Republic, Haiti, Jamaica, Leeward Islands, Netherland Antilles, Puerto Rico, Tortuga Island,
UK Virgin Islands, U.S. Virgin Islands, Windward Islands; 4) North America: Canada,
Greenland, Mexico, U.S.A. First order administrative divisions (departments, provinces,
states) for all most diverse countries are interpolated in the locality names. A list of species
by first order administrative divisions is provided for all countries treated. The most diverse
country is Brazil, with 855 species of Laniatores, followed by Venezuela with 328 species.
An exhaustive list of the depository institutions of the type material with curators and
contacts addresses is given.
Goniosomatine harvestmen have strongly armed pedipalps, generally large bodies and, commonly, very long legs (sometimes more than 20cm), 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-Leito, 1922); G. unicolor (Mello-Leito, 1932); G. carum (Mello-Leito, 1936); Serracutisoma proximum (Mello-Leito, 1922); S. banhadoae (Soares Soares, 1947); S. molle (Mello-Leito, 1933); S. thalassinum (Simon, 1879); S. catarina (Machado, Pinto-da-Rocha Ramires, 2002); S. inerme (Mello-Leito, 1927); S. spelaeum (Mello-Leito, 1933); Heteromitobates inscriptus (Mello-Leito, 1922); H. albiscriptus (Mello-Leito, 1932); Mitogoniella modesta (Perty, 1833); and M. badia (Koch, 1839). Reestablished combinations include: Mitogoniella indistincta Mello-Leito, 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-Leito, 1932 (and its species, G. viridans Mello-Leito, 1932) is removed from Goniosomatinae and considered incertae sedis.
Pseudotrogulus Roewer 1932, described from Amazonia as a cranaine Gonyleptidae, is newly reported from the Brazilian Atlantic Forest, where it is represented by the type species, P. telluris and Pseudotrogulus mirim new species, herein described. These, along with the two monotypic genera Multumbo Roewer 1927, and Piassagera Roewer 1928, are the only Cranainae recorded from the Brazilian Atlantic Forest. A review of these three presumed cranaine genera suggests they are only superficially similar to the Cranainae, and are instead related to the Gonyleptinae. therefore they are newly referred to this subfamily. in the same way, Stygnobates Mello-Leitõ 1927, also from the Brazilian Atlantic Forest, and currently included in the Andean Stygnicranainae, is shown to be more closely related to the progonyleptoidellinae. a closer relationship among Gonyleptinae, Hernandariinae, Progonyleptoidellinae, Caelopyginae and Sodreaninae is suggested, based on genital morphology.
A cladistic analysis of the genera of Mitobatinae and species of Promitobates Roewer, is presented. Only one equally most parsimonious tree was obtained (L=273, CI=0.36; RI=0.73) from a data matrix of 77 equally weighted characters. Mitobatinae is recovered as a monophyletic group, with low Bremer support, sister to ("Pachylinae" + Bourguyiinae). Despirus parvulus, Roewer is sister to the remaining mitobatines, which are divided into two major clades: [1] one that possesses a roughly rectangular body, and males and females with coxa and trochanter IV without large apophyses, including the genera Ischnotherus Kury, Encheiridium Kury, Metamitobates Roewer, Ruschia Mello-Leitão, Mitobatula Roewer, and Mitobates Sundevall; and [2] one that possesses the body roughly pyriform, with evident sexual dimorphism of coxa and trochanter IV, in which males have a large apophysis on coxa IV and tubercles on trochanter IV, including the genera Discocyrtoides Mello-Leitão, Longiperna Roewer, Neoancistrotus Mello-Leitão, and Promitobates. A taxonomic review of Promitobates and a new classification of its species, supported by results of the cladistic analysis, are presented. Based on these results P. intermedius (Mello-Leitão) comb. n., and P. nigripes (Mello-Leitão) comb. n., are revalidated
Longiperna Roewer, 1929 is revised and new records of distribution are presented for the Brazilian Costal Atlantic Rain Forest (from Rio de Janeiro to Santa Catarina states). The following new synonymies are established: Longiperna concolor (Mello-Leitao, 1923) = L. zonata Mello-Leitao, 1935 and L. heliaca B. Soares, 1942; Longiperna coxalis (Roewer, 1943) = L. areolata B. Soares, 1944; Longipema insperata (Soares & Soares, 1947) = L. paranensis Soares & Soares, 1947 and L. curitibana Kury, 2003. Two new species are described: Longipema kuryi sp. nov. (type-locality: Brazil, Rio de Janeiro, Parati); Longipema trembao sp. nov. (type-locality: Brazil, Minas Gerais, Belo Horizonte).
Two new species of troglomorph Pachylinae are described from caves in a xeric biome of eastern Brazil. Both are assigned to extant genera, known from epigean environment – Discocyrtus Holmberg, 1878 and Eusarcus Perty, 1833. These are the first records of Opiliones from the caves “Gruna do Brejo & Verruga” and “Caverna Pedra Furada,” Sistema Lapa Doce in the Brazilian state of Bahia. Both species show moderate troglomorph features such as partial depigmentation and elongation of appendages.
Harvestmen show a high degree of endemism in the Atlantic Rain Forest (eastern coast of Brazil). This biome shows the highest diversity of harvestmen inhabiting Brazil; 2/3 of the species are found in this area. Most of the species are distributed in a few thousand square kilometers, almost always within one mountain range. The similarities of 26 localities were studied, including sites from the Brazilian savanna, using data from recent collections (more than 8,000 specimens) and published data. A cluster analysis using Sørensen´s Coefficient indicated a high degree of endemism of species of harvestmen (similarity indexes below 0.5). It resulted in six main clusters related to the large mountain ranges and near sites. A high variation in richness was observed; 4–64 species per locality. The distribution of 84 species of four recently reviewed subfamilies of Gonyleptidae (Goniosomatinae, Caelopyginae, Progonyleptoidellinae and Sodreaninae) was studied. Eleven areas of endemism, with 3–14 endemic species each, were proposed. A primary Brooks Parsimony Analysis showed a possible first vicariant event splitting the fauna of two northern areas from the rest, and a second event splitting the fauna of southern areas (until 24°35″S) from those areas related to certain mountain ranges in the central Atlantic Rain Forest. The vicariant events were related to the uplifting of the Serra do Mar and the Serra da Mantiqueira, and the appearance of large rivers and climatic changes.
Aim To use the method of parsimony analysis of endemism to identify areas of endemism for passerine birds in the Atlantic Forest, South America, and to compare the locations of these areas with areas previously identified for birds as well as other taxa.
Location The Atlantic Forest, eastern South America.
Methods We analysed a matrix composed of the presence (1) or absence (0) of 140 endemic species in 24 quadrats of 1 × 1 degree distributed along the Atlantic Forest to find the most parsimonious area cladogram.
Results Fourteen most parsimonious cladograms were found and then summarized in a single consensus tree. Four areas of endemism were identified: Pernambuco, Central Bahia, Coastal Bahia, and Serra do Mar.
Main conclusions Avian areas of endemism in the Atlantic Forest have significant generality, as they are highly nonrandom and congruent with those of other groups of organisms. A first hypothesis about the historical relationships among the four areas of avian endemism in the Atlantic Forest is delineated. There is a basal dichotomy among areas of endemism in the Atlantic Forest, with Pernambuco forming a northern cluster and Coastal Bahia, Central Bahia and Serra do Mar comprising a southern cluster. Within the southern cluster, Central Bahia and Serra do Mar are more closely related to each other than to Coastal Bahia.
Heteropachylinae Kury, 1994 is currently considered an early lineage of Gonyleptidae Sundevall, 1833 and includes small harvestmen that occur in the Atlantic Forests, mainly in the Northeast Region of Brazil. The species of Heteropachylinae were spread in different subfamilies until the establishment of the group. As a result of that, until now there were no comparative works on the subfamily in the literature. In this study, the first systematic review for Heteropachylinae was carried out and the first phylogenetic hypothesis for their taxa is proposed, by means of a cladistic approach. A total of 102 morphological characters was coded for 21 terminal taxa: 11 species of Heteropachylinae, two species from the genus Mangaratiba Mello-Leitão, 1940 (considered as Heteropachylinae before this paper), seven other gonyleptids, and one cosmetid. The data were analysed using heuristic search algorithms, resulting in one most parsimonious tree (280 steps, consistency index = 0.47, retention index = 0.63). A parsimony analysis under implied weights was also performed, resulting in three trees. According to the results, Heteropachylinae is a monophyletic and well-supported group if Mangaratiba is excluded. This genus should be returned to where it was originally described – Pachylinae. The subfamily distribution is restricted from Ceará to Espírito Santo Brazilian states. Pseudopucrolia Roewer, 1912 is considered a senior synonym of Melloa Roewer, 1930, Thaumatoleptes Roewer, 1930, and Tribunosoma Roewer, 1943. Two species of Heteropachylus Roewer, 1913, Heteropachylus ramphonotus sp. nov. and Heteropachylus gracilis sp. nov., from Bahia and Espírito Santo Brazilian states, are newly described. Heteropachylus is considered a senior synonym of Chavesincola Soares & Soares, 1946 and Aesotrinoma H. Soares, 1977. A new genus is also described for a new species from Bahia and Espírito Santo Brazilian states, Magnispina neptunus gen. et sp. nov. Lectotypes are designated for the following species: Pseudopucrolia incerta (Mello-Leitão, 1928) comb. nov., Pseudopucrolia discrepans (Roewer, 1943) comb.nov., Pseudopucrolia rugosa (Roewer, 1930) comb. nov., Heteropachylus inexpectabilis (Soares & Soares, 1946) comb. nov. and Heteropachylus spiniger Roewer, 1913.© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, 437–483.
Biogeographical affinities among cloud forests in the Neotropical region were studied through a track approach, by constructing
generalised tracks based on the results of a parsimony analysis of endemicity (PAE). Distributional data on 946 genera and
1,266 species of vascular plants (Pteridophyta, angiosperms, and gymnosperms) from 26 cloud forest patches from Colombia,
Costa Rica, Cuba, Honduras, Jamaica, Mexico, Peru, Puerto Rico, and Venezuela were analysed; and four localities from eastern
and western United States were also included as outgroups. The track analysis identified six generalised tracks: a first one
that includes the majority of the cloud forests of Mexico, Central America, the Antilles, and northern Colombia; a second
one that includes southern Mexico and northern Central America; a third one that includes the mountains in northwestern South
America; a fourth one that includes the mountains in southwestern South America; and two others in western and eastern United
States. It is concluded that the Neotropical cloud forests are closely related and that those of the Caribbean subregion exhibit
complex relationships, which could be due to the complex tectonic history of the area.
Summary: Goniosoma includes large and conspicuous species of harvestmen that occur exclusively in the Brazilian Atlantic forest. This paper reports on the parental activities, defensive behavior, and gregariousness of five species of Goniosoma, and also summarizes the published biological data for the genus. The behavioral patterns within the genus are discussed and a hypothesis for the emergence of gregariousness in the group is presented. Several Goniosoma live inside caves, mainly as trogloxenes. Although species of the genus comprise only 5% of the harvestmen fauna in Brazil, they account for 20.6% of all harvestmen species recorded in Brazilian caves. In addition to several morphological and physiological features that may favor the occupation of caves, species of Goniosoma also present subsocial behavior, which may confer a special advantage in this particular environment. Female protection is crucial for egg survival since predators may consume entire batches in a single night. Most species of Goniosoma form dense diurnal aggregations ranging from three to 200 individuals (mainly subadults and adults of both sexes). In a hypothetical scenario for the evolution of sociality in the Goniosoma, physiological constraints acting on individual harvestmen would lead to a behavioral response for the selection of sites with appropriate microclimatic conditions. The lack of cannibalism among adults and subadults may favor tolerance towards conspecifics, and may represent a pre-adaptation to gregariousness. The derived functions of gregarious behavior may include strengthening of the defensive signal through the collective release of a repugnant secretion, the dilution effect, and promptness in fleeing a predator attack as a consequence of the additional alarm role of scent gland secretions. These defensive functions may overcome the costs of group living and may be responsible for the maintenance of gregariousness in Goniosoma.
Tropical lowland areas have often been seen as the centres of terrestrial species proliferation, but recent evidence suggests that young species may be more frequent in montane areas. Several montane speciation modes have been proposed, but their relative frequencies and predominant evolutionary sequence remain unclear because so few biogeographic and phylogenetic studies have tested such questions. I use morphological data to generate a phylogenetic hypothesis for all 11 species of the riodinid butterfly genus Ithomiola (Riodininae: Mesosemiini: Napaeina). These species are shown here to be all strictly geographically and elevationally allo- or parapatrically distributed with respect to their closest relatives in lowland and montane regions throughout the Neotropics. The overwhelming pattern in Ithomiola is of repeated upward parapatric speciation across an elevational gradient, and the genus appears to provide the clearest example to date of vertical montane speciation. All of the young derived species are montane and all of the old basal species are confined to the lowlands, supporting the hypothesis of montane regions largely as 'species pumps' and lowland regions as 'museums'. Possible reasons for the post-speciation maintenance of parapatric ranges in Ithomiola are discussed.
To investigate the past spatial dynamics of Atlantic Forest (AF)‘island-like’ enclaves in northeastern Brazil, we employed ecological niche modeling (ENM)using a set of georeferenced records of 13 woody plant species. ENMs were constructed using the Ensemble forecasting approach and were projected into past climatic conditions for the last interglacial period (LIG, 120 kyr), last glacial maximum (LGM, 21 kyr), and the Middle Holocene (MH, 6 kyr). Our results suggest an expansion of wetter forests during the LGM into areas currently covered by the Caatinga seasonally dry woodlands, with recent retraction to the current distribution. Central AF islands located south of the São Francisco River underwent a different history compared to Northern AF islands north of this river: the former were mostly connected to coastal AF since the LIG (with a very recent separation), whereas the latter presented a more dynamic historical distribution. Results reveal contrasting spatiotemporal histories of forest instability and isolation across the various enclaves, supporting three main biogeographic hypotheses: i)moderate connectivity with coastal AF and a recent population bottleneck in the Araripe and Pernambuco/Paraíba enclaves, ii)low connectivity to, and long-term isolation from, other enclaves, as well as recent population bottleneck, in the northernmost North Ceará enclaves; and iii)high connectivity with coastal AF and recent population expansion in the southernmost Chapada Diamantina enclaves. Future comparative phylogeography studies will largely aid in assessing the herein proposed biogeographic scenarios during the highly dynamic recent history of the AF enclaves.
Version 1.5 of the computer program TNT completely integrates landmark data into phylogenetic analysis. Landmark data consist of coordinates (in two or three dimensions) for the terminal taxa; TNT reconstructs shapes for the internal nodes such that the difference between ancestor and descendant shapes for all tree branches sums up to a minimum; this sum is used as tree score. Landmark data can be analysed alone or in combination with standard characters; all the applicable commands and options in TNT can be used transparently after reading a landmark data set. The program continues implementing all the types of analyses in former versions, including discrete and continuous characters (which can now be read at any scale, and automatically rescaled by TNT). Using algorithms described in this paper, searches for landmark data can be made tens to hundreds of times faster than it was possible before (from T to 3T times faster, where T is the number of taxa), thus making phylogenetic analysis of landmarks feasible even on standard personal computers.
A new species of the Bokermannohyla circumdata group, B. diamantina sp. nov., is described from Serra dos Barbados, Municipality of Abaíra, State of Bahia, Brazil. This locality belongs to Chapada Diamantina, a semiarid region in central Bahia. This is the first record of a species of the B. circumdata group from the Brazilian semiarid. Bokermannohyla diamantina is diagnosed by the medium size (snout-vent length 47.0-51.7 mm in adult specimens); head length 2.8-2.9 times smaller than snout-vent length; tympanum medium-sized, its diameter 1.3-1.5 times smaller than eye diameter (tympanum diameter/snout-vent length 0.07); thigh and tibia lengths 0.48-0.52 and 0.49-0.51 of snout-vent length, respectively; dorsum medium to blackish brown, with weak transverse dark brown bars; flanks, anterior, and posterior surfaces of thighs with dark transverse brown stripes, lacking additional thin stripes between them; distinct vocal slits in adult males; advertisement call with 2-4 notes, and dominant frequency from 0.39-0.56 kHz.
Abstract
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.
The complex geography of the Neotropical montane system is a natural laboratory for population divergence. Understanding which geographic barriers (lowland barriers, arid river valleys, and montane barriers above the tree line separate these regions of endemism) are instrumental in promoting and maintaining population divergence is an important step in preserving genetic diversity and endemism within the region. Here, I analyze patterns of genetic differentiation between 16 predefined regions of endemism for 43 co-distributed zoogeographic species complexes of Neotropical montane forest birds. The analysis shows that lowland barriers generate the highest levels of genetic differentiation, while barriers above the tree line in the Andes show the least. Within the Andes, arid river valleys promote population divergence to varying degrees. The Río Marañón shows the greatest effect, but the Río Apurímac and Río Quinimari are also associated with extensive genetic differentiation, while genetic divergence across other river valleys is generally weak. Most barriers are associated with a wide span of divergence times, supporting a protracted history of dispersal postdating barrier formation. If the goal is to maintain genetic diversity, preservation of populations within each region of endemism would help to ensure the continued survival of evolutionarily distinct lineages within species. Considering the alarming rate of deforestation in Neotropical montane regions, preservation of suitable tracts of montane forest is needed within each region of endemism, with special emphasis placed on endemism regions separated by lowland barriers and by deep intermontane river valleys.
Abstract Aim To examine how the genetic diversity of selected taxa of forest-dwelling small mammals is distributed between and within the major rain forest domains of Amazonia and Atlantic Forest and the intervening interior forests of Brazil, as inferred by the relationships between gene genealogies and geography. I also addressed the historical importance of the central Brazilian forests in connecting Amazon and Atlantic Forest populations of rodents and marsupials.
Methods I evaluated variation in the mitochondrial cytochrome b gene to estimate the levels of sequence divergence between those taxa occurring throughout the Amazon, Atlantic Forest, and forests in the Cerrado and Caatinga regions. I inferred the hierarchical relationships between haplotypes, populations and formal taxa using the cladistic approach of maximum parsimony. I compared areas and the clades identified by superimposing cladograms on the geographical distribution of samples. The degree of concordance both in phylogeny and the depth of the nodes in these phylogenies, in addition to patterns of geographical distribution of clades, permitted me to make inferences on how, when and where the taxa differentiated.
Results Sequence similarity is often greater between samples from the Atlantic Forest and either Amazon or central Brazilian forests than it is within each of the two rain forest domains. The Atlantic Forest clades are either not reciprocally monophyletic or are the sister group to all the other clades. There is some indication of northern and southern components in the Atlantic Forest. Given the geographical distribution of clades and the relatively deep levels of divergence, the central Brazilian area does not behave as a separate region but is complementary to either Amazon or Atlantic Forest. Patterns of area relationships differ across taxa, suggesting that different processes and/or historic events affected the diversification within each lineage.
Main conclusions The Amazon and the Atlantic forests are not exclusive in terms of their small mammal faunas; both overlap broadly with taxa occurring in gallery forests and dry forests in central Brazil. Central Brazilian forests are an integral part of the evolutionary scenario of lowland small mammals, playing an important role as present and past habitats for rain forest species. Therefore, representatives from this area should always be included in analyses of the evolutionary history of lowland rain forest faunas. The incongruence of branching patterns among areas is in agreement with recent results presented for Neotropical passerine birds and indicates that a single hypothesis of Neotropical area relationships is unlikely. These findings reinforce the idea that speciation in the Neotropics will not be explained by any single model of vicariance or climatic changes.
The arachnids of the order Opiliones (harvestmen) produce substances used in defense. In the present paper, we analyzed 22 species of Gonyleptidae to explore the use of defensive substances in taxonomy and evolutionary biology. Thirty-seven different compounds were detected, 18 of which were preliminarily identified. These compounds were mapped onto a phylogenetic tree showing the relationships within the Gonyleptidae. Data from Cosmetidae were used as an outgroup. Five ketones and six alkyl phenols were reported for the first time in harvestmen.
It is generally accepted that accentuated global climatic cycles since the Plio-Pleistocene (2.8 Ma ago) have caused the intermittent fragmentation of forest regions into isolated refugia thereby providing a mechanism for speciation of tropical forest biota contained within them. However, it has been assumed that this mechanism had its greatest effect in the species rich lowland regions. Contrary evidence from molecular studies of African and South American forest birds suggests that areas of recent intensive speciation, where mostly new lineages are clustered, occur in discrete tropical montane regions, while lowland regions contain mostly old species. Two predictions arise from this finding. First, a species phylogeny of an avian group, represented in both lowland and montane habitats, should be ordered such that montane forms are represented by the most derived characters. Second, montane speciation events should predominate within the past 2.8 Ma. In order to test this model I have investigated the evolutionary history of the recently radiated African greenbuls (genus Andropadus), using a molecular approach. This analysis finds that montane species are a derived monophyletic group when compared to lowland species of the same genus and recent speciation events (within the Plio-Pleistocene) have exclusively occurred in montane regions. These data support the view that montane regions have acted as centres of speciation during recent climatic instability.
Although many studies have documented the effect of glaciation on the evolutionary history of Northern Hemisphere flora and fauna, this study is the first to investigate how the indirect aridification of Africa caused by global cooling in response to glacial cycles at higher latitudes has influenced the evolutionary history of an African montane bird. Mitochondrial DNA sequences from the NADH 3 gene were collected from 283 individual Starred Robins (Pogonocichla stellata, Muscicapoidea). At least two major vicariant events, one that separated the Albertine Rift from all but the Kenyan Highlands around 1.3-1.2 Myrs BP, and another that separated the Kenyan Highlands from the northern Eastern Arc, and the northern Eastern Arc from the south-central Eastern Arc between 0.9 and 0.8 Myrs BP appear to underlie much of the observed genetic diversity and structure within Starred Robin populations. These dates of divergence suggest a lack of recurrent gene flow; although the Albertine Rift and south-central Eastern Arc share haplotypes, based on coalescent analyses this can confidently be accounted for by ancestral polymorphism as opposed to recurrent gene flow. Taken collectively, strong evidence exists for recognition of four major ancestral populations: (1) Kenyan Highlands (subspecies keniensis), (2) Albertine Rift (ruwenzori), (3) northern Eastern Arc (helleri), and (4) south-central Eastern Arc, Ufipa and the Malawi Rift (orientalis). The estimated divergence times cluster remarkably around one of the three estimated peaks of aridification in Africa during the Plio-Pleistocene centred on 1 Myrs BP. Further, time to most recent common ancestor (TMRCA) estimates (1.7-1.6 Myrs BP) of gene divergence between the Albertine Rift and the other montane highlands corresponds closely with a second estimated peak of aridification at about 1.7 Myrs BP. Collectively, these results suggest that aridification of Africa in response to glaciation at higher latitudes during the Pleistocene has had a profound influence on montane speciation in east and central Africa.
The arachnids of the order Opiliones (harvestmen) have a pair of scent glands opening at the sides of the body, the substances of which are used in defense. Several types of behavioral, morphological and chemical defensive mechanisms have been assigned to the order as a whole, although some of these tactics were restricted to particular groups. Only around 25 species have been studied from this perspective so far. In the present paper, we analyzed 33 species (mostly from the largest harvestmen family, the Gonyleptidae) aiming at recognizing the usefulness of the defensive characters in taxonomy and evolutionary biology. We observed the morphology of the gland opening (ozopore) area and the defensive behavior, and their relationship, and mapped these traits on an available phylogenetic hypothesis of relationship within Gonyleptidae. As outgroups, we analyzed Cosmetidae and Stygnidae. Combining the observed behavioral characters of the emission of defensive secretion (near the ozopore, with liquid displacement through an integumentary groove, or in form of a jet) with the morphological types of the gland opening (direction of the integumentary dome that surrounds the gland opening, presence of two openings and the relationship between their sizes, and presence of a V-shaped cut at the anterior opening), we recognized eight patterns. In addition, we could examine the evolution of such traits within Gonyleptidae.
Description of Mitogoniella mucuri sp
- L N Ázara
- M B Dasilva
- R L Ferreira
Ázara, L.N., DaSilva, M.B. & Ferreira, R.L. (2013) Description of Mitogoniella mucuri sp. nov. (Opiliones: Gonyleptidae) and
considerations on polymorphic traits in the genus and Gonyleptidae. Zootaxa, 3786 (1), 69-81.
https://doi.org/10.11646/zootaxa.3736.1.3
Die Arachniden getreu nach der Natur abgebildet und beschrieben
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Koch, C.L. (1839a) Die Arachniden getreu nach der Natur abgebildet und beschrieben. Band 7. Parts 1-5. C. H. Zeh'schen
Buchhandlung, Nuremberg, 106 pp.
Annals and Magazine of Natural History
- C F Mello-Leitão
Mello-Leitão, C.F. (1922) Some new Brazilian Gonyleptidae. Annals and Magazine of Natural History, 9, 329-348.
https://doi.org/10.1080/00222932208632683
Opiliões Laniatores do Brasil. Archivos do Museu Nacional do Rio de Janeiro
- C F Mello-Leitão
Mello-Leitão, C.F. (1923) Opiliões Laniatores do Brasil. Archivos do Museu Nacional do Rio de Janeiro, 24, 107-197.
Novos Gonyleptidae do Brasil meridional. Arquivos da Escola Superior de Agricultura e Medicina Veterinária
- C F Mello-Leitão
Mello-Leitão, C.F. (1933) Novos Gonyleptidae do Brasil meridional. Arquivos da Escola Superior de Agricultura e Medicina
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A propósito de alguns opiliões novos
- C F Mello-Leitão
Mello-Leitão, C.F. (1935) A propósito de alguns opiliões novos. Memórias do Instituto Butantan, 9, 369-411.
Notas sobre opiliões
- C F Mello-Leitão
Mello-Leitão, C.F. (1936) Notas sobre opiliões. Boletim do Museu Nacional do Rio de Janeiro, 12 (3-4), 1-41.
Sete gêneros e vinte e oito espécies de Gonyleptidae. Arquivos de Zoologia do Estado de São Paulo
- C F Mello-Leitão
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1 (1), 1-52.