Proechimys e Trinomys (Rodentia: Echimyidae): diferenças no sincrânio em dois gêneros de ratos de espinho na região Neotropical

As coleções científicas têm um papel muito importante em salvaguardar registro da biodiversidade. Neste estudo, utilizamos crânios e mandíbulas de roedores obtidos em três museus e uma Universidade do Brasil. O nosso objetivo foi descrever as diferenças morfológicas entre os dois gêneros de ratos de espinho Proechimys e Trinomys. Para isso, foram fotografados um total de 612 indivíduos de ambos os gêneros. Utilizamos métodos de morfometria geométrica com a plotagem de marcos anatômicos bidimensionais para descrever a forma e tamanho do sincrânio. Nossos resultados demonstraram que não há diferença estatística de tamanho do sincrânio entre os gêneros Proechimys e Trinomys. Porém, há diferença de forma: as espécies de Trinomys apresentaram bulas auditivas proporcionalmente menores, um processo jugal relativamente mais proeminente e um crânio relativamente mais robusto do que as espécies de Proechimys, enquanto as espécies de Proechimys apresentam um crânio relativamente mais delgado e afilado do que as espécies de Trinomys. Este trabalho adiciona informações referentes às diferenças já estabelecidas para os gêneros Proechimys e Trinomys. Além disso, demonstra a importância das coleções científicas para a descrição da biodiversidade

Phenotypic plasticity in Heliconius erato (Lepidoptera: Nymphalidae) mandibles induced by different host plants (Passifloraceae)

El objetivo fue analizar si el consumo de diferentes plantas influenciaba el tamaño/forma de las mandíbulas de larvas de lepidópteros oligófagas y si dicho consumo ocasionaba tasas de crecimiento diferenciadas entre el músculo aductor mandibular y las mandíbulas durante su ontogénesis. Para esto se usaron larvas de Heliconius erato criadas en cuatro plantas hospederas naturales, a las que se les tomaron medidas de longitud de cápsula cefálica, tamaño mandibular y área del músculo aductor. Para detectar eventuales diferencias de forma mandibular se empleó microscopía electrónica y morfometria geométrica, y para posibles cambios alométricos se realizaron comparaciones entre curvas de crecimiento mandíbula vs. músculo aductor de los individuos criados en las cuatro plantas hospederas. Se encontró que no existen diferencias de tamaño entre las mandíbulas ni en músculos aductores de individuos criados en las diferentes plantas pero la forma mandibular fue significativamente diferente, mostrando una relación directa con la planta hospedera usada como alimento. Generalmente, la mandíbula creció proporcionalmente más que el músculo aductor, cuyo crecimiento fue estable. El cambio más significativo fue el alargamiento de los dientes incisivos en mandíbulas de larvas criadas con Passiflora suberosa. Se sugiere que esto se debe a que las hojas de esta especie poseen mayor dureza comparadas con las otras y que el desarrollar dientes largos facilitaría su corte y consumo.This study aimed to test whether the consumption of different host plants influences the size and shape of mandibles and associated muscles in oligophagic insects throughout ontogeny. Larvae of Heliconius erato phyllis (Lepidoptera: Nymphalidae) were reared on four species of passion vines (Passifloraceae) and had the size of their head caspules, mandibles and associated adductor muscles measured afterward. Corresponding effects on shape and potential allometries in mandibles were also evaluted using geometric morphometric analysis based on scanning electron micrographs. Growth in mandibular size was greater compared to that of associated adductor muscles. There was no host plant effect on the sizes of either mandibles or adductor muscles. However, the shapes of mandibles was significantly different among passion vine species used as larval food. Greater differences in shape associated with wider teeth were obtained for Passiflora suberosa, whose leaves are tougher compared to those of P. misera, P. caerulea, and P. actinia; and, among which there was no difference in manbilular shape. Thus, it was inferred that the larval stage of this Heliconius species is able to adjust its mandible shape according to the toughness of host plants; a phenotypic plasticity mechanism that can be supposed to have evolved in order to facilitate processing these kinds of leaves

Leaching of carbon from native and non-native leaf litter of subtropical riparian forests

We evaluated the leaching of dissolved organic and inorganic carbon (DOC and DIC, respectively) from leaf litter of plant species of the native Atlantic Forest and of Pinus elliottii (Engelm.) and Eucalyptus grandis (Hill ex Maiden) forests. We carried out experiments in which leaves from each forest system (native, E. grandis, and P. elliottii) were placed in reactors with sterile water. After 1/4 h, 1 h, 4 h, 12 h, 24 h, and 48 h of incubation, we quantified the concentrations of DIC and DOC of each leachate. The greatest quantity of DIC was leached in the native forest system, whereas the greatest quantity of DOC was leached in the E. grandis forest system. With respect to the period of year, the greatest quantity of DIC was leached in autumn, whereas that of DOC was leached in the summer. Our results demonstrated that the replacement of native species in riparian zones, by non-native forest monocultures, could alter the chemical composition of the water. Finally, the results presented in this paper reinforce the need for a more careful look at ecological processes

Estudo da variação morfológica craniana entre quatro blocos populacionais de Ctenomys lami (Rodentia, Ctenomydae) através de morfometria geométrica

Este estudo analisa a variação na forma do crânio de Ctenomys lami, em relação a machos e fêmeas, diferentes blocos populacionais, diferentes números diplóides, cariótipos e em relação aos pares cromossômicos 1 e 2. Foram utilizados 90 crânios de espécimes adultos, 36 machos e 54 fêmeas, todos provenientes da região da Coxilha das Lombas, Rio Grande do Sul, Brasil. Esta espécie apresenta sete citótipos diferentes distribuídos em quatro blocos populacionais, sendo o bloco A com 2n = 54, 55a e 56a; bloco B com 2n = 57 e 58; bloco C com 2n = 54 e 55a; e o bloco D com 2n = 55b e 56b. A variação morfométrica foi estudada nas vistas dorsal, ventral e lateral dos crânios, sendo determinados marcos anatômicos em cada uma delas. Os marcos foram posicionados com o programa TPSDig, totalizando 14 marcos para a vista dorsal, 14 para a ventral e 15 para a lateral. Para as análises de morfometria geométrica foi utilizado o método de sobreposição de Procrustes com as coordenadas dos marcos para diferentes vistas foram feitas análises de componentes principais (PCA), análises de variáveis canônicas (CVA), análise das distâncias de Procrustes entre as formas médias. Os resultados confirmam a presença de diferentes blocos populacionais com formas específicas para cada um deles, variação da forma em relação aos diferentes números cromossômicos e presença de dimorfismo sexual para a espécie, sendo este mais intenso no bloco B. Estes dados nos levam a sugerir que existe associação entre variação cromossômica e variação morfológica.This study examine the shape variation in the skull of Ctenomys lami, in relation to males and females, different population blocks, different diploid numbers, karyotypes and in relation to chromosomal pairs 1 and 2. We used 90 skulls of adults specimens (36 males and 54 females), all of them deriving from Coxilha das Lombas, Rio Grande do Sul, Brazil. This species have seven different cytotypes distributed in four population blocks, block A with 2n = 54, 55a and 56a; block B with 2n = 57 and 58; block C with 2n = 54 and 55a; and block D with 2n = 55b and 56b. The morphometric variation was studied in the dorsal, ventral and lateral views of the skull. We established landmarks in each view. Landmarks were located with TPSDig software, 14 in the dorsal view, 14 in the ventral view and 15 for lateral view. For the geometric morphometric analysis we used Procrustes superimposition method based in the coordinates of landmarks for different views. We made Principal Component Analysis (PCA), Canonical Variate Analysis (CVA), comparisons between distances between means shape. The results confirm the presence of different population blocks with specific shapes for each one, shape variation in relation to different chromosome number and existence of sexual dimorphism for this species, been more intense in block B. This data led us to suggest the existence of association between chromosome variation and morphological variation

Evolução na forma e tamanho do crânio no gênero Ctenomys (Rodentia: Ctenomydae)

Ctenomys é um gênero de roedores subterrâneos endêmico ao sul da região Neotropical, que compreende cerca de 60 espécies conhecidas como “tuco-tucos”. Este gênero é um exemplo de evolução rápida, que tem uma alta diversidade cariotípica com número diplóide variando de 2n = 10 até 2n = 70. Com exceção de algumas espécies, a variação morfológica quantitativa entre os tuco-tucos é pouco conhecida. Recentemente, estudos usando citocromo b propuseram relações filogenéticas para Ctenomys e apoiam oito grandes grupos. O objetivo deste estudo é investigar neste gênero, os padrões de evolução morfológica na forma e no tamanho do crânio, assim como diferenças entre sexos, em nível macro e microevolutivo, usando uma abordagem de morfometria geométrica. Em nível macroevolutivo, analisamos padrões de variação em 47 espécies do gênero Ctenomys e a variação dentro e entre os grupos mendocinus e torquatus. Em nível microevolutivo, analisamos mais detalhadamente as espécies C. minutus e C. lami. Ambas ocorrem na planície costeira do sul do Brasil. A primeira espécie tem 15 cariótipos (2n = 42, 43, 44, 45, 46a, 46b, 47a, 47b, 48a, 48b, 48c, 49a, 49b, 50a e 50b), o maior polimorfismo cromossômico conhecido para o gênero Ctenomys. C. lami também apresenta uma alta variação cariotípica, com números diplóides de 2n = 54 até 2n = 58. Baseado em diferentes frequências de rearranjos cromossômicos ao longo de sua distribuição geográfica, quatro blocos populacionais foram previamente propostos. Em nível macroevolutivo, a diversificação na forma do crânio parece estar estruturada geograficamente. Encontramos gradientes tanto leste-oeste quanto norte-sul, crânios mais robustos no norte e crânios mais gráceis nas espécies do sul. A mandíbula se mostrou menos variável na forma do que o crânio. Os fenogramas gerados com distâncias de Mahalanobis mostraram um forte sinal filogenético para o grupo torquatus e similaridade morfológica entre os dois grupos Bolivianos e entre os grupos mendocinus e Patagônico. O grupo torquatus é muito congruente na forma e pouco varia em tamanho. Uma relação de ancestralidade comum entre os grupos Patagônico e mendocinus, baseada na forma do crânio, é reforçada pela mesma morfologia de espermatozóide assimétrico encontrada nos dois grupos. Em nível microevolutivo, a forma do crânio não parece obedecer ao mesmo padrão. Em C. minutus a variação morfológica é estruturada espacialmente por um modelo de isolamento pela distância, por barreiras geográficas antigas (paleo-canais) e provavelmente por uma inversão cromossômica. Os rearranjos cromossômicos envolvendo fusões e fissões robertsonianas parecem não ser um fator primário que promova a estruturação na variação do crânio. A diferença de habitats também foi um dos fatores detectados como estruturante. Além disso, nós encontramos deriva, seleção diversificadora e seleção estabilizadora atuando na variação da forma do crânio nos quatro blocos populacionais cromossômicos de C. lami. O padrão de seleção diversificadora envolve estruturas relacionadas à escavação, que são compatíveis com modelos biomecânicos propostos para Ctenomys. O padrão de seleção estabilizadora envolve um ajuste das proporções da bula timpânica, entre a capacidade auditiva e a abertura mandibular. De forma geral, nossos achados sugerem que a evolução do crânio de Ctenomys é resultado principalmente de processos históricos regidos pela biologia da espécie em níveis macroevolutivos. Porém, em níveis microevolutivos, a combinação de diversas forças evolutivas e processo históricos produzem padrões mais variáveis do que o esperado e pouco relacionados com a distribuição geográfica.Ctenomys is a genus of subterranean rodents endemic to the southern Neotropical region, which comprise about 60 living species known as “tuco-tucos”. This genus is an example of rapid evolution, which have high karyotypic diversity with diploid numbers varying from 2n = 10 to 2n = 70. To the exception of a few species, quantitative morphological variation among tuco-tucos is poorly known. Recently, studies using cytochrome b proposed phylogentic relationships for Ctenomys and support eight great groups. The goal of this study is to investigate in this genus, the morphological evolution patterns in skull shape and size variation between sexes, in macro and microevolutionary levels, using a geometric morphometrics approach. In macroevolutionay level, we analyze the variation pattern in 47 species of the genus Ctenomys and the variation within and between mendocinus and torquatus groups. In microevolutionary level we analyze in more details C. lami and C. minutus. Both occurs in coastal plain of southern Brazil. The first species has 15 karyotypes (2n = 42, 43, 44, 45, 46a, 46b, 47a, 47b, 48a, 48b, 48c, 49a, 49b, 50a and 50b), the highest intraspecific chromosomal polymorphism known for the genus Ctenomys. C. lami also shows high karyotypic variation, with diploid numbers ranging from 2n = 54 to 2n = 58. Based on different chromosome rearrangement frequencies along its geographical distribution, four population blocks were previously proposed. In macroevolutionary level we found that skull evolution appear geographically structured. We found west-east and north-south gradients, skulls more robust in north and skulls more gracile in south species. The mandible is less variable in shape than skull. The phenograms of Mahalanobis distances show strong phylogenetic signal only for torquatus-group and morphological similarity between two Bolivian groups, and between mendocinus and Patagonico groups. The torquatus-group is very congruent in shape and have low variation in size. Common ancestral relationship between Patagonico and mendocinus groups based in skull shape is reinforced by the same asymmetry in sperm morphology found in both groups. In microevolutionary level, the skull shape does not follow the same pattern. For C. minutus the morphological variation is spatial structured by the isolation-by-distance model by ancient barriers (paleochannels) and probably by a chromosomal inversion. Chromosomal rearrangements involving robertsonians fusions and fissions appear not be a primary factor that promote the structure in skull shape. The habitats difference also a factor in structure shape change. Moreover, we found drift, diversifying selection and stabilizing selection acting on variation in skull shape in four chromosomal population blocks of C. lami. Diversifying patterns involve structures related to digging, that are compatible with biomechanical models proposed for Ctenomys. Stabilizing patterns involve a compromise in the proportions of the auditory bulla, between hearing ability and the mandibular opening. In general, our findings suggested that Ctenomys skull evolution is a result of historical factors conducted by species biology in macroevolutionary level. However, in microevolutionary level the combination of many forces and historical process producing more variable patterns than expected and weak related to geographical distribution

Estudo da variação morfológica craniana entre quatro blocos populacionais de Ctenomys lami (Rodentia, Ctenomydae) através de morfometria geométrica

Este estudo analisa a variação na forma do crânio de Ctenomys lami, em relação a machos e fêmeas, diferentes blocos populacionais, diferentes números diplóides, cariótipos e em relação aos pares cromossômicos 1 e 2. Foram utilizados 90 crânios de espécimes adultos, 36 machos e 54 fêmeas, todos provenientes da região da Coxilha das Lombas, Rio Grande do Sul, Brasil. Esta espécie apresenta sete citótipos diferentes distribuídos em quatro blocos populacionais, sendo o bloco A com 2n = 54, 55a e 56a; bloco B com 2n = 57 e 58; bloco C com 2n = 54 e 55a; e o bloco D com 2n = 55b e 56b. A variação morfométrica foi estudada nas vistas dorsal, ventral e lateral dos crânios, sendo determinados marcos anatômicos em cada uma delas. Os marcos foram posicionados com o programa TPSDig, totalizando 14 marcos para a vista dorsal, 14 para a ventral e 15 para a lateral. Para as análises de morfometria geométrica foi utilizado o método de sobreposição de Procrustes com as coordenadas dos marcos para diferentes vistas foram feitas análises de componentes principais (PCA), análises de variáveis canônicas (CVA), análise das distâncias de Procrustes entre as formas médias. Os resultados confirmam a presença de diferentes blocos populacionais com formas específicas para cada um deles, variação da forma em relação aos diferentes números cromossômicos e presença de dimorfismo sexual para a espécie, sendo este mais intenso no bloco B. Estes dados nos levam a sugerir que existe associação entre variação cromossômica e variação morfológica.This study examine the shape variation in the skull of Ctenomys lami, in relation to males and females, different population blocks, different diploid numbers, karyotypes and in relation to chromosomal pairs 1 and 2. We used 90 skulls of adults specimens (36 males and 54 females), all of them deriving from Coxilha das Lombas, Rio Grande do Sul, Brazil. This species have seven different cytotypes distributed in four population blocks, block A with 2n = 54, 55a and 56a; block B with 2n = 57 and 58; block C with 2n = 54 and 55a; and block D with 2n = 55b and 56b. The morphometric variation was studied in the dorsal, ventral and lateral views of the skull. We established landmarks in each view. Landmarks were located with TPSDig software, 14 in the dorsal view, 14 in the ventral view and 15 for lateral view. For the geometric morphometric analysis we used Procrustes superimposition method based in the coordinates of landmarks for different views. We made Principal Component Analysis (PCA), Canonical Variate Analysis (CVA), comparisons between distances between means shape. The results confirm the presence of different population blocks with specific shapes for each one, shape variation in relation to different chromosome number and existence of sexual dimorphism for this species, been more intense in block B. This data led us to suggest the existence of association between chromosome variation and morphological variation

Phenotypic plasticity in Heliconius erato (Lepidoptera: Nymphalidae) mandibles induced by different host plants (Passifloraceae).

This study aimed to test whether the consumption of different host plants influences the size and shape of mandibles and associated muscles in oligophagic insects throughout ontogeny. Larvae of Heliconius erato phyllis (Lepidoptera: Nymphalidae) were reared on four species of passion vines (Passifloraceae) and had the size of their head caspules, mandibles and associated adductor muscles measured afterward. Corresponding effects on shape and potential allometries in mandibles were also evaluted using geometric morphometric analysis based on scanning electron micrographs. Growth in mandibular size was greater compared to that of associated adductor muscles. There was no host plant effect on the sizes of either mandibles or adductor muscles. However, the shapes of mandibles was significantly different among passion vine species used as larval food. Greater differences in shape associated with wider teeth were obtained for Passiflora suberosa, whose leaves are tougher compared to those of P. misera, P. caerulea, and P. actinia; and, among which there was no difference in manbilular shape. Thus, it was inferred that the larval stage of this Heliconius species is able to adjust its mandible shape according to the toughness of host plants; a phenotypic plasticity mechanism that can be supposed to have evolved in order to facilitate processing these kinds of leaves

Geographic variation in skull shape of the water rat Scapteromys tumidus (Cricetidae, Sigmodontinae): isolation-by-distance plus environmental and geographic barrier effects?

ABSTRACT The geographic variation in skull size and shape of the swamp rat Scapteromys tumidus was examined in samples from eight geographic clusters in almost of its distribution in southern Brazil and Uruguay. For analysis we used two-dimensional geometric morphometric methods for dorsal, ventral and lateral views of the skull. The geometric descriptors showed no significant differences in skull size between geographic clusters, while differences in shape were highly significant. We found a significant and moderate correlation between geographic and morphological distances, corroborating the isolation-by-distance model. Samples from the Rio Grande do Sul central coastal plain were the most differentiated, segregating completely from all other samples in canonical variate analysis for the dorsal view. The most visible variable regions in skull were the zygomatic arch (mainly the squamosal root of zygomatic) and the lateral braincase borders. Once correlation between geographic and morphological distances were not strong, it is possible that other factors (environmental heterogeneity and/or geographic barriers) may are acting in S. tumidus skull differentiation

Plasticidad fenotípica en mandíbulas de Heliconius erato (Lepidoptera: Nymphalidae) inducida por diferentes plantas hospederas (Passifloraceae)

This study aimed to test whether the consumption of different host plants influences the size and shape of mandibles and associated muscles in oligophagic insects throughout ontogeny. Larvae of Heliconius erato phyllis (Lepidoptera: Nymphalidae) were reared on four species of passion vines (Passifloraceae) and had the size of their head caspules, mandibles and associated adductor muscles measured afterward. Corresponding effects on shape and potential allometries in mandibles were also evaluted using geometric morphometric analysis based on scanning electron micrographs. Growth in mandibular size was greater compared to that of associated adductor muscles. There was no host plant effect on the sizes of either mandibles or adductor muscles. However, the shapes of mandibles was significantly different among passion vine species used as larval food. Greater differences in shape associated with wider teeth were obtained for Passiflora suberosa, whose leaves are tougher compared to those of P. misera, P. caerulea, and P. actinia; and, among which there was no difference in manbilular shape. Thus, it was inferred that the larval stage of this Heliconius species is able to adjust its mandible shape according to the toughness of host plants; a phenotypic plasticity mechanism that can be supposed to have evolved in order to facilitate processing these kinds of leaves.El objetivo fue analizar si el consumo de diferentes plantas influenciaba el tamaño/forma de las mandíbulas de larvas de lepidópteros oligófagas y si dicho consumo ocasionaba tasas de crecimiento diferenciadas entre el músculo aductor mandibular y las mandíbulas durante su ontogénesis. Para esto se usaron larvas de Heliconius erato criadas en cuatro plantas hospederas naturales, a las que se les tomaron medidas de longitud de cápsula cefálica, tamaño mandibular y área del músculo aductor. Para detectar eventuales diferencias de forma mandibular se empleó microscopía electrónica y morfometria geométrica, y para posibles cambios alométricos se realizaron comparaciones entre curvas de crecimiento mandíbula vs. músculo aductor de los individuos criados en las cuatro plantas hospederas. Se encontró que no existen diferencias de tamaño entre las mandíbulas ni en músculos aductores de individuos criados en las diferentes plantas pero la forma mandibular fue significativamente diferente, mostrando una relación directa con la planta hospedera usada como alimento. Generalmente, la mandíbula creció proporcionalmente más que el músculo aductor, cuyo crecimiento fue estable. El cambio más significativo fue el alargamiento de los dientes incisivos en mandíbulas de larvas criadas con Passiflora suberosa. Se sugiere  que esto se debe  a que las hojas de esta especie poseen mayor dureza comparadas con las otras y que el desarrollar dientes largos facilitaría su corte y consumo

Skull shape and size variation within and between mendocinus and torquatus groups in the genus Ctenomys (Rodentia: Ctenomyidae) in chromosomal polymorphism context

Abstract We tested the association between chromosomal polymorphism and skull shape and size variation in two groups of the subterranean rodent Ctenomys. The hypothesis is based on the premise that chromosomal rearrangements in small populations, as it occurs in Ctenomys, produce reproductive isolation and allow the independent diversification of populations. The mendocinus group has species with low chromosomal diploid number variation (2n=46-48), while species from the torquatus group have a higher karyotype variation (2n=42-70). We analyzed the shape and size variation of skull and mandible by a geometric morphometric approach, with univariate and multivariate statistical analysis in 12 species from mendocinus and torquatus groups of the genus Ctenomys. We used 763 adult skulls in dorsal, ventral, and lateral views, and 515 mandibles in lateral view and 93 landmarks in four views. Although we expected more phenotypic variation in the torquatus than the mendocinus group, our results rejected the hypothesis of an association between chromosomal polymorphism and skull shape and size variation. Moreover, the torquatus group did not show more variation than mendocinus. Habitat heterogeneity associated to biomechanical constraints and other factors like geography, phylogeny, and demography, may affect skull morphological evolution in Ctenomys