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.     

When does diversity fit null model predictions? Scale and range size mediate the mid‐domain effect

Aim  Recently, a flurry of studies have focused on the extent to which geographical patterns of diversity fit mid-domain effect (MDE) null models. While some studies find strong support for MDE null models, others find little. We test two hypotheses that might explain this variation among studies: small-ranged groups of species are less likely than large-ranged species to show mid-domain peaks in species richness, and mid-domain null model predictions are less robust for smaller spatial extents than for larger spatial extents.
Location  We analyse data sets from elevational, riverine, continental and other domains from around the world.
Methods  We use a combination of Spearman rank correlations and binomial tests to examine whether differences within and among studies and domains in the predictive power of MDE null models vary with spatial scale and range size.
Results  Small-ranged groups of species are less likely to fit mid-domain predictions than large-ranged groups of species. At large spatial extents, diversity patterns of taxonomic groups with large mean range sizes fit MDE null model predictions better than did diversity patterns of groups with small mean range sizes. MDE predictions were more explanatory at larger spatial extents than at smaller extents. Diversity patterns at smaller spatial extents fit MDE predictions poorly across all range sizes. Thus, MDE predictions should be expected to explain patterns of species richness when ranges and the scale of analysis are both large.
Main conclusions  Taken together, the support for these hypotheses offers a more sophisticated model of when MDE predictions should be expected to explain patterns of species richness, namely when ranges and the scale of analysis are both large. Thus the circumstances in which the MDE is important are finite and apparently predictable.     

The mid-domain effect in ectomycorrhizal fungi: range overlap along an elevation gradient on Mount Fuji,Japan

Mid-domain effect (MDE) models predict that the random placement of species'' ranges within a bounded geographical area leads to increased range overlap and species richness in the center of the bounded area. These models are frequently applied to study species-richness patterns of macroorganisms, but the MDE in relation to microorganisms is poorly understood. In this study, we examined the characteristics of the MDE in richness patterns of ectomycorrhizal (EM) fungi, an ecologically important group of soil symbionts. We conducted intensive soil sampling to investigate overlap among species ranges and the applicability of the MDE to EM fungi in four temperate forest stands along an elevation gradient on Mount Fuji, Japan. Molecular analyses using direct sequencing revealed 302 EM fungal species. Of 73 EM fungal species found in multiple stands, 72 inhabited a continuous range along the elevation gradient. The maximum overlap in species range and the highest species richness occurred at elevations in the middle of the gradient. The observed richness pattern also fit within the 95% confidence interval of the mid-domain null model, supporting the role of the MDE in EM fungal richness. Deviation in observed richness from the mean of the mid-domain null estimation was negatively correlated with some environmental factors, including precipitation and soil C/N, indicating that unexplained richness patterns could be driven by these environmental factors. Our results clearly support the existence of microbial species'' ranges along environmental gradients and the potential applicability of the MDE to better understand microbial diversity patterns.     

Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains

Aim  This research aims to understand the factors that shape elevational diversity gradients and how those factors vary with spatial grain. Specifically, we test the predictions of the species–productivity hypothesis, species–temperature hypothesis, the metabolic theory of ecology and the mid-domain effects null model. We also examine how the effects of productivity and temperature on richness depend on spatial grain.
Location  Deciduous forests along an elevational gradient in Great Smoky Mountains National Park, USA.
Methods  We sampled 22 leaf litter ant assemblages at three spatial grains, from 1-m² quadrats to 50 × 50 m plots using Winkler samplers.
Results  Across spatial grains, warmer sites had more species than did cooler sites, and primary productivity did not predict ant species richness. We found some support for the predictions of the metabolic theory of ecology, but no support for the mid-domain effects null model. Thus, our data are best explained by some version of a species–temperature hypothesis.
Main conclusions  Our results suggest that temperature indirectly affects ant species diversity across spatial grains, perhaps by limiting access to resources. Warmer sites support more species because they support more individuals, thereby reducing the probability of local extinction. Many of our results from this elevational gradient agree with studies at more global scales, suggesting that some mechanisms shaping ant diversity gradients are common across scales.     

The mid-domain effect applied to elevational gradients: species richness of small mammals in Costa Rica

Aim The objective of this study was to comprehensively document and examine the alpha and gamma patterns of species richness in non-volant, small mammals (rodents, shrews and mouse opossums) along a tropical elevational gradient. These data were used to determine the support for existing hypotheses of species richness encompassing mid-domain null models, as well as climatic, and community overlap hypotheses. Location Field studies were conducted along a Caribbean slope of the Río Peñas Blancas watershed in the north-eastern region of Costa Rica between 750 and 1850 m at 10 sampling sites. Methods Species richness and abundances of small mammals were surveyed for four seasons including three temporal replicates at each of five elevational sites: late wet season (2000), early wet season (2001), and dry season (2002), and one spatial replicate at five different sites within the same elevations during the late wet season (2001). Species richness at elevations below 700 m was compiled from specimen records from 23 US national and international collections. Predictions of a null model based solely on geometric constraints were examined using a Monte Carlo simulation program, Mid-Domain Null. Results In 16,900 trap nights, 1561 individuals from 16 species were captured. Both alpha and gamma species richness peaked at mid-elevation between 1000 and 1300 m, with richness declining both at higher and lower elevations. Most of the empirical curves of species richness occur within 95% prediction curves of the mid-domain model, although deviations from the null model exist. Regression of the empirical richness on the null model predictions explained nearly half of the variation observed (r² = 0.45, P = 0.002). Main conclusions The geometric constraints of montane topography appear to influence the diversity pattern of small mammals, although climatic conditions including an intermediate rainfall and temperature regime, and distance from the persistent cloud cap also are correlated with the pattern of species richness. The predictions of productivity, and community overlap hypotheses are not supported with the empirical data.     

Climate and history explain the species richness peak at mid-elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions

Aim  We studied elevational species richness patterns of Schizothorax fishes and identified the roles of ecological and evolutionary factors in shaping the patterns of elevational diversity.
Location  The Tibetan Plateau and its adjacent regions.
Methods  We assembled distribution and altitude data for all Schizothorax species using the literature. We merged ecological and evolutionary approaches to test the relationships between species richness and ecological factors (climate, area, the mid-domain effect) or evolutionary factors (diversification rates and time of colonization).
Results  We found that species richness of Schizothorax fishes peaked at mid-elevations. Rainfall, area, the mid-domain effect and diversification rate were weak predictors of the richness pattern. Temperature showed a nonlinear relationship with species richness. Temperature and time of colonization were the most important variables in explaining the elevational diversity pattern.
Main conclusion  Our findings indicate that the time-for-speciation effect and niche conservatism play important roles in variation of species richness.     

Process-based models of species distributions and the mid-domain effect

Null models that place species ranges at random within a bounded geographical domain produce hump-shaped species richness gradients (the "mid-domain effect," or MDE). However, there is debate about the extent to which these models are a suitable null expectation for effects of environmental gradients on species richness. Here, I present a process-based framework for modeling species distributions within a bounded geographical domain. Analysis of null models consistent with the mid-domain hypothesis shows that MDEs are indeed likely to be ubiquitous consequences of geographical domain boundaries. Comparing the probability distributions of range locations for the process-based and randomization-based models reveals that randomization models probably overestimate the contribution of MDEs to empirical patterns of species richness, but it also indicates that other testable predictions from randomization models are likely to be robust. I also show how this process-based framework can be extended beyond null models to incorporate effects of environmental gradients within the domain. This study provides a first step toward an ecological theory of species distributions in geographical space that can incorporate both "geometric constraints" and effects of environmental gradients, and it shows how such a theory can inform our understanding of species richness gradients in nature.     

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.     

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.     

Challenges in the application of geometric constraint models

Discerning the processes influencing geographical patterns of species richness remains one of the central goals of modern ecology. Traditional approaches to exploring these patterns have focused on environmental and ecological correlates of observed species richness. Recently, some have suggested these approaches suffer from the lack of an appropriate null model that accounts for species ranges being constrained to occur within a bounded domain. Proponents of these null geometric constraint models (GCMs), and the mid-domain effect these models produce, argue their utility in identifying meaningful gradients in species richness. This idea has generated substantial debate. Here we discuss what we believe are the three major challenges in the application of GCMs. First, we argue that there are actually two equally valid null models for the random placement of species ranges within a domain, one of which actually predicts a uniform distribution of species richness. Second, we highlight the numerous decisions that must be made to implement a GCM that lead to marked differences in the predictions of the null model. Finally, we discuss challenges in evaluating the importance of GCMs once they have been implemented.     

Latitudinal gradients in diversity: real patterns and random models

Mid-domain models have been argued lo 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.     

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

The "mid-domain 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.     

Using habitat models to map diversity: pan-African species richness of ticks (Acari: Ixodida)

Aims To show how logistic regression models for individual species can be used to produce improved estimates of species richness at a continental scale; to present these data for African ticks (Acari: Ixodida); and to address the question of whether there is a latitudinal gradient in tick species richness. Location Africa. Methods A database of 34,060 collection records for African ticks is used to produce a pan‐African map of known tick species richness at 0.25 × 0.25‐degree resolution. The likely distributions of seventy‐three species are then estimated from environmental factors using logistic regression, and localities where there is a suitably high probability of occurrence for a given species are added to the original data for that species. These augmented data are combined to produce a map of the predicted pan‐African distribution of tick species richness. The relationship of species richness to latitude is considered along a transect placed across some of the more extensively collected areas. Results Maps of known and predicted pan‐African tick species richness are presented, and deficiencies in the available data are highlighted. Correlations using both known and predicted estimates of tick species richness suggest that ticks follow similar species richness patterns to those described for African mammals and birds, with a latitudinal gradient and highest species richness in east equatorial Africa. Tick species ranges are log‐normally distributed. Main conclusions Carefully constructed probability surfaces offer a more powerful approach to mapping species ranges than simple presence‐absence maps. Such models are a useful extension to current biogeographical methods and have a wide range of potential applications in ecology, epidemiology and conservation. Tick species richness at a continental scale follows similar trends to those reported for mammals and birds.     

The mid-domain effect and species richness patterns:what have we learned so far?

If species' ranges are randomly shuffled within a bounded geographical domain free of environmental gradients, ranges overlap increasingly toward the center of the domain, creating a "mid-domain" peak of species richness. This "mid-domain effect" (MDE) has been controversial both in concept and in application. Empirical studies assess the degree to which the evolutionary, ecological, and historical processes that undeniably act on individual species and clades produce geographical patterns that resemble those produced by MDE models. MDE models that resample empirical range size frequency distributions (RSFDs) balance the risk of underestimating and overestimating the role of MDE, whereas theoretical RSFDs are generally biased toward underestimating MDE. We discuss the inclusion of nonendemic species in MDE models, rationales for setting domain limits, and the validity of one- and two-dimensional MDE models. MDE models, though null models, are not null hypotheses to be simplistically rejected or accepted. They are a means of estimating the expected effect of geometric constraints within the context of multiple causality. We call for assessment of MDE on an equal statistical footing with other candidate explanations for richness gradients. Although some critics have categorically dismissed MDE, an overview of the 21 MDE studies published to date reveals a substantial signature of MDE in natural patterns and justifies continued work.     

The mid-domain effect: geometric constraints on the geography of species richness

Geographic patterns of species richness are influenced by many factors, but the role of shared physiographical and physiological boundaries in relation to range-size distributions has been surprisingly neglected, in spite of the fact that such geometric constraints lead to mid-domain richness peaks even without environmental gradients (the mid-domain effect). Relying on null models, several recent studies have begun to quantify this problem using simulated and empirical data. This approach promises to transform how we perceive geographic variation in diversity, including the long unresolved latitudinal gradient in species richness. The question is not whether geometry affects such patterns, but by how much.     

Environmental and geometric drivers of small mammal diversity along elevational gradients in Utah

The mechanisms shaping patterns of biodiversity along spatial gradients remain poorly known and controversial. Hypotheses have emphasized the importance of both environmental and spatial factors. Much of the uncertainty about the relative role of these processes can be attributed to the limited number of comparative studies that evaluate multiple potential mechanisms. This study examines the relative importance of six variables: temperature, precipitation, productivity, habitat heterogeneity, area, and the mid-domain effect on patterns of species richness for non-volant small mammals along four neighboring mountain ranges in central Utah. Along each of these elevational gradients, a hump-shaped relationship of richness with elevation is evident. This study evaluates whether the processes shaping this common pattern are also common to all gradients. Model selection indicates that no one factor or set of factors best explains patterns of species richness across all gradients, and drivers of diversity may vary seasonally. These findings suggest that commonality in the pattern of species richness, even among elevational gradients with a similar biogeographic history and fauna, cannot be attributed to a simple universal explanation.     

The role of fire in terrestrial vertebrate richness patterns

Productivity is strongly associated with terrestrial species richness patterns, although the mechanisms underpinning such patterns have long been debated. Despite considerable consumption of primary productivity by fire, its influence on global diversity has received relatively little study. Here we examine the sensitivity of terrestrial vertebrate biodiversity (amphibians, birds and mammals) to fire, while accounting for other drivers. We analyse global data on terrestrial vertebrate richness, net primary productivity, fire occurrence (fraction of productivity consumed) and additional influences unrelated to productivity (i.e., historical phylogenetic and area effects) on species richness. For birds, fire is associated with higher diversity, rivalling the effects of productivity on richness, and for mammals, fire's positive association with diversity is even stronger than productivity; for amphibians, in contrast, there are few clear associations. Our findings suggest an underappreciated role for fire in the generation of animal species richness and the conservation of global biodiversity.     

A null model for species richness gradients: bounded range overlap of butterflies and other rainforest endemics in Madagascar

Species richness has classically been thought to increase from the poles towards the Equator, and from high elevations down to sea-level. However, the largest radiation of butterflies in Madagascar, the subtribe Mycalesina (c. 67 spp.) does not exhibit such a monotonic pattern, either for empirical records or for interpolated species ranges. Instead, summation of mycalesine ranges generates a domed curve of species richness values approximately symmetric around mid latitudes within the island, a pattern most smoothly exhibited by the wider ranging and better known species, and a less symmetric curve peaking near mid elevations. Hotspots for the summation of 1183 species ranges and seven out of the ten groups of insects and vertebrates analysed (butterflies, cicindelid and enariine melonthid beeties, ctenuchiine moths, chameleons, frogs, birds, lemurs, tenrecs, and rodents) also occur at both mid latitudes and elevations. The most strongly parabolic pattern is shown by animals (637 spp.) whose ranges are confined to the highly linear rainforest biome. This rainforest species richness curve is resilient in shape even after controlling for particular effects of area and irregular sample effort. In sharp contrast, at least eight different environmental parameters for the rainforest biome tend to increase monotonically towards the northern, more tropical, boundary, a trend evident only in species richness gradients of more narrow-ranging species. The one-dimensional latitudinal species richness curves and hotspots observed in fact best reflect overall the geometric predictions of a null model for ranked range size partitions of the regional species pool. This analytical model is based on the uniform probability distribution, and assumes that species ranges are constrained by the position of biome or island boundaries. The same logarithmic equations applied iteratively to longitude also accurately predict hotspots for more realistic species ranges containing gaps, as shown for two-dimensional species richness patterns for the Madagascan rainforest dataset. Bio-geographic and conservation implications of the bounded range overlap concept are discussed.     

Biogeographic patterns of the East African coastal forest vertebrate fauna

The archipelago-like coastal forest of East Africa is one of the highest priority ecosystems for biodiversity conservation worldwide. Here we investigate patterns of species richness and biogeographic distribution among birds, mammals and reptiles of these forests, using distribution data obtained from recently published reviews and information collated by the WWF Eastern Africa Coastal Forest Ecoregion Programme. Birds and mammals species were divided into forest specialists and generalists, and forest specialist reptiles into ‘coastal’ and ‘forest’ endemics. The species richness of birds and generalist mammals increased with area, and is probably a result of area-dependent extinction. Only in birds, however, species richness increased with decreasing isolation, suggesting possible isolation-dependent colonization. Forest diversity, associated to altitudinal range, is important for specialist birds and mammals, whose species richness increased with wider altitudinal range. The number of relict coastal endemic and forest endemic reptiles was higher in forests with wider altitudinal ranges and on relatively higher altitude, respectively. Such forests have probably provided a suitable (and perhaps stable) environment for these species through time, thus increasing their persistence. Parsimony analysis of distributions (PAD) and cluster analyses showed geographical distance and general ecological similarity among forests as a determinant factor in bird distribution patterns, with compositional similarity decreasing with increasing inter-forest distance. Compositional similarity patterns of mammals among the forests did not show a strong geographical correspondence or a significant correlation with inter-forest distance, and those of reptiles were not resolved, with very low similarity levels among forest faunas. Our results suggest that the relative importance (and causal relationship) of forest attributes affecting the distribution of the East African coastal forest vertebrate fauna varies depending on life history traits such as dispersal ability and forest specialization. The groupings in PAD are partly congruent with some of the previous classifications of areas of endemism for this region, supporting the ‘naturalness’ of these regions.     

Differentiation between populations of a termite in eastern Africa: implications for biogeography

Aim  African forests are divided by an arid corridor which runs from the Horn of Africa to the Namib Desert. Several forest species occur in the forests of eastern Africa as well as in the Guineo-Congolian forest block. We evaluate the possibility that such species may have crossed the arid corridor along a route through the Kenyan Highlands and down the eastern drainages during climatologically favourable periods in the past.
Locations  Eastern Africa, Ivory Coast.
Methods  We used the termite species Schedorhinotermes lamanianus (Sjöstedt). This species occurs in lowland forests and woodland throughout Africa south of the Sahara. We sampled termites from 12 populations. We evaluated the differentiation between populations using amplified fragment length polymorphisms as well as morphometrical measurements.
Results  Genetic and morphometrical analysis demonstrated substantial differentiation between populations west and east of the arid corridor in Kenya. To the east of this corridor we found an increase of morphological distance with geographical distance. Schedorhinotermes lamanianus occurs not only along the coast but also at isolated locations (e.g. ground-water forests in foothills) within the arid hinterland.
Main conclusions  We interpret these populations as remnants of a wider distribution during wet climatic periods. At these times, populations of S. lamanianus were apparently able to establish along extensive gallery forests protruding into the arid belt of the Kenyan hinterland. There have been no connections between populations of this species east and west of the arid corridor across the Kenyan Highlands.