RESUMO

Abstract: The Mid-Domain Effect (MDE) and the Rapoport (RE) effect are two biogeographical theories that make predictions about biogeogaphic patterns. MDE predicts higher richness in the central portions of a gradient if it is within a bounded domain. RE predicts a positive relation between altitude and species range size along an altitudinal gradient. Our aim was to document the distribution of spider species richness along an altitudinal gradient in the Brazilian Amazon, and to test the influence of MDE and RE on the diversity patterns. Our study was conducted at the Pico da Neblina (Amazonas state, Brazil), and we sampled spiders at six different altitudes using two methods: nocturnal hand sampling and a beating tray. We obtained 3,140 adult spiders from 39 families, sorted to 529 species/morphospecies. Richness declined continuously with an altitude increase, but the fit with the MDE richness estimates was very weak and was not significant. Range size was not related to altitude, i. e., no RE. Finally, the abundance distribution within each species range varied more specifically, which prevented the occurrence of a RE at the community level. The influence of MDE was extremely low, a consequence of our community characteristics, formed mostly by small range size species. Short and medium range species were located at all altitudes, preventing a significant relation between range size and altitude. The distribution of abundance within a species range varied specifically and do not support a RE hypothesis.

Resumo: O Efeito do Domínio Central (MDE) e o Efeito Rapoport (ER) são duas teorias biogeografias que fazem previsões sobre a distribuição da diversidade ao longo de gradientes. O MDE prevê maior riqueza nas porções centrais de um gradiente, se este estiver dentro de um domínio fechado. O ER prevê uma relação positiva entre altitude e tamanho da distribuição ao longo do gradiente altitudinal. Nosso objetivo foi o de registrar a distribuição de uma comunidade de aranhas ao longo de um gradiente altitudinal na Amazônia Brasileira, e testar se há uma influência do EDC e do ER sobre os padrões de diversidade da comunidade. Nosso estudo foi feito no Parque Nacional do Pico da Neblina (AM, Brasil), e nós amostramos aranhas em seis altitudes diferentes. Nós coletamos 3.140 exemplares adultos de 39 famílias, que foram divididos em 529 espécies/morfoespécies. A riqueza declinou com o aumento de altitude, mas o padrão não mostrou ajuste com as previsões feitas pelo EDC. O tamanho da distribuição altitudinal também não esteve relacionado ao previsto pelo ER. Por fim, a distribuição de abundância ao longo da distribuição altitudinal das espécies variou de maneira específica, o que impediu a ocorrência de um ER nos padrões da comunidade. A influência do EDC sobre os padrões observados foi baixíssima, uma consequência de características de nossa comunidade, já que esta é formada por espécies com pequena distribuição altitudinal. Espécies de distribuição altitudinal médias e grandes ocorreram em todas as partes do gradiente o que impediu a ocorrência de um ER. Por fim, o ER também não foi observado na distribuição de abundância das espécies ao longo do gradiente, já que essa variou de maneira específica.

RESUMO

The Mid-Domain Effect (MDE) and the Rapoport (RE) effect are two biogeographical theories that make predictions about biogeogaphic patterns. MDE predicts higher richness in the central portions of a gradient if it is within a bounded domain. RE predicts a positive relation between altitude and species range size along an altitudinal gradient. Our aim was to document the distribution of spider species richness along an altitudinal gradient in the Brazilian Amazon, and to test the influence of MDE and RE on the diversity patterns. Our study was conducted at the Pico da Neblina (Amazonas state, Brazil), and we sampled spiders at six different altitudes using two methods: nocturnal hand sampling and a beating tray. We obtained 3,140 adult spiders from 39 families, sorted to 529 species/morphospecies. Richness declined continuously with an altitude increase, but the fit with the MDE richness estimates was very weak and was not significant. Range size was not related to altitude, i. e., no RE. Finally, the abundance distribution within each species range varied more specifically, which prevented the occurrence of a RE at the community level. The influence of MDE was extremely low, a consequence of our community characteristics, formed mostly by small range size species. Short and medium range species were located at all altitudes, preventing a significant relation between range size and altitude. The distribution of abundance within a species range varied specifically and do not support a RE hypothesis.

O Efeito do Domínio Central (MDE) e o Efeito Rapoport (ER) são duas teorias biogeografias que fazem previsões sobre a distribuição da diversidade ao longo de gradientes. O MDE prevê maior riqueza nas porções centrais de um gradiente, se este estiver dentro de um domínio fechado. O ER prevê uma relação positiva entre altitude e tamanho da distribuição ao longo do gradiente altitudinal. Nosso objetivo foi o de registrar a distribuição de uma comunidade de aranhas ao longo de um gradiente altitudinal na Amazônia Brasileira, e testar se há uma influência do EDC e do ER sobre os padrões de diversidade da comunidade. Nosso estudo foi feito no Parque Nacional do Pico da Neblina (AM, Brasil), e nós amostramos aranhas em seis altitudes diferentes. Nós coletamos 3.140 exemplares adultos de 39 famílias, que foram divididos em 529 espécies/morfoespécies. A riqueza declinou com o aumento de altitude, mas o padrão não mostrou ajuste com as previsões feitas pelo EDC. O tamanho da distribuição altitudinal também não esteve relacionado ao previsto pelo ER. Por fim, a distribuição de abundância ao longo da distribuição altitudinal das espécies variou de maneira específica, o que impediu a ocorrência de um ER nos padrões da comunidade. A influência do EDC sobre os padrões observados foi baixíssima, uma consequência de características de nossa comunidade, já que esta é formada por espécies com pequena distribuição altitudinal. Espécies de distribuição altitudinal médias e grandes ocorreram em todas as partes do gradiente o que impediu a ocorrência de um ER. Por fim, o ER também não foi observado na distribuição de abundância das espécies ao longo do gradiente, já que essa variou de maneira específica.

RESUMO

The Atlantic Forest is an excellent case study for the elevational diversity of birds, and some inventories along elevational gradients have been carried out in Brazil. Since none of these studies explain the patterns of species richness with elevation, we herein review all Brazilian studies on bird elevational diversity, and test a geometric constraint null model that predicts a unimodal species-altitude curve, the Mid-domain Effect (MDE). We searched for bird inventories in the literature and also analysed our own survey data using limited-radius point counts along an 800 m elevational gradient in the state of São Paulo, Brazil. We found 10 investigations of elevational diversity of Atlantic Forest birds and identified five different elevational patterns: monotonic decreasing diversity, constant at low elevations, constant at low elevations but increasing towards the middle, and two undescribed patterns for Atlantic Forest birds, trough-shaped and increasing diversity. The average MDE fit was low (r² = 0.31) and none of the MDE predictions were robust across all gradients. Those studies with good MDE model fits had obvious sampling bias. Although it has been proposed that the MDE may be positively associated with the elevational diversity of birds, it does not fit the Brazilian Atlantic Forest bird elevational diversity.

RESUMO

The Atlantic Forest is an excellent case study for the elevational diversity of birds, and some inventories along elevational gradients have been carried out in Brazil. Since none of these studies explain the patterns of species richness with elevation, we herein review all Brazilian studies on bird elevational diversity, and test a geometric constraint null model that predicts a unimodal species-altitude curve, the Mid-domain Effect (MDE). We searched for bird inventories in the literature and also analysed our own survey data using limited-radius point counts along an 800 m elevational gradient in the state of São Paulo, Brazil. We found 10 investigations of elevational diversity of Atlantic Forest birds and identified five different elevational patterns: monotonic decreasing diversity, constant at low elevations, constant at low elevations but increasing towards the middle, and two undescribed patterns for Atlantic Forest birds, trough-shaped and increasing diversity. The average MDE fit was low (r² = 0.31) and none of the MDE predictions were robust across all gradients. Those studies with good MDE model fits had obvious sampling bias. Although it has been proposed that the MDE may be positively associated with the elevational diversity of birds, it does not fit the Brazilian Atlantic Forest bird elevational diversity.

RESUMO

The Atlantic Forest is an excellent case study for the elevational diversity of birds, and some inventories along elevational gradients have been carried out in Brazil. Since none of these studies explain the patterns of species richness with elevation, we herein review all Brazilian studies on bird elevational diversity, and test a geometric constraint null model that predicts a unimodal species-altitude curve, the Mid-domain Effect (MDE). We searched for bird inventories in the literature and also analysed our own survey data using limited-radius point counts along an 800 m elevational gradient in the state of São Paulo, Brazil. We found 10 investigations of elevational diversity of Atlantic Forest birds and identified five different elevational patterns: monotonic decreasing diversity, constant at low elevations, constant at low elevations but increasing towards the middle, and two undescribed patterns for Atlantic Forest birds, trough-shaped and increasing diversity. The average MDE fit was low (r² = 0.31) and none of the MDE predictions were robust across all gradients. Those studies with good MDE model fits had obvious sampling bias. Although it has been proposed that the MDE may be positively associated with the elevational diversity of birds, it does not fit the Brazilian Atlantic Forest bird elevational diversity.

RESUMO

Patterns of species richness along latitudinal, elevational and depth gradients often exhibit a mid-gradient peak. Similar patterns with a mid-domain peak in richness are produced, as a result of geometric constraints on species distributions, by models that randomize species range size and placement over a bounded gradient. Proponents of these so-called mid-domain models argue that they provide an appropriate null hypothesis for examining species richness patterns along spatial gradients. Furthermore, some claim that because these models seem to explain a large proportion of the large-scale spatial variation in richness, geometric constraints on species distribution are in fact the cause of these patterns. A critical examination of model assumptions reveals that some are unrealistic, conceptually flawed or internally inconsistent. Additionally, tests of mid-domain models have suffered from methodological deficiencies derived from arbitrariness and circularity in the definition of domain boundaries, collapsing two-dimensional (2-D) patterns into a single dimension (1-D), and the use of interpolated ranges, all of which can bias test results in favour of the models. Tests have also been statistically naive by using fairly insensitive measures of deviation between observed and predicted patterns and ignoring the increased probability of Type I error that can result in analyses of spatially autocorrelated data. In spite of this, a review of the empirical evidence indicates that most studies do not show a high degree of concordance between observed and predicted species richness patterns, particularly in 2-D. Additionally, spatial patterns of variation in range size and species turnover do not unequivocally support mid-domain models. Thus, the models do not adequately describe observed species richness gradients and thus fail to explain them. Although mid-domain models have served a useful purpose in drawing attention to the need to clarify the null expectation in the study of species richness gradients, their use appears to be severely limited by difficulties associated with the treatment of ranges, boundary definitions and a lack of clarity regarding the extent to which the observed data should be used to generate the null patterns.