An analysis of the floristic composition and diversity of Amazonian forests including those of the Guiana Shield

A large number of newly published and unpublished hectare plots in Amazonia and the Guiana Shield area allow an analysis of family composition and testing of hypotheses concerning alpha-diversity in the south American rain forest. Using data from 94 plots the family-level floristic patterns in wet tropical South America are described. To test diversity patterns, 268 plots are used in this large area. Contrary to a widely held belief, western Amazonian plots are not necessarily the most diverse. Several central Amazonian plots have equal or even higher tree diversity. Annual rainfall is not a good estimator for tree diversity in the Amazonia area and Guiana shield. Plots in the Guiana Shield area (and eastern Amazonia) usually have lower diversity than those in central or western Amazonia. It is argued that this is not because of low rainfall or low nutrient status of the soil but because of the small area of the relatively isolated rain forest area in eastern Amazonia and the Guiana Shield. The low diversity on nutrient-poor white sand soils in the Amazon basin is not necessarily due to their Low nutrient status but is, at least partly, caused by their small extent and fragmented nature.

Tometes camunani (Characiformes: Serrasalmidae), a new species of phytophagous fish from the Guiana Shield, rio Trombetas basin, Brazil

Uma espécie nova de Serrasalmidae, Tometes camunani, é descrita para as drenagens superiores da bacia do rio Trombetas, estado do Pará, Brasil. A espécie nova distingue-se dos congêneres pela presença de uma ligeira concavidade no neurocrânio na altura do frontal (vs. concavidade ausente, perfil dorsal do neurocrânio reto). Também pode ser adicionalmente distinguido dos seus congêneres por possuir dentes com a cúspide central mais alta e cume agudo (vs. cúspide central mais baixa e com cume arredondado em T. trilobatus), a boca terminal (vs. boca orientada para cima em T. lebaili), e 12-26 espinhos pré-pélvicos (vs. 0-9 em T. makue). A espécie nova é estritamente reofílica, como as demais espécies de Tometes, e ocorre exclusivamente nas zonas encachoeiradas dos rios de escudo, biótopos complexos, frágeis e ameaçados por ações antropogênicas. Uma chave de identificação para as espécies do grupo Myleus é apresentada.

Multiple Quaternary Refugia in the Eastern Guiana Shield Revealed by Comparative Phylogeography of 12 Frog Species

The Guiana Shield (GS) is one of the most pristine regions of Amazonia and biologically one of the richest areas on Earth. How and when this massive diversity arose remains the subject of considerable debate. The prevailing hypothesis of Quaternary glacial refugia suggests that a part of the eastern GS, among other areas in Amazonia, served as stable forested refugia during periods of aridity. However, the recently proposed disturbance-vicariance hypothesis proposes that fluctuations in temperature on orbital timescales, with some associated aridity, have driven Neotropical diversification. The expectations of the temporal and spatial organization of biodiversity differ between these two hypotheses. Here, we compare the genetic structure of 12 leaf-litter inhabiting frog species from the GS lowlands using a combination of mitochondrial and nuclear sequences in an integrative analytical approach that includes phylogenetic reconstructions, molecular dating, and Geographic Information System methods. This comparative and integrated approach overcomes the well-known limitations of phylogeographic inference based on single species and single loci. All of the focal species exhibit distinct phylogeographic patterns highlighting taxon-specific historical distributions, ecological tolerances to climatic disturbance, and dispersal abilities. Nevertheless, all but one species exhibit a history of fragmentation/isolation within the eastern GS during the Quaternary with spatial and temporal concordance among species. The signature of isolation in northern French Guiana (FG) during the early Pleistocene is particularly clear. Approximate Bayesian Computation supports the synchrony of the divergence between northern FG and other GS lineages. Substructure observed throughout the GS suggests further Quaternary fragmentation and a role for rivers. Our findings support fragmentation of moist tropical forest in the eastern GS during this period when the refuge hypothesis would have the region serving as a contiguous wet-forest refuge.

Cladistic analysis of the Stygninae and description of a new species of Protimesius Roewer, 1913 (Opiliones: Stygnidae)

A new species of Stygnidae is described from the state of Bahia, Brazil. Protimesius bahiensis sp. nov. can be distinguished from the remaining species of the genus by the combination of: male femur IV unarmed and cylindrical; male patella IV with a row of large dorsal acute tubercles, increasing in size distally and male tibia IV with one mesodistal tubercle; ventral plate of the penis with three pairs of distal curved setae and one pair of intermediate setae, smaller than the rest. A cladistic analysis of the subfamily is presented. Stygninae is divided in two groups of genera: (Ricstygnus, Stygnus, Sickesia), with a wide distribution and (Pickeliana (Protimesius (Phareus (Stenophareus (Auranus (Verrucastygnus, Stenostygnoides)))))), associated to the Guiana Shield, Amazon basin and Northeastern Brazil. The monophyly of Protimesius is supported by the apex of pedipalpal tibia sockets bifid (homoplastically present in Verrucastygnus and Stenostygnoides) and by the presence of scopulae with non-spatulated hairs.

Phylogenetic position of Dendropsophus gaucheri (Lescure and Marty 2000) highlights the need for an in-depth investigation of the phylogenetic relationships of Dendropsophus (Anura: Hylidae)

Dendropsophus gaucheri is a recently described species which inhabits open areas of the eastern part of the Guiana Shield and is currently assigned to the D. parviceps species group based on the presence of a subocular cream spot. Herein we investigate its phylogenetic position including material from the type locality and newly documented populations from Suriname and Brazil based on mtDNA sequences. The species, as well as D. riveroi which is assigned to the D. minimus species group, were recovered nested within the D. microcephalus species group which implies the paraphyly of the three Dendropsophus species groups. Such result, along with other evidences, highlights the need for a thorough revision of the genus. The genetic distances among D. gaucheri samples studied are low confirming their conspecificity and suggesting recent connections among populations from open areas currently isolated by rainforest in the lowlands of the Guiana Shield.

THE RIO APA CRATON IN MATO GROSSO DO SUL (BRAZIL) AND NORTHERN PARAGUAY: GEOCHRONOLOGICAL EVOLUTION, CORRELATIONS AND TECTONIC IMPLICATIONS FOR RODINIA AND GONDWANA

The Rio Apa cratonic fragment crops out in Mato Grosso do Sul State of Brazil and in northeastern Paraguay. It comprises Paleo-Mesoproterozoic medium grade metamorphic rocks, intruded by granitic rocks, and is covered by the Neoproterozoic deposits of the Corumbi and Itapocurni Groups. Eastward it is bound by the southern portion of the Paraguay belt. In this work, more than 100 isotopic determinations, including U-Pb SHRIMP zircon ages, Rb-Sr and Sm-Nd whole-rock determinations, as well as K-Ar and Ar-Ar mineral ages, were reassessed in order to obtain a complete picture of its regional geological history. The tectonic evolution of the Rio Apa Craton starts with the formation of a series of magmatic arc complexes. The oldest U-Pb SHRIMP zircon age comes from a banded gneiss collected in the northern part of the region, with an age of 1950 +/- 23 Ma. The large granitic intrusion of the Alumiador Batholith yielded a U-Pb zircon age of 1839 +/- 33 Ma, and from the southeastern part of the area two orthogneisses gave zircon U-Pb ages of 1774 +/- 26 Ma and 1721 +/- 25 Ma. These may be coeval with the Alto Terere metamorphic rocks of the northeastern corner, intruded in their turn by the Baia das Garcas granitic rocks, one of them yielding a zircon U-Pb age of 1754 +/- 49 Ma. The original magmatic protoliths of these rocks involved some crustal component, as indicated by the Sm-Nd TDm model ages, between 1.9 and 2.5 Ga. Regional Sr isotopic homogenization, associated with tectonic deformation and medium-grade metamorphism occurred at approximately 1670 Ma, as suggested by Rb-Sr whole rock reference isochrons. Finally, at 1300 Ma ago, the Ar work indicates that the Rio Apa Craton was affected by widespread regional heating, when the temperature probably exceeded 350 degrees C. Geographic distribution, age and isotopic signature of the fithotectonic units suggest the existence of a major suture separating two different tectonic domains, juxtaposed at about 1670 Ma. From that time on, the unified Rio Apa continental block behaved as one coherent and stable tectonic unit. It correlates well with the SW corner of the Amazonian Craton, where the medium-grade rocks of the Juruena-Rio Negro tectonic province, with ages between 1600 and 1780 Ma, were reworked at about 1300 Ma. Looking at the largest scale, the Rio Apa Craton is probably attached to the larger Amazonian Craton, and the actual configuration of southwestern South America is possibly due to a complex arrangement of allochthonous blocks such as the Arequipa, Antofalla and Pampia, with different sizes, that may have originated as disrupted parts of either Laurentia or Amazonia, and were trapped during later collisions of these continental masses.

Phylogenetic position of Dendropsophus gaucheri (Lescure and Marty 2000) highlights the need for an in-depth investigation of the phylogenetic relationships of Dendropsophus (Anura: Hylidae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Reinforcing and expanding the predictions of the disturbance vicariance hypothesis in Amazonian harlequin frogs: A molecular phylogenetic and climate envelope modelling approach

The disturbance vicariance hypothesis (DV) has been proposed to explain speciation in Amazonia, especially its edge regions, e. g. in eastern Guiana Shield harlequin frogs (Atelopus) which are suggested to have derived from a cool-adapted Andean ancestor. In concordance with DV predictions we studied that (i) these amphibians display a natural distribution gap in central Amazonia; (ii) east of this gap they constitute a monophyletic lineage which is nested within a pre-Andean/western clade; (iii) climate envelopes of Atelopus west and east of the distribution gap show some macroclimatic divergence due to a regional climate envelope shift; (iv) geographic distributions of climate envelopes of western and eastern Atelopus range into central Amazonia but with limited spatial overlap. We tested if presence and apparent absence data points of Atelopus were homogenously distributed with Ripley's K function. A molecular phylogeny (mitochondrial 16S rRNA gene) was reconstructed using Maximum Likelihood and Bayesian Inference to study if Guianan Atelopus constitute a clade nested within a larger genus phylogeny. We focused on climate envelope divergence and geographic distribution by computing climatic envelope models with MaxEnt based on macroscale bioclimatic parameters and testing them by using Schoener's index and modified Hellinger distance. We corroborated existing DV predictions and, for the first time, formulated new DV predictions aiming on species' climate envelope change. Our results suggest that cool-adapted Andean Atelopus ancestors had dispersed into the Amazon basin and further onto the eastern Guiana Shield where, under warm conditions, they were forced to change climate envelopes. © 2010 The Author(s).

Systematics of spiny-backed treefrogs (Hylidae: Osteocephalus): An Amazonian puzzle

Spiny-backed tree frogs of the genus Osteocephalus are conspicuous components of the tropical wet forests of the Amazon and the Guiana Shield. Here, we revise the phylogenetic relationships of Osteocephalus and its sister group Tepuihyla, using up to 6134 bp of DNA sequences of nine mitochondrial and one nuclear gene for 338 specimens from eight countries and 218 localities, representing 89% of the 28 currently recognized nominal species. Our phylogenetic analyses reveal (i) the paraphyly of Osteocephalus with respect to Tepuihyla, (ii) the placement of 'Hyla' warreni as sister to Tepuihyla, (iii) the non-monophyly of several currently recognized species within Osteocephalus and (iv) the presence of low (<1%) and overlapping genetic distances among phenotypically well-characterized nominal species (e.g. O. taurinus and O. oophagus) for the 16S gene fragment used in amphibian DNA barcoding. We propose a new taxonomy, securing the monophyly of Osteocephalus and Tepuihyla by rearranging and redefining the content of both genera and also erect a new genus for the sister group of Osteocephalus. The colouration of newly metamorphosed individuals is proposed as a morphological synapomorphy for Osteocephalus. We recognize and define five monophyletic species groups within Osteocephalus, synonymize three species of Osteocephalus (O. germani, O. phasmatus and O. vilmae) and three species of Tepuihyla (T. celsae, T. galani and T. talbergae) and reallocate three species (Hyla helenae to Osteocephalus, O. exophthalmus to Tepuihyla and O. pearsoni to Dryaderces gen. n.). Furthermore, we flag nine putative new species (an increase to 138% of the current diversity). We conclude that species numbers are largely underestimated, with most hidden diversity centred on widespread and polymorphic nominal species. The evolutionary origin of breeding strategies within Osteocephalus is discussed in the light of this new phylogenetic hypothesis, and a novel type of amplexus (gular amplexus) is described. © 2013 The Norwegian Academy of Science and Letters.

Diversidade de espécies no complexo Monodelphis brevicaudata (Didelphimorphia:Didelphidae), inferida por dados moleculares e morfológicos

O complexo de espécies M. brevicaudata possui distribuição reconhecida para o Norte da América do sul e compreende três espécies descritas ‐ M. brevicaudata, M. glirina, e M. palliolata ‐ e duas não descritas, reconhecidas em estudos prévios. A delimitação de espécies baseada somente em caracteres morfológicos é complicada, de forma que diversos táxons nominais já foram associados ao grupo e diversos arranjos taxonômicos foram propostos. Os poucos estudos baseados em dados moleculares que incluíram espécimes do complexo brevicaudata revelaram altas taxas de divergência genética. Este trabalho buscou elucidar a sistemática do complexo de espécies M. brevicaudata através do estudo dos padrões de variação morfológica e genética. Para tal, desenvolvemos análises filogenéticas baseadas em dois genes mitocondriais: citocromo b e 16 S rDNA. Adicionalmente, estudamos a morfologia externa e craniana dos espécimes, investigando a existência de congruência entre a variação genética e morfológica. As análises morfológicas foram, em geral, congruentes com as moleculares, as quais indicaram os mesmos clados em todas as análises filogenéticas. Foram formalmente reconhecidas nove espécies para o complexo. Monodelphis brevicaudata, M. palliolata e M. glirina são consideradas espécies válidas; M. touan é revalidado da sinonímia de M. brevicaudata e duas espécies novas são descritas e nomeadas; a espécie M. domestica provou ser intimamente relacionada a espécimes do grupo brevicaudata, sendo aqui considerada como integrante do referido grupo; duas espécies reconhecidas como distintas permanecem sem uma descrição formal; M. maraxina é sinonimizada com M. glirina. Foi observado dimorfismo sexual para as espécies estudadas, sendo que para as duas espécies estatisticamente testadas (teste T de student), M. glirina e M. sp. nov. “Trombetas”, os machos apresentaram crânios significativamente maiores que as fêmeas. Rios de grande porte parecem ter participado na diferenciação genética e estruturação filogeográfica das espécies. O padrão filogeográfico encontrado sugere ao menos dois centros de diversificação para o grupo, um no escudo das Guianas, envolvendo as espécies ao norte do rio Amazonas, e outro no escudo brasileiro, envolvendo M. glirina e M. domestica.

Deformação das rochas siliciclásticas paleoproterozoicas do Grupo Araí como exemplo das reativações de falhas do embasamento, Serra do Tepequém, Roraima, norte do Brasil

As rochas siliciclásticas da Serra do Tepequém são correlacionadas aos Grupos Araí e Suapi, pertencentes ao Supergrupo Roraima de idade paleoproterozoica. A análise estrutural destas rochas revelou que o acamamento exibe arranjos com mergulhos preferencialmente para SE e NW, individualizados em domínios limitados por zonas de falhas oblíquas sinistrais com rejeitos normais e inversos, com direção NE-SW. Essa estruturação é formada por dobras forçadas quilométricas do tipo kink bands e chevrons. O novo arcabouço geométrico observado é característico de um ambiente de deformação de nível crustal raso a médio. A história tectônica é controlada por reativações dos planos de fraqueza preexistentes nas tramas antigas, dúcteis, do embasamento. Esse modelo diverge dos regionais prévios para a região, os quais consideram as dobras existentes como produtos de ambiente dúctil sob tectônica colisional. Os resultados evidenciam a importância da presença de estruturas antigas do embasamento, relacionado ao Escudo das Guianas, como controladoras da geometria das rochas da Serra do Tepequém, em ambiente rúptil.

A new distinctively banded species of Pseudolithoxus (Siluriformes: Loricariidae) from the upper Orinoco River

Pseudolithoxus kelsorum is described as a new species based on type material from the upper Orinoco in Amazonas State, Venezuela. Pseudolithoxus kelsorum is diagnosed from all other Pseudolithoxus by having dark brown to black base color with eight to 11 (usually nine) light yellow vertical or oblique transversal bands between orbits and caudal fin, bands wide and rarely but sometimes incomplete or contorted as swirls (vs. dark brown to black base color with 18 or more thin, light yellow, frequently contorted transversal bands between orbits and caudal fin in P. tigris; black base color with small white spots in P. anthrax and P. nicoi; and light brown base color with dark brown to black spots in P. dumus). Distributions of P. kelsorum and other Ancistrini taxa support the presence of a zoogeographic filter limiting fish distributions across a reach of the Orinoco River between the Ventuari-Orinoco confluence and the Maipures rapids.

Taxonomic revision of extant Doras Lacepede, 1803 (Siluriformes: Doradidae) with descriptions of three new species

Extant Doras are newly diagnosed among Doradidae by the unique combination of maxillary barbels long and fimbriate; mesethmoid with anterior lateral margins converging towards narrow tip; single anterior cranial fontanel contained largely within frontals and anteriorly by mesethmoid (posterior cranial fontanel occluded); anterior nuchal plate wide, pentaganol or roughly hexagonal, sharing distinct lateral suture with epioccipital and isolating supraoccipital from middle nuchal plate; nuchal foramina absent; coracoid process short, posterior tip falling well short of that of postcleithral process; dentary with acicular teeth; and skin immediately ventral to postcleithral process perforated with conspicuous pores. One fossil species, dagger D. dioneae, and two nominal extant species, D. carinatus and D. micropoeus, are recognized as valid and the latter two redescribed. Three additional extant species, D. phlyzakion, D. higuchii and D. zuanoni, are newly described from the middle Amazon and tributaries, lower Amazon tributaries and rio Araguaia (Tocantins drainage), respectively. Doras phlyzakion and D. zuanoni form a monophyletic group that is found in lowland, lentic habitats, and is characterized by multiple conspicuous pores in skin on breast and abdomen, a trait unique among doradids and rare if not unique among all catfishes. The remaining extant species, D. carinatus, D. higuchii and D. micropoeus, with uncertain relationships, are found in upland, lotic habitats. The occurrence of D. carinatus in the Orinoco basin suggests a historical link between right-bank tributaries of the lower Orinoco (e.g., Caroni) draining the western Guiana Shield and more eastern rivers (e.g., Cuyuni-Essequibo) that drain the Shield directly into the Atlantic Ocean. A key to extant species is provided, a neotype is designated for Silurtis carinatus Linnaeus 1766, and Mormyropsis Miranda Ribeiro, 1911, is placed in the synonymy of Doras Lacepede, 1803.

Tree height integrated into pantropical forest biomass estimates

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- andWeibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (?40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8Mgha?1 (range 6.6 to 112.4) to 8.0Mgha?1 (?2.5 to 23.0). For all plots, aboveground live biomass was ?52.2 Mgha?1 (?82.0 to ?20.3 bootstrapped 95%CI), or 13%, lower when including H estimates, with the greatest relative reductions in estimated biomass in forests of the Brazilian Shield, east Africa, and Australia, and relatively little change in the Guiana Shield, central Africa and southeast Asia. Appreciably different stand structure was observed among regions across the tropical continents, with some storing significantly more biomass in small diameter stems, which affects selection of the best height models to reduce uncertainty and biomass reductions due to H. After accounting for variation in H, total biomass per hectare is greatest in Australia, the Guiana Shield, Asia, central and east Africa, and lowest in eastcentral Amazonia, W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if tropical forests span 1668 million km2 and store 285 Pg C (estimate including H), then applying our regional relationships implies that carbon storage is overestimated by 35 PgC (31?39 bootstrapped 95%CI) if H is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree H is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of tropical carbon stocks and emissions due to deforestation.