Mid-domain Model Research Articles

Amphibian community structure along elevation gradients in eastern Nepal Himalaya

BackgroundSpecies richness and composition pattern of amphibians along elevation gradients in eastern Nepal Himalaya are rarely investigated. This is a first ever study in the Himalayan elevation gradient, the world’s highest mountain range and are highly sensitive to the effects of recent global changes. The aim of the present study was to assess amphibian community structure along elevation gradients and identify the potential drivers that regulate community structures. Amphibian assemblages were sampled within 3 months in both 2014 and 2015 (from May to July) using nocturnal time constrained and acoustic aids visual encounter surveys. In total, 79 transects between 78 and 4200 m asl were sampled within 2 years field work. A combination of polynomial regression, generalized linear models, hierarchical partitioning and canonical correspondence analysis were used to determine the effects of elevation and environmental variables on species richness, abundance, and composition of amphibian communities.ResultsSpecies richness and abundance declined linearly with increasing elevation, which did not support the Mid-Domain Model. Among all the environmental variables, elevation, surface area and humidity were the best predictors of species richness, abundance and composition of amphibians. The majority of amphibian species had narrow elevation ranges. There was no significant correlation between species range size and elevation gradients. However, body size significantly increased along elevation gradients, indicating that Bergmann’s rule is valid for amphibians in eastern Nepal Himalaya.ConclusionsThis study indicates that eastern Nepal Himalaya is a hotspot in amphibian diversity, and it should be served as a baseline for management and conservation activities.

Effects of neutrality, geometric constraints, climate, and habitat quality on species richness and composition of Atlantic Forest small-mammals

Aim To compare the fit of models of climate, habitat quality, neutral processes, and geometric constraints to species richness and composition of small mammal assemblages. Location The South American Atlantic Forest biome. Methods Using neutral models and mid-domain effect models, we simulated species spread in a spatially explicit array of grid cells representing the Atlantic Forest domain. We compared empirical patterns of species richness and composition with predictions of the neutral and mid-domain effect models. We also modeled individual species responses to climatic conditions and forest integrity, a measure of habitat quality. Results Habitat quality was the single best predictor of local species richness (α-diversity), but was a poor predictor of local species composition and of the decay in species similarity with distance (β-diversity). The neutral and mid-domain models generated very similar predictions, and were better predictors of species composition than of species richness. Climate variables were also strongly associated with overall species composition, but not with species richness. Main Conclusions The species richness of small-mammal assemblages in the Atlantic Forest is best explained by variation in habitat quality. In contrast, the composition of small-mammal assemblages is best explained by models of limited dispersal (neutral and mid-domain) and effects of climate on local species composition. Collectively, these results suggest that regional patterns of species richness may be uncoupled from patterns of species composition. Both species richness and composition should be considered when evaluating the predictions of neutral and mid-domain effect models, and of correlations of community structure with climatic or habitat variables.

Climatic and geometric constraints as driving factors of butterfly species richness along a Neotropical elevational gradient

We tested the effects of temperature, humidity and geographical constraints upon butterfly species richness along an elevational gradient covering an altitude ranging from 117 to 3,104 m above sea level (m. a.s.l.), in Southern Mexico. Ten transect sites were sampled 219 times from May 2010 to May 2011, along the elevational gradient to estimate range and population abundance of butterfly species. The effects of temperature, humidity and geometric constraints (mid-domain effects) on species richness along the study gradient were assessed using ordinary least squares regression. A total of 7,005 specimens representing 193 species were recorded. Species richness was relatively higher at elevations between 117 and 1,000 m. a.s.l. with an observed decline in richness values as elevation increased. Butterfly species richness along the study environmental gradient was predominantly determined by climatic constraints, rather than geometric constraints—a mid-domain model fit well only for large-ranged Pieridae species. Temperature and humidity explained the variation species richness for the entire butterfly community and for the three families evaluated; however the effect of predictor variables varied according to the measure of species richness and taxonomic family. This discrepancy in the response of butterfly richness to temperature, humidity and geometric constraints emphasizes the need to evaluate the response of different taxa to elevational gradients, to establish general patterns that help us to prioritize conservation measures that reduce population declines and local extinctions predicted by climate change in highly diverse tropical mountain ecosystems.

Geographical patterns in species richness of the benthic polychaetes in the continental shelf of the Gulf of California, Mexican Pacific

The present study is the first attempt to describe meso-scale patterns in the species richness of polychaetes along the Gulf of California, which stretches from about 23°N to 31°N. We examine herein the spatial changes in species distribution and explore the overlapping of species’ ranges towards the centre of the Gulf, to test whether the mid-domain effect (MDE) could explain an expected mid-domain peak in species richness. The faunal composition and the latitudinal range of 244 species of polychaetes recorded along the continental shelf of the Gulf of California were analysed in latitude bands of 1°. The species composition changes around the Gulf’s archipelago (~29°N), and the highest values of species richness are found at the 25° (197 species) and 26° (193 species) of latitude. Although the species richness pattern could be described by a parabolic shape, the regional trend was not strongly consistent with the peak of diversity at 27°N (176–191 species) predicted by the mid-domain effect: the random sorting of species’ ranges within spatial domain does not explain satisfactorily the geographical patterns of diversity. Nevertheless, a partial contribution of MDE to these natural patterns of diversity could be detected, and the increase in species richness towards middle latitudes was basically determined by species with distribution ranges larger than 6°. The low level of significance between the empirical species richness pattern and the mid-domain model prediction for polychaetes in the Gulf does not restrict their use as a model for exploring the randomness of the diversity patterns.

A Phenological Mid-Domain Analysis of Non-Native and Native Species Recruitment Richness

A mid-domain model was used to examine differences in patterns of species recruitment diversity between native and non-native species groups within shallow sub-tidal epifaunal communities of southern New England. A mid-domain model is a stochastic null model that predicts species richness patterns within a bounded domain. Deviations from the mid-domain model were examined to assess: 1) if each species group’s recruitment diversity differed from the mid-domain model; and 2) if there were differences in the patterns of deviation between groups. In general, the shape of mid-domain null was correlated to empirical patterns of recruitment diversity (i.e., mid-summer peaks in recruitment diversity) although among-year variation occurred in both the pattern of recruitment diversity and the level of fit. Akiake’s information criterion was used to determine if the mid-domain effect contributed to linear regression model predictions of phenological species recruitment patterns. We found distinct differences between the level of fit between native and non-native species groups. Native species did not significantly differ from the mid-domain null in five of seven years examined, while non-native species were significantly different in all years but two. Non-native species also showed a much weaker correlation to the mid-domain model and the seasonal water temperature signal. Collectively, this evidence supports the hypothesis that native and non-native species have different factors controlling the timing of their recruitment. Mid-domain nulls proved to be useful in the analysis of species richness patterns along temporal axes and provide a valuable baseline for further analysis.

One, two and three-dimensional geometric constraints and climatic correlates of North American tree species richness

The 'mid-domain effect' (MDE) has received much attention recently as a candidate explanation for patterns in species richness over large geographic areas. Mid-domain models generate a central peak in richness when species ranges are randomly placed within a bounded geographic area (i.e. the domain). The most common terrestrial mid-domain models published to date have been 1-D latitude or elevation models and 2-D latitude-longitude models. Here, we test 1-D, 2-D and 3-D mid-domain models incorporating latitude, longitude and elevation, and assess independent and concurrent effects of geometric constraints and climatic variables on species richness of North American trees. We use both the traditional 'global' regression models as well as geographically weighted regressions ('local' models) to examine local variation in the contribution of MDE and climatic variables to species richness across the domain. Our results show that in some dimensions the contribution of MDE to patterns of species richness can be quite substantial, and we show that in most cases a combination of MDE and climate predicted empirical species richness best in both local and global models. For the North American domain, MDE in the elevation dimension is clearly important in describing patterns of empirical species richness. We also show that the assumption of stationarity in global models is not met in the North American domain and that results of these models mask complex patterns in both the effect of MDE on richness and the response of species richness to climate. In particular we show the increased explanatory role of MDE in predicting species richness as domain edges are approached. Our results support the hypothesis that geometric constraints contribute to species richness patterns and we suggest the mid-domain effect should be considered alongside more traditional environmental correlates in understanding patterns of species diversity.

Fish diversity patterns in the Mediterranean Sea: deviations from a mid-domain model

Many ecological drivers have been proposed to explain the spatial distribution of species richness (productivity, energy, climate), but historical and paleogeographical contingencies may also be important. All these hypotheses make the implicit assumption that species richness would be similar everywhere in absence of any direct environmental variations; this is known as the mid-domain effect (MDE). The Mediterranean Sea is an 'archetypal situation' where we would expect this effect, so we used the MDE as a null model for fish richness and analyzed the deviations. Temperature, primary production and distance from both the Gibraltar Strait and Suez Canal have been included in our models as predictors. To obtain our objective we (1) built a database containing the geographic distributions of 619 Mediterranean fishes, (2) implemented 1- and 2-dimensional MDE models, and (3) carried out spatial autoregressive models on MDE deviations. It appeared that in 1 dimension, the only true MDE was displayed longitudinally by the large-ranged endemic species. In 2 dimensions, the MDE accounted for 21% of variation in fish richness distribution on the continental shelf. However, we showed that neglecting the MDE when relating fish richness to envi- ronmental predictors is misleading. In fact, minimum temperature and primary production negatively influenced fish species richness per se, but had a positive effect on deviations from MDE predictions. Finally, our study suggested that by analyzing the deviations from MDE predictions and taking into account spatial autocorrelation we developed a more rigorous assessment of the factors influencing diversity patterns.

Local and regional species diversity of benthic Isopoda (Crustacea) in the deep Gulf of Mexico

Abstract Recent studies of deep-sea faunas considered the influence of mid-domain models in the distribution of species diversity and richness with depth. In this paper, I show that separating local diversity from regional species richness in benthic isopods clarifies mid-domain effects in the distribution of isopods in the Gulf of Mexico. Deviations from the randomised implied species ranges can be informative to understanding general patterns within the Gulf of Mexico. The isopods from the GoMB study contained 135 species, with a total of 156 species including those from an earlier study. More than 60 species may be new to science. Most families of deep-sea isopods (suborder Asellota) were present, although some were extremely rare. The isopod family Desmosomatidae dominated the samples, and one species of Macrostylis (Macrostylidae) was found in many samples. Species richness for samples pooled within sites ranged from 1 to 52 species. Because species in pooled samples were highly correlated with individuals, species diversity was compared across sites using the expected species estimator ( n =15 individuals, ES 15 ). Six depth transects had idiosyncratic patterns of ES 15 , and transects with the greatest short-range variation in topography, such as basins and canyons, had the greatest short-range disparity. Basins on the deep slope did not have a consistent influence (i.e., relatively higher or lower than surrounding areas) on the comparative species diversity. ES 15 of all transects together showed a weak mid-domain effect, peaking around 1200–1500 m, with low values at the shallowest and deepest samples (Sigsbee Abyssal Plain); no longitudinal (east–west) pattern was found. The regional species pool was analyzed by summing the implied ranges of all species. The species ranges in aggregate did not have significant patterns across longitudes, and many species had broad depth ranges, suggesting that the isopod fauna of the Gulf of Mexico is well dispersed. The summed ranges, as expected, had strong mid-domain patterns, contrasting with the local species richness estimates. The longitudinal ranges closely matched a randomized pattern (species ranges placed randomly, 1000 iterations), with significant deviations in the east attributable to lower sampling effort. The depth pattern, however, deviated from the mid-domain model, with a bimodal peak displaced nearly 500 m shallower than the mode of the randomized distribution. The deviations from random expectation were significantly positive above 1600 m and negative below 2000 m, with the result that regional species richness peaked between 800 and 1200 m, and decreased rapidly at deeper depths. The highest species richness intervals corresponded to the number of individuals collected. Residuals from a regression of the deviations on individual numbers, however, still deviated from the randomized pattern. In this declining depth-diversity pattern, the Gulf of Mexico resembles other partially enclosed basins, such as the Norwegian Sea, known to have suffered geologically recent extinction events. This displaced diversity pattern and broad depth ranges implicate ongoing re-colonization of the deeper parts of the Gulf of Mexico. The Sigsbee Abyssal Plain sites could be depauperate for historical reasons (e.g., one or more extinction events) rather than ongoing ecological reasons (e.g., low food supply).

Three-dimensional mid-domain predictions: geometric constraints in North American amphibian, bird, mammal and tree species richness patterns

The effect (MDE) has received much attention as a candidate explanation for patterns in species richness over large geographic areas. Mid-domain models generate a central peak in richness when species ranges are placed randomly within a bounded geographic area (i.e. the domain). Until now, domain limits have been described mostly in one-dimension, usually latitude or elevation, and only occasionally in two-dimensions. Here we test 1-D, 2-D and, for the first time, 3-D mid-domain models and assess the effects of geometric constraints on species richness in North American amphibian, bird, mammal and tree species. Using spatially lagged simultaneous autoregressive models, empirical richness was predicted quite well by the mid-domain predictions and the spatial autoregressive term (45–92% R2). However, our results show that empirical species richness peaks do deviate from those of the MDE predictions in 3 dimensions. Variation explained (R2) by MDE predictions generally increased with increasing mean range size of the different biotic groups (from amphibian, to tree, mammal and finally bird data), and decreased with increasing dimensions being accounted for in the models. The results suggest geometric constraints alone can explain much of the variation in species richness with elevation, specifically with respect to the larger-range taxa, birds and mammals. Our analysis addresses many of the recent methodological criticisms directed at studies testing the MDE, and our results support the hypothesis that species diversity patterns are influenced by geometric constraints.

TESTS OF THE MID-DOMAIN HYPOTHESIS: A REVIEW OF THE EVIDENCE

Geographic variation of species richness is strongly correlated with environmental gradients. However, random arrangement of species distributions within a bounded domain can also theoretically produce richness gradients without underlying environmental gradients. This mid-domain effect (MDE) could serve as the null hypothesis against which to test effects of environmental variables, or as a component of a multivariate explanation of species-richness patterns. Recent reviews have concluded that there is a substantial MDE signature in observed geographical patterns of richness, based on correlations between observed patterns of richness and the predictions of mid-domain models. However, the mid-domain hypothesis makes additional powerful predictions about how richness should vary through space, and about the slope of the relationship between predicted and observed richness. Very few studies have tested these more powerful MDE predictions. Here, we reexamine the published mid-domain literature for agreement between observed patterns of richness and MDE predictions. We find that 50 of 53 published studies of MDEs showed significant deviations from the predictions of mid-domain models. When observed richness is correlated with MDE predictions, there are nearly always strongly collinear environmental gradients (e.g., in the Americas, climatic favorability and MDE-predicted richness are both maximal in the middle). Interpolation in sparsely sampled data can also give rise to spurious, apparently strong, mid-domain effects (e.g., the classic study of the Madagascan rain forest). We conclude that observed broad-scale patterns of species richness are not consistent with the mid-domain hypothesis.

The missing Madagascan mid‐domain effect

Species richness varies enormously across geographical gradients, a well-known phenomenon for which there are many hypothesized explanations. One recent hypothesis uses null models to demonstrate that random re-distribution of species' ranges within a given domain leads to a 'mid-domain effect' (MDE): increasing species richness towards the centre of the area. Madagascar is especially well-suited for empirical evaluation of mid-domain models by virtue of its large endemic fauna and its clearly defined boundaries. Lees et al. [Biol. J. Linn. Soc.67 (1999) 529] observed patterns of species richness consistent with MDEs in the Madagascan rainforest (a slim, north-south belt). In this study, we test one-dimensional and two-dimensional mid-domain model predictions for the birds and mammals of the entire island of Madagascar. When only latitudinal extents of species' distribution are considered, patterns of richness in Madagascar show an MDE. However, this pattern disappears for both taxa after accounting for the tendency of latitudinal bands nearer the middle of the country to be larger. Two-dimensional mid-domain model predictions of species richness are qualitatively opposite to observed patterns. Instead, island-wide spatial gradients of species richness in Madagascar relate strongly to patterns of primary productivity and amount of remaining natural habitat. Earlier work that showed a mid-domain peak within the rainforest biome (effectively after controlling for climate and natural habitat) seems likely to have reflected methodological artefacts. The classic case in which MDEs should occur is, in fact, inconsistent with the mid-domain hypothesis.

ELEVATIONAL SPECIES RICHNESS PATTERNS EMERGE FROM MULTIPLE LOCAL MECHANISMS IN HIMALAYAN WOODY PLANTS

We compared woody plant species distributions across nested spatial scales (local scale to entire Western Himalaya) and explored landscape scale patterns in detail to obtain inferences about the elevational gradient in species richness. Distribution data were compiled for 1100 species in the Western Himalaya, and primary data, comprising 123 species and 47 000 individuals, were collected for a landscape. Correlates of diversity were examined for the five spatial scales, and for different biogeographic groups at the landscape scale. The results indicate multiple mechanisms both within and across scales. At the landscape scale, though the mechanisms explaining unimodal species richness patterns were hard to separate, the underlying correlates of biogeographic groups were more distinct; temperate species richness followed mid-domain model predictions, and showed a nonlinear relationship with temperature, whereas tropical species richness tracked temperature and area. Simulations demonstrated that models with varying assumptions, while resulting in monotonic, unimodal, or multimodal patterns at local scales, could all lead to unimodal patterns at regional scales when multiple local replicates are aggregated, with a peak in the major ecotone. The turnover or successive accumulation of marginal species in ecotones potentially explains the mid-elevational peak in this zone. Landscape scale primary data on distribution and abundance could therefore be critical to understanding key aspects of macroecological patterns.

Range size in mid-domain models of species diversity

Geographical patterns of species diversity have been examined using mid-domain null models, in which the ranges of individual species are simulated by randomly arranging them on a bounded one- or two-dimensional continent. These models have shown that structured patterns in the geographical distribution of biodiversity can arise even under a fully stochastic procedure. In particular, mid-domain models have demonstrated that the random generation of ranges of different sizes and locations can produce a gradient of species diversity similar to the one found in real assemblages, with a peak at the middle of a continent. A less explored feature of mid-domain models is the pattern of range-size frequency distribution. Numerical simulations have provided some insights about the geographic pattern of average range size, but no exploration of the shape of range-size frequency distributions has been carried out. Here I present analytical and numerical models that generate explicit predictions for patterns of range size under the assumptions of mid-domain models of species diversity. Some generalizations include: (1) Mid-domain models predict no geographic gradient of average range size; the mean range size of species occurring at any point on a continent is constant (0.5 of the extent of the continent in the one-dimensional model, 0.25 of the area of the continent in the two-dimensional case); (2) Variance in range size is lowest at the middle of a continent and highest near the corners of a square-shaped continent; (3) The range-size frequency distribution is highly right-skewed at any point of a continent, but the skewness is highest near the corners. Despite their alleged weaknesses, mid-domain models are adequate null models against which real-world patterns can be contrasted.

INDO-PACIFIC BIODIVERSITY OF CORAL REEFS: DEVIATIONS FROM A MID-DOMAIN MODEL

Understanding the nature and causes of global gradients in species richness is a perennial ecological problem, and recent work has highlighted the need to assess these gradients relative to an appropriate statistical expectation. This paper examines latitudinal and longitudinal gradients in species richnesses of corals and reef fishes in the Indo-Pacific domain and compares them with gradients predicted by a mid-domain model in which geographic domains are located at random between the latitudinal and longitudinal boundaries of this region. We test for significant differences between observed and predicted species-richness patterns, and we identify regions that are enriched or depauperate in species, relative to expectation. In addition, we move beyond previous mid-domain analyses by directly comparing observed spatial distributions of geographic ranges with those predicted by a mid-domain model. This comparison indicates precisely how species-richness anomalies are produced by nonrandomness in the distribution of species ranges. For both corals and fishes, large and statistically significant differences exist between observed latitudinal and longitudinal species-richness gradients and those predicted by mid-domain models. Longitudinally, species richness is markedly higher than predicted along the African coast and, to a lesser extent, within the Indo-Australian Archipelago (IAA), and it is markedly lower than expected in the eastern Pacific. Latitudinally, species richness becomes increasingly higher than predicted as one moves from the equator to the tropical margins; then it becomes sharply lower than predicted beyond the tropics. Unexpectedly, differences between observed and predicted spatial distributions of range endpoints and midpoints reveal a pattern of nonrandomness that is highly congruent with the hypothesis that gyres in the Indian and Pacific Oceans, with the IAA forming a porous boundary between them, have a major influence on Indo-Pacific species-richness patterns. Our analyses indicate that the perspective offered by a focus on explaining nonrandomness in the location of geographic ranges (rather than explaining why species numbers vary in space) is likely to dramatically alter our assessments of alternative explanations for global species-richness gradients.

Mid-domain models of species richness gradients: assumptions, methods and evidence.

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.

Latitudinal gradients in diversity: real patterns and random models

Mid-domain models have been argued to provide a default explanation for the best known spatial pattern in biodiversity, namely the latitudinal gradient in species richness. These models assume no environmental gradients, but merely a random latitudinal association between the size and placement of the geographic ranges of species. A mid-domain peak in richness is generated because when the latitudinal extents of species in a given taxonomic group are bounded to north and south, perhaps by a physical constraint such as a continental edge or perhaps by a climatic constraint such as a critical temperature or precipitation threshold, then the number of ways in which ranges can be distributed changes systematically between the bounds. In addition, such models make predictions about latitudinal variation in the latitudinal extents of the distributions of species, and in beta diversity (the spatial turnover in species identities). Here we test how well five mid-domain models predict observed latitudinal patterns of species richness, latitudinal extent and beta diversity in two groups of birds, parrots and woodpeckers, across the New World. Whilst both groups exhibit clear gradients in richness and beta diversity and the general trend in species richness is acceptably predicted (but not accurately, unless substantial empirical information is assumed), the fit of these models is uniformly poor for beta diversity and latitudinal range extent. This suggests either that, at least for these data, as presently formulated mid-domain models are too simplistic, or that in practice the mid-domain effect is not significant in determining geographical variation in diversity.

Overlap, Regularity, and Flowering Phenologies

Previous articleNext article No AccessNotes and CommentsOverlap, Regularity, and Flowering PhenologiesBlaine J. ColeBlaine J. Cole Search for more articles by this author PDFPDF PLUS Add to favoritesDownload CitationTrack CitationsPermissionsReprints Share onFacebookTwitterLinkedInRedditEmail SectionsMoreDetailsFiguresReferencesCited by The American Naturalist Volume 117, Number 6Jun., 1981 Published for The American Society of Naturalists Article DOIhttps://doi.org/10.1086/283784 Views: 7Total views on this site Citations: 31Citations are reported from Crossref Copyright 1981 The University of ChicagoPDF download Crossref reports the following articles citing this article:Susanne S. Renner Placing the Time of Leaf Emergence in an Evolutionary Context, The American Naturalist 189, no.22 (Dec 2016): ii–iv.https://doi.org/10.1086/690020Ingi Agnarsson, Nicholas J. 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