RESUMO
The ecology and evolution of Caribbean anoles are well described, yet little is known about mainland anole species. Lack of phylogenetic information limits our knowledge about species boundaries, morphological evolution, and the biogeography of anoles in South America. To help fill this gap, we provide an updated molecular phylogeny of the Dactyloa (Dactyloidae), with emphasis on the punctata species group. By sampling understudied Amazonian taxa, we (i) assess the phylogenetic placement of the 'odd anole', D. dissimilis; (ii) infer the relationships of the proboscis-bearing D. phyllorhina, testing the hypothesis of independent nasal appendage evolution within the anole radiation; and (iii) examine genetic and dewlap color variation in D. punctata and D. philopunctata. Combining multiple nuclear loci with a review of the fossil record, we also (iv) estimate divergence times within the pleurodont iguanian clade of lizards, including Amazonian representatives of Dactyloa and Norops (Dactyloidae) and of Polychrus (Polychrotidae). We recover the five Dactyloa clades previously referred to as the aequatorialis, heteroderma, latifrons, punctata and roquet species groups, as well as a sixth clade composed of D. dissimilis and the non-Amazonian D. neblinina and D. calimae. We find D. phyllorhina to be nested within the punctata group, suggesting independent evolution of the anole proboscis. We consistently recover D. philopunctata nested within D. punctata, and report limited genetic divergence between distinct dewlap phenotypes. The most recent common ancestor of Dactyloa, Anolis and Norops dates back to the Eocene. Most Amazonian taxa within both Dactyloa and Norops diverged in the Miocene, but some diversification events were as old as the late Eocene and late Oligocene. Amazonian Polychrus diverged in the Pliocene. Our findings have broad implications for anole biogeography, disputing recent suggestions that modern dactyloid genera were present in the Caribbean region during the Cretaceous.
Assuntos
Evolução Biológica , Lagartos/classificação , Filogenia , Animais , Teorema de Bayes , Fósseis , Funções Verossimilhança , Lagartos/anatomia & histologia , Lagartos/genética , Modelos Genéticos , Fenótipo , Análise de Sequência de DNA , América do SulRESUMO
Escape by Anolis lizards is influenced by microhabitats and fight initiation distance increases with predation risk. Differences in microhabitat use among ecomorphs affect escape behavior, but only two studies have reported ecomorphological differences in flight initiation distance among Greater Antillean species. I studied effects of predation risk and microhabitats on escape behavior by conducting field experiments using two species of anoles, Anolis lineatopus and A. grahami, on the campus of the University of the West Indies at Mona, Jamaica. Because ecomorphological variation of anoles has evolved independently within each island of the Greater Antilles, but relationships between ecomorphs and escape behaviors are poorly known, I characterized microhabitat use and escape tactics, and determined relationships between flight initiation distance and two risk factors, habituation to human presence and perch height, in Anolis lineatopus, a trunk-ground anole and A. grahami, a trunk-crown anole. Sample sizes for A. lineatopus and A. grahami were 214 and 93, for microhabitat use and escape destinations, 74 and 34 for human presence and 125 and 34 for perch height. The two species occurred in similar microhabitats and exhibited similar escape tactics, but exhibited key differences expected for their ecomorphs. Both species were sighted frequently on the ground and on trees, but A. lineatopus were more frequently on ground and were perched lower than A. grahami. Both species escaped from ground to trees and when on trees hid on far sides and escaped without changing climbing direction with equal frequency. The frequency of fleeing upward was greater for A. grahami than A. lineatopus. Both species exhibited habituation by having shorter flight initiation distances in areas with more frequent exposure to people. In both species flight initiation distance increased as perch height decreased because, lizards had to climb farther to be out of reach when perched lower. The relationship between flight initiation distance and perch height may apply to other anole ecomorphs that flee upward when low perched on trees. Rev. Biol. Trop. 58 (4): 1199-1209. Epub 2010 December 01.
El escape de las largarijas Anolis está influenciado por el microhábitat y la distancia de iniciación de escape incrementa el riesgo de depredación. Las diferencias en el uso de microhábitats entre ecomorfos afecta el comportamiento de escape, pero sólo dos estudios han reportado diferencias ecomorfológicas en la distancia de iniciación de escape entre las especies de las Antillas Mayores. Se estudió el efecto de riesgo de depredación y la influencia del microhábitat en el comportamiento de escape, mediante la realización de experimentos de campo con Anolis lineatopus y A. grahami, en el campus de la Universidad West Indies en Mona, Jamaica. Debido a que las variaciones ecomorfológicas de Anolis han evolucionado independientemente en cada isla de las Antillas Mayores, la relación entre ecomorfos y el comportamiento de escape son pobremente conocidos. Se caracteriza el uso del microhábitat y las tácticas de escape, se determinan las relaciones entre la distancia de iniciación de escape y los dos factores de riesgo (habituación a presencia humana y altura a la que se posan) de Anolis lineatopus, una lagartija que habita en troncos-tierra y A. grahami, una lagartija de troncos-partes más altas. Los tamaños de muestra para A. lineatopus y A. grahami fueron: 214 y 93, para uso del microhábitat y destinos de escape 74, para presencia humana 34 y para perchas altas 125 y 34. Las dos especies se presentan en microhábitats similares y mostraron tácticas de escape parecidas, pero exhibieron diferencias claves esperadas para sus ecomorfos. Ambas especies fueron vistas con frecuencia en el suelo y en los árboles, pero A. lineatopus fue encontrada más frecuentemente en el suelo y debajo de A. grahami. Ambas especies escaparon del suelo a esconderse en los árboles y huían con igual frecuencia sin cambiar de dirección. La frecuencia de huir hacia arriba fue mayor para A grahami. Ambas especies mostraron habituación al tener distancias más cortas de iniciación de escape en zonas con exposición frecuente a la gente y la distancia de iniciación de escape incrementa cuando la altura de la percha disminuye, porque las lagartijas tienden a subir más al estar fuera de nuestro alcance cuando se posan en la parte baja. La relación entre la distancia de iniciación de escape y altura de la percha puede aplicar a otros ecomorfos de Anolis que huyen hacia arriba cuando están posados en las partes bajas de los árboles.
Assuntos
Animais , Humanos , Ecossistema , Reação de Fuga/fisiologia , Lagartos/fisiologia , Jamaica , Comportamento Predatório , Especificidade da EspécieRESUMO
Among a series of eight specimens of Anolis phyllorhinus obtained in the northern part of the state of Mato Grosso, Brazil, one was determined to be the first female known for the species and the first so far reported for proboscid anoles. The female lacks the characteristic male proboscis and, excepting a few consistent differences, is morphologically similar to females of Anolis punctatus, a species closely related and broadly sympatric with A. phyllorhinus. Males and females of the new series are described and new data on their ecology and behavior presented. The new data are consistent with the hypotheses that the male proboscis probably evolved as a sexual signal and that visual signals for intra and interspecific recognition are redundant in anoles.
Em uma série de 8 exemplares de Anolis phyllorhinus coletados em Aripuanã, no norte do estado de Mato Grosso encontra-se a primeira fêmea da espécie, que também vem a ser a primeira fêmea conhecida para os anolis dotados de apêndice nasal. A fêmea não apresenta a probóscide característica dos machos e, exceto por pequenas diferenças, embora consistentes, é morfologicamente muito semelhante às fêmeas de Anolis punctatus, espécie relacionada e amplamente simpátrica com Anolis phyllorhinus. Descrevem-se os machos e a fêmea da nova série, acrescidos de informações obtidas no campo sobre sua ecologia e comportamento. Os novos dados, concordam com hipóteses anteriores sugerindo que a probóscide do macho possívelmente evoluiu como um sinal sexual e que os sinais visuais para reconhecimento intra e interespecífico são redundantes no gênero Anolis.
RESUMO
Among a series of eight specimens of Anolis phyllorhinus obtained in the northern part of the state of Mato Grosso, Brazil, one was determined to be the first female known for the species and the first so far reported for proboscid anoles. The female lacks the characteristic male proboscis and, excepting a few consistent differences, is morphologically similar to females of Anolis punctatus, a species closely related and broadly sympatric with A. phyllorhinus. Males and females of the new series are described and new data on their ecology and behavior presented. The new data are consistent with the hypotheses that the male proboscis probably evolved as a sexual signal and that visual signals for intra and interspecific recognition are redundant in anoles.
Em uma série de 8 exemplares de Anolis phyllorhinus coletados em Aripuanã, no norte do estado de Mato Grosso encontra-se a primeira fêmea da espécie, que também vem a ser a primeira fêmea conhecida para os anolis dotados de apêndice nasal. A fêmea não apresenta a probóscide característica dos machos e, exceto por pequenas diferenças, embora consistentes, é morfologicamente muito semelhante às fêmeas de Anolis punctatus, espécie relacionada e amplamente simpátrica com Anolis phyllorhinus. Descrevem-se os machos e a fêmea da nova série, acrescidos de informações obtidas no campo sobre sua ecologia e comportamento. Os novos dados, concordam com hipóteses anteriores sugerindo que a probóscide do macho possívelmente evoluiu como um sinal sexual e que os sinais visuais para reconhecimento intra e interespecífico são redundantes no gênero Anolis.