Termites (Isoptera: Kalotermitidae,Rhinotermitidae, Termitidae) of Ecuador

Termites are an abundant and diverse group in the Neotropics with about 500 species representing 83 genera. The paucity of the termite fauna recorded from Ecuador is due, in part, to a lack of deliberate surveys. We revise the termite fauna of Ecuador and raise the number of species from 25 species to 72 based on our recent termite surveys. Of the 72 species, 18 could not be conclusively identified and are likely new species. Given the limited area that has been covered in surveys of the Ecuadorian termite fauna, there are undoubtedly many more species to be recorded for Ecuador, primarily in the eastern lowland areas, cloud forests on both the eastern and western slopes of the Andes, and the Amazonian lowland forests.     

Integrating alpha,beta, and phylogenetic diversity to understand anuran fauna along environmental gradients of tropical forests in western Ecuador

The study of current distribution patterns of amphibian species in South America is of particular interest in areas such as evolutionary ecology and conservation biology. These patterns could be playing an important role in biological interactions, population size, and connectivity, and potential extinction risk in amphibians. Here, we tested the effects of spatial and environmental factors on the variation, turnover, and phylogenetic diversity of anuran amphibian species in tropical forests of western Ecuador. Data for presence/absence of 101 species of 34 genera and 10 families registered in 12 sites (nested in four biogeographic units) were obtained through fieldwork, museum collections, and literature records. We examined the influence of geographical, altitudinal, temperature, and precipitation distances on differences in anuran composition between sites. We found significant positive correlations among all of these variables with anuran distribution. The greatest alpha diversity (species richness) was found in the Equatorial Chocó biogeographic unit. Equatorial Pacific biogeographic unit could act as a transition zone between the Equatorial Chocó and Equatorial Tumbes. The western Andes (Western Cordillera biogeographic unit) was the most dissimilar and exhibited a higher species turnover rate than the other biogeographic units. Our results suggest that precipitation and elevation play a key role in maintaining the diversity of amphibian species in western Ecuador.     

Rediscovery of ‘lost’ treefrogs from the Oaxacan highlands of Mexico

The Oaxacan highlands of Mexico boast an exceptional diversity of stream-breeding treefrogs (Hylidae). Recent surveys in these regions have reported dramatic declines for entire stream-breeding communities, and many endemic species have not been seen in several decades and are feared to be extinct. In 2007 we conducted a rapid survey in two focal regions that were surveyed in 1970 and 2000; both prior to and subsequent to reported population declines. We report on extant populations of six species missing from recent surveys following declines, one of which (Plectrohyla thorectes) had not been seen in over 28 years and was feared to be extinct. We document new populations of Exerodonta abdivita, a species previously known only from its type locality and currently listed as data deficient. We make comparisons with historical records and note changes in species composition and abundance with recent work. Differences in species compositions among studies may reflect population declines and rebounds over relatively short periods. However, we argue that aspects of species natural history in combination with the variable nature of brief surveys may account for some of the observed differences, and are important factors to consider when inferring species declines.     

Upward range extension of Andean anurans and chytridiomycosis to extreme elevations in response to tropical deglaciation

High-alpine life forms and ecosystems exist at the limits of habitable environments, and thus, are especially sensitive to environmental change. Here we report a recent increase in the elevational limit of anurans following glacial retreat in the tropical Peruvian Andes. Three species have colonized ponds in recently deglaciated terrain at new record elevations for amphibians worldwide (5244–5400 m). Two of these species were also found to be infected with Batrachochytrium dendrobatidis ( Bd ), an emerging fungal pathogen causally associated with global amphibian declines, including the disappearance of several Latin American species. The presence of this pathogen was associated with elevated mortality rates of at least one species. These results represent the first evidence of upward expansion of anurans to newly available habitat brought about by recent deglaciation. Furthermore, the large increase in the upper limit of known Bd infections, previously reported as 4112 m in Ecuador, to 5348 m in this study, also expands the spatial domain of potential Bd pathogenicity to encompass virtually all high elevation anuran habitats in the tropical Andes.     

Ecological and Geographical Analysis of the Distribution of the Mountain Tapir (Tapirus pinchaque) in Ecuador: Importance of Protected Areas in Future Scenarios of Global Warming

In Ecuador, Tapirus pinchaque is considered to be critically endangered. Although the species has been registered in several localities, its geographic distribution remains unclear, and the effects of climate change and current land uses on this species are largely unknown. We modeled the ecological niche of T. pinchaque using MaxEnt, in order to assess its potential adaptation to present and future climate change scenarios. We evaluated the effects of habitat loss due by current land use, the ecosystem availability and importance of Ecuadorian System of Protected Areas into the models. The model of environmental suitability estimated an extent of occurrence for species of 21,729 km² in all of Ecuador, mainly occurring along the corridor of the eastern Ecuadorian Andes. A total of 10 Andean ecosystems encompassed ~98% of the area defined by the model, with herbaceous paramo, northeastern Andean montane evergreen forest and northeastern Andes upper montane evergreen forest being the most representative. When considering the effect of habitat loss, a significant reduction in model area (~17%) occurred, and the effect of climate change represented a net reduction up to 37.86%. However, the synergistic effect of both climate change and habitat loss, given current land use practices, could represent a greater risk in the short-term, leading to a net reduction of 19.90 to 44.65% in T. pinchaque’s potential distribution. Even under such a scenarios, several Protected Areas harbor a portion (~36 to 48%) of the potential distribution defined by the models. However, the central and southern populations are highly threatened by habitat loss and climate change. Based on these results and due to the restricted home range of T. pinchaque, its preference for upland forests and paramos, and its small estimated population size in the Andes, we suggest to maintaining its current status as Critically Endangered in Ecuador.     

The diversity of cloud forest birds on the eastern and western slopes of the Ecuadorian Andes: a latitudinal and comparative analysis with implications for conservation

We studied the avifaunas of five cloud forest localities at 3000–3350 mas1 on the eastern slope of the Andes of Ecuador A method standardised for area, altitude and effort was the basis for obtaining data on species richness and abundances Richness and abundances (diversity) were unchanged along the limned latitudinal gradient (530 km) studied, whereas there was a considerable turnover in community structure between localities This result has important implications for ranking conservation priorities the total community richness and the dominance/evenness component of diversity does not provide a basis for making priorities along this limited latitudinal gradient Instead, species compositions and species referred to threatened categories (e g, endemics, restricted–range, IUCN threatened/near–threatened, CITES) will be the most meaningful data for making priorities for conservation We compared our results with similar results from the Andean western slope of Ecuador, obtained with the same method and standardised for time of the year The number of species was significantly higher on the eastern slope than on the western slope, a characteristic that can be explained from differences in extension of adjacent source areas and geohistory However, because the species compositions on the two slopes are rather different, it is not recommended to use the difference in richness to rank conservation priorities Abundances were quite similar on both sides of the Andes Contrary to the general belief, the montane avifaunas investigated have tropical and not temperate species richness which agrees with the tropical clutch sizes found among montane birds The avian richness found on the eastern slope of Ecuador near the Equator is definitely among the highest m the world for this altitude     

Where are the survivors? Tracking relictual populations of endangered frogs in Costa Rica

Aim We used abiotic environmental variables and historical locality records to infer distributions of endangered anuran species of Costa Rica to promote efficient strategies for future amphibian surveys. Location Costa Rica. Methods We used a Maximum Entropy Algorithm (Maxent) to predict potential distribution maps for 17 species of endangered anurans and create a consensus map of species richness. We compared the environmental conditions from localities where relictual amphibian populations were recently rediscovered with the conditions across their historical range to evaluate the possibility that these relictual populations might occur in specific climatic conditions that could explain their persistence. We used a multicriteria analysis considering the following factors: the intersection zones between the consensus map, conservation areas, potential Batrachochytrium dendrobatidis (Bd) distribution, collecting effort and areas within the precipitation range at which reappearances had occurred to locate sites for future surveys. Results The resulting predictions suggest that suitable areas for the highest number of species occur between 1300 and 2500 m.a.s.l and are concentrated along the Pacific slopes of the Cordillera de Talamanca and Cordillera Volcánica Central. Around 45% of the high potential richness area is under protection. Relictual populations of declined species seem to persist mainly in highly humid localities (2500–3500 mm of mean annual precipitation). Around 240 km² has an ideal environment for the rediscovery of relictual populations. The multicriteria analysis showed that around 0.5% of the Costa Rican territory should be surveyed exhaustively for frogs. Main conclusions Many of the potential refugia we identified here have not been surveyed since 2000, the areas identified by the best model predictions correspond well with the localities of the relictual populations recently reported. We suggest future surveys of missing amphibian species should focus on these areas. The discovery of populations of endangered species can be used to propose conservation areas.     

Frogs in the spotlight: a 16‐year survey of native frogs and invasive toads on a floodplain in tropical Australia

Although widespread declines in anuran populations have attracted considerable concern, the stochastic demographics of these animals make it difficult to detect consistent trends against a background of spatial and temporal variation. To identify long‐term trends, we need datasets gathered over long time periods, especially from tropical areas where anuran biodiversity is highest. We conducted road surveys of four anurans in the Australian wet–dry tropics on 4637 nights over a 16‐year period. Our surveys spanned the arrival of invasive cane toads (Rhinella marina), allowing us to assess the invader's impact on native anuran populations. Our counts demonstrate abrupt and asynchronous shifts in abundance and species composition from one year to the next, not clearly linked to rainfall patterns. Typically, periods of decline in numbers of a species were limited to 1–2 years and were followed by 1‐ to 2‐year periods of increase. No taxa showed consistent declines over time, although trajectories for some species showed significant perturbations coincident with the arrival of toads. None of the four focal frog species was less common at the end of the study than at the beginning, and three of the species reached peak abundances after toad arrival. Survey counts of cane toads increased rapidly during the initial stage of invasion but have subsequently declined and fluctuated. Distinguishing consistent declines versus stochastic fluctuations in anuran populations requires extensive time‐series analysis, coupled with an understanding of the shifts expected under local climatic conditions. This is especially pertinent when assessing impacts of specific perturbations such as invasive species.     

Patterns of plant species endemism in Ecuador

Distribution patterns of plant species endemic to Ecuador and adjacent parts of southern Colombia and northern Peru are analysed on the basis of information in the Flora of Ecuador. A total of 827 restricted-range species were found, many of which are known from extremely small areas, often only one or a few localities. A total of 27% of the species treated in the Flora of Ecuador are endemic to that country. The overall proportion of endemic and restricted-range species is greater in the Andes than in the lowland areas on either side of these mountains; particularly the southern Andes appears to be very rich in endemic species. Spatial analysis of distribution data results in the recognition of 15 floristic elements and 18 geographical endemism regions in Ecuador, the characteristics of which are discussed. Comparison with distribution patterns of restricted-range bird species show a general correspondence, with the main difference that birds tend to be more widely distributed than plants along the Andes. Comparison of the results with the location of national parks and other protected areas shows that the endemic floras in the northern and eastern parts of the country are much better protected than those of the southern and western parts.     

Historical biogeography and post-glacial recolonization of South American temperate rain forest by the relictual marsupial Dromiciops gliroides

Aim Long‐term climatic variation has generated historical expansions and contractions of species ranges, with accompanying fragmentation and population bottlenecks, which are evidenced by spatial variation in genetic structure of populations. We examine here hypotheses concerning dispersal and vicariance in response to historical geoclimatic change and potential isolation produced by mountains and water barriers. Location The temperate rain forest of southern South America, which is distributed from coastal Chile, including the large continental island of Chiloé, across the Andes into Argentina. Methods We investigated our hypotheses in the phylogenetically and biogeographically relictual marsupial Dromiciops gliroides. We examined 56 specimens, which resulted from field samples and museum study skins from 21 localities. We evaluated the influence of two major barriers, the Andean cordillera and the waterway between the mainland and the large island of Chiloé, by performing Bayesian and maximum‐likelihood phylogenetic analyses on sequences of 877 base pairs of mitochondrial DNA. We further tested the contribution of the proposed geographical barriers using analysis of molecular variance (amova ). We also evaluated the responses of populations to historical north–south shifts of habitat associated with glacial history and sea‐level change. Results Our analyses revealed a phylogeny with three clades, two of which are widespread and contain nearly all the haplotypes: a northern clade (36–39° S) and a southern clade (40–43° S). These two clades contain forms from both sides of the Andes. Within the southern clade, island and mainland forms were not significantly differentiated. Tests of recent demographic change revealed that southern populations have experienced recent expansion, whereas northern populations exhibit long‐term stability. The direction of recent gene flow and range expansion is predominantly from Chile to Argentina, with a modest reciprocal exchange across the Andes. Recent gene flow from the island of Chiloé to the mainland is also supported. Main conclusions The genetic structure of contemporary D. gliroides populations suggests recent gene flow across the Andes and between the mainland and the island of Chiloé. Differences in demographic history that we detected between northern and southern populations have resulted from historical southward shifts of habitat associated with glacial recession in South America. Our results add to a growing literature that demonstrates the value of genetic data to illuminate how environmental history shapes species range and population structure.     

Phylogeography of a morphologically diverse Neotropical montane species, the Common Bush-Tanager (Chlorospingus ophthalmicus)

The Common Bush-Tanager (Chlorospingus ophthalmicus) is distributed in Neotropical cloud-forests from Mexico to Argentina and contains 25 subspecies divided into eight subspecies groups based on biogeography, eye coloration, presence of a postocular spot and chest band. All of Central America is occupied by a single subspecies group; whereas the Andes are believed to be occupied by seven additional subspecies groups. We used five mitochondrial genes to investigate the phylogeography and possible species limits of the ophthalmicus complex. A total of 14 monophyletic lineages were uncovered within the ophthalmicus complex, including three clades currently classified as separate species (C. semifuscus, inornatus and tacarcunae). Divergence estimates for these clades date between 0.8 and 5.2 million years ago (Ma). Contrary to expectations based on morphological diversity, phylogeographic structure was greatest in Mexico and Central America and weakest in the Andes. Morphological and genetic divergences were not significantly correlated and most morphologically defined subspecies groups were not supported. Our evidence suggests the ophthalmicus complex originated in Mexico ca. 6.0 Ma (million years ago) and spread south into the Andes ca. 4.7 Ma before the completion of the Isthmus of Panama. Three genetically divergent lineages of ophthalmicus that formed in the Andes possess a complex checkerboard distribution, with a single lineage represented by disjunct populations from Venezuela and the southern Andes, while intervening populations in Ecuador and Central Peru form two genetically and morphologically divergent lineages.     

High levels of cryptic species diversity uncovered in Amazonian frogs

One of the greatest challenges for biodiversity conservation is the poor understanding of species diversity. Molecular methods have dramatically improved our ability to uncover cryptic species, but the magnitude of cryptic diversity remains unknown, particularly in diverse tropical regions such as the Amazon Basin. Uncovering cryptic diversity in amphibians is particularly pressing because amphibians are going extinct globally at an alarming rate. Here, we use an integrative analysis of two independent Amazonian frog clades, Engystomops toadlets and Hypsiboas treefrogs, to test whether species richness is underestimated and, if so, by how much. We sampled intensively in six countries with a focus in Ecuador (Engystomops: 252 individuals from 36 localities; Hypsiboas: 208 individuals from 65 localities) and combined mitochondrial DNA, nuclear DNA, morphological, and bioacoustic data to detect cryptic species. We found that in both clades, species richness was severely underestimated, with more undescribed species than described species. In Engystomops, the two currently recognized species are actually five to seven species (a 150-250% increase in species richness); in Hypsiboas, two recognized species represent six to nine species (a 200-350% increase). Our results suggest that Amazonian frog biodiversity is much more severely underestimated than previously thought.     

Morphological abnormalities in anurans from southern Brazil

In recent years, the progressive decline in populations of anuran amphibians has left many species increasingly vulnerable to extinction. The potential role of morphological abnormalities as a mechanism of this decline has attracted attention in many recent studies. We investigated the occurrence and frequency of morphological abnormalities in anurans from northern of the state of Rio Grande do Sul, Brazil. We sampled 1674 individuals from three different environments – forest, farmland and peri‐urban areas – between 2001 and 2017. We recorded 89 individuals with morphological abnormalities, with 12 different types of abnormality in 19 anuran species. We recorded nine types of abnormality in forest and seven in the farmland, with brachydactyly being the most common abnormality in both areas. Only three abnormalities were recorded in the peri‐urban area, and all three were equally frequent. Studies of morphological alterations provide important insights into the potential effects of environmental impacts on anuran populations and may be essential for the planning of effective conservation strategies.     

Molecular Markers and Conservation of Plant Species in the Latin-America: The Case of Phaedranassa viridiflora (Amaryllidaceae)

Phaedranassa viridiflora (Amaryllidaceae) is an endemic and endangered plant restricted to the Northern Andes in Ecuador. It is known in three locations where it is sympatric with other Phaedranassa species. Phaedranassa viridiflora is the only species of the genus with yellow flowers. We analyzed 13 microsatellite loci to elucidate the genetic structure of the populations of P. viridiflora. Our results provided the first evidence of natural hybridization in the genus (between P. viridiflora and P. dubia in the Pululahua crater of northern Ecuador). The central and southern populations did not show hybridization. Genetic diversity was the highest in the Pululahua population. Central and southern populations have a higher proportion of clones than Pululahua. Bayesian and cluster analysis suggest that the yellow flower type evolved at least three times along the Ecuadorean Andes. In contrast to other Phaedranassa species, Phaedranassa viridiflora shows lower genetic diversity, which is likely related to a vegetative reproductive strategy.     

The role of nurse plants in the establishment of shrub seedlings in the semi-arid subtropical Andes

In the nurse plant syndrome, or nurse association, seedlings (beneficiaries) are associated with adult shrubs/trees (benefactors). This phenomenon has been documented in several regions of the planet. Abiotic stress amelioration (one mechanism of facilitation) is one of the causes of this association. Most of the studies addressing the nurse syndrome have been conducted on spatial scales of a few hectares and have focused on only one or a few species. Moreover, there is an almost complete lack of studies addressing the incidence and characteristics of the nurse phenomenon in the arid Andes of South America. We undertook a first approximation to the study of facilitation in these ecosystems. The study was conducted at local and regional scales and involved the assessment of the spatial distribution of juveniles (seedlings and saplings) of 51 populations of 16 shrub and 12 cactus species in relation to shrub cover at 20 localities of the Prepuna (subtropical Andes of Bolivia and Argentina, 20–26°S). In terms of spatial distribution, the juveniles of most of the populations of shrubs studied were distributed both under the shrubs and in open spaces, thereby showing an apparent indifference to microhabitat. Globose and opuntioid cacti were preferentially distributed below the canopies of shrubs and were usually more associated with the dominant shrub species, which stood out as better potential nurses. The pattern was consistent throughout the region, including the more mesic and arid localities. The fact that Prepuna woody species are capable of establishing in open spaces would confer this region a greater resilience. Our findings further suggest that community dynamics in arid and semi-arid environments are more variable than previously thought.     

Comparative phylogeography of short-tailed bats (Carollia: Phyllostomidae)

This is the first study of comparative phylogeography involving closely related species of Neotropical bats of the family Phyllostomidae. We compared patterns of geographical variation within the five species of fruit-eating bats currently recognized in the genus Carollia using the complete mitochondrial cytochrome-b gene. Our results suggest that the combined effect of the uplift of the Andes and the Panamanian land bridge has been as important for bats as for terrestrial mammals in shaping present-day biodiversity in the New World tropics. Species in this genus can be arranged in two highly supported clades, with a deep subdivision within each that corresponds well to differences across the Andes. We found three congruent phylogeographical patterns across species in this genus. First, the closer relationship between samples from western Ecuador and those from Central America, compared with populations east of the Andes in C. brevicauda, C. castanea and C. perspicillata. Second, the likelihood of a similar timing in South America for the arrival and diversification of C. brevicauda and C. perspicillata from their Central America ancestors. Third, the expansion of C. perspicillata and C. sowelli into northwestern Central America in the relatively recent past. Using a molecular clock, with rates ranging from 2.3 to 5% per 10(6) years, diversification within Carollia would have occurred over the last 1-4.5 Myr. These estimates agree well with the last rise of the Northern Andes and the Panama isthmus.     

Divergence with gene flow and fine-scale phylogeographical structure in the wedge-billed woodcreeper, Glyphorynchus spirurus, a Neotropical rainforest bird

Determining the relative roles of vicariance and selection in restricting gene flow between populations is of central importance to the evolutionary process of population divergence and speciation. Here we use molecular and morphological data to contrast the effect of isolation (by mountains and geographical distance) with that of ecological factors (altitudinal gradients) in promoting differentiation in the wedge-billed woodcreeper, Glyphorynchus spirurus , a tropical forest bird, in Ecuador. Tarsus length and beak size increased relative to body size with altitude on both sides of the Andes, and were correlated with the amount of moss on tree trunks, suggesting the role of selection in driving adaptive divergence. In contrast, molecular data revealed a considerable degree of admixture along these altitudinal gradients, suggesting that adaptive divergence in morphological traits has occurred in the presence of gene flow. As suggested by mitochondrial DNA sequence data, the Andes act as a barrier to gene flow between ancient subspecific lineages. Genome-wide amplified fragment length polymorphism markers reflected more recent patterns of gene flow and revealed fine-scale patterns of population differentiation that were not detectable with mitochondrial DNA, including the differentiation of isolated coastal populations west of the Andes. Our results support the predominant role of geographical isolation in driving genetic differentiation in G. spirurus , yet suggest the role of selection in driving parallel morphological divergence along ecological gradients.     

Biogeographic area relationships of lowland Neotropical rainforest based on raw distributions of vertebrate groups

Hypotheses of the historic biogeography of Neotropical anurans inhabiting lowland forests were generated using Parsimony Analysis of Endemicity. In order to establish comparisons with the biogeographical patterns of other vertebrates, previous cladistic analyses reported in the literature (for lizards and primates) were extended and reanalysed to match the geographical scope of the anuran analysis. Cladistic analysis of the distribution of 335 anuran species at 14 localities showed two regions that form a basal dichotomy: (1) Central America + Choco and (2) Amazon Basin + Brazilian Atlantic Forest. This result is interpreted as the first vicariance event that separated lowland Neotropical rainforests into Cis-Andean (east from the Andes) and Trans-Andean (west from the Andes) areas. Within the Cis-Andean localities, the earliest separation occurred between the Amazon Basin and the Brazilian Atlantic Forest. Within the Amazon Basin, three distinctive clusters are defined: (1) Belem, (2) Guianan Region, and (3) Upper Amazon Basin. Data sets on the distribution of anurans, lizards, and mammals have strong cladistic signal. Strong congruence exists among the area cladograms of anurans, lizards, and primates. All of them have, or at least did not conflict with: (1) a basal separation between Cis- and Trans-Andean regions, (2) a Central American clade, (3) the Choco Region is sister to the Central American clade, (4) an Amazon Basin clade, (5) an Upper Amazon Basin clade, and (6) a Guianan clade. The area cladograms are dichotomous and therefore do not support biogeographic theories that hypothesize simultaneous isolations of biotas in the Neotropics.     

Cryptic diversity and deep divergence in an upper Amazonian leaflitter frog, <Emphasis Type="Italic">Eleutherodactylus ockendeni</Emphasis>

Background  

The forests of the upper Amazon basin harbour some of the world's highest anuran species richness, but to date we have only the sparsest understanding of the distribution of genetic diversity within and among species in this region. To quantify region-wide genealogical patterns and to test for the presence of deep intraspecific divergences that have been documented in some other neotropical anurans, we developed a molecular phylogeny of the wide-spread terrestrial leaflitter frog Eleutherodactylus ockendeni (Leptodactylidae) from 13 localities throughout its range in Ecuador using data from two mitochondrial genes (16S and cyt b; 1246 base pairs). We examined the relation between divergence of mtDNA and the nuclear genome, as sampled by five species-specific microsatellite loci, to evaluate indirectly whether lineages are reproductively isolated where they co-occur. Our extensive phylogeographic survey thus assesses the spatial distribution of E. ockendeni genetic diversity across eastern Ecuador.