Keyword: Rodentia

Los análisis de egagrópilas y su impacto en el conocimiento de los micromamíferos de la Patagonia

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Los micromamíferos representan uno de los componentes más importantes de las comunidades de vertebrados terrestres neotropicales. Una metodología muy utilizada para analizar la composición de sus ensambles y los parámetros que los caracterizan (i.e. riqueza, diversidad) es el análisis de egagrópilas de aves rapaces. En Argentina, uno de los pioneros en utilizar esta metodología fue Elio Massoia en la década de 1960. Desde entonces, los trabajos basados en el estudio de egagrópilas para obtener información sobre micromamíferos fueron en aumento. El objetivo de este trabajo es documentar cómo esta metodología cambió nuestro conocimiento sobre los micromamíferos patagónicos. Se revisó la bibliografía publicada sobre análisis de egagrópilas y estudios enfocados en micromamíferos para la Patagonia argentina. El material recuperado de las egagrópilas ha permitido analizar la variabilidad intraespecífica de especies poco representadas en colecciones de museos o difíciles de capturar mediante trampas. Esta herramienta cambió de forma extraordinaria nuestra percepción sobre la biogeografía y distribuciones geográficas de muchas especies patagónicas (e.g. Calomys musculinus, Galea leucoblephara, Graomys griseoflavus, Lestodelphys halli, Loxodontomys micropus, Microcavia australis, Notiomys edwardsii, Paynomys macronyx, Thylamys pallidior). Asimismo, permitió reconsiderar las categorías de conservación de algunas especies que se creían raras y que demostraron solamente estar subrepresentadas en los muestreos tradicionales (i.e. N. edwardsii, L. halli, Euneomys chinchilloides e Irenomys tarsalis). Finalmente, también nos ha permitido reconstruir la estructura de los ensambles y los cambios que han atravesado desde al menos el Pleistoceno Tardío, documentando retracciones regionales, variaciones dramáticas en las frecuencias relativas y cambios en la dominancia de especies en los conjuntos actuales como consecuencia del impacto antrópico. El análisis de egagrópilas de aves rapaces representa una herramienta poderosa, especialmente en áreas extensas que serían difíciles de prospectar mediante otros métodos.

The analysis of owl pellets and their impact on knowledge of micromammals from Patagonia. Micromammals represent one of the most important components of the Neotropical terrestrial vertebrate communities. A widely used methodology to analyze the composition of their assemblages and the parameters that characterize them (i.e., richness, diversity) is the analysis of pellets produced by birds of prey. In Argentina, one of the pioneers in using this methodology was Elio Massoia. From that moment on, the studies based on the study of pellets to obtain information on small mammals were on the rise. The objective of this work is to document how this methodology changed our knowledge about Patagonian micromammals. We reviewed the published bibliography on pellet analysis and studies focused on small mammals for Argentinian Patagonia. The material recovered from pellets has made it possible to analyze the intraspecific variability of species poorly represented in museum collections or diffcult to capture using traps. This tool also dramatically changed our perception of the biogeography and geographical distributions of many Patagonian species (e.g., Calomys musculinus, Galea leucoblephara, Graomys griseoflavus, Lestodelphys halli, Loxodontomys micropus, Microcavia australis, Notiomys edwardsii, Paynomys macronyx, Thylamys pallidior). Likewise, it allowed reconsidering the conservation categories of some species that were believed to be rare (i.e., N. edwardsii, L. halli, Euneomys chinchilloides, and Irenomys tarsalis). Finally, it has also allowed us to reconstruct the structure of the assemblages and the changes on them since at least the Late Pleistocene, documenting regional retractions, dramatic variations in relative frequencies, and changes in the dominance of species in recent assemblages as a consequence of the anthropic impact. Pellet analysis of birds of prey represents a powerful tool, especially in large areas that would be diffcult to survey by other methods.

Reply to: Teta et al. (2020): on the distinction and availability of the new taxa proposed by Agnolin et al. 2019

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In a recent work Teta and collaborators conclude that all taxonomic decisions and new taxa established by Agnolin et al. should be considered invalid. It is not our aim to analyze point by point the critics and observations made by Teta et al., some of which we recognize are correct. In the present contribution we emphasize on the epistemological and philosophical flaws made by those authors. Teta et al. sustain their critics almost on authority criterion, acting as judge and jury on what works and what does not in mammalogy.We are convinced that without discussion there is a danger that any scientific discipline can be transformed in a dogma.

Respuesta a: Teta et al. (2020): Sobre la distinción y disponibilidad de los nuevos taxones propuestos por Agnolin et al. 2019. En una contribución reciente Teta y colaboradores concluyen que la totalidad de las decisiones taxonómicas y los nuevos taxones establecidos por Agnolin et al. deben ser considerados inválidos. No es nuestro fin analizar punto por punto las críticas y observaciones hechas por Teta et al., algunas de las cuales reconocemos son correctas. En la presente contribución enfatizamos en los errores epistemológicos y filosóficos cometidos por esos autores. Estamos convencidos de que sin discusión existe el peligro de que cualquier disciplina científica pueda ser transformada en un dogma.

Efecto de la condición social de alojamiento en cautiverio sobre comportamientos individuales: Calomys musculinus como especie modelo ecológico

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Numerosos estudios experimentales realizados en varias especies de mamíferos, pero principalmente en ratones y ratas de laboratorio, demostraron que el aislamiento social afecta profundamente los comportamientos individuales. Este estudio fue diseñado para comprender cómo los comportamientos agresivos y exploratorios de machos de Calomys musculinus, sexualmente maduros, podrían verse afectados por las condiciones de alojamiento. El estudio se realizó en bioterio durante marzo y mayo de 2018. Durante una semana, 35 machos fueron alojados en condición solitaria y 30 en condición agrupada (3 individuos por caja). Para estudiar el comportamiento de exploración se utilizó una cámara de desplazamiento. Para el estudio de agresividad, en una arena circular se realizaron enfrentamientos diádicos de 5 minutos de duración cada uno de ellos, entre individuos provenientes del mismo tratamiento. Los resultados obtenidos en este estudio experimental apoyaron la hipótesis planteada: los machos de C. musculinus alojados individualmente fueron más exploradores y menos agresivos que los provenientes de la condición agrupada. La mayor agresividad de los machos de C. musculinus alojados en grupo se vio reflejada por la intensidad de las aproximaciones ofensivas, ataques o persecuciones exhibidos durante los enfrentamientos intrasexuales. Nuestros resultados constituyen un aporte importante para la interpretación de futuros estudios en C. musculinus que requieran tanto del alojamiento transitorio de individuos, como del establecimiento de colonias fundadoras de breve permanencia en bioterio, de la conformación de una colonia bioterio o de la constitución de grupos experimentales.

Effect of the social status of housing in captivity on individual behaviors: Calomys musculinus as species ecological model. Several experimental studies conducted in various mammalian species, but mainly in laboratory mice and rats, demonstrated that social isolation deeply affects individual behaviors. This study was designed to understand how aggressive and exploratory behaviors of sexually mature Calomys musculinus males could be affected by housing conditions. The study was conducted in laboratory during March and May 2018. During one week, 35 males were housed in solitary condition and 30 in grouped condition (3 individuals per box). A displacement camera was used to study the scanning behavior. For the study of aggressiveness, dyadic confrontations of 5 minutes each were carried out in a circular arena, between individuals from the same treatment. The results obtained in this experimental study supported our hypothesis: C. musculinus males housed alone were more exploratory and less aggressive than those from the grouped condition. The greater aggressiveness of the C. musculinus males housed in a group was reflected by the intensity of the offensive approaches, attacks or persecutions exhibited during intra-sexual trials. Our results constitute an important contribution for the interpretation of future studies in C. musculinus that require both the transitory accommodation of individuals, the establishment of founder colonies of short stay in the bioterium, the formation of a bioterium colony, or the constitution of experimental groups.

Small mammals in restinga areas of north-eastern Brazil

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The restingas are an important component of the Atlantic Forest, affected by anthropic impacts along the Brazilian coast. The mastofauna that occurs in these environments is little known, especially in the north-east of the country. This study characterized the richness, abundance and composition of small mammal species in arboreal restinga areas in the state of Sergipe, north-eastern Brazil. Sampling was performed monthly, from September 2017 to August 2018. An effort composed of 8188 trap-nights with Sherman traps and 1782 trap-nights with pitfall traps resulted in the capture of 120 individuals from 10 species: four marsupials (Didelphis albiventris, Marmosa demerarae, Marmosa murina, and Marmosops incanus) and six rodents (Calomys sp., Cerradomys vivoi, Oligoryzomys sp., Phyllomys blainvilii, Rhipidomys mastacalis, and Trinomys sp.). Phyllomys blainvilii is a new record for the state. The observed richness corresponds to 73% of that estimated by Jackknife 1 (13.6 ± 1.5 species). Marmosops incanus and R. mastacalis were the most abundant, corresponding to 84% of the individuals captured. The community composition differs from the restingas of other regions of Brazil, but the majority of the species correspond to a subset of those found in the Atlantic Forest of the region. The locality presented the greatest richness of small mammals in areas of the Atlantic Forest of Sergipe, reinforcing the importance of the preservation of these areas and contributing to understanding the mastofauna in the restingas. This study also presents an updated list for this group in the state.

Pequenos mamíferos em áreas de restinga no nordeste do Brasil. As restingas são um componente importante da Mata Atlântica, sendo afetadas por impactos antrópicos ao longo da costa brasileira. A mastofauna que ocorre nesses ambientes ainda é pouco conhecida, principalmente no nordeste do país. O presente estudo caracterizou a riqueza, abundância e composição de pequenos mamíferos em uma fitofisionomia de restinga arbórea em Sergipe, nordeste do Brasil. A amostragem foi realizada mensalmente de setembro de 2017 a agosto de 2018. O esforço de 8188 armadilhas-noite e 1782 pitfalls-noite resultou na captura de 120 indivíduos pertencentes a 10 espécies: quatro marsupiais (Didelphis albiventris, Marmosa demerarae, Marmosa murina e Marmosops incanus) e seis roedores (Calomys sp., Cerradomys vivoi, Oligoryzomys sp., Phyllomys blainvilii, Rhipidomys mastacalis e Trinomys sp.). O registro de P. blainvilii é inédito para o estado. A riqueza observada corresponde a 73% da estimada por Jackknife 1 (13.6 ± 1.5 espécies). Marmosops incanus e R. mastacalis foram as espécies mais abundantes, correspondendo a 84% dos indivíduos capturados. A composição da comunidade difere das restingas de outras regiões do Brasil, mas a maioria das espécies corresponde a um subconjunto das encontradas nas áreas de Mata Atlântica da região. A localidade apresentou a maior riqueza de pequenos mamíferos em áreas da Mata Atlântica de Sergipe, reforçando a importância da preservação dessas áreas e contribuindo para o entendimento sobre a mastofauna que ocorre nas restingas. Este estudo apresenta assim uma lista atualizada de pequenos mamíferos para o estado.

On the distinction and availability of the new taxa proposed by Agnolin et al. 2019

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Recently, Agnolin et al. (2019) described 14 new species of mammals, including 12 rodents, one bat, and one carnivore, and one new subspecies of rodent. In addition, these authors proposed several other nomenclatorial acts: some nominal forms were removed from synonymies and hypothesized as distinct species, at the time that three new genera, one subtribe, and one tribe of mammals were also named. We reviewed the merits of all nomenclatorial acts proposed by Agnolin at al. (2019) and concluded that all 14 new species and the new subspecies, as well as those forms removed from synonymies, should be treated as synonyms of already known species. We suggest the same regarding the three new supraspecific taxa presented by Agnolin et al., two of which are not available as they fail to comply with the provisions of the International Code of Zoological Nomenclature. We end this contribution criticizing the way that mammal taxonomy was approached by Agnolin et al. (2019).

Sobre la distinción y disponibilidad de los nuevos taxones propuestos por Agnolin et al. 2019. Recientemente, Agnolin et al. (2019) describieron 14 especies nuevas de mamíferos, incluyendo 12 roedores, un murciélago y un carnívoro, y una nueva subespecie de roedor. Además, estos autores propusieron varios otros actos nomenclatoriales: algunas formas nominales se eliminaron de las sinonimias y se hipotetizaron como especies distintas; se nombraron tres nuevos géneros, una subtribu y una tribu de mamíferos. Revisamos todos los actos nomenclatoriales propuestos por Agnolin et al. (2019) y concluimos que las 14 nuevas especies y la nueva subespecie, así como las formas eliminadas de las sinonimias, deben tratarse como sinónimos de especies ya conocidas. Sugerimos lo mismo con respecto a los tres nuevos taxones supraespecíficos presentados por Agnolin et al. (2019), de los cuales dos no están disponibles ya que no cumplen con las disposiciones del Código Internacional de Nomenclatura Zoológica. Terminamos esta contribución criticando la forma en que Agnolin et al. (2019) realizaron su abordaje taxonómico.

Use of forest fragments and agricultural matrices by small mammals in southeastern Brazil

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The use of matrix by species that naturally inhabit forest fragments can be an important instrument to restore connectivity in fragmented landscapes. The type and structure of the matrix are determinant for this connectivity, however, few studies have explored this theme. In this study, we evaluated the community of small mammals within forest remnants and within two types of matrices (coffee plantation and pasture). In all, 11 species of small mammals were captured, being Akodon montensis, Oligoryzomys nigripes and Calomys cerqueirai the most abundant in fragments, coffee plantation and pasture, respectively. The results show that the composition and structure of the community in the two matrix types are distinct from those found in fragments, and the coffee plantation matrix has greater abundance and greater richness of small mammals than the pasture matrix.

Uso de fragmentos florestais e matrizes agrícolas por pequenos mamíferos no sudeste do Brasil. O uso da matriz por espécies que naturalmente habitam manchas florestais pode ser um instrumento importante para restaurar a conectividade em paisagens fragmentadas. O tipo e a estrutura da matriz são determinantes para esta conectividade, porém, poucos estudos exploram este tema. Neste estudo, nós avaliamos a comunidade de pequenos mamíferos dentro de fragmentos florestais e também dentro de dois tipos de matrizes (plantação de café e pasto). No total foram capturadas 11 espécies de pequenos mamíferos, sendo Akodon montensis, Oligoryzomys nigripes e Calomys cerqueirai as mais abundantes nos fragmentos, plantação de café e pasto, respectivamente. Os resultados mostram que a composição e a estrutura da comunidade nos dois tipos de matrizes foram diferentes daquelas encontradas nos fragmentos, e que a matriz de café possui maior abundância e riqueza de espécies que a matriz de pasto.

Richness, endemism and conservation of sigmodontine rodents in Argentina

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Sigmodontine rodents, with 86 genera and ~430 living species, constitute one of the most successful radiations of Neotropical mammals. In this contribution, we studied the distributional ranges of 108 sigmodontine species in Argentina. Our objectives were (i) to establish geographical patterns of species richness and endemism, and (ii) to evaluate the regional conservation status of these taxa. We constructed a minimum convex polygon for each species, using information from literature and biological collections. Individual maps were superimposed on a map of Argentina divided into cells of 25 km on each side. For each cell, we calculated the species richness, which varied between 1 and 21 species, and its degree of endemism, which fluctuated between 0.001 and 3.28. There were 30 species of sigmodontine rodents distributed almost exclusively in Argentina, most of them restricted to forested areas (Southern Andean Yungas) or to arid and semiarid environments (High and Low Monte and Patagonian Steppe). Areas with high species richness and endemism scores corresponded grossly with the Southern Andean Yungas, the Humid Chaco plus the Paraná flooded savannas, the Alto Paraná Atlantic forests plus the Araucaria moist forests, the High Monte and the ecotone between the Patagonian steppe and the Valdivian temperate forests. A reassessment of the conservation status of sigmodontine rodents distributed in Argentina retrieved 2 extinct species, 7 endangered, 7 vulnerable, 6 near threatened and 13 data deficient. These numbers suggest a much more serious situation than the expressed by previous evaluations, highlighting the urgent need to establish conservation measures for the protection of this group.

Riqueza, endemismo y conservación de roedores sigmodontinos en Argentina. Los roedores sigmodontinos, con 86 géneros y ~430 especies vivientes, constituyen una de las radiaciones más exitosas de mamíferos neotropicales. En esta contribución estudiamos los rangos de distribución de 108 especies de sigmodontinos en Argentina. Nuestros objetivos fueron (i) establecer patrones geográficos de riqueza de especies y endemismo y (ii) evaluar el estado de conservación regional de estos taxones. Construimos un polígono convexo mínimo para cada especie, utilizando información de la literatura y colecciones biológicas. Los mapas individuales fueron superpuestos en un mapa de Argentina dividido en celdas de 25 km de lado. Para cada celda calculamos la riqueza de especies, que varió entre 1 y 21, y su grado de endemismo, que fluctuó entre 0.001 y 3.28. Hubo 30 especies de roedores sigmodontinos distribuidos casi exclusivamente en Argentina, la mayoría de ellos restringidos a áreas boscosas (Yungas andinas del sur) o a ambientes áridos y semiáridos (Monte alto y bajo y Estepa Patagónica). Las áreas con mayor riqueza de especies y valores más altos de endemismo se correspondieron groseramente con las Yungas andinas del sur, el Chaco húmedo más las Sabanas inundadas de Paraná, el Bosque Atlántico del Alto Paraná más los Bosques húmedos de araucaria, el Monte alto y el ecotono entre la Estepa Patagónica y los Bosques templados valdivianos. Una reevaluación del estado de conservación de los roedores sigmodontinos distribuidos en Argentina recuperó 2 especies extintas, 7 en peligro, 7 vulnerables, 6 casi amenazadas y 13 con datos deficientes. Estas cifras sugieren una situación mucho más grave que la expresada en evaluaciones anteriores, destacando la necesidad urgente de establecer medidas de conservación para la protección de este grupo.

Terrestrial mammals of the Jequitinhonha River Basin, Brazil: a transition area between Atlantic Forest and Cerrado

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Here we present an extensive survey for non-volant mammals along the margins of the Jequitinhonha River, which represents a contact area between the Cerrado and Atlantic Forest biomes. Our data were obtained after 13 trapping expeditions (from 2005 to 2012) on both banks of the river, from its source to its mouth, enriched by data obtained from a literature review and voucher specimens deposited in six mammal collections. All collected specimens were identified through their karyotypes and/or morphology. We obtained data for 75 localities (34 at the rigth margin, 42 at the left margin; 30 in the Cerrado and 45 in the Atlantic Forest). We obtained voucher records for 91 species, ca. 17% of all known Brazilian terrestrial non-volant mammals. Our results include range extensions for six species (Monodelphis scalops, Thylamys velutinus, Calomys mattevii, Wiedomys pyrrhorhinos, Galea spixii, Kerodon rupestris), and karyotype information for 26 species (11 marsupials and 15 rodents), with four new karyotypes. While most species have wide geographical range in both biomes, 26 were recorded only in the Cerrado and 19 were recorded only in the Atlantic Forest. Recorded species represent 28.7% (6.3% endemic) of the species known from the Cerrado and 21.5% (22.2% endemic) of those from the Atlantic Forest. The richness of the fauna recorded in this region supports its importance for the understanding of mammal biogeography, diversity and evolution.

Mamíferos terrestres da bacia do Rio Jequitinhonha, Brasil: uma área de transição entre a Floresta Atlântica e o Cerrado. Aqui apresentamos uma extensa pesquisa de mamíferos não-voadores ao longo das margens do Rio Jequitinhonha, região biogeográfica interessante que representa uma área de contato entre dois biomas brasileiros (Cerrado e Mata Atlântica), ainda pouco conhecidos para a fauna de mamíferos. Nossos dados foram obtidos após 13 expedições de captura (de 2005 a 2012), em ambas as margens do rio, da nascente até à foz, enriquecida por dados obtidos a partir de amostras de material depositado em seis coleções de mamíferos. Todos os espécimes coletados foram identificados através de seus cariótipos ou morfologia. Foram obtidos dados de 75 localidades (34 na margem direita, 41 à esquerda; 30 no Cerrado e 45 na Floresta Atlântica). Obtivemos registros de 91 espécies de mamíferos não-voadores, ca. de 17% de todos os mamíferos terrestres brasileiros conhecidos. Nossos resultados incluem extensões de distribuição de seis espécies (Monodelphis scalops, Thylamys velutinus, Calomys mattevii, Wiedomys pyrrhorhinos, Galea spixii, Kerodon rupestris), e informações de cariótipo de 26 espécies (11 marsupiais e 15 de roedores), com quatro novos cariótipos. Enquanto a maioria das espécies têm ampla distribuição geográfica, 26 ocorreram apenas no Cerrado e 19 apenas na Mata Atlântica. Foram registradas de 28.7% (6.3% endêmicas) de todas as espécies conhecidas para o Cerrado e 21.5% (22.2% endêmicas) da Floresta Atlântica. A riqueza da fauna registrada nesta região apoia a sua importância da compreensão na diversidade, biogeografia e evolução dos mamíferos.

Variación morfológica de Oligoryzomys longicaudatus magellanicus (Rodentia: Sigmodontinae) al oeste de los Campos de Hielo Patagónico Sur en Chile

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Oligoryzomys es uno de los géneros de roedores más ampliamente distribuido en el territorio chileno. Se describe la variabilidad morfológica de individuos provenientes de localidades al oeste de Campos de Hielo Patagónico Sur entre los 48° S y 51° S. Análisis uni y multivariados realizados sobre la base de mediciones del cuerpo, mandíbula y cráneo no mostraron diferencias significativas por sexo, pero sí por procedencia geográfica. Los individuos insulares presentaron valores mayores para casi todos los caracteres evaluados, lo que podría corresponderse con la ausencia de predadores y competidores. Morfológica y distribucionalmente, los individuos analizados son asignados a la subespecie Oligoryzomys longicaudatus magellanicus del extremo sur del continente.

Morphological variation of Oligoryzomys longicaudatus magellanicus (Rodentia: Sigmodontinae) west of the Southern Patagonian Ice Fields in Chile. Oligoryzomys is one of the most widely distributed mammal genera in Chile. We describe morphological variation among specimens from west of the Southern Patagonian Ice Fields, between 48° S and 51° S. Univariate and multivariate analysis showed no significant differences between sexes, but uncovered significant geographical variation. Island specimens had higher values for almost all characters evaluated, which could correspond to the absence of predators and competitors. Based on morphological and distributional information, we suggest that individuals analyzed belong to the subspecies Oligoryzomys longicaudatus magellanicus.

Variación morfológica cualitativa y cuantitativa en Abrothrix longipilis (Cricetidae, Sigmodontinae)

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Se estudió la taxonomía alfa de Abrothrix longipilis (Waterhouse, 1837), uno de los abrotriquinos más ampliamente distribuidos y abundantes en las porciones australes de Argentina y Chile. Se realizó un análisis morfológico cualitativo (6 caracteres discretos) y cuantitativo (18 medidas cráneo-dentarias) de 477 especímenes adultos, agrupados primero según un criterio geográfico y luego según su pertenencia a los clados definidos por un estudio filogeográfico previo basado en 2 marcadores moleculares. Los análisis multivariados (análisis de componentes principales y análisis discriminantes «libres de tamaño») indican que A. longipilis es un complejo que incluye por lo menos 2 especies: A. longipilis stricto sensu, aquí restringida al centro de Chile, y A. hirta (Thomas, 1895), distribuida en el sur de Chile y sudoeste de Argentina. Las poblaciones de A. hirta mostraron una correspondencia moderada entre los patrones filogeográficos y de variación fenotípica. Preliminarmente, puede considerarse que el tamaño en A. hirta se vincula más estrechamente con las características del ambiente (con los individuos más grandes en áreas boscosas, de mayor productividad primaria y los más pequeños en ambientes áridos, de menor productividad) antes que con su pertenencia a los distintos clados. En este escenario, la divergencia morfológica podría estar mayormente determinada por la selección diferencial a lo largo de un gradiente ambiental extremo, aun en presencia de flujo génico (modelo de divergencia con flujo génico).

Qualitative and quantitative morphological variation in Abrothrix longipilis (Cricetidae, Sigmodontinae). We studied the alpha taxonomy of Abrothrix longipilis (Waterhouse, 1837), one of the most widely distributed and abundant Abrotrichini along the southern portion of Argentina and Chile. We conducted a qualitative (6 discrete characters) and quantitative (18 skull and dental measurements) morphological analysis of 477 adult specimens, grouped first by geographical criteria and later by membership to the clades defined in a previous phylogeographical study based on 2 molecular markers. Multivariate analyses (principal component analysis, discriminant analyses) suggested that in its current conception A. longipilis is a species-complex that includes at least two nominal taxa: A. longipilis stricto sensu, here restricted to Central Chile, and A. hirta (Thomas, 1895), distributed in southern Chile and southwestern Argentina. Populations of A. hirta showed a moderate correspondence between phylogeographic and phenotypic patterns. Tentatively, we considered that general size in A. hirta is mostly linked to environmental conditions (with larger individuals in forested areas, under high primary productivity conditions and the smaller ones in arid environments of low productivity) rather than to their inclusion in different clades. Under this scenario, morphological differentiation was probably driven by differential selection across a sharp ecological gradient, in spite of gene flow (divergence-with-gene-flow model).