Elevational and seasonal patterns of plant pollinator networks in two highland tropical ecosystems in Costa Rica.

Many plant species in high montane ecosystems rely on animal pollination for sexual reproduction, however, our understanding of plant-pollinator interactions in tropical montane habitats is still limited. We compared species diversity and composition of blooming plants and floral visitors, and the structure of plant-floral visitor networks between the Montane Forest and Paramo ecosystems in Costa Rica. We also studied the influence of seasonality on species composition and interaction structure. Given the severe climatic conditions experienced by organisms in habitats above treeline, we expected lower plant and insect richness, as well as less specialized and smaller pollination networks in the Paramo than in Montane Forest where climatic conditions are milder and understory plants are better protected. Accordingly, we found that blooming plants and floral visitor species richness was higher in the Montane Forest than in the Paramo, and in both ecosystems species richness of blooming plants and floral visitors was higher in the rainy season than in the dry season. Interaction networks in the Paramo were smaller and more nested, with lower levels of specialization and modularity than those in the Montane Forest, but there were no seasonal differences within either ecosystem. Beta diversity analyses indicate that differences between ecosystems are likely explained by species turnover, whereas within the Montane Forest differences between seasons are more likely explained by the rewiring of interactions. Results indicate that the decrease in species diversity with elevation affects network structure, increasing nestedness and reducing specialization and modularity.

Efecto de la actividad truchícola sobre la concentración de coliformes totales y Escherichia coli en aguas de la cuenca alta del río Savegre, Costa Rica

Introducción: La acuacultura de truchas ha incrementado gradualmente en las tierras altas de Costa Rica. Las aguas residuales de esta actividad son descartadas directamente en los ríos, sin tratamientos previos. Como consecuencia, la actividad truchícola puede contaminar severamente el agua de los ríos con bacterias que pueden afectar la salud humana. Objetivo: Evaluar la contaminación bacteriana por la acuacultura de truchas en el río Savegre, Costa Rica. Métodos: Contamos los coliformes totales y Escherichia coli de muestras mensuales (2015-2018) en tres secciones del proyecto de acuacultura más grande de la cuenca alta del río. Recolectamos las muestras en la entrada de los estanques para las truchas, a la salida, y 200 m hacia abajo. Resultados: Encontramos menos coliformes totales y E. coli en el agua recolectada justo en la salida del agua de los estanques. El número de coliformes totales fue mayor en el 2016 y 2017, y de E. coli en el 2016. Conclusiones: Conteos de coliformes y de E. coli es muy alto en el río, pero inesperadamente, su número disminuye en el agua residual descartada de los estanques. Podría ser que el mucus producido por las truchas o sustancias liberadas del musgo que cubre la pared de los estanques reduzca el crecimiento de bacterias, como se ha sido sugerido en otros estudios. La contaminación del río parece venir de otras fuentes.

Introduction: The trout aquacultural activity has gradually increased in Costa Rican highlands. Residual waters from this activity are discarded directly in the rivers without any previous treatment process. Consequently, this activity could severely contaminate the river with bacteria that can affect human health. Objective: To evaluate bacterial contamination from trout aquaculture on Río Savegre, Costa Rica. Methods: We counted total coliforms and Escherichia coli from monthly samples (2015-2018) at three sections of the largest aquacultural development in the upper drainage of the river. We collected samples at the fish ponds entrance, exit and 200 m downwards. Results: We found fewer total coliforms and E. coli in the water collected just at the exit of the fish ponds. We counted more total coliforms in 2016 and 2017, and more E. coli in 2016. Conclusions: Coliform and E. coli counts are high in the river, but, unexpectedly, low in the water discarded from the fish tanks. Perhaps the mucus produced by the trouts or substances released by mosses on the fish tank walls reduce bacterial growth, as suggested by other studies. River pollution appears to come from other sources.

A potential consequence for urban birds' fitness: Exposed anthropogenic nest materials reduce nest survival in the clay-colored thrush.

Urban bird dwellers survive and reproduce in highly urbanized ecosystems. Some individuals adjust to these novel conditions by changing natural nesting materials for artificial materials, thus making nests more conspicuous in the environment. The consequences of using artificial materials for nesting remain poorly understood, especially from a nest-predator perspective. We studied if exposed artificial materials on bird nests affect the daily survival rate in a common dweller species, the clay-colored thrush (Turdus grayi). We used previously collected nests with a different area of exposed artificial materials and placed them with clay eggs on the main campus of the Universidad de Costa Rica. During 12 days of the reproductive season, we monitored the nests using trap cameras located in front of each nest. We found that nest survival decreased as the proportion of exposed artificial materials increased in the nest and, unexpectedly, that the principal predators were conspecifics. Thus, artificial materials used in the outer layer of nests make them more susceptible to predation. The use of artificial materials likely reduce reproductive success and population size in urban clay-colored thrush, but further field experiments measuring the effect of waste in birds' nests on the reproductive success of urban birds are needed.

Blooming plant species diversity patterns in two adjacent Costa Rican highland ecosystems.

The Costa Rican Paramo is a unique ecosystem with high levels of endemism that is geographically isolated from the Andean Paramos. Paramo ecosystems occur above Montane Forests, below the permanent snow level, and their vegetation differs notably from that of adjacent Montane Forests. We compared the composition and beta diversity of blooming plant species using phenological data from functional plant groups (i.e., insect-visited, bird-visited and insect + bird-visited plants) between a Paramo and a Montane Forest site in Costa Rica and analyzed seasonal changes in blooming plant diversity between the rainy and dry seasons. Species richness was higher in the Montane Forest for all plant categories, except for insect-visited plants, which was higher in the Paramo. Beta diversity and blooming plant composition differed between both ecosystems and seasons. Differences in species richness and beta diversity between Paramo and the adjacent Montane Forest are likely the result of dispersal events that occurred during the last glacial period and subsequent isolation, as climate turned to tropical conditions after the Pleistocene, and to stressful abiotic conditions in the Paramo ecosystem that limit species establishment. Differences in blooming plant composition between both ecosystems and seasons are likely attributed to differential effects of climatic cues triggering the flowering events in each ecosystem, but phylogenetic conservatism cannot be discarded. Analyses of species composition and richness based on flowering phenology data are useful to evaluate potential floral resources for floral visitors (insects and birds) and how these resources change spatially and temporarily in endangered ecosystems such as the Paramo.

Genetic structure in neotropical birds with different tolerance to urbanization.

Gene flow in birds can be affected by urbanization depending on natural history traits and adaptability to habitat change. Contrasting results can be expected when comparing species with opposite resilience to urbanization. In this study, we assessed genetic diversity and structure for two bird species, the urban avoider white-eared ground-sparrow, Melozone leucotis, and the urban dweller house wren Troglodytes aedon. We used seven microsatellite loci and sampled five locations with differing levels of urbanization in Costa Rica. We found considerably higher genetic structure in white-eared ground-sparrows than in house wrens. Circuit theory analyses proved a higher isolation from urban resistance for the white-eared ground-sparrow than for house wrens. These results support that urbanization is a significant barrier for gene flow in urban avoiders, in contrast to urban dweller species that showed little to no impact. Differences could be attributed to a higher plasticity in habitat and nesting site preferences in the house wren, and significant dispersal limitation for the white-eared ground-sparrow. These results emphasize the need for conservation strategies towards white-eared ground-sparrows and other urban avoider species whose habitat and connectivity have been reduced by the recent urban expansion.

Human waste used as nesting material affects nest cooling in the clay-colored thrush.

The internal temperature of nests largely depends on the materials used in their construction because the characteristics of each material affect the isolation of nest walls. In urban environments, the availability of natural materials for nest building decreases, while the availability of artificial materials increases. Therefore, many urban bird species use more artificial materials for nest building inside cities, which may affect the thermal properties of the nest. We conducted an experiment to measure the effect of artificial materials included as part of the nest structure, on nest thermoregulation. We used as a model, nests of the clay-colored thrush (Turdus grayi), an urban bird species that have been reported using artificial nest materials. In our experiment, we measured how variation in artificial materials mass affects the nest cooling rate in a climate-controlled room. We found that artificial materials increased the cooling rate of clay-colored thrush nests, compared with nests with only natural materials. This result is especially relevant because showed a negative direct effect of the use of artificial material for nest building in birds. Considering that the availability of artificial material is increasing in urban areas, while natural material is decreasing, it is expected that the negative effect of using artificital material for nest construction would increase in the clay-colored thrush and other city bird species.

Complex behavioral plasticity is not reduced in spiderlings with miniature brains.

The brains of smaller animals are smaller than those of their larger relatives, but it is not clear whether their adaptive behavioral flexibility is more limited. Previous interspecific comparisons found that aspects of web construction behavior of very small orb weaving spiders (0.005 mg) were no less precise than those of much larger related orb weavers (30 mg), but the behaviors tested were relatively simple. Here we perform a more sensitive intraspecific test involving the multiple behavioral adjustments of orb web designs made by Leucauge argyra to confinement in very small spaces. Web adjustments of spiderlings as small as ~0.1 mg were compared to previously published observations of ~80 mg conspecific adults. Spiderlings in constrained spaces made all of the complex adjustments made by adults in at least seven independent web design variables, and their adjustments were no less precise. Rough estimates based on previously published data on total brain volumes and the mean diameters of neuron cell bodies suggested that spiderlings and adult females of Leucauge may have similar numbers of neurons, due to spiderlings having smaller neurons and a greater percentage of body tissues dedicated to the brain. We speculate that this neural similarity may explain why L. argyra spiderlings showed no behavioral deficits compared with adults.

The effect of noise variation over time and between populations on the fine spectrotemporal characteristics of different vocalization types.

Noise affects the recognition of acoustic signals by masking information. To compensate for increased noise, individuals often increase the minimum frequency of their vocalization to reduce noise interference. Our goal was to analyze the effect of noise on the characteristics of different bird vocalizations, through a comparative study of vocalizations on the same bird species. We analyzed the effects of noise variation on the fine spectrotemporal characteristics of calls, songs, and duets of White-eared Ground-sparrows (Melozone leucotis) across three populations over a three-year period. We recorded vocalizations and noise levels simultaneously from 41 territories between 2012 and 2014. We measured the duration, minimum, maximum, and maximum amplitude frequency, and counted the number of songs elements for each vocalization recorded. As we predicted, noise influences the minimum frequency of song, but did not have an effect on the fine spectrotemporal characteristics for calls and duets. We did, however, find that low and high frequency of calls and duets increased with noise-population-year interaction. Our results suggest that differences in noise values at each population were inadequate to observe changes in vocalization characteristics. In conclusion, evaluating responses to different noise levels on different vocalization types for the same species expands our understanding of the flexibility of birds to adjust vocalizations in response to anthropogenic noise.

Do Different Methods Yield Equivalent Estimations of Brain Size in Birds?

The ratio of brain size to body size (relative brain size) is often used as a measure of relative investment in the brain in ecological and evolutionary studies on a wide range of animal groups. In birds, a variety of methods have been used to measure the brain size part of this ratio, including endocranial volume, fixed brain mass, and fresh brain mass. It is still unclear, however, whether these methods yield the same results. Using data obtained from fresh corpses and from published sources, this study shows that endocranial volume, mass of fixed brain tissue, and fresh mass provide equivalent estimations of brain size for 48 bird families, in 19 orders. We found, however, that the various methods yield significantly different brain size estimates for hummingbirds (Trochilidae). For hummingbirds, fixed brain mass tends to underestimate brain size due to reduced tissue density, whereas endocranial volume overestimates brain size because it includes a larger volume than that occupied by the brain.

Allometry of reaching and contact structures in Kukulcania hibernalis (Araneae: Filistatidae).

Large part of the morphological diversity observed across taxa is attributed to the effect of sexual selection; and the static allometry of these structures vary largely from highly positive to negative, depending on their function, and position on the animal's body. In arthropods, information of how sexually selected contact and reaching male structures use during courtship scale on body size is scarce. We tested two complementary hypotheses: the reaching structure hypothesis and the contact-function hypothesis, in the spider Kukulcania hibernalis. We used the length of the proximal segments of the male pedipalp to test the reaching structure hypothesis, and claw features to test the contact-function hypothesis. Our results support both hypotheses. Small males have disproportionally longer pedipalps (highly negative allometry) than large males, increasing the probability of small-bodied males to inseminate even large females. We also found that both distal contact and noncontact homologous structures scaled shallow (slope < 1) on body size, but allometry was significantly shallower for contact than for noncontact distal structures, providing support to the contact-function hypothesis, and allowing teasing apart the effect of sexual selection on distal contact structures with dual functions.

Asociación entre la morfología alar y el uso del hábitat en seis especies de palomas (Columbidae) neotropicales

Los factores biomecánicos, el comportamiento e historia natural de las especies pueden influenciar la forma y dimensión del ala en las aves. Por lo tanto, la morfología de las alas de cada especie podría estar adaptada al ambiente en el que habitan. En palomas (familia Columbidae) existe un gradiente de especies que tienden a volar mucho y otras que vuelan muy poco. El objetivo de esta investigación es determinar si la forma de las alas de seis especies de palomas se asocia con su capacidad de vuelo. Predecimos que las especies Patagioenas flavirostris y P. nigrirostris que tienden a volar largas distancias presentan alas relativamente largas y puntiagudas; mientras que Geotrygon montana que se desplaza principalmente caminando presenta alas relativamente anchas y redondeadas. Especies con capacidad de vuelo intermedio (Leptotila verreauxi, L. cassini y Zenaida asiatica) van a presentar alas con características intermedias. Estimamos la carga alar, proporción de la forma, conicidad alar, largo relativo y el área alar promedio para cada especie. Realizamos un análisis discriminante para determinar cuál combinación de variables explica mejor la diferencia morfológica de las alas entre las seis especies y probamos que tan robusto es este modelo mediante una prueba binomial. El modelo clasificó correctamente el 57 % de los individuos en su propia especie. Por lo tanto, encontramos que el tipo de desplazamiento predominante en cada una de las seis especies de Columbidae analizadas, se asocia con la morfología alar: alas adaptadas para vuelos en zonas abiertas en P. flavirostris y P. nigrirostris, alas adaptadas para vuelos muy maniobrables en hábitats densos en G. montana, y alas con características intermedias en L. verreauxi, L. cassini y Z. asiatica.

Biomechanics, behavior, and natural history influence wing dimension and shape. Wing design often correlates with features of the habitat in which each species is found. Doves and pigeons (Columbidae family) range from long-distance fliers (e.g., canopy and open area species) to very short-distance fliers (e.g., species adapted to dense understory forests) and such variation makes this group fit to test the association between flying habits and wing morphology. Our objective in this study is to determine whether the wing morphology (shape and dimensions) of six dove species is associated to their flying capability. We predict that the long-distance fliers Patagioenas flavirostris and P. nigrirostris will have long and sharp wings; while the very short-distance flier Geotrygon montana will have broad and rounded wings. Other species (e.g., Leptotila verreauxi, L. cassini and Zenaida asiatica) whose flying capability fits in between these two will have wings with intermediate morphological features. We measured the wing disc loading, shape ratio, the ratio of mean to maximum wing chord, relative wing length, and wing area for each species. We conducted a discriminant function analysis to compare which variables explain better the differences in wing morphology across the six species, and used a binomial test to evaluate the power of the model. The model correctly classified 57 % of individuals within their own species. The flying capability is associated with the wing morphology of the six Columbidae species; with a wing design for long-distance fliers in P. flavirostris and P. nigrirostris, wing design for maneuvering in dense habitats in G. montana, and wings with an intermediate design in L. verreauxi, L. cassini and Z. asiatica.

The forgotten habitats in conservation: early successional vegetation

Conservation efforts in terrestrial environments have focused on preserving patches of natural habitats and restoring disturbed habitats, with the main goal of transforming them into forests or habitats that resemble the original conditions. This approach tends to overlook the importance of conserving early successional vegetation (e.g., riverside vegetation, natural regeneration, young secondary forests), which often includes a large number of species (e.g., plants and animals) associated with or restricted to these habitats. In this paper we want to bring to attention the importance of preserving early successional vegetation, and to encourage scientists to investigate, e.g., the diversity, distribution, and species interactions occurring in these habitats. To address these goals, we focus on two main objectives: (1) to identify the common types of early successional vegetation in the Costa Rican Central Valley; and (2) to use some case studies to draw attention to the importance that such areas have as reservoirs of a large portion of the diversity unique to early successional stages. We first include an example to show the diversity of plants in small forest patches immersed in a large urbanized area. We provide general information on the insects that occur in early successional vegetation in urban areas, and in further detail examples of butterflies. Additionally, we provide examples of birds and mammals that are restricted to early successional vegetation, and how the reduction of this vegetation type affects species conservation. Finally, we encourage scientists to investigate these early successional habitats, particularly those species exclusive to early successional stages. Special attention should be paid to endemic species and those with a restricted distribution. Information of this type will make conservation of the diversity contained in these habitats possible.

Los esfuerzos de conservación en ambientes terrestres se han centrado principalmente en la preservación de ambientes naturales y la restauración de diferentes hábitats, con la meta principal de transformar estos ambientes en bosques maduros o hábitats que asemejen las condiciones originales. Este enfoque tiende a pasar por alto la importancia de conservar la vegetación de estados de regeneración temprana (e.g., vegetación riparia, regeneración natural, bosque secundario joven), la cual incluye un gran número de especies (e.g., plantas y animales) asociadas o restringidas a estos hábitats. Con este artículo queremos llamar la atención sobre la importancia de preservar áreas cubiertas con vegetación de sucesión temprana, e instar a científicos y naturalistas a investigar, e.g., la diversidad, distribución, e interacciones entre las especies presentes en estos ambientes. Para apoyar esta meta, nos enfocamos en dos objetivos principales: (1) identificar los tipos más comunes de vegetación pionera en el Valle Central de Costa Rica; y (2) utilizar algunos casos de estudio para llamar la atención sobre la importancia que tales áreas tienen como reservorio de gran parte de la diversidad, mucha de la cual es única de los estados de sucesión temprana. Primero se incluye un ejemplo particular en el cual se muestra la diversidad de plantas en pequeños fragmentos de bosque y matorral inmersos en una gran área urbanizada. Después se presenta una revisión general de los insectos que habitan en la vegetación de sucesión temprana en áreas urbanas, para luego discutir en mayor detalle ejemplos de mariposas. Además, proporcionamos ejemplos de especies de aves y mamíferos que están restringidos a vegetación de sucesión temprana, y cómo la reducción de este ambiente afecta su conservación. Finalmente, instamos a los científicos de diferentes áreas a investigar los diversos procesos ecológicos e interacciones biológicas inherentes a los estados de regeneración temprana. Especial atención requieren aquellas especies exclusivas o endémicas de estos ambientes. Sin esta información es imposible conservar la diversidad de estos hábitats.

How many larvae and nymphs of Amblyomma spp. ticks (Ixodida: Ixodidae) are needed to detect the presence of Rickettsia (Rickettsiales: Rickettsiaceae)?

16. Tick-borne rickettsiosis is becoming a common emergent disease in many tropical countries, and the rapid detection of this disease could reduce the impact on wild life and public health. Pools of larvae and nymphs, and individual adult ticks are used for detection of Rickettsia species (Rickettsiaceae), but the number of larvae and nymphs in the pools likely affect the probability of detection of Rickettsia spp. We tested whether the number of larvae or nymphs of Amblyomma (Ixodidae) included in the pools affected the probability of detection. The number of larvae in each pool did not affect the probability of detecting Rickettsia spp., but the probability of detection increased rapidly with the number of nymphs. In this study, 20 nymphs are enough to approach a probability of 1 of detecting Rickettsia.

20. La rickettsiosis transmitida por garrapatas es una enfermedad emergente cada vez más común en muchos países tropicales, por lo que su rápida detección podría reducir su impacto negativo en la salud humana. Grupos (pozos) de larvas y ninfas de garrapatas, así como individuos adultos son usados para detectar la presencia de diferentes especies de Rickettsia. Sin embargo, el número de larvas y ninfas incluidos en estos grupos probablemente afecta la detección de Rickettsia spp. Nosotros probamos si el número de larvas o ninfas incluidas en estos grupos afecta la probabilidad de detección. Encontramos que el número de larvas incluidas en cada grupo no afectó la probabilidad de detección de Rickettsia spp., pero la probabilidad de detección aumenta rápidamente con el número de ninfas incluidas en los grupos. En este estudio, 20 ninfas por grupo fueron suficientes para tener una probabilidad cercana a 1 de detectar Rickettsia.

Effect of El Niño and La Niña on abundance of frugivorous and nectarivorous terrestrial birds in three tropical forests

El Niño and La Niña climatic oscillations have dramatic effects on population dynamics and community structure of different animals. For marine birds, El Niño phenomenon drastically increases their mortality and reduces their reproductive success. In terrestrial ecosystems, the lack of long-term longitudinal data limits our understanding of the impact of El Niño and La Niña on bird populations and communities. We analyzed changes in abundance of frugivorous (large, medium, and small) and nectarivorous birds on three tropical forest types (lowland, premontane and montane) during El Niño, La Niña and neutral years using data from 16 Christmas' Bird Counts in Costa Rica. Abundance of large and medium size frugivores increased during La Niña events, and the abundance of nectarivores during El Niño in the lowland forest, but neither of these events affected the abundance of small frugivores. In the montane forest, abundance of all four bird groups increased during El Niño, but decreased during La Niña events. Abundance of large, and small frugivores increased in the premontane forest during La Niña events, but other bird groups were not affected by La Niña. The abundance of small frugivores also increased during El Niño. We hypothesize that changes in abundance of frugivores and nectarivores during La Niña and El Niño events is probably correlated with fluctuations in food resources as it has been reported for other terrestrial tropical ecosystems.

Las oscilaciones climáticas causadas por el fenómeno de El Niño y La Niña tienen efectos drásticos sobre la dinámica de las poblaciones y la estructura de las comunidades. Para aves marinas, el fenómeno de El Niño incrementa drásticamente su mortalidad y reduce su éxito reproductivo. En ecosistemas terrestres, la falta de datos a largo plazo limita nuestro entendimiento sobre el impacto de El Niño y La Niña sobre la dinámica de las poblaciones y comunidades de aves. Nosotros analizamos los cambios en la abundancia de aves frugívoras (grandes, medianos y pequeños) y nectarívoras en tres tipos de bosque tropical (lluvioso de tierras bajas, premontano y montano) durante los años de El Niño, La Niña, y años neutrales usando datos de al menos 16 conteos de navidad (Christmas' Bird Counts) en Costa Rica. La abundancia de frugívoros grandes y medianos incrementó durante los años de La Niña, y la abundancia de nectarívoros durante el Niño en el bosque lluvioso de tierras bajas, pero ninguno de estos eventos afectó la abundancia de frugívoros pequeños en este bosque. En el bosque montano, la abundancia de los cuatro grupos de aves incrementó durante los años de El Niño, pero disminuyó durante los años de La Niña. La abundancia de frugívoros grandes y pequeños aumentó en el bosque pre-montano durante los años de La Niña; sin embargo, la abundancia de los frugívoros medianos y los nectarívoros no fue afectada por los eventos por La Niña o El Niño. La abundancia de frugívoros pequeños también aumentó durante El Niño en este bosque. Proponemos la hipótesis de que los cambios en la abundancia de las aves frugívoras y nectarívoras reflejan fluctuaciones en el recurso alimentario, como ha sido reportado en otros ecosistemas terrestres tropicales.

Effect of barriers and distance on song, genetic, and morphological divergence in the highland endemic Timberline Wren (Thryorchilus browni, Troglodytidae).

Populations may become isolated by distance, geographic barriers or both. Isolated populations often diverge in behavioral, morphological and genetic traits as a result of reduced inter-population gene flow. Highland species commonly present naturally fragmented distributions that confine populations to the highest mountain peaks, isolated by mountain passes and distance. The endemic Timberline Wren (Thryorchilus browni) inhabits the highlands of the Talamanca mountain range, including western Panama, and the highest peak in the Central Volcanic mountain range of Costa Rica. Using microsatellites and song recordings we studied the effect of a geographic barrier and distance on song, genetic and morphological divergence among four populations in Costa Rica. A lowland mountain pass resulted in the largest genetic, vocal, and morphological (bill length) differences among populations, likely due to reduce the gene flow. Cultural drift and assortative mating by females selecting songs from their own population likely accentuates the effect of isolation and limited gene flow between populations. This pattern of population divergence has been found in other Neotropical highland birds, but over larger geographical scales. We conclude that mountain passes and distance both reduce gene flow between populations in recently-isolated highland species with restricted distributions.

Morphological adaptations for relatively larger brains in hummingbird skulls.

A common allometric pattern called Haller's Rule states that small species have relatively larger brains and eyes than larger species of the same taxonomic group. This pattern imposes drastic structural changes and energetic costs on small species to produce and maintain a disproportionate amount of nervous tissue. Indeed, several studies have shown the significant metabolic costs of having relatively larger brains; however, little is known about the structural constraints and adaptations required for housing these relatively larger brains and eyes. Because hummingbirds include the smallest birds, they are ideal for exploring how small species evolve morphological adaptations for housing relatively larger brain and eyes. We here present results from a comparative study of hummingbirds and show that the smallest species have the lowest levels of ossification, the most compact braincases, and relatively larger eye sockets, but lower eye/head proportion, than larger species. In contrast to Passerines, skull ossification in hummingbirds correlates with body and brain size but not with age. Correlation of these skull traits with body size might represent adaptations to facilitate housing relatively larger brain and eyes, rather than just heterochronic effects related to change in body size. These structural changes in skull traits allow small animals to accommodate disproportionately larger brains and eyes without further increasing overall head size.

Mating and aggressive behaviour of Brachyrhaphis olomina (Cyprinodontiformes: Poeciliidae).

Despite the increasing interest in the use of intromittent male genitalia and coercive mating behaviour in poeciliids, detailed studies of the mating behaviour of most species in this family are lacking. We describe here the mating and aggressive behaviours of Brachyrhaphis olomina, and correlate them with the condition of the female's ovum and embryos (immature, mature and pregnant). B. olomina performed a wide range of aggressive (sidle spread, tail beating, coordinate) and mating behaviours (approximation, touch, lateral display, touch-lateral display). Some behaviours (e.g. tail beating) are shared with other poeciliids, but two sexual behaviours (touch and lateral display) and one aggressive (coordinate) behaviour may be unique to B. olomina and were not reported in a previous study. Differences in male behaviour when paired with a female with mature ovum (more mating displays, no agonistic movements) suggest that males detect the female's reproductive condition from some distance. The distinctive nature of mating behaviour in B. olomina highlights the importance of studying different species to have a better understanding of the evolution of mating and aggressive behaviours in poeciliids. Digital video images related to the article are available at http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo01a, http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo02a and http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo03a.

Effect of fragmentation on the Costa Rican dry forest avifauna.

Deforestation and changes in land use have reduced the tropical dry forest to isolated forest patches in northwestern Costa Rica. We examined the effect of patch area and length of the dry season on nestedness of the entire avian community, forest fragment assemblages, and species occupancy across fragments for the entire native avifauna, and for a subset of forest dependent species. Species richness was independent of both fragment area and distance between fragments. Similarity in bird community composition between patches was related to habitat structure; fragments with similar forest structure have more similar avian assemblages. Size of forest patches influenced nestedness of the bird community and species occupancy, but not nestedness of assemblages across patches in northwestern Costa Rican avifauna. Forest dependent species (species that require large tracts of mature forest) and assemblages of these species were nested within patches ordered by a gradient of seasonality, and only occupancy of species was nested by area of patches. Thus, forest patches with a shorter dry season include more forest dependent species.

Song plasticity over time and vocal learning in clay-colored thrushes.

Songbirds have been traditionally classified into close-ended or open-ended learning species according to the length of the sensitive period during which birds are able to memorize new vocalizations. Closed-ended learners are generally not capable of changing their song after the first year of life, while open-ended learners show song plasticity as adults. A few Turdus species have been suggested to be open-ended learners, but no long-term study has been conducted to investigate their song plasticity over time. We analyzed the songs of clay-colored thrushes, T. grayi, over four successive breeding seasons to assess song plasticity in their syllable repertoires within and between breeding seasons. A total of 16,262 syllables were classified through visual inspection of spectrograms and multidimensional scaling analysis based on spectrogram correlations. On average, 563 ± 153 (SD) syllables per male per breeding season were analyzed. Male repertoire size was 9-20 syllable types. Males were capable of modifying their syllable repertoire between the initial and final periods of the breeding season. Song plasticity within breeding seasons may be associated with imitation between neighboring males, suggesting song learning in males that were ≥2 years old. This short-term plasticity is not enough, however, to explain the high proportion of change (mean = 65 % syllable types) in repertoire composition between breeding seasons in adult males. Song plasticity resulting from annual changes in repertoire composition could be explained by open-ended learning, but another mechanism, extended memory and re-expression, could also explain long-term plasticity. Experimental studies controlling the acoustic environment are needed to determine which mechanism is responsible for such a high level of song plasticity.

Sexual behavior, cannibalism, and mating plugs as sticky traps in the orb weaver spider Leucauge argyra (Tetragnathidae).

Unpublished field observations in Leucauge argyra, a tropical orb weaver spider, suggest the occurrence of conspicuous mating plugs that could reduce or prevent remating attempts. Otherwise, the sexual behavior of this species remains unknown. The aims of this study were to describe the courtship behavior and copulation in L. argyra and investigate mating plug formation in this species. Fourteen virgin females and 12 plugged females were exposed to up to three males and checked for mating plug formation. Of the 12 virgins that copulated, nine produced plugs (five immediately after copulation), and the five plugged females that copulated produced another mating plug immediately after copulation. We did not detect the transfer of any male substance during copulation but observed a whitish liquid emerging from female genital ducts. Plug formation was positively associated with male twanging during courtship. One virgin and four plugged females cannibalized males. In seven trials with virgins and in three trials with plugged females, the male's palp adhered to a substance that emerged from female genital ducts and spread on her genital plate. The male had to struggle energetically to free his glued palp; two of these males were cannibalized while trying to release their palps. Females seem to determine copulation duration by altering the timing of mating plug formation and through sexual cannibalism. This is the first case reported of a mating plug as a sticky trap for males.