Areas of endemism and spatial diversification of the Muscidae (Insecta: Diptera) in the Andean and Neotropical regions

Aim We present a biogeographical analysis of the areas of endemism and areas of diversification in the Muscidae. This analysis searched for geographical patterns in the Muscidae to reconstruct elements of the evolutionary biogeographical history of this insect family. Location Andean and Neotropical regions. Method We constructed a geographic database of 728 species from the literature and museum specimens. Areas of endemism were established by parsimony analysis of endemicity (PAE) based on grids of two different sizes: 5° (550 × 550 km) and 2° (220 × 220 km). Areas of diversification were delimited by track analysis that also included phylogenetic information. This process was independently applied to 11 genera. For each genus, we plotted generalized tracks generated by sister species on a map. When these generalized tracks supported inter‐generic nodes they were manually contoured and inferred to be areas of diversification for the Muscidae. Results Thirteen endemic areas were found using the 5° grid, and eight endemic areas resulted from the 2° grid. Ten areas were in agreement with previous studies, and 11 were new. Amazonian and Atlantic areas of diversification agreed with previous areas for the genus Polietina, and new areas of diversification were found in Panama and in central Chile. Main conclusions Six spatial patterns in the Muscidae were identified: (1) areas of endemism in both Pampa and Puna provinces were established with species whose distributions had not previously been analysed; (2) a new area of endemism was established in extreme southern South America, in Tierra del Fuego; (3) two new areas of diversification, which include Panama and central Chile, were identified; (4) a spatial association was identified between the separation of Chiloe Island from the continent and the diversification in Andean species; (5) a north–south track axis and latitudinal node intervals were identified, interpreted as spatial responses to glaciation or glacial retreat in the Andes; and (6) a spatial coincidence of areas of endemism, of diversification and high species richness in the Muscidae was discovered. The analysis of a complete database and the recognition of areas of diversification are extremely important in elucidating novel biogeographical patterns, which will in turn contribute to a better understanding of the geographical patterns of evolution in the Muscidae.     

Tropical niche conservatism as a historical narrative hypothesis for the Neotropics: a case study using the fly family Muscidae

Aim We evaluate the extent to which the tropical conservatism hypothesis can explain the evolutionary development of the Muscidae. Furthermore, we compare the geographical patterns of muscid phylogenetic structure with biogeographical regions that have been identified for Neotropical insects. Location Central and South America. Methods We modelled the geographic distributions of 658 species using Maxent and 19 environmental variables. A generic‐level supertree of the Muscidae was assembled using matrix representation with parsimony and used to map the geographic pattern of mean root distance (MRD), a metric of the relative evolutionary development of assemblages. Regression models (ordinary least squares and regression trees) were used to examine temperature and other environmental correlates of MRD to explore potential environmental drivers of muscid diversification. We used the regression tree results to recognize variable intervals that best explained MRD, and these intervals were mapped to recognize and compare with biogeographical regions of Neotropical insects. Results The geographic pattern of MRD was consistent with the tropical conservatism hypothesis: species in genera that diversified relatively early, as measured by their distance from the tree root, dominate lowland tropical South America, whereas species in genera that diversified more recently occupy extra‐tropical areas, sub‐Antarctic areas and the Andean highlands. Temperature was the strongest correlate of MRD. Three biogeographical regions were recognized and they coincided with two regions known for insects. Main conclusions Evolutionary responses of muscid flies to post‐Eocene climate change taking the form of an expansion of a tropical group into regions with colder climates may be fundamental to explaining their distribution in the Neotropics. Our biogeographical regions delimited by temperature and the phylogenetic metric, surrogates of the tropical conservatism hypothesis, were very similar to general insect patterns, supporting the ‘tropical origin and evolutionary response to climate cooling’ as a broadly based historical narrative for the Neotropics.     

Evolutionary islands in the Andes: persistence and isolation explain high endemism in Andean dry tropical forests

Aim The tropical Andes are a world biodiversity hotspot. With diverse biomes and dramatic, geologically recent mountain uplift, they offer a system to study the relative contributions of geological and biome history to species richness. There are preliminary indications that historical species assembly in the Andes has been influenced by physiographical heterogeneity and that distinct biomes have evolved in relative isolation despite physical proximity. Here we test this ‘Andean biotic separation hypothesis’ by focusing on the low‐elevation, seasonally dry tropical forest (SDTF) biome to determine whether patterns of plant diversification within the SDTF differ from those in mid‐ and high‐elevation biomes. Location Tropical Andes, South America. Methods Densely sampled time‐calibrated phylogenies for five legume genera (Amicia, Coursetia, Cyathostegia, Mimosa and Poissonia) containing species endemic to the Andean SDTF biome were used to investigate divergence times and levels of geographical structure. Geographical structure was measured using isolation‐by‐distance methods. Meta‐analysis of time‐calibrated phylogenies of Andean plant groups was used to compare the pattern and tempo of endemic species diversification between the major Andean biomes. Results Long‐term persistence of SDTF in the Andes is suggested by old stem ages (5–27 Ma) of endemic genera/clades within genera, and deep divergences coupled with strong geographical structure among and within species. Comparison of species diversification patterns among different biomes shows that the relatively old, geographically confined pattern of species diversification in SDTF contrasts with the high‐elevation grasslands that show rapid and recent radiations driven by ecological opportunities. Main conclusions The SDTF biome has a long history in the Andes. We suggest that the diverse SDTF flora has been assembled gradually over the past c. 19 Ma from lineages exhibiting strong phylogenetic niche conservatism. These patterns suggest that Andean SDTFs have formed stable and strongly isolated ‘islands’ despite the upheavals of Andean uplift. Indeed, the Andean SDTFs may represent some of the most isolated and evolutionarily persistent continental plant communities, similar in many respects to floras of remote oceanic islands.     

Areas of endemism and distribution patterns for Neotropical Piper species (Piperaceae)

Aim The study aimed to establish areas of endemism and distribution patterns for Neotropical species of the genus Piper in the Neotropical and Andean regions by means of parsimony analysis of endemicity (PAE) and track‐compatibility analysis. Location The study area includes the Neotropical region and the Northern Andean region (Páramo‐Punan subregion). Methods We used distribution information from herbarium specimens and recent monographic revisions for 1152 species of Piper from the Neotropics. First, a PAE was attempted in order to delimit the areas of endemism. Second, we performed a track‐compatibility analysis to establish distribution patterns for Neotropical species of Piper. Terminology for grouping Piper is based on recent phylogenetic analyses. Results The PAE yielded 104 small endemic areas for the genus Piper, 80 of which are in the Caribbean, Amazonian and Paranensis subregions of the Neotropical region, and 24 in the Páramo‐Punan subregion of the Andean region. Track‐compatibility analysis revealed 26 generalized tracks, one in the Páramo‐Punan subregion (Andean region), 19 in the Neotropical region, and six connecting the Andean and Neotropical regions. Both the generalized tracks and endemic areas indicate that distribution of Piper species is restricted to forest areas in the Andes, Amazonia, Chocó, Central America, the Guayana Shield and the Brazilian Atlantic coast. Main conclusions Piper should not be considered an Andean‐centred group as it represents two large species components with distributions centred in the Amazonian and Andean regions. Furthermore, areas of greater species richness and/or endemism are restricted to lowland habitats belonging to the Neotropical region. The distribution patterns of Neotropical species of Piper could be explained by recent events in the Neotropical region, as is the case for the track connecting Chocó and Central America, where most of the species rich groups of the genus are found. Two kinds of event could explain the biogeography of a large part of the Piper taxa with Andean–Amazonian distribution: pre‐Andean and post‐Andean events.     

Neotropical diversification: the effects of a complex history on diversity within the poison frog genus Dendrobates

Aim We study the Neotropical poison frogs of the genus Dendrobates Wagler, 1830 in order to clarify their phylogenetic relationships and biogeographical history. The genus Dendrobates is an excellent taxon for examining patterns of Neotropical diversification as the four major species groups appear to correspond roughly to distinct geographical regions: (1) trans‐Andean, (2) Andean foreland, (3) Brazilian Shield and (4) Guianan Shield/Central America. In order to test the agreement of five of the most prominent hypotheses of Amazonian diversification, phylogenetic patterns were examined for agreement with patterns predicted by these hypotheses. Location Central and South America Methods The phylogenetic relationships of the genus Dendrobates were examined from novel and existing (GenBank) sequences of four mitochondrial loci totalling c. 1400 bp from 40 specimens of 22 different species using maximum parsimony and Bayesian methods. Results were compared with traditional taxonomic arrangements by means of SH tests. Phylogenetic relationships and genetic distances were used to test the adequacy of various diversification hypotheses. Results Phylogenetic analyses support the restructuring of two species groups of Dendrobates and the creation of a new species group. Statistical tests of the traditional taxonomic arrangement indicate a significantly bad fit to the molecular data. This restructuring has important implications for the understanding of the historical biogeography of Dendrobates. Biogeographical patterns within this genus suggest that a complex interaction of biotic and abiotic factors since the Eocene have produced the diversity observed today. Main conclusions The current classification of Dendrobates into discrete species groups does not accurately reflect evolutionary history. Data presented here strongly support a monophyletic Brazilian Shield lineage whose members have previously been split among the quinquevittatus and tinctorius groups. Furthermore, previous attempts at elucidating the historical biogeography of this genus were compromised by incomplete sampling and conclusions drawn from a paraphyletic ingroup. Our findings demonstrate a role for numerous hypotheses of diversification (e.g. river, refuge, disturbance–vicariance) in the history of Dendrobates, supporting previous warnings about the dangers of over‐simplification in the study of Neotropical diversification.     

Rediscovery and Mitochondrial Relationships of the Dendrobatid Frog Colostethus humilis Suggest Parallel Colonization of the Venezuelan Andes by Poison Frogs

Colostethus humilis Rivero, ‘1978’1980 was rediscovered near the type locality. The species is redescribed, and data on its advertisement call are provided for the first time. Its morphology and colour pattern differ from other Andean dendrobatids. It shows morphological and bio-acoustic similarities to species of Colostethus from the Amazonian lowlands. A phylogenetic analysis of sequences of the mitochondrial 16S rRNA gene confirmed that C. humilis is related to Amazonian lowland Colostethus, rather than to other Andean poison frogs of the genera Nephelobates and Mannophryne. This indicates that several groups of basal dendrobatids independently have colonised the Venezuelan Andes.     

Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae)

Euptychiina is the most species‐rich subtribe of Neotropical Satyrinae, with over 450 known species in 47 genera (14 monotypic). Here, we use morphological characters to examine the phylogenetic relationships within Euptychiina. Taxonomic sampling included 105 species representing the majority of the genera, as well as five outgroups. A total of 103 characters were obtained: 45 from wing pattern, 48 from genitalia and 10 from wing venation. The data matrix was analysed using maximum parsimony under both equal and extended implied weights. Euptychiina was recovered as monophyletic with ten monophyletic genera, contrasting previous DNA sequence‐based phylogenies that did not recover the monophyly of the group. In agreement with sequence‐based hypotheses, however, three main clades were recognized: the ‘Megisto clade’ with six monophyletic and three polyphyletic genera, the ‘Taygetis clade’ with nine genera of which three were monophyletic, and the ‘Pareuptyhia clade’ with four monophyletic and two polyphyletic genera. This is the first morphology‐based phylogenetic hypothesis for Euptychiina and the results will be used to complement molecular data in a combined analysis and to provide critical synapomorphies for clades and genera in this taxonomically confused group.     

Out of Amazonia again and again: episodic crossing of the Andes promotes diversification in a lowland forest flycatcher

Most Neotropical lowland forest taxa occur exclusively on one side of the Andes despite the availability of appropriate habitat on both sides. Almost all molecular phylogenies and phylogenetic analyses of species assemblages (i.e. area cladograms) have supported the hypothesis that Andean uplift during the Late Pliocene created a vicariant barrier affecting lowland lineages in the region. However, a few widespread plant and animal species occurring in lowland forests on both sides of the Andes challenge the generality of this hypothesis. To understand the role of the Andes in the history of such organisms, we reconstructed the phylogeographic history of a widespread Neotropical flycatcher (Mionectes oleagineus) in the context of the other four species in the genus. A molecular phylogeny based on nuclear and mitochondrial sequences unambiguously showed an early basal split between montane and lowland Mionectes. The phylogeographic reconstruction of lowland taxa revealed a complex history, with multiple cases in which geographically proximate populations do not represent sister lineages. Specifically, three populations of M. oleagineus west of the Andes do not comprise a monophyletic clade; instead, each represents an independent lineage with origins east of the Andes. Divergence time estimates suggest that at least two cross-Andean dispersal events post-date Andean uplift.     

Evolutionary history and precipitation seasonality shape niche overlap in Neotropical bat–plant pollination networks

Species interactions are one dimension of a niche. Niche overlap arises when two species share an interaction partner. In pollination systems, environmental and biotic factors affect niche overlap. Here, we explored the effects of climate seasonality, plant and bat richness, morphological traits, and phylogenetic distance in shaping the niche overlap of Neotropical bat–plant pollination networks. We examined a dataset of 22 bat–plant pollination networks in the Neotropical region. We measured niche overlap in bats and plants with the Morisita-Horn index (Ĉ_H) and used a SAR model to test the relationships between niche overlap and both abiotic and biotic factors. We found a lower niche overlap among bats in communities composed of phylogenetically distant bat species. Moreover, plant and bat overlap was lower in regions with higher precipitation seasonality. Our results indicate that climate seasonality and bat evolutionary history drive niche overlap in Neotropical bat–plant pollination interactions. These findings suggest that a higher precipitation seasonality promotes the emergence of temporal modules, which reduces niche overlap, likely due to seasonal species phenologies. Furthermore, the method used to record the interactions affects the degree of niche overlap. Interactions recorded with pollen samples tend to have higher niche overlap than direct observations. The responses of morphological traits and phylogenetic distances in bat niche overlap were uncoupled, suggesting an effect of historical processes independent of morphological traits. Our findings reinforce the importance of evolutionary history and ecological processes in imprinting patterns of interaction niche overlap.     

Phylogeny of the Neotropical killifish family Rivulidae (Cyprinodontiformes, Aplocheiloidei) inferred from mitochondrial DNA sequences

Phylogenetic relationships of 70 taxa representing 68 species of the Neotropical killifish family Rivulidae were derived from analysis of 1516 nucleotides sampled from four different segments of the mitochondrial genome: 12S rRNA, 16S rRNA, cytochrome oxidase I, and cytochrome b. The basal bifurcation of Cynolebiatinae and Rivulinae (Costa, 1990a,b) is supported; however, Terranatos, Maratecoara, and Plesiolebias are rivulins, not cynolebiatins. These three genera, along with the other recognized annual rivulin genera, form a monophyletic clade. Austrofundulus, Rachovia, Renova, Terranatos, and 3 species of the genus Pterolebias, all from northeastern South America, form a monophyletic clade excluding other species of Pterolebias. Pterolebias as presently understood is clearly polyphyletic. Trigonectes and Moema are supported as sister groups but do not form a monophyletic group with the genera Neofundulus and Renova as previously proposed. The suite of adaptations necessary for an annual life history has clearly been lost several times in the course of rivulid evolution. Also revealed is a considerable increase in substitution rate in most annual lineages relative to the nonannual Rivulus species. The widespread and speciose genus Rivulus is paraphyletic, representing both basal and terminal clades within the Rivulidae. Previous hypotheses regarding the vicariant origin of Greater Antillean Rivulus species are supported. Most rivulid clades show considerable endemism; thus, detailed analysis of rivulid phylogeny and distribution will contribute robust hypotheses to the clarification of Neotropical biogeography.     

Global diversity of water mites (Acari,Hydrachnidia; Arachnida) in freshwater

The Hydrachnidia (water mites) represent the most important group of the Arachnida in fresh water. Over 6,000 species have been described worldwide, representing 57 families, 81 subfamilies and more than 400 genera. The article analyzes extant water mite diversity and biogeography. Data on distribution and species richness of water mites are substantial but still far from complete. Many parts of the world are poorly investigated, Oriental and Afrotropical regions in particular. Moreover, information among different freshwater habitats is unbalanced with springs and interstitial waters disproportionately unrepresented. Therefore, more than 10,000 species could be reasonably expected to occur in inland waters worldwide. Based on available information, the Palaearctic region represents one of the better investigated areas with the highest number of species recorded (1,642 species). More than 1,000 species have been recorded in each of the Neotropical (1,305 species) and Nearctic regions (1,025 species). Known species richness is lower in Afrotropical (787 species) and Australasian (694 species) regions, and lowest in the Oriental region (554 species). The total number of genera is not correlated with species richness and is distinctly higher in the Neotropical (164 genera); genus richness is similar in the Palaearctic, Nearctic and Australasian regions (128–131 genera) and is lower in the Afrotropical and Oriental regions with 110 and 94 genera, respectively. A mean number of about three genera per family occur in the Palaeartic, Nearctic and Oriental while an average of more than four genera characterizes the families of Australasian and Afrotropical regions and more than five genera those of the Neotropical. Australasian fauna is also characterized by the highest percentage of endemic genera (62%), followed by Neotropical (50.6%) and Afrotropical (47.2%) regions. Lower values are recorded for the Palaearctic (26.9%), Oriental (24.4%) and Nearctic (21.4%). The Palaearctic and Nearctic have the highest faunistic similarity, some minor affinities are also evident for the generic diversification of Holarctic and Oriental families. The faunas of Southern Hemisphere bioregions are more distinct and characterized by the presence of ancient Gondwanan clades with a regional diversification particularly evident in the Neotropics and Australasia. This scenario of water mite diversity and distribution reflect the basic vicariance pattern, isolation, phylogenetic diversification, recent climatic vicissitudes and episodes of dispersal between adjacent land masses together with extant ecological factors can be evoked to explain distribution patterns at a global scale. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Tracing the diversification history of a Neogene rodent invasion into South America

We investigated spatial patterns of evolutionary relatedness and diversification rates to test hypotheses about the historical biogeographic processes underlying the radiation of Neotropical rats and mice (Sigmodontinae, ~400 species). A negative correlation between mean phylogenetic distance and diversification rates of rodent assemblages reveals a pattern of species co‐occurrence in which assemblages of closely related species are also the fastest diversifying ones. Subregions of the Neotropics occupied by distantly related species that are on average more slowly diversifying include Central America, northern South America, and the Atlantic forest. In southern South America, recent species turnover appears to have been higher. Ancestral locations for the main tribes of sigmodontines were also estimated, suggesting eastern South America and the Amazonian lowlands were colonized before some central Andean regions, even though the latter are now centers of species richness for these rodents. Moreover, a past connection between the tropical Andes and the Atlantic Forest is suggested by our results, highlighting a role for a hypothetical arc connecting the two biomes, which would have impacted many other groups of organisms. Whether rapid, recent speciation in some regions is related to Quaternary climatic fluctuations and the young age of sigmodontines (~12.7 Ma crown age) or instead to intrinsic traits of these rodents remains an open question. If the former is true, we hypothesize that contrasting trends will characterize older Neotropical clades.     

INVESTIGATING PROCESSES OF NEOTROPICAL RAIN FOREST TREE DIVERSIFICATION BY EXAMINING THE EVOLUTION AND HISTORICAL BIOGEOGRAPHY OF THE PROTIEAE (BURSERACEAE)

Andean uplift and the collision of North and South America are thought to have major implications for the diversification of the Neotropical biota. However, few studies have investigated how these geological events may have influenced diversification. We present a multilocus phylogeny of 102 Protieae taxa (73% of published species), sampled pantropically, to test hypotheses about the relative importance of dispersal, vicariance, habitat specialization, and biotic factors in the diversification of this ecologically dominant tribe of Neotropical trees. Bayesian fossil‐calibrated analyses date the Protieae stem at 55 Mya. Biogeographic analyses reconstruct an initial late Oligocene/early Miocene radiation in Amazonia for Neotropical Protieae, with several subsequent late Miocene dispersal events to Central America, the Caribbean, Brazil's Atlantic Forest, and the Chocó. Regional phylogenetic structure results indicate frequent dispersal among regions throughout the Miocene and many instances of more recent regional in situ speciation. Habitat specialization to white sand or flooded soils was common, especially in Amazonia. There was one significant increase in diversification rate coincident with colonization of the Neotropics, followed by a gradual decrease consistent with models of diversity‐dependent cladogenesis. Dispersal, biotic interactions, and habitat specialization are thus hypothesized to be the most important processes underlying the diversification of the Protieae.     

Different evolutionary histories underlie congruent species richness gradients of birds and mammals

Aim The global species richness patterns of birds and mammals are strongly congruent. This could reflect similar evolutionary responses to the Earth’s history, shared responses to current climatic conditions, or both. We compare the geographical and phylogenetic structures of both richness gradients to evaluate these possibilities. Location Global. Methods Gridded bird and mammal distribution databases were used to compare their species richness gradients with the current environment. Phylogenetic trees (resolved to family for birds and to species for mammals) were used to examine underlying phylogenetic structures. Our first prediction is that both groups have responded to the same climatic gradients. Our phylogenetic predictions include: (1) that both groups have similar geographical patterns of mean root distance, a measure of the level of the evolutionary development of faunas, and, more directly, (2) that richness patterns of basal and derived clades will differ, with richness peaking in the tropics for basal clades and in the extra‐tropics for derived clades, and that this difference will hold for both birds and mammals. We also explore whether alternative taxonomic treatments for mammals can generate patterns matching those of birds. Results Both richness gradients are associated with the same current environmental gradients. In contrast, neither of our evolutionary predictions is met: the gradients have different phylogenetic structures, and the richness of birds in the lowland tropics is dominated by many basal species from many basal groups, whereas mammal richness is attributable to many species from both few basal groups and many derived groups. Phylogenetic incongruence is robust to taxonomic delineations for mammals. Main conclusions Contemporary climate can force multiple groups into similar diversity patterns even when evolutionary trajectories differ. Thus, as widely appreciated, our understanding of biodiversity must consider responses to both past and present climates, and our results are consistent with predictions that future climate change will cause major, correlated changes in patterns of diversity across multiple groups irrespective of their evolutionary histories.     

Classification, diversity, and distribution of Chilean Asteraceae: implications for biogeography and conservation

This paper provides a synopsis of the Chilean Asteraceae genera according to the most recent classification. Asteraceae is the richest family within the native Chilean flora, with a total of 121 genera and c . 863 species, currently classified in 18 tribes. The genera are distributed along the whole latitudinal gradient in Chile, with a centre of richness at 33°–34° S. Almost one-third of the genera show small to medium-small ranges of distribution, while two-thirds have medium-large to large latitudinal ranges of distribution. Of the 115 mainland genera, 46% have their main distribution in the central Mediterranean zone between 27°–37° S. Also of the mainland genera, 53% occupy both coastal and Andean environments, while 33% can be considered as strictly Andean and 20% as strictly coastal genera. The biogeographical analysis of relationships allows the distinction of several floristic elements and generalized tracks: the most marked floristic element is the Neotropical, followed by the antitropical and the endemic element. The biogeographical analysis provides important insights into the origin and evolution of the Chilean Asteraceae flora. The presence of many localized and endemic taxa has direct conservation implications.     

Epiphytism and pollinator specialization: drivers for orchid diversity?

Epiphytes are a characteristic component of tropical rainforests. Out of the 25,000 orchid species currently known to science, more than 70% live in tree canopies. Understanding when and how these orchids diversified is vital to understanding the history of epiphytic biomes. We investigated whether orchids managed to radiate so explosively owing to their predominantly epiphytic habit and/or their specialized pollinator systems by testing these hypotheses from a statistical and phylogenetic standpoint. For the first approach, species numbers of 100 randomly chosen epiphytic and terrestrial genera were compared. Furthermore, the mean number of pollinators per orchid species within the five subfamilies was calculated and correlated with their time of diversification and species richness. In the second approach, molecular epiphytic orchid phylogenies were screened for clades with specific suites of epiphytic adaptations. Epiphytic genera were found to be significantly richer in species than terrestrial genera both for orchids and non-orchids. No evidence was found for a positive association between pollinator specialization and orchid species richness. Repeated associations between a small body size, short life cycle and specialized clinging roots of twig epiphytes in Bulbophyllinae and Oncidiinae were discovered. The development of twig epiphytism in the first group seems repeatedly correlated with speciation bursts.     

Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions

Abstract.  1. Patterns of simuliid species richness were examined over a variety of scales at 532 stream sites in the Nearctic (394) and Neotropical (138) regions. In Nearctic streams, species richness of immature blackflies both within and across ecoregions and over two seasons was examined. Stream variables at each site included seston, width, depth, velocity, discharge, conductivity, pH, dissolved oxygen, water temperature, dominant streambed-particle size, canopy cover, and riparian vegetation. These variables were subjected to a principal component analysis and derived principal components were related back to richness, using regression analysis. At the level of the stream reach, richness was not highly correlated with single-point measurements of stream conditions.
2. Using data from both Nearctic and Neotropical sites, the effect of regional richness on local richness was examined. As regional richness increased, local diversity reached an asymptote in which further increases in regional richness were not matched by increases in local richness. Hence, simuliid communities are best described as saturated (type II) communities, consistent with the current view of lotic communities as non-equilibrium systems.
3. The well-established pattern of greater species richness in tropical regions was not observed in this study. To the contrary, blackfly richness is higher in temperate streams than in tropical streams at both local and regional scales.     

The impact of parsimony weighting schemes on inferred relationships among toucans and neotropical barbets (Aves: piciformes)

The development of new schemes for weighting DNA sequence data for phylogenetic analysis continues to outpace the development of consensus on the most appropriate weights. The present study is an exploration of the similarities and differences between results from 22 character weighting schemes when applied to a study of barbet and toucan (traditional avian families Capitonidae and Ramphastidae) phylogenetic relationships. The dataset comprises cytochrome b sequences for representatives of all toucan and Neotropical barbet genera, as well as for several genera of Paleotropical barbets. The 22 weighting schemes produced conflicting patterns of relationship among taxa, often with conflicting patterns each receiving strong bootstrap support. Use of multiple weighting schemes helped to identify the source within the dataset (codon position, transitions, transversions) of the various putative phylogenetic signals. Importantly, some phylogenetic hypotheses were consistently supported despite the wide range of weights employed. The use of phylogenetic frameworks to summarize the results of these multiple analyses proved very informative. Relationships among barbets and toucans inferred from these data support the paraphyly of the traditional Capitonidae. Additionally, these data support paraphyly of Neotropical barbets, but rather than indicating a relationship between Semnornis and toucans, as previously suggested by morphological data, most analyses indicate a basal position of Semnornis within the Neotropical radiation. The cytochrome b data also allow inference of relationships among toucans. Supported hypotheses include Ramphastos as the sister to all other toucans, a close relationship of Baillonius and Pteroglossus with these two genera as the sister group to an (Andigena, Selenidera) clade, and the latter four genera as a sister group to Aulacorhynchus.     

Hidden generic diversity in Neotropical birds: molecular and anatomical data support a new genus for the "Scytalopus"indigoticus species-group (Aves: Rhinocryptidae)

The genus Scytalopus is a species-rich and taxonomically complicated component of the Neotropical avian family Rhinocryptidae. Probably because Scytalopus is a superficially uniform assemblage, its monophyly has not been seriously questioned. We investigated phylogenetic relationships of a representative set of species in the genus using nuclear and mitochondrial DNA sequences as well as anatomical data, and provided the first test of its presumed monophyly by including in the analyses its hypothesized closest relatives (the genera Myornis, Eugralla, and Merulaxis) as well as most rhinocryptid genera. We found strong support for the paraphyly of the genus Scytalopus, with the Scytalopus indigoticus species-group forming a clade with Merulaxis. A well-supported clade including the genera Eugralla, Myornis, and the remaining Scytalopus was also recovered. Because these results were recovered independently and with strong support using mitochondrial and nuclear data, and were entirely consistent with anatomical data, we erect a new genus for the S.indigoticus species-group. These findings illustrate the importance of formally testing hypotheses of monophyly even for well-accepted groups of Neotropical birds.     

Climate change and the future restructuring of Neotropical anuran biodiversity

Climate change is likely to impact multiple dimensions of biodiversity. Species range shifts are expected and may drive changes in the composition of species assemblages. In some regions, changes in climate may precipitate the loss of geographically restricted, niche specialists and facilitate their replacement by more widespread, niche generalists, leading to decreases in β-diversity and biotic homogenization. However, in other regions climate change may drive local extinctions and range contraction, leading to increases in β-diversity and biotic heterogenization. Regional topography should be a strong determinant of such changes as mountainous areas often are home to many geographically restricted species, whereas lowlands and plains are more often inhabited by widespread generalists. Climate warming, therefore, may simultaneously bring about opposite trends in β-diversity in mountainous highlands versus relatively flat lowlands. To test this hypothesis, we used species distribution modelling to map the present-day distributions of 2669 Neotropical anuran species, and then generated projections of their future distributions assuming future climate change scenarios. Using traditional metrics of β-diversity, we mapped shifts in biotic homogenization across the entire Neotropical region. We used generalized additive models to then evaluate how changes in β-diversity were associated with shifts in species richness, phylogenetic diversity and one measure of ecological generalism. Consistent with our hypothesis, we find increasing biotic homogenization in most highlands, associated with increased numbers of generalists and, to a lesser extent, losses of specialists, leading to an overall increase in alpha diversity, but lower mean phylogenetic diversity. In the lowlands, biotic heterogenization was more common, and primarily driven by local extinctions of generalists, leading to lower α-diversity, but higher mean phylogenetic diversity. Our results suggest that impacts of climate change on β-diversity are likely to vary regionally, but will generally lead to lower diversity, with increases in β-diversity offset by decreases in α-diversity.