An integrative approach to understanding diversity patterns and assemblage rules in Neotropical bats.

Understanding the mechanisms shaping species composition of assemblages is critical for incorporating ecological and evolutionary perspectives into biodiversity conservation. Thus, we quantified the relative support of community assembly mechanisms by assessing how species richness relates to the functional and phylogenetic biodiversity of Neotropical bat assemblages. We assessed the association of functional diversity for functional categories and phylogenetic diversity with species richness for 20 assemblages of Neotropical bats. In addition, we contrasted functional and phylogenetic diversity against null models to determine the mechanisms that structure the assemblages. We hypothesize functional/phylogenetic overdispersion for high species sites and a positive relationship between those dimensions of diversity and richness. Functional divergence increased with species richness, indicating that the variability in ecological attributes among abundant bats increases as the assemblages contain more species. Taxa were more distantly related as richness increases, but distances among closely related species remained constant. We found a consistent tendency of clustering of functional traits in site assemblages, particularly in abundant species. We proposed competition between clades as a possible mechanism modulating the community structure in Neotropical bat assemblages. Our results suggest that decreasing overlap in functional traits between abundant species could promote coexistence with rare species that can buffer ecosystem function due to species loss.

Time partitioning among neotropical frugivorous bats: effects of phylogeny, body size, and abundance / Distribución del tiempo entre murciélagos frugívoros neotropicales: efectos de la filogenia, el tamaño corporal y la abundancia

Abstract Introduction: Resource partitioning by promoting coexistence is essential to determine species richness and composition in natural communities. However, the partitioning of time has been questioned as a mechanism that promotes the coexistence of ecologically similar species. Objective: To determine the importance of the partitioning of time as a mechanism that promotes coexistence, we compared the activity patterns of tropical frugivorous bats. Methods: We captured bats with mist nets from sunset to sunrise in three study sites (tropical dry forest, wet forest, and rainforest) to calculate activity patterns of the species using Kernel density estimation. We used the superposition coefficient (Δ1) to compare activity patterns between (1) bat assemblages of study sites, (2) frugivorous species in the same site, and (3) populations of the same species among different sites. To determine whether the overlap in the activity patterns was related to the ecological similarity of species, we evaluated the association between Δ1 and similarity in abundances and body mass and phylogenetic closeness. Results: We found geographical variations in the overall activity patterns of the assemblages of the three localities. Likewise, we found variations in activity patterns between species at each study site and between populations in different study sites. Overlap in activity patterns tended to decrease as species were phylogenetically more closely related and similar in abundance and body size. Conclusions: Our results provide empirical support for the role of temporal segregation in activity patterns as a mechanism that promotes the coexistence of ecologically similar species in nature.

Resumen Introducción: La partición de recursos es esencial para determinar la riqueza y composición de especies en las comunidades naturales. Sin embargo, la segregacion temporal ha sido cuestionada como un mecanismo que promueve la coexistencia de especies ecológicamente similares. Objetivo: Con el fin de determinar la importancia de la segregación en los patrones de actividad, como mecanismo que promueve la coexistencia; se compararon los patrones de actividad de murciélagos frugívoros neotropicales. Métodos: Se capturaron murciélagos con redes de niebla desde el atardecer hasta el amanecer en tres localidades (bosque seco tropical, bosque muy húmedo y bosque lluvioso submontano) con el fin de calcular los patrones de actividad de las especies empleando la estimación de densidad Kernel. Se utilizó el coeficiente de superposición (Δ) para comparar patrones de actividad entre: (1) ensamblajes de murciélagos de los sitios de estudio, (2) especies frugívoras en el mismo sitio, y (3) poblaciones de la misma especie en diferentes sitios. Resultados: Se encontraron variaciones geográficas en los patrones de actividad de los ensamblajes en las tres localidades. Asimismo, se encontraron variaciones en los patrones de actividad entre especies en cada sitio y entre poblaciones en diferentes sitios. Determinamos que la superposición en los patrones de actividad disminuyó a medida que las especies estaban más relacionadas filogenéticamente, similares en abundancia y tamaño corporal; lo que sugiere una tendencia hacia la segregación temporal de especies ecológicamente similares. Conclusión: Estos resultados proveen apoyo empírico del papel de la segregación temporal en los patrones de actividad como un mecanismo que promueve la coexistencia de especies ecológicamente similares en la naturaleza.

Comparative mitogenome phylogeography of two anteater genera (Tamandua and Myrmecophaga; Myrmecophagidae, Xenarthra): Evidence of discrepant evolutionary traits.

The species within Xenarthra (sloths, anteaters, and armadillos) are quintessential South American mammals. Of the three groups, Vermilingua (anteaters) contains the fewest extant and paleontological species. Here, we sampled and sequenced the entire mitochondrial genomes (mitogenomes) of two Tamandua species (Tamandua tetradactyla and Tamandua mexicana) (n=74) from Central and South America, as well as Myrmecophaga tridactyla (n=41) from South America. Within Tamandua, we detected three different haplogroups. The oldest (THI) contained many specimens with the T. tetradactyla morphotype (but also several with the T. mexicana morphotype) and originated in southeastern South America (currently Uruguay) before moving towards northern South America, where the THII haplogroup originated. THII primarily contained specimens with the T. mexicana morphotype (but also several with the T. tetradactyla morphotype) and was distributed in Central America, Colombia, and Ecuador. THI and THII yielded a genetic distance of 4%. THII originated in either northern South America or "in situ" in Central America with haplogroup THIII, which consisted of ~50% T. mexicana and 50% T. tetradactyla phenotypes. THIII was mostly located in the same areas as THII, i.e., Central America, Ecuador, and Colombia, though mainly in the latter. The three haplogroups overlapped in Colombia and Ecuador. Thus, T. tetradactyla and T. mexicana were not reciprocally monophyletic. For this reason, we considered that a unique species of Tamandua likely exists, i.e., T. tetradactyla. In contrast to Tamandua, M. tridactyla did not show different morphotypes throughout its geographical range in the Neotropics. However, two very divergent genetic haplogroups (MHI and MHII), with a genetic distance of ~10%, were detected. The basal haplogroup, MHI, originated in northwestern South America, whereas the more geographically derived haplogroup, MHII, overlapped with MHI, but also expanded into central and southern South America. Thus, Tamandua migrated from south to north whereas Myrmecophaga migrated from north to south. Our results also showed that temporal mitochondrial diversification for Tamandua began during the Late Pliocene and Upper Pleistocene, but for Myrmecophaga began during the Late Miocene. Furthermore, both taxa showed elevated levels of mitochondrial genetic diversity. Tamandua showed more evidence of female population expansion than Myrmecophaga. Tamandua experienced population expansion ~0.6-0.17 million years ago (Mya), whereas Myrmecophaga showed possible population expansion ~0.3-0.2 Mya. However, both taxa experienced a conspicuous female decline in the last 10 000-20 000 years. Our results also showed little spatial genetic structure for both taxa. However, several analyses revealed higher spatial structure in Tamandua than in Myrmecophaga. Therefore, Tamandua and Myrmecophaga were not subjected to the same biogeographical, geological, or climatological events in shaping their genetic structures.

Elevation does not strongly influence interspecific variation in body size of small Tropical endotherms / La elevación no influencia fuertemente la variación interespecífica en el tamaño corporal de los endotermos tropicales pequeños

Introduction: Body size is an essential trait for endotherms to face the physiological requirements of cold, so there is a tendency to large body size at high altitudes and latitudes, known as Bergmann's rule. However, the validity of this ecomorphological rule to small-bodied endotherms across altitudinal gradients is poorly known. Objective: To understand the effects of environmental variation on body size, we assessed whether interspecific variation in body size of small tropical endotherms follows Bergmann's rule along tropical altitudinal gradients. Methods: We compiled data on elevational ranges and body masses for 133 species of hummingbirds of Colombia. We then assessed the association between body mass and mid-point of the altitudinal distribution using phylogenetic generalized least squares (PGLS) analyses under different evolutionary models. Results: We found a decelerating rate of evolution for body size since the Early Burst model of evolution provided a better fit to body mass data. For elevational range, we found a slow and constant rate since Pagel's lambda model provided a better fit to the mid-point of the altitudinal distribution data. Besides, phylogenetic regression analysis indicated that body mass and the altitudinal range of hummingbirds are associated through the phylogeny, with a positive but slight association (R2= 0.036). Conclusions: We found that body mass and altitude of hummingbirds are positively related, which is in agreement with expectations under Bergmann's rule. However, this association was weaker than expected for small and non-passerine birds like hummingbirds. Thus, our results suggest that environmental changes across altitudinal gradients do not strongly influence body mass in small tropical endotherms as hummingbirds.

Introducción: El tamaño corporal es un rasgo importante para determinar la respuesta de los endotermos a los requerimientos que exigen las zonas frías, por lo cual se espera una tendencia hacia el incremento del tamaño corporal al aumentar la altitud y la latitud. Sin embargo, se conoce poco acerca de la validez de esta regla ecomorfológica, conocida como la regla de Bergmann, para endotermos pequeños en gradientes altitudinales tropicales. Objetivo: Con el fin de entender los efectos de la variación ambiental sobre el tamaño corporal, se evaluó sí la variación interespecífica en la masa corporal de endotermos tropicales pequeños se ajusta a la regla de Bergmann a lo largo de gradientes de elevación. Métodos: Se compilaron datos sobre los rangos de distribución altitudinal y los tamaños corporales de 133 especies de colibríes en Colombia. Posteriormente, se evaluó la asociación entre la masa corporal y el punto medio de distribución altitudinal de los colibríes mediante análisis de mínimos cuadrados generalizados filogenéticos (PGLS) bajo diferentes modelos evolutivos. Resultados: La evolución de la masa corporal se ajustó mejor a un modelo de evolución Early Burst, mientras que el rango de elevación al modelo evolutivo lambda de Pagel; lo que indica que la tasa de evolución es desacelerada para el tamaño del cuerpo, mientras es lenta y constante para el rango de elevación. Además, el análisis de regresión filogenética indica que la masa corporal y el rango de elevación están positiva y ligeramente asociados (R2 = 0.036). Conclusiones: De acuerdo con lo esperado por la regla de Bergmann, los resultados indican que los colibríes tienden a ser más grandes a mayores altitudes. Sin embargo, esta asociación es más débil de lo esperado para aves no paseriformes de tamaño pequeño como los colibríes.Por lo tanto, los resultados sugieren que las variaciones ambientales a lo largo de gradientes de elevación no tienen una influencia fuerte sobre el tamaño corporal de endotermos pequeños como los colibríes.

Taxonomic identity of an insular population of sac-winged bat Saccopteryx (Chiroptera: Emballonuridae)

Las poblaciones insulares frecuentemente difieren de las continentales, en morfológica y comportamiento. Consecuentemente, utilizar características externas puede dificultar la identificación taxonómica de especies en las islas. En el Parque Nacional Natural Gorgona, particularmente, ha existido controversia sobre la identidad de varias especies de murciélagos residentes. Nuestro objetivo fue determinar la identidad taxonómica de la población de Saccopteryx de esta isla, que tiene la particularidad de tener actividad diurna. Evaluamos las diferencias entre las poblaciones mediante morfometría (tradicional y geométrica) y llamados de ecolocalización. Además, debido a que las especies difieren en su sistema de apareamiento, evaluamos la conformación de los grupos en los refugios. Pese a la variación en la morfometría craneal y los llamados de ecolocalización entre las poblaciones, la población de Gorgona fue más similar a S. leptura que a S. bilineata. Las conformaciones de los grupos en los refugios sugieren un sistema monógamo como en S. leptura. Por ende, concluimos que la especie de la isla es S. leptura. Adicionalmente, la historia biogeográfica y el aislamiento geográfico de la isla sugieren que esta población puede ser diferente de sus contrapartes continentales debido al efecto fundador, deriva genética y/o adaptaciones a las condiciones ambientales locales.

Insular populations can differ from their continental counterparts since they are subject to particular pressures, biotic and abiotic, that can lead to variations in morphology and behavior; thus, the use of external characters may complicate taxonomic identification of insular populations. Even though the bat assemblage of Gorgona Island is a conservation target, there has been controversy about the taxonomic identity of bats from this Colombian National Natural Park. Specifically, our goal was to determine the taxonomic identity of the population of the sac-winged bat (genus Saccopteryx) resident in Gorgona, which have the peculiarity of having diurnal activity. Thus, we assessed differences among populations using morphometrics (traditional and geometric) and echolocation calls. Additionally, since the species of Saccopteryx involved have different mating systems, we assessed roosting association of Gorgona population as a surrogate for mating system. Despite the variation in skull morphometrics and echolocation calls among populations, Gorgona population was more similar to S. leptura than to S. bilineata. Besides, evolutionary allometry was significant, but size explained a small proportion of shape differences among populations. Finally, roosting association of Gorgona population suggests a monogamous mating system such as in S. leptura. We conclude that Saccopteryx from Gorgona corresponds to S. leptura. In addition, biogeographic history and geographic isolation of the island suggest that this population may have differentiated from their mainland counterparts through founder effect, genetic drift and/or adaptation to the local biotic and abiotic environmental conditions.

Demography of population recovery: survival and fidelity of peregrine falcons at various stages of population recovery.

Factors influencing vital demographic rates and population dynamics can vary across phases of population growth. We studied factors influencing survival and fidelity of peregrine falcons in south Scotland-north England at two stages of population growth: when the population was recovering from pesticide-related declines and density was low, and when it had largely recovered from pesticide effects and density was high. Fidelity was higher for: adults and subadults than for juveniles, females than for males, and juveniles and adults during the low-density than during the high-density study period. Survival was age specific, with lower survival for juveniles than for older birds (juveniles, 0.600 ± SE 0.063; subadults, 0.811 ± 0.058; adults, 0.810 ± 0.034). Furthermore, there was some evidence that survival was generally lower for all age classes during the low-density period than during the high-density period, possibly due to a chronic, persistent effect of organochlorine pesticides as the population recovered. Evidence for a density-dependent effect on survival was weak, but a negative effect of density on fidelity of juveniles (dispersing age class) during the recovery phase suggests density-dependent dispersal when the population was increasing. Our results show how population density can influence demographic parameters differently and how such influences can vary across phases of population growth.

Murciélagos de cola corta (Carollia: Phyllostomidae) del Parque Nacional Natural Gorgona (Colombia) y sus implicaciones biogeográficas / Short-tailed bats (Carollia: Phyllostomidae) from Gorgona National Natural Park (Colombia) and its biogeographical implications

Taxonomy of short-tailed bats (Phyllostomidae: Carollia) has been unclear due to the extreme morphological similarity among species and the significant morphological variation within species. The identity of the Carollia species in Gorgona National Natural Park (Colombian Pacific) has been controversial due to the high morphological similarity between Carollia perspicillata and C. brevicauda. C. perspicillata is common in lowlands, and more likely to be the Gorgona species than C. brevicauda, while geological evidence suggests that common between 1 500 and 3 000m. We recorded eight measurements from jaws and skulls of confirmed C. perspicillata (24 individuals) and C. brevicauda (23) and 35 individuals captured in Gorgona. Discriminant analyses showed that, contrary to expectations based on current altitudinal distributions, the Gorgona population is morphologically closer to C. brevicauda. Biological evidence suggests that Gorgona was connected in the past to mainland South America. Gorgona may be the highest part of a now submerged mountain range (“Cordillera de la Costa”) that was part of the continent during the Pleistocene glaciations. Consequently, I hypothesize that C. brevicauda colonized Gorgona overland during last Pleistocene glaciations. Rev. Biol. Trop. 62 (Suppl. 1): 435-445. Epub 2014 February 01.

La identidad de la especie de Carollia (Chiroptera: Phyllostomidae) en el Parque Nacional Natural Gorgona ha sido controvertida debido a la alta similitud entre C. perspicillata y C. brevicauda. C. perspicillata es la especie con mayor probabilidad de habitar en la isla debido a que actualmente se distribuye en las tierras bajas de la costa pacífica colombiana, mientras C. brevicauda es una especie predominantemente de tierras altas. Con el objetivo de resolver la controversia acerca de la especie del género Carollia que habita la isla, se realizaron análisis morfométricos con base en característica de cráneos y mandíbulas de especímenes de las dos especies y del PNN Gorgona. Los resultados evidenciaron que, contrario a lo esperado, esta población insular es morfológicamente más similar a C. brevicauda. Evidencia biológica para diferentes grupos de organismos sugiere que Gorgona puede haber estado conectada a los andes y las zonas bajas del pacífico del sur de Colombia-norte de Ecuador en el pasado. Por otra parte, existe evidencia geológica que muestra que Gorgona es la parte más alta de una cordillera (Cordillera de la Costa) que se encuentra sumergida actualmente, pero que probablemente fue parte del continente durante el Pleistoceno. Por lo tanto, con base en esta evidencia biológica y geológica se plantea la hipótesis de una ruta de colonización de C. brevicauda a Gorgona desde los Andes a través de la Cordillera de la Costa. Este proceso pudo facilitarse por el descenso marino durante las glaciaciones del Pleistoceno.