Lowland Species Research Articles

Fitness of natural willow hybrids in a pioneer mosaic hybrid zone.

Hybrid fitness is an important parameter to predict the evolutionary consequences of a hybridization event and to characterize hybrid zones. We studied fitness parameters of F1 and later‐generation hybrids between the lowland species Salix purpurea and the alpine S. helvetica that have recently emerged during colonization of an alpine glacier forefield. Fruit production (number of capsules per catkin and fruit set) did not differ between hybrids and parents, but the number of seeds per capsule of F1 hybrids was slightly lower than that of later‐generation hybrids and of the parents. Germination rates and seedling growth were tested on three substrates (pH 4.5, 7.0, and 8.0). Germination rates of seeds collected from F1 hybrids were lower on acid and neutral substrates, but equal at pH 8.0 compared to all other groups, while the seeds from later‐generation hybrids performed as well as the parents on all three substrates. In seedling growth, the colonizer S. purpurea performed better than all other taxa on all three substrates, while hybrids resembled the subalpine species S. helvetica. Results suggest that endogenous selection acts against F1 hybrids, but favors fitter genotypes in later‐generation hybrids. Exogenous selection via soil pH appears to be weak during seedling establishment. The pioneer vegetation on the glacier forefield may offer sufficient niche space for hybrid seedlings. Owing to the relatively high fitness of the hybrids and the scattered distribution of hybrids and parental individuals on the glacier forefield, this hybrid zone can be assigned to a mosaic model, probably facilitating gene flow and introgression between the parental species. As establishment of the hybrid zone appears to be linked to a colonization process, we propose to call it a pioneer mosaic hybrid zone.

Habitat and conservation status of molinillo (Magnolia sambuensis) and laurel arenillo (Magnolia katiorum), two endangered species from the lowland, Colombia

The conservation of the Magnoliaceae family is considered a priority worldwide, especially in Colombia, where all of its 36 species are seriously endangered while frustratingly little is known about their ecology. We therefore, assessed some ecological aspects of two lowland forest species: laurel arenillo ( Magnolia katiorum) and molinillo ( M. sambuensis). The first is an endemic species of the Urabá region in northwestern Colombia, and the second is distributed from Panama to Colombia along the Chocó Biogeographical region. Both species have a very low tree density and are found in forest stands that are disturbed by the timber exploitation of valuable woody species. The trees of both species had an average height of 26 m and shared similar environmental conditions such as soil (25°C) and air (28°C) temperatures, soil moisture (113%), and slope (25%). A redundancy analysis showed that each Magnolia species grows in a different community. Laurel arenillo was not a clear dominant species in a more diverse forest community, and was positively associated to altitude, longitudinal slope, and soil humidity. In contrast, molinillo was positively associated to air temperature and transversal slope and was negatively associated with longitudinal slope. This species was the most dominant in its community, which was less diverse than that of laurel arenillo. The last species should be considered a priority for conservation while the conservation status of molinillo should be further reviewed. Both species displayed large, fragrant, eye-catching flowers throughout most of the year.

Dual colonization of the Palaearctic from different regions in the Afrotropics by Senecio

AbstractAimInvestigation of the geographical origin of Senecio and the colonization of the Palaearctic including the relationship of life‐history strategy and elevational distribution in the source area and the colonized area.LocationWorldwide with a focus on the Afrotropic and the Palaearctic.MethodsSampling focused on adding species from the Afrotropic to existing datasets of Senecio. Two nuclear markers and three chloroplast markers were amplified and sequenced. Maximum likelihood and Bayesian inference were used to infer phylogeny, divergence times, biogeographical history and life‐history strategy evolution.ResultsSenecio originated most likely during the Mid to Late Miocene in western southern Africa (WSA). Colonization of the Palaearctic probably started from two different areas. One lineage of mostly annual lowland species seems to have originated in WSA, whereas the other lineage of perennial mountain species originated in mountainous areas of eastern southern Africa or tropical/subtropical East Africa.Main conclusionsSenecio evolved in arid or semi‐arid regions in WSA during a period of ongoing aridification and broad‐scale biome changes at the onset of a winter rainfall regime in the Greater Cape Floristic Region. Life‐history strategy and elevational distribution of the two Palaearctic lineages did not change between the source areas and the colonized areas.

SpeciesGeoCoder: Fast Categorization of Species Occurrences for Analyses of Biodiversity, Biogeography, Ecology, and Evolution.

Understanding the patterns and processes underlying the uneven distribution of biodiversity across space constitutes a major scientific challenge in systematic biology and biogeography, which largely relies on effectively mapping and making sense of rapidly increasing species occurrence data. There is thus an urgent need for making the process of coding species into spatial units faster, automated, transparent, and reproducible. Here we present SpeciesGeoCoder, an open-source software package written in Python and R, that allows for easy coding of species into user-defined operational units. These units may be of any size and be purely spatial (i.e., polygons) such as countries and states, conservation areas, biomes, islands, biodiversity hotspots, and areas of endemism, but may also include elevation ranges. This flexibility allows scoring species into complex categories, such as those encountered in topographically and ecologically heterogeneous landscapes. In addition, SpeciesGeoCoder can be used to facilitate sorting and cleaning of occurrence data obtained from online databases, and for testing the impact of incorrect identification of specimens on the spatial coding of species. The various outputs of SpeciesGeoCoder include quantitative biodiversity statistics, global and local distribution maps, and files that can be used directly in many phylogeny-based applications for ancestral range reconstruction, investigations of biome evolution, and other comparative methods. Our simulations indicate that even datasets containing hundreds of millions of records can be analyzed in relatively short time using a standard computer. We exemplify the use of SpeciesGeoCoder by inferring the historical dispersal of birds across the Isthmus of Panama, showing that lowland species crossed the Isthmus about twice as frequently as montane species with a marked increase in the number of dispersals during the last 10 million years.

Herbivory and nutrient limitation protect warming tundra from lowland species' invasion and diversity loss.

Herbivory and nutrient limitation can increase the resistance of temperature-limited systems to invasions under climate warming. We imported seeds of lowland species to tundra under factorial treatments of warming, fertilization, herbivore exclusion and biomass removal. We show that warming alone had little impact on lowland species, while exclusion of native herbivores and relaxation of nutrient limitation greatly benefitted them. In contrast, warming alone benefitted resident tundra species and increased species richness; however, these were canceled by negative effects of herbivore exclusion and fertilization. Dominance of lowland species was associated with low cover of tundra species and resulted in decreased species richness. Our results highlight the critical role of biotic and abiotic filters unrelated to temperature in protecting tundra under warmer climate. While scarcity of soil nutrients and native herbivores act as important agents of resistance to invasions by lowland species, they concurrently promote overall species coexistence. However, when these biotic and abiotic resistances are relaxed, invasion of lowland species can lead to decreased abundance of resident tundra species and diminished diversity.

Do thermoregulatory costs limit altitude distributions of Andean forest birds?

Summary Along tropical mountains, species often occupy narrow altitude ranges. Numerous biotic and abiotic factors have been proposed as determinants of altitude occupancy. We measured several aspects of thermal physiology of 215 bird species across a 2·6‐km altitude gradient in the Peruvian Andes. We predicted that highland species would show adaptation to the colder high‐altitude climate and that energy costs of thermoregulation might limit upslope dispersal of lowland natives. We found reductions in thermal conductance, body temperature and lower critical temperature in highland birds compared with lowland species. These combine to make highland natives more resistant to heat loss. We did not find convincing evidence that acute thermal limits or energy costs of thermoregulation constrained altitude distributions. Heat‐budget models predicted low‐to‐moderate long‐term costs at native altitudes. Costs increased for lowland natives modelled in the highland climate, but for all but a few species, costs remained within putative expenditure limits. Although we did not test heat tolerances, we measured all species at temperatures similar to the hottest air temperatures at the lowland site. There was no evidence that high lowland temperatures preclude downslope movements of highland birds. While thermal tolerances probably do not directly determine altitude occupancy by most species, the additional energy cost of thermoregulation experienced by lowland species moving upslope may trade off against investment in important life‐history components such as breeding, and thereby affect altitude range limits. A Lay Summary is available for this article.

Reproductive ecology of a Tibetan frog Nanorana parkeri (Anura: Ranidae)

ABSTRACTEndemic to the Tibetan Plateau, Nanorana parkeri figures among the highest altitude amphibians in the world. The present work, which was conducted in an alpine marsh at 4300 m altitude, is the first report of the species’ reproductive ecology. The breeding season, as indicated by the presence of amplectant pairs and fresh eggs in ponds, lasted from early Mary to early July. The ratios of adult male to female were 1.3 in hibernation ponds and 2.3 in spawning ones. Males were smaller than females. Amplexus was typically axillary (81.7%), with a few exceptional amplexus in which a male clasped another male or object rather than gravid females. Scramble competition occurred in 6.8% of the normal amplexus located. The body sizes of amplexed couples correlated positively. Deposited eggs were attached to submerged plants solitarily (60.5%) or in small clusters (39.5%), and the scattered arrangement of individual eggs could be a strategy to ensure embryonic development in oxygen-poor environments. Clutch size averaged 189 [standard deviation (SD) = 64], and correlated positively with female body size; egg size averaged 2.18 mm (SD = 0.10), independent of female body size; both traits within a clutch were negatively related with each other even controlling for female body size. Relatively few and large eggs laid by the Tibetan frogs, compared with those by lowland aquatic-spawning anuran species, should be adaptive to the harsh high-altitude conditions. Larvae hatched from eggs after 16.4 days on average (SD = 5.8). Froglets emerged between late July and early August, but some tadpoles were present in winter. These findings may help to understand the evolution of amphibians in extreme environments and to protect these remote species.

Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru.

We examined whether variations in photosynthetic capacity are linked to variations in the environment and/or associated leaf traits for tropical moist forests (TMFs) in the Andes/western Amazon regions of Peru. We compared photosynthetic capacity (maximal rate of carboxylation of Rubisco (Vcmax ), and the maximum rate of electron transport (Jmax )), leaf mass, nitrogen (N) and phosphorus (P) per unit leaf area (Ma , Na and Pa , respectively), and chlorophyll from 210 species at 18 field sites along a 3300-m elevation gradient. Western blots were used to quantify the abundance of the CO2 -fixing enzyme Rubisco. Area- and N-based rates of photosynthetic capacity at 25°C were higher in upland than lowland TMFs, underpinned by greater investment of N in photosynthesis in high-elevation trees. Soil [P] and leaf Pa were key explanatory factors for models of area-based Vcmax and Jmax but did not account for variations in photosynthetic N-use efficiency. At any given Na and Pa , the fraction of N allocated to photosynthesis was higher in upland than lowland species. For a small subset of lowland TMF trees examined, a substantial fraction of Rubisco was inactive. These results highlight the importance of soil- and leaf-P in defining the photosynthetic capacity of TMFs, with variations in N allocation and Rubisco activation state further influencing photosynthetic rates and N-use efficiency of these critically important forests.

Climate, habitat associations and the potential distributions of Neotropical birds: Implications for diversification across the Andes

En contraste con la visión tradicional de que el levantamiento de los Andes impulsó la diversificación biótica causando vicarianza en varios grupos con distribuciones que antes eran continuas, investigaciones recientes sugieren que eventos de dispersión a través de los Andes sucedidos después de los principales episodios de levantamiento fueron catalizadores principales de la especiación en aves neotropicales, y que la habilidad de los linajes para persistir y dispersarse en el paisaje predice fuertemente los patrones de especiación. Sin embargo, se desconoce cuándo y dónde sucedieron dichos eventos de dispersión, y el papel de las fluctuaciones climáticas y el consecuente desplazamiento de la vegetación como promotores de la dispersión a través de los Andes en distintos momentos no ha sido estudiado. Empleamos modelos de la distribución potencial de especies basados en datos de clima actual e histórico para examinar escenarios de dispersión a través de los Andes en 41 especies de aves neotropicales de tierras bajas con diferentes afinidades de hábitat. Nuestros resultados indican que la conectividad ecológica que favorecería la dispersión a través de los Andes en el presente es mayor en pasos bajos de la cordillera del sur de Ecuador y el norte de Perú que en pasos bajos de Colombia, y este patrón espacial parece haberse mantenido en cuatro momentos diferentes de los últimos 130,000 años. También encontramos que aunque algunas áreas actualmente no serían propicias para la dispersión de las aves a través de los Andes, bajo condiciones climáticas del pasado (i.e. durante períodos más fríos y secos) presentaron condiciones climáticas sustancialmente más idóneas para permitir la conectividad ecológica de poblaciones a través de la cordillera. No encontramos diferencias consistentes en la conectividad ecológica presente y pasada a través de los Andes entre especies de diferentes tipos de hábitat. Sugerimos que los valles andinos impulsan la diversificación evolutiva no solo porque aíslan las poblaciones de especies de montaña, sino porque permiten la dispersión episódica de especies de las tierras bajas. Nuestros modelos permiten hacer predicciones sobre flujo genético que pueden ser evaluadas en estudios futuros que examinen patrones de intercambio genético a escala fina usando herramientas de genética del paisaje. © Acad. Colomb. Cienc. Ex. Fis. Nat. 2016.

Hybridization as a threat in climate relict Nuphar pumila (Nymphaeaceae)

Field studies and conceptual work on hybridization-mediated extinction risk in climate relicts are extremely rare. Nuphar pumila (Nymphaeaceae) is one of the most emblematic climate relicts in Europe with few isolated populations in the Alpine arc. The extent of introgression with related lowland and generalist species Nuphar lutea has never been studied using molecular methods. All biogeographical regions where N. pumila naturally occurs in the neighbourhood of the Alpine arc were sampled and studied using nuclear microsatellite markers. Furthermore, we used forward-in-time simulations and Approximate Bayesian Computation to check whether an introgression scenario fits with the observed admixture patterns and estimated the demographic parameters associated with this process. Our study confirms ongoing hybridization between N. pumila and N. lutea and validates it by the use of population models. More than 40 % of investigated N. pumila individuals were admixed and hybrids were found in over 60 % of studied populations. The introgression is bidirectional and is most likely a result of very recent gene flow. Our work provides strong evidence for rapid extinction risk and demographic swamping between specialized climatic relicts and closely related generalists. The remaining pure populations of N. pumila are rare in the Alpine arc and deserve high conservation priority.

Spread of Amphibian Chytrid Fungus across Lowland Populations of Túngara Frogs in Panamá.

Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd), is an emergent infectious disease partially responsible for worldwide amphibian population declines. The spread of Bd along highland habitats (> 500 meters above sea level, m a.s.l.) of Costa Rica and Panamá is well documented and has been linked to amphibian population collapses. In contrast, data are scarce on the prevalence and dispersal of Bd in lowland habitats where amphibians may be infected but asymptomatic. Here we describe the spread (2009 to 2014) of Bd across lowland habitats east of the Panamá Canal (< 500 m a.s.l.) with a focus on the Túngara frog (Physalaemus [Engystomops] pustulosus), one of the most common and abundant frog species in this region. Highland populations in western Panamá were already infected with Bd at the start of the study, which was consistent with previous studies indicating that Bd is enzootic in this region. In central Panamá, we collected the first positive samples in 2010, and by 2014, we detected Bd from remote sites in eastern Panamá (Darién National Park). We discuss the importance of studying Bd in lowland species, which may serve as potential reservoirs and agents of dispersal of Bd to highland species that are more susceptible to chytridiomycosis.

Species delimitation, phylogeny and evolutionary demography of co-distributed, montane frogs in the southern Brazilian Atlantic Forest

The Brazilian Atlantic Forest (BAF) is recognized as one of the world’s biodiversity hotspots, with even more species per unit of area than the Amazon, however the mechanisms that led to such astonishing diversity are yet to be fully understood. In this study, we investigate the diversification of two co-distributed frog genera associated with montane areas of southern BAF: Melanophryniscus (Bufonidae) and Brachycephalus (Brachycephalidae). Species delimitation methods using mitochondrial and nuclear loci supported the existence of a remarkable number of highly endemic species in each genus, most of which occupy only one or a few adjacent mountaintops. Their timing of diversification was highly congruent, supporting recent speciation events within the past 600 thousand years. Extended Bayesian skyline plots indicate that most populations have remained relatively stable in size across the evolutionary past, with recent growth after 0.15My, suggesting that the drastic changes found in previous studies on lowland frog species were not shared by these montane taxa. These results are consistent with the existence of a montane refugium in southern BAF, allowing species persistence through the climatic shifts experienced along the BAF during the Quaternary.

Cryptic diversity in Afro-tropical lowland forests: The systematics and biogeography of the avian genus Bleda

Recent investigations of distributional patterns of Afro-tropical lowland forest species have demonstrated to some degree our overall lack of understanding involving historical diversification patterns. Traditionally, researchers have relied upon two hypotheses, each of which views the lowland forest of Africa in differing roles. The Pleistocene Forest Refuge Hypothesis (PFRH) posits that biogeographic patterns of avian lowland species are explained via allopatric speciation during forest fragmentation cycles in the Pleistocene epoch (c. 1.8Ma–11,700Ka). The Montane Speciation Hypothesis (MSH) countered by suggesting that lowland forests are “evolutionary museums” where species, which originally evolved in montane forest refuge centers, remained without further diversification. Furthermore, investigations have largely regarded widespread, avian species which lack phenotypic variability as biogeographically “uninformative”, with regards to historical biogeographic patterns. To test the tenets of these ideas, we investigated the systematics and biogeography of the genus Bleda, whose constituent species are restricted to lowland forest and are lacking in phenotypic variation. Using extracted DNA from 179 individuals, we amplified two mitochondrial genes and three nuclear loci and utilized Bayesian phylogenetic methods and molecular clock dating to develop a time-calibrated phylogeny of Bleda. We used LaGrange to develop an ancestral area reconstruction for the genus. Haplotype networks for three species were generated using Network. We recovered the four currently recognized species of Bleda, plus a monophyletic B. ugandae, a current sub-species which may warrant full species status. We found that the origins of the genus Bleda are estimated to be in the Upper Guinean forests of West Africa, dating to the Miocene (c. 7.5Ma), while the speciation events for the rest of the genus are dated to the Pliocene (c. 5–1.8Ma). Our analyses recovered discrete and highly differentiated geographic structuring of genetic diversity in West and Central Africa in three of five species, with many of the diversification events dating to the Pleistocene. The biogeographic patterns observed in Bleda can be explained through a combination of isolation via forest refuges during the Plio–Pleistocene and riverine barriers limiting secondary contact after forest expansion. We find evidence for the PFRH as a driver of intra-specific diversity, but conclude that it does not facilitate an explanation for speciation in the genus Bleda. The “evolutionary museum” concept furnished by the MSH is countered by our evidence of in situ diversification in the lowland forests of Africa. Additionally, our results provide strong evidence of the value of seemingly “uninformative” widespread avian taxa for revealing complex patterns of forest diversity. Overall, our study highlights that past researchers have both underestimated the amount of diversity found in lowland forests and failed to understand the complexity of historical forces shaping that diversity. Gaining a better understanding of lowland forest diversity and the historical factors which have shaped it will crucial in determining conservation tactics in the near future.

East Asian Lobelioideae and ancient divergence of a giant rosette Lobelia in Himalayan Bhutan

AbstractLobelioideae comprise about 1200 species in 30 genera worldwide and include famous cases of parallel evolution in the “giant rosette” lobelias occurring in East Africa, Hawaii, and Bhutan. To date, no phylogeny has included the Himalayan giant lobelia, and its age and phylogenetic distance from the Hawaiian and African giant species (the latter in the same section as the Bhutan species) are therefore unknown. Asian Lobelioideae in general have received little attention, partly because of the difficult access to material. Here we use nuclear and plastid markers of Lobelioideae (with worldwide sampling) to test the monophyly of the section that includes most Asian lobelias, and we then use a reduced concatenated matrix to resolve the position of Lobelia nubigena from Bhutan and of the likewise cold‐adapted lowland species L. sessilifolia. Most sequences come from herbarium material. Lobelia nubigena diverged from its Asian lowland relatives 13.8 (12.6–15) million years ago (about the time of the Bhutan Himalaya orogeny), and in an unexpected twist, L. sessilifolia is the sister species to a huge clade (ca. 265 species) that comprises African, South American, Asian, and the entire Hawaiian Lobelioideae radiation. The African and Hawaiian high‐elevation lobelias in our sample are younger than L. nubigena, although the time when gigantism evolved in this species cannot be precisely inferred. Our results support the previously hypothesized Asian origin of (1) the Hawaiian Lobelioideae, (2) Southeast Asian islands lobelias, and (3) the African Lobelia‐Rynchopetalum group, which also reached South America.

Mountain roads shift native and non‐native plant species' ranges

Roads are known to act as corridors for dispersal of plant species. With their variable microclimate, role as corridors for species movement and reoccurring disturbance events, they show several characteristics that might influence range dynamics of both native and non‐native species. Previous research on plant species ranges in mountains however seldom included the effects of roads. To study how ranges of native and non‐native species differ between roads and adjacent vegetation, we used a global dataset of plant species composition along mountain roads. We compared average elevation and range width of species, and used generalized linear mixed models (GLMMs) to compile their range optimum and amplitude. We then explored differences between roadside and adjacent plots based on a species’ origin (native vs non‐native) and nitrogen and temperature affinity. Most non‐native species had on average higher elevational ranges and broader amplitudes in roadsides. Higher optima for non‐native species were associated with high nitrogen and temperature affinity. While lowland native species showed patterns comparable to those in non‐native species, highland native species had significantly lower elevational ranges in roadsides compared to the adjacent vegetation. We conclude that roadsides indeed change the elevational ranges of a variety of species. These changes are not limited to the expansion of non‐native species along mountain roads, but also include both upward and downward changes in ranges of native species. Roadsides may thus facilitate upward range shifts, for instance related to climate change, and they could serve as corridors to facilitate migration of alpine species between adjacent high‐elevation areas. We recommend including the effects of mountain roads in species distribution models to fine‐tune the predictions of range changes in a warming climate.

Facing the Heat: Thermoregulation and Behaviour of Lowland Species of a Cold-Dwelling Butterfly Genus, Erebia.

Understanding the potential of animals to immediately respond to changing temperatures is imperative for predicting the effects of climate change on biodiversity. Ectothermic animals, such as insects, use behavioural thermoregulation to keep their body temperature within suitable limits. It may be particularly important at warm margins of species occurrence, where populations are sensitive to increasing air temperatures. In the field, we studied thermal requirements and behavioural thermoregulation in low-altitude populations of the Satyrinae butterflies Erebia aethiops, E. euryale and E. medusa. We compared the relationship of individual body temperature with air and microhabitat temperatures for the low-altitude Erebia species to our data on seven mountain species, including a high-altitude population of E. euryale, studied in the Alps. We found that the grassland butterfly E. medusa was well adapted to the warm lowland climate and it was active under the highest air temperatures and kept the highest body temperature of all species. Contrarily, the woodland species, E. aethiops and a low-altitude population of E. euryale, kept lower body temperatures and did not search for warm microclimates as much as other species. Furthermore, temperature-dependence of daily activities also differed between the three low-altitude and the mountain species. Lastly, the different responses to ambient temperature between the low- and high-altitude populations of E. euryale suggest possible local adaptations to different climates. We highlight the importance of habitat heterogeneity for long-term species survival, because it is expected to buffer climate change consequences by providing a variety of microclimates, which can be actively explored by adults. Alpine species can take advantage of warm microclimates, while low-altitude grassland species may retreat to colder microhabitats to escape heat, if needed. However, we conclude that lowland populations of woodland species may be more severely threatened by climate warming because of the unavailability of relatively colder microclimates.

Aboveground and belowground competition affect seedling performance and allometry in a tropical monsoon forest

Distinguishing the relative effects of above- and belowground competition can improve our understanding of the forces shaping community assembly in different ecosystems. In this study, we investigated the impacts of above- and belowground competition on seedling growth and allometry in a tropical monsoon forest (TMF) on Hainan Island, China. Four common dominant canopy tree species from three forest types (Schima superba in secondary forest, Peltophorum tonkinense and Vatica mangachapoi in lowland rainforest, and Terminalia nigrovenulosa in TMF) were planted in different treatments (understory vegetation removal and trenching to decrease above- and belowground competition, respectively). Three species had greater relative growth rates in height in the vegetation removal, but not in the trenching treatment. Vegetation removal reduced mortality rates for all species and increased biomass in three of the four. Trenching alone did not affect biomass for any species. Vegetation removal affected seedling allometry, resulting in higher leaf mass fraction in the lowland rainforest species P. tonkinense and V. mangachapoi. The secondary forest species, S. superba, did not successfully establish in plots with intact understory because of aboveground competition. The highly drought-adapted traits of T. nigrovenulosa allowed its seedlings to establish successfully in TMF, where it is a canopy dominant. Our results demonstrate that aboveground competition is more important than belowground competition in structuring seedling communities in TMF, because light availability largely limits seedling establishment in this ecosystem.

Amphibians of varzea environments and floating meadows of the oxbow lakes of the Middle Purus River, Amazonas, Brazil

Despite the historical knowledge on amphibians of the Purus River basin, basic information on assemblages is fragmented, with gaps in knowledge existing at various scales, which limits conservation strategies. This study provides information on the composition, richness and abundance of the amphibian fauna in varzea environments and floating meadows of the oxbow lakes of the Middle Purus River between the Boca do Acre and Pauiní municipalities, Amazonas, Brazil. We sampled six oxbow lakes using forty-seven 200-meter transects, distributed among the "floating meadows," "high varzea" and "low varzea," from April to January 2014. We recorded 59 species, with the family Hylidae being predominant. This amphibian fauna represents approximately 19% of the species known for the Amazon, 28% for Amazonas State and 45% of the species recorded so far in the Purus River, increasing the richness of the basin to 132 species. Eight species were considered rare, and 29 are endemic to the Amazon. This study adds to the knowledge on the amphibian species of the Amazonian lowlands, including the expansion of known distributions, as well as increases the knowledge of several species that are locally rare, endemic and/or that are data deficient regarding distribution and ecology.

Using chromosomal data in the phylogenetic and molecular dating framework: karyotype evolution and diversification in Nierembergia (Solanaceae) influenced by historical changes in sea level.

Karyotype data within a phylogenetic framework and molecular dating were used to examine chromosome evolution in Nierembergia and to infer how geological or climatic processes have influenced in the diversification of this solanaceous genus native to South America and Mexico. Despite the numerous studies comparing karyotype features across species, including the use of molecular phylogenies, to date relatively few studies have used formal comparative methods to elucidate chromosomal evolution, especially to reconstruct the whole ancestral karyotypes. Here, we mapped on the Nierembergia phylogeny one complete set of chromosomal data obtained by conventional staining, AgNOR-, C- and fluorescent chromosome banding, and fluorescent insitu hybridisation. In addition, we used a Bayesian molecular relaxed clock to estimate divergence times between species. Nierembergia showed two major divergent clades: a mountainous species group with symmetrical karyotypes, large chromosomes, only one nucleolar organising region (NOR) and without centromeric heterochromatin, and a lowland species group with asymmetrical karyotypes, small chromosomes, two chromosomes pairs with NORs and centromeric heterochromatin bands. Molecular dating on the DNA phylogeny revealed that both groups diverged during Late Miocene, when Atlantic marine ingressions, called the 'Paranense Sea', probably forced the ancestors of these species to find refuge in unflooded areas for about 2Myr. This split agrees with an increased asymmetry and heterochromatin amount, and decrease in karyotype length and chromosome size. Thus, when the two Nierembergia ancestral lineages were isolated, major divergences occurred in chromosomal evolution, and then each lineage underwent speciation separately, with relatively minor changes in chromosomal characteristics.

A new species of Microhyla (Anura: Microhylidae) from Sri Lanka: an integrative taxonomic approach.

Species boundaries of Microhyla rubra of India and Sri Lanka were assessed using the following criteria: genetic barcoding, morphology, and vocalization. We use a ca. 500 bp fragment of the 16S rRNA mitochondrial gene and show that there is an uncorrected pairwise distance of 2.7-3.2% between the Indian and Sri Lankan populations of M. rubra. We show that they are different in several call characteristics such as, dominant frequency, call duration, call rise time and pulse rate. Morphologically, the Sri Lankan population can be distinguished from the typical M. rubra described from southern India, by a combination of characters: body size, skin texture, and feet dimensions. We recognize the population from Sri Lanka as a new species, Microhyla mihintalei sp. nov., a widely distributed lowland species with an elevational distribution of up to 500 m a.s.l.