Actividad reproductiva y dieta de Hemidactylus frenatus (Sauria: Gekkonidae) en el norte de Colombia

A pesar de su éxito como especie invasora, existe poca información sobre los aspectos ecológicos del gecónido Hemidactylus frenatus en Colombia. En el presente estudio se determinó el tamaño de madurez, dimorfismo sexual, actividad reproductiva y composición de la dieta de una población de esta especie en una localidad urbana del Norte de Colombia. Se realizaron once muestreos desde septiembre de 2011 hasta agosto de 2012 en edificaciones del municipio de Sincelejo. Se capturaron 264 ejemplares, 133 machos adultos, 112 hembras adultas y 19 juveniles. Los machos alcanzaron la madurez sexual a un tamaño (longitud hocico-cloaca) menor (35.7 mm) que las hembras (42.7 mm), además alcanzan mayor tamaño y tienen cabezas y bocas proporcionalmente más grandes que éstas. Los machos fueron reproductivos durante todo el año; aunque el volumen testicular varió significativamente entre los muestreos, esta variación no estuvo asociada con el tamaño corporal ni con la precipitación en el área de estudio. Durante todos los muestreos se encontraron hembras adultas reproductivas. Las hembras tienen un tamaño de nidada invariable de dos huevos y no existen diferencias entre oviductos respecto al diámetro y peso de los huevos. La dieta de H. frenatus es variada, siendo los dípteros, hemípteros y formícidos las presas de mayor importancia relativa. Los adultos de ambos sexos consumen un volumen y número similar de presas. Así, la población de H. frenatus estudiada tiene una actividad reproductiva continua y un comportamiento dietario generalista y oportunista. Las condiciones climáticas, la disponibilidad ambiental de presas en el área de estudio y las características intrínsecas de esta especie parecen ser responsables de su abundancia y éxito colonizador en esta y otras localidades.

Despite its success as an invasive species, little is known about the ecological aspects of the gekkonid lizard Hemidactylus frenatus in Colombia. In the present study the size at maturity, sexual dimorphism, reproductive activity, and diet composition of a population of this species in an urban locality of Northern Colombia were determined. We conducted eleven samplings from September 2011 to August 2012 in buildings of the municipality of Sincelejo. A total of 264 specimens H. frenatus were captured, 112 were adult females, 133 adult males and 19 juveniles. Males reach sexual maturity at a smaller size (snout-vent length) than females (males: 35.7 mm; females: 42.7 mm), also they are larger and have proportionally larger heads and mouths than females. Males were reproductive throughout the year; although testicular volume varied significantly between samples, this variation was not associated with body size and precipitation in the study area. Reproductive adult females were found during all the sampling period. Females have an invariable clutch size of two eggs and we found no differences in the diameter and weight of eggs in each oviduct. The diet of H. frenatus is varied, with Diptera, Hemiptera and Formicidae being the prey types with the greatest relative importance values. Individuals of both sexes consume a similar volume and number of prey. Thus, the studied population of H. frenatus has continuous reproductive activity and a generalist-opportunistic feeding behavior. The climatic conditions of the study area, environmental availability of prey and intrinsic features of this species appear to be responsible for their abundance and colonizing success in this and other localities.

Actividad reproductiva y dieta de Hemidactylus frenatus (Sauria: Gekkonidae) en el norte de Colombia

A pesar de su éxito como especie invasora, existe poca información sobre los aspectos ecológicos del gecónido Hemidactylus frenatus en Colombia. En el presente estudio se determinó el tamaño de madurez, dimorfismo sexual, actividad reproductiva y composición de la dieta de una población de esta especie en una localidad urbana del Norte de Colombia. Se realizaron once muestreos desde septiembre de 2011 hasta agosto de 2012 en edificaciones del municipio de Sincelejo. Se capturaron 264 ejemplares, 133 machos adultos, 112 hembras adultas y 19 juveniles. Los machos alcanzaron la madurez sexual a un tamaño (longitud hocico-cloaca) menor (35.7 mm) que las hembras (42.7 mm), además alcanzan mayor tamaño y tienen cabezas y bocas proporcionalmente más grandes que éstas. Los machos fueron reproductivos durante todo el año; aunque el volumen testicular varió significativamente entre los muestreos, esta variación no estuvo asociada con el tamaño corporal ni con la precipitación en el área de estudio. Durante todos los muestreos se encontraron hembras adultas reproductivas. Las hembras tienen un tamaño de nidada invariable de dos huevos y no existen diferencias entre oviductos respecto al diámetro y peso de los huevos. La dieta de H. frenatus es variada, siendo los dípteros, hemípteros y formícidos las presas de mayor importancia relativa. Los adultos de ambos sexos consumen un volumen y número similar de presas. Así, la población de H. frenatus estudiada tiene una actividad reproductiva continua y un comportamiento dietario generalista y oportunista. Las condiciones climáticas, la disponibilidad ambiental de presas en el área de estudio y las características intrínsecas de esta especie parecen ser responsables de su abundancia y éxito colonizador en esta y otras localidades.(AU)

Despite its success as an invasive species, little is known about the ecological aspects of the gekkonid lizard Hemidactylus frenatus in Colombia. In the present study the size at maturity, sexual dimorphism, reproductive activity, and diet composition of a population of this species in an urban locality of Northern Colombia were determined. We conducted eleven samplings from September 2011 to August 2012 in buildings of the municipality of Sincelejo. A total of 264 specimens H. frenatus were captured, 112 were adult females, 133 adult males and 19 juveniles. Males reach sexual maturity at a smaller size (snout-vent length) than females (males: 35.7 mm; females: 42.7 mm), also they are larger and have proportionally larger heads and mouths than females. Males were reproductive throughout the year; although testicular volume varied significantly between samples, this variation was not associated with body size and precipitation in the study area. Reproductive adult females were found during all the sampling period. Females have an invariable clutch size of two eggs and we found no differences in the diameter and weight of eggs in each oviduct. The diet of H. frenatus is varied, with Diptera, Hemiptera and Formicidae being the prey types with the greatest relative importance values. Individuals of both sexes consume a similar volume and number of prey. Thus, the studied population of H. frenatus has continuous reproductive activity and a generalist-opportunistic feeding behavior. The climatic conditions of the study area, environmental availability of prey and intrinsic features of this species appear to be responsible for their abundance and colonizing success in this and other localities.(AU)

[Reproductive ecology of Geobatrachus walkeri (Anura: Strabomantidae)]. / Ecología reproductiva de Geobatrachus walkeri (Anura: Srabomantidae).

Geobatrachus walkeri is a frog that belongs to a monotypic genus and is endemic to the Sierra Nevada de Santa Marta, Colombia. The species inhabits different microhabitats between 2 000 to 3 500m altitude, including the leaf litter of a pine plantation. To understand its reproductive ecology, we conducted eight frog samplings, covering the rainy and dry seasons, and two habitat types (secondary native forest and pine plantation) during 2010-2011. For this study, we also included data obtained from five previous similar samplings undertaken during 2008-2009. The pine leaf litter was the main microhabitat where frogs were found; we heard choruses of six-ten calling males during all sampled months, and observed the frogs having diurnal and nocturnal activity. Regardless of the year of study, the population consisted of neonates recruited several times of the year, a large number of juveniles with a wide range of body sizes, and fewer adults with a narrower range of body size. The histological analyses of the gonads showed that the size at maturity was near 18mm SVL for males and females, and those adult males and females were reproductive active during all sampling months, suggesting a continuous reproductive activity pattern. However, during the dry season, the seminiferous tubules showed a drastically diminished spermatic epithelium although containing abundant luminal spermatozoa, which suggest a reduction in the sperm production at the end of this season. Similarly, frogs of all age categories were significantly more abundant during the early dry season, whereas were significantly less abundant with the advancement of the dryness suggesting that the intensity of the dry season could temporally stop the activity and reproduction of this population.

Ecología reproductiva de Geobatrachus walkeri (Anura: Srabomantidae) / Reproductive ecology of Geobatrachus walkeri (Anura: Strabomantidae)

Geobatrachus walkeri is a frog that belongs to a monotypic genus and is endemic to the Sierra Nevada de Santa Marta, Colombia. The species inhabits different microhabitats between 2 000 to 3 500m altitude, including the leaf litter of a pine plantation. To understand its reproductive ecology, we conducted eight frog samplings, covering the rainy and dry seasons, and two habitat types (secondary native forest and pine plantation) during 2010-2011. For this study, we also included data obtained from five previous similar samplings undertaken during 2008-2009. The pine leaf litter was the main microhabitat where frogs were found; we heard choruses of six-ten calling males during all sampled months, and observed the frogs having diurnal and nocturnal activity. Regardless of the year of study, the population consisted of neonates recruited several times of the year, a large number of juveniles with a wide range of body sizes, and fewer adults with a narrower range of body size. The histological analyses of the gonads showed that the size at maturity was near 18mm SVL for males and females, and those adult males and females were reproductive active during all sampling months, suggesting a continuous reproductive activity pattern. However, during the dry season, the seminiferous tubules showed a drastically diminished spermatic epithelium although containing abundant luminal spermatozoa, which suggest a reduction in the sperm production at the end of this season. Similarly, frogs of all age categories were significantly more abundant during the early dry season, whereas were significantly less abundant with the advancement of the dryness suggesting that the intensity of the dry season could temporally stop the activity and reproduction of this population.

Geobatrachus walkeri es una rana que pertenece a un género monotípico y es endémica de la Sierra Nevada de Santa Marta, Colombia. La especie habita en diferentes microhábitats entre los 2 000 y 3 500m de altitud, incluyendo la hojarasca de una plantación de pinos. Para entender su ecología reproductiva se realizaron ocho muestreos de individuos, que abarcaron las estaciones de lluvias y de sequía y dos tipos de hábitats (bosque secundario y plantación de pino) durante 2010-2011. También se incluyeron los datos de cinco muestreos similares obtenidos previamente durante el período 2008-2009. El principal microhábitat donde las ranas se encontraron fue la hoja-rasca de pino, allí escuchamos coros de seis-diez machos durante todos los meses muestreados, y observamos que las ranas tienen actividad diurna y nocturna. Sin importar el año de estudio, la población se compone de neonatos reclutados en varios meses del año, un gran número de juveniles con un amplio rango de tamaños corporales, y un menor número de adultos con un rango más reducido de tamaño corporal. Los análisis histológicos de las gónadas mostraron que el tamaño a la madurez se adquiere cerca de los 18mm LRC para machos y hembras, y que los machos y hembras adultos fueron reproductivos durante todo el muestreo, lo que sugiere un patrón continuo de actividad reproductiva. Sin embargo, durante la estación seca, los túbulos seminíferos presentan un epitelio espermático drásticamente disminuido, aunque contienen abundantes espermatozoides luminales, lo que sugiere una reducción en la producción de espermatozoides en el final de la estación seca. Del mismo modo, las ranas de todas las categorías de edad fueron significativamente más abundantes durante el inicio de la estación seca, mientras que fueron significativamente menos abundantes con el avance de la sequedad, lo que sugiere que una mayor intensidad de la temporada de sequía podría detener temporalmente la actividad y la reproducción en esta población.

[Relative abundance and microhabitat use by the frog Geobatrachus walkeri (Anura: Strabomantidae) in two habitats of Sierra Nevada de Santa Marta, Colombia]. / Abundancia relativa y uso de microhábitat de la rana Geobatrachus walkeri (Anura: Strabomantidae) en dos hábitats en Sierra Nevada de Santa Marta, Colombia.

Geobatrachus walkeri belongs to a monotypic frog genus endemic to the San Lorenzo area, Sierra Nevada de Santa Marta, Colombia. This species has been categorized as endangered because of its small distribution area and the decline in the extent and quality of its habitat. It inhabits two forest types with different composition and structure, the native secondary forest and a pine plantation (dominated by Pinus patula). To compare the relative abundance and microhabitat use of this species in these habitat types, 30 quadrants/environment were distributed randomly. The individual number, microhabitat use and other aspects of its natural history were registered using visual encounter surveys in both sites, including non-sampled areas in the quadrants. The relative abundance of frogs was significantly different between habitats and among seasons. The highest abundance of G. walkeri relative to the total area was found in the pine plantation, being 2.3 times higher than in the natural forest. More frogs were significantly found during the rainy season; nevertheless, active individuals were also found during the dry season. Significant differences were found in the microhabitat use with respect to the forest type and season. The most frequently microhabitat used in the two forest types was the pine leaf-litter; besides, in the native forest, the microhabitat occupied more frequently presented medium and large size stones. Geobatrachus walkeri is a successful species in pine plantations, associated permanently to its leaf-litter environment where it seems to develop its entire life cycle. The clear modifications in the soils and water, derived from the introduction of the pine plantation in this area, seem not to have negatively affected the conservation and successful maintenance of this species.

Abundancia relativa y uso de microhábitat de la rana Geobatrachus walkeri (Anura: Strabomantidae) en dos hábitats en Sierra Nevada de Santa Marta, Colombia

Relative abundance and microhabitat use by the frog Geobatrachus walkeri (Anura:Strabomantidae) in two habitats of Sierra Nevada de Santa Marta, Colombia. Geobatrachus walkeri belongs to a monotypic frog genus endemic to the San Lorenzo area, Sierra Nevada de Santa Marta, Colombia. This species has been categorized as endangered because of its small distribution area and the decline in the extent and quality of its habitat. It inhabits two forest types with different composition and structure, the native secondary forest and a pine plantation (dominated by Pinus patula). To compare the relative abundance and microhabitat use of this species in these habitat types, 30 quadrants/environment were distributed randomly. The individual number, microhabitat use and other aspects of its natural history were registered using visual encounter surveys in both sites, including non-sampled areas in the quadrants. The relative abundance of frogs was significantly different between habitats and among seasons. The highest abundance of G. walkeri relative to the total area was found in the pine plantation, being 2.3 times higher than in the natural forest. More frogs were significantly found during the rainy season; nevertheless, active individuals were also found during the dry season. Significant differences were found in the microhabitat use with respect to the forest type and season. The most frequently microhabitat used in the two forest types was the pine leaf-litter; besides, in the native forest, the microhabitat occupied more frequently presented medium and large size stones. Geobatrachus walkeri is a successful species in pine plantations, associated permanently to its leaf-litter environment where it seems to develop its entire life cycle. The clear modifications in the soils and water, derived from the introduction of the pine plantation in this area, seem not to have negatively affected the conservation and successful maintenance of this species. Rev. Biol. Trop. 59 (2): 907-920. Epub 2011 June 01.

La rana Geobatrachus walkeri pertenece a un género monotípico y endémico del área de San Lorenzo, Sierra Nevada de Santa Marta, Colombia. La especie habita en el bosque secundario natural y en una plantación de pino (dominada por Pinus patula). Para comparar la abundancia relativa y el uso del microhábitat en estos dos hábitat y durante los períodos de máxima y mínima precipitación, se distribuyeron aleatoriamente 30 cuadrantes en cada bosque sobre los que se registraron el número de individuos y los microhábitats ocupados. Además se reconocieron otros aspectos de la historia natural mediante muestreo por encuentros casuales (VES), se incluye en los dos hábitats áreas no muestreadas por los cuadrantes. La mayor abundancia de ranas se obtuvo en la plantación de pino y durante la época de lluvias. El microhábitat más usado fue la hojarasca de pino. Geobatrachus walkeri es una especie exitosa en las plantaciones de pino, asociada permanentemente a su hojarasca en donde parece desarrollar todo su ciclo de vida. Las claras modificaciones derivadas de la introducción de las plantaciones de pino en el suelo y el agua, parecen no haber afectado la supervivencia y mantenimiento exitoso de esta especie categorizada como en peligro de extinción.

The morphology and post-hatching development of the skull of Bolitoglossa nicefori (Caudata: Plethodontidae): developmental implications of recapitulation and repatterning.

The cranial morphology of the direct-developing salamander Bolitoglossa nicefori and its post-hatching development are described and compared with that of other urodeles. Four stages of cranial development are defined on the basis of conspicuous events that occur during post-hatching ontogeny. The adult skull morphology of B. nicefori is similar to that of other plethodontids; however, some regions show interspecific variation. The post-hatching ontogeny of the skull and the stage of ossification observed in the hatchlings of B. nicefori show two important ontogenetic features: (1) a mosaic of early larval, metamorphic and post-metamorphic skull features in hatchlings, and (2) absence of characteristic larval elements in skull and hyoid apparatus. The distinctive stage of ossification in the hatchlings of B. nicefori could be caused by heterochronic changes in the ossification sequence, compared to the ontogeny of metamorphic salamanders. The possible heterochronic changes and the absence of larval traits are perhaps due to ontogenetic repatterning, yet without an obvious impact on the adult skull morphology (absence of morphological novelties). This might indicate a compartmentalized development. Further studies should be performed in order to establish the possible occurrence of recapitulatory patterns or ontogenetic repatterning in the skull morphogenesis of B. nicefori during its embryonic development.

How does diet influence the reproductive seasonality of tropical freshwater fish?: A case study of a characin in a tropical mountain river

Seasonal breeding of tropical freshwater fish may be synchronized with periods of high food consumption. We explored this hypothesis by studying the relationship between diet and reproductive activity of Creagrutus guanes (Teleostei, Characidae). Our results showed that C. guanes had a generalist and omnivorous diet dominated by aquatic insects (mainly Diptera larvae) and seeds. Creagrutus guanes did not show intersexual or ontogenetic variation in diet. Peaks of feeding activity during rainy months were not synchronized with breeding in dry months. Our results do not support the hypothesis that the reproductive season has to be synchronized with high food consumption. We discussed the hypothesis fat reserves may be an important factor for the desynchronization of peaks of feeding and reproduction as explanation of seasonal breeding of this species.(AU)

A reprodução sazonal de peixes tropicais de água doce pode estar sincronizada com períodos de alto consumo de alimento. Nós examinamos esta hipótese estudando a relação entre dieta e atividade reprodutiva de Creagrutus guanes (Teleostei, Characidae). Nossos resultados mostraram que C. guanes teve uma dieta generalista e onívora, com predominância de insetos aquáticos (principalmente larva de Diptera) e sementes. Creagrutus guanes não apresentou variação intersexual ou ontogenética na dieta. Picos de atividade alimentar durante os meses chuvosos não foram sincronizados com atividade reprodutiva nos meses secos. Nossos resultados não sustentam a hipótese de que a estação reprodutiva está sincronizada com o consumo elevado de alimento. A reserva de gordura pode ser um fator importante para a dessincronização de picos de alimentação e reprodução (a reserva de gordura pode influenciar a ausência de sincronia entre os picos de alimentação e reprodução). São apresentadas hipóteses alternativas para a explicação de desova sazonal da espécie(AU)

How does diet influence the reproductive seasonality of tropical freshwater fish?: A case study of a characin in a tropical mountain river

Seasonal breeding of tropical freshwater fish may be synchronized with periods of high food consumption. We explored this hypothesis by studying the relationship between diet and reproductive activity of Creagrutus guanes (Teleostei, Characidae). Our results showed that C. guanes had a generalist and omnivorous diet dominated by aquatic insects (mainly Diptera larvae) and seeds. Creagrutus guanes did not show intersexual or ontogenetic variation in diet. Peaks of feeding activity during rainy months were not synchronized with breeding in dry months. Our results do not support the hypothesis that the reproductive season has to be synchronized with high food consumption. We discussed the hypothesis fat reserves may be an important factor for the desynchronization of peaks of feeding and reproduction as explanation of seasonal breeding of this species.(AU)

A reprodução sazonal de peixes tropicais de água doce pode estar sincronizada com períodos de alto consumo de alimento. Nós examinamos esta hipótese estudando a relação entre dieta e atividade reprodutiva de Creagrutus guanes (Teleostei, Characidae). Nossos resultados mostraram que C. guanes teve uma dieta generalista e onívora, com predominância de insetos aquáticos (principalmente larva de Diptera) e sementes. Creagrutus guanes não apresentou variação intersexual ou ontogenética na dieta. Picos de atividade alimentar durante os meses chuvosos não foram sincronizados com atividade reprodutiva nos meses secos. Nossos resultados não sustentam a hipótese de que a estação reprodutiva está sincronizada com o consumo elevado de alimento. A reserva de gordura pode ser um fator importante para a dessincronização de picos de alimentação e reprodução (a reserva de gordura pode influenciar a ausência de sincronia entre os picos de alimentação e reprodução). São apresentadas hipóteses alternativas para a explicação de desova sazonal da espécie(AU)