The mid-domain effect: It’s not just about space

Ecologists and biogeographers have long sought to understand how and why diversity varies across space. Up until the late 20th century, the dominant role of environmental gradients and historical processes in driving geographical species richness patterns went largely undisputed. However, almost 20 years ago, Colwell & Hurtt (1994) proposed a radical reappraisal of ecological gradient theory that called into question decades of empirical and theoretical research. That controversial idea was later termed the ‘the mid-domain effect’: the simple proposition that in the absence of environmental gradients, the random placement of species ranges within a bounded domain will give rise to greatest range overlap, and thus richness, at the center of the domain (Colwell & Lees, 2000) (Fig. 1a). The implication of this line of reasoning is that the conventional null model of equal species richness regardless of latitude, elevation or depth should be replaced by one where richness peaks at some midpoint in geographical space. Our intention here is to draw attention to a neglected, yet important manifestation of the mid-domain effect, namely the application of mid-domain models (also referred to as geometric constraint models) to non-spatial domains. If individual species have ranges that exist not just in geographical space but also in environmental factors, such as temperature, rainfall, pH, productivity or disturbance, shouldn’t we also expect mid-domain richness peaks along non-spatial gradients? A mid-domain model applied to non-spatial gradients predicts the maximum potential richness for every value of an environmental factor. As with spatial mid-domain models, realized richness would probably be less, but the limits to richness are still predicted to be hump-shaped. Indeed, hump-shaped relationships emerge with remarkably high frequency across various non-spatial gradients. For instance, two of ecology’s most fundamental, albeit controversial theories – the productivity–diversity relationship and the intermediate disturbance hypothesis – predict mid-domain peaks in species richness. However, the potential of non-spatial mid-domain models has gone largely ignored

Mammal community structure through the Paleocene-Eocene Thermal Maximum

Human-mediated species invasion and climate change are leading to global extinctions and are predicted to result in the loss of important axes of phylogenetic and functional diversity. However, the long-term robustness of modern communities to invasion is unknown, given the limited timescales over which they can be studied. Using the fossil record of the Paleocene- Eocene Thermal Maximum (PETM; ~ 56 Ma) in North America, we evaluate mammalian community-level response to a rapid global warming event (5° to 8°C) and invasion by three Eurasian mammalian orders and by species undergoing northward range shifts. We assembled a database of 144 species body sizes and created a time-scaled composite phylogeny. We calculated the phylogenetic and functional diversity of all communities before, during, and after the PETM. Despite increases in the phylogenetic diversity of the regional species pool, phylogenetic diversity of mammalian communities remained relatively unchanged, a pattern that is invariant to the tree dating method, uncertainty in tree topology, and resolution. Similarly, body size dispersion and the degree of spatial taxonomic turnover of communities remained similar across the PETM. We suggest that invasion by new taxa had little impact on Paleocene- Eocene mammal communities because niches were not saturated. Our findings are consistent with the numerous studies of modern communities that record little change in community-scale richness despite turnover in taxonomic composition during invasion. What remains unknown is whether long-term robustness to biotic and abiotic perturbation are retained by modern communities given global anthropogenic landscape modification

Biotic interchange has structured Western Hemisphere mammal communities

Aim.— Many hypotheses posit that species-rich tropical communities are dominated by speciesspecies interactions, apparent as competitive exclusion or character displacement, whereas species-poor temperate communities are dominated by species-environment interactions. Recent studies demonstrate a strong influence of macroevolutionary and biogeographic factors. We simultaneously test for the effects of species interactions, climate, and biotic interchange on Western Hemisphere mammal communities using a phylogenetic and functional diversity approach. Location.— Western Hemisphere. Time period.— Modern Major taxa studied.— Mammalia Methods.— Using Western Hemisphere mammal distributional and body mass data, we calculate body mass dispersion, phylogenetic diversity (Net Relatedness Index), and assemblage-averaged rates of co-occurrence (Checkerboard scores) in 100 km by 100 km grid cells under an equal area projection. We model body mass dispersion as a function of phylogenetic diversity, cooccurrence rates, and species richness, as well as mean annual temperature and precipitation. We infer rates of dispersal among the temperate and tropical zones of the Western Hemisphere using phylogenetic methods. Results.— The dispersion of Northern Temperate mammal body masses is higher than null communities and shows correlated change with climate, consistent with resource competition and environmental filtering. Conversely, the dispersion of tropical and Southern Temperate mammal body masses are lower than and not differentiable from null expectations, respectively, suggesting a limited role of species-species and species-environment interactions at the grain of our analysis. Low tropical body mass dispersion and phylogenetic evenness are best explained by the high rates of faunal mixing. High rates of dispersal might also explain the similarity in community structure between the Southern temperate and tropical zones. Main Conclusions.— Mammal community assembly processes differ among the temperate and tropical zones of the Western Hemisphere and faunal mixture during dispersal events such as the Great American Biotic Interchange (Pliocene ~3 Ma) may have been important in structuring Western Hemisphere mammal communities. 9 (!) supplementary files attached below

The accelerating influence of humans on mammalian macroecological patterns over the late Quaternary

The transition of hominins to a largely meat-based diet ~1.8 million years ago led to the exploitation of other mammals for food and resources. As hominins, particularly archaic and modern humans, became increasingly abundant and dispersed across the globe, a temporally and spatially transgressive extinction of large-bodied mammals followed; the degree of selectivity was unprecedented in the Cenozoic fossil record. Today, most remaining large-bodied mammal species are confined to Africa, where they coevolved with hominins. Here, using a comprehensive global dataset of mammal distribution, life history and ecology, we examine the consequences of “body size downgrading” of mammals over the late Quaternary on fundamental macroecological patterns. Specifically, we examine changes in species diversity, global and continental body size distributions, allometric scaling of geographic range size with body mass, and the scaling of maximum body size with area. Moreover, we project these patterns toward a potential future scenario in which all mammals currently listed as vulnerable on the IUCN\u27s Red List are extirpated. Our analysis demonstrates that anthropogenic impact on earth systems predates the terminal Pleistocene and has grown as populations increased and humans have become more widespread. Moreover, owing to the disproportionate influence on ecosystem structure and function of megafauna, past and present body size downgrading has reshaped Earth\u27s biosphere. Thus, macroecological studies based only on modern species yield distorted results, which are not representative of the patterns present for most of mammal evolution. Our review supports the concept of benchmarking the “Anthropocene” with the earliest activities of Homo sapiens

Evidence for Trait-Based Dominance in Occupancy among Fossil Taxa and the Decoupling of Macroecological and Macroevolutionary Success

Biological systems provide examples of differential success among taxa, from ecosystems with a few dominant species (ecological success) to clades that possess far more species than sister clades (macroevolutionary success). Macroecological success, the occupation by a species or clade of an unusually high number of areas, has received less attention. If macroecological success reflects heritable traits, then successful species should be related. Genera composed of species possessing those traits should occupy more areas than genera with comparable species richness that lack such traits. Alternatively, if macroecological success reflects autapomorphic traits, then generic occupancy should be a by-product of species richness among genera and occupancy of constituent species. We test this using Phanerozoic marine invertebrates. Although temporal patterns of species and generic occupancy are strongly correlated, inequality in generic occupancy typically is greater than expected. Genus-level patterns cannot be explained solely with species-level patterns. Within individual intervals, deviations between the observed and expected generic occupancy correlate with the number of lithological units (stratigraphic formations), particularly after controlling for geographic range and species richness. However, elevated generic occupancy is unrelated to or negatively associated with either generic geographic ranges or within-genus species richness. Our results suggest that shared traits among congeneric species encourage shortterm macroecological success without generating short-term macroevolutionary success. A broad niche may confer high occupancy but does not necessarily promote speciation. Supplemental files attached below (.zip

Evidence for Trait-Based Dominance in Occupancy among Fossil Taxa and the Decoupling of Macroecological and Macroevolutionary Success

Biological systems provide examples of differential success among taxa, from ecosystems with a few dominant species (ecological success) to clades that possess far more species than sister clades (macroevolutionary success). Macroecological success, the occupation by a species or clade of an unusually high number of areas, has received less attention. If macroecological success reflects heritable traits, then successful species should be related. Genera composed of species possessing those traits should occupy more areas than genera with comparable species richness that lack such traits. Alternatively, if macroecological success reflects autapomorphic traits, then generic occupancy should be a by-product of species richness among genera and occupancy of constituent species. We test this using Phanerozoic marine invertebrates. Although temporal patterns of species and generic occupancy are strongly correlated, inequality in generic occupancy typically is greater than expected. Genus-level patterns cannot be explained solely with species-level patterns. Within individual intervals, deviations between the observed and expected generic occupancy correlate with the number of lithological units (stratigraphic formations), particularly after controlling for geographic range and species richness. However, elevated generic occupancy is unrelated to or negatively associated with either generic geographic ranges or within-genus species richness. Our results suggest that shared traits among congeneric species encourage shortterm macroecological success without generating short-term macroevolutionary success. A broad niche may confer high occupancy but does not necessarily promote speciation. Supplemental files attached below (.zip

Evidence for Trait-Based Dominance in Occupancy among Fossil Taxa and the Decoupling of Macroecological and Macroevolutionary Success

Biological systems provide examples of differential success among taxa, from ecosystems with a few dominant species (ecological success) to clades that possess far more species than sister clades (macroevolutionary success). Macroecological success, the occupation by a species or clade of an unusually high number of areas, has received less attention. If macroecological success reflects heritable traits, then successful species should be related. Genera composed of species possessing those traits should occupy more areas than genera with comparable species richness that lack such traits. Alternatively, if macroecological success reflects autapomorphic traits, then generic occupancy should be a by-product of species richness among genera and occupancy of constituent species. We test this using Phanerozoic marine invertebrates. Although temporal patterns of species and generic occupancy are strongly correlated, inequality in generic occupancy typically is greater than expected. Genus-level patterns cannot be explained solely with species-level patterns. Within individual intervals, deviations between the observed and expected generic occupancy correlate with the number of lithological units (stratigraphic formations), particularly after controlling for geographic range and species richness. However, elevated generic occupancy is unrelated to or negatively associated with either generic geographic ranges or within-genus species richness. Our results suggest that shared traits among congeneric species encourage shortterm macroecological success without generating short-term macroevolutionary success. A broad niche may confer high occupancy but does not necessarily promote speciation. Supplemental files attached below (.zip

Evidence for Trait-Based Dominance in Occupancy among Fossil Taxa and the Decoupling of Macroecological and Macroevolutionary Success

Biological systems provide examples of differential success among taxa, from ecosystems with a few dominant species (ecological success) to clades that possess far more species than sister clades (macroevolutionary success). Macroecological success, the occupation by a species or clade of an unusually high number of areas, has received less attention. If macroecological success reflects heritable traits, then successful species should be related. Genera composed of species possessing those traits should occupy more areas than genera with comparable species richness that lack such traits. Alternatively, if macroecological success reflects autapomorphic traits, then generic occupancy should be a by-product of species richness among genera and occupancy of constituent species. We test this using Phanerozoic marine invertebrates. Although temporal patterns of species and generic occupancy are strongly correlated, inequality in generic occupancy typically is greater than expected. Genus-level patterns cannot be explained solely with species-level patterns. Within individual intervals, deviations between the observed and expected generic occupancy correlate with the number of lithological units (stratigraphic formations), particularly after controlling for geographic range and species richness. However, elevated generic occupancy is unrelated to or negatively associated with either generic geographic ranges or within-genus species richness. Our results suggest that shared traits among congeneric species encourage shortterm macroecological success without generating short-term macroevolutionary success. A broad niche may confer high occupancy but does not necessarily promote speciation. Supplemental files attached below (.zip

Changes in the diet and body size of a small herbivorous mammal (hispid cotton rat, \u3ci\u3eSigmodon hispidus\u3c/i\u3e) following the late Pleistocene megafauna extinction

The catastrophic loss of large-bodied mammals during the terminal Pleistocene likely led to cascading effects within communities. While the extinction of the top consumers probably expanded the resources available to survivors of all body sizes, little work has focused on the responses of the smallest mammals. Here, we use a detailed fossil record from the southwestern United States to examine the response of the hispid cotton rat Sigmodon hispidus to biodiversity loss and climatic change over the late Quaternary. In particular, we focus on changes in diet and body size. We characterize diet through carbon (δ13C) and nitrogen (δ15N) isotope analysis of bone collagen in fossil jaws and body size through measurement of fossil teeth; the abundance of material allows us to examine population level responses at millennial scale for the past 16 ka. Sigmodon was not present at the cave during the full glacial, first appearing at ~16 ka after ice sheets were in retreat. It remained relatively rare until ~12 ka when warming tempera­tures allowed it to expand its species range northward. We find variation in both diet and body size of Sigmodon hispidus over time: the average body size of the population varied by ~20% (90–110 g) and mean δ13C and δ15N values ranged between −13.5 to −16.5‰ and 5.5 to 7.4‰ respectively. A state–space model suggested changes in mass were influenced by diet, maximum temperature and community structure, while the modest changes in diet were most influenced by community structure. Sigmodon maintained a fairly similar dietary niche over time despite contemporaneous changes in climate and herbivore community composition that followed the megafauna extinc­tion. Broadly, our results suggest that small mammals may be as sensitive to shifts in local biotic interactions within their ecosystem as they are to changes in climate and large-scale biodiversity loss

Ecological interactions disrupted by habitat alteration in the Neotropics

Ecological interactions help determine the distribution of species across landscapes and play crucial roles in ecosystem services such as pollination, seed dispersal, and pest control1. Human disturbances, particularly habitat alteration, have the potential to modify or erase ecological interactions2,3 and so jeopardise the processes they control. While examples of interactions becoming rewired under human influence have been recorded, studies of this process for speciose assemblages at regional to continental scales are uncommon4 and obstructed by logistical difficulties2. The consequences for ecological communities and people are therefore poorly understood. Here we show that human habitat alteration is associated with a decrease in the spatial aggregation of Neotropical bat pairs and bird pairs that share similar dietary requirements. We find that groups of species pairs with similar vs. different diets have positive spatial associations on average, but pairs within dietary guilds have stronger associations than pairs with disparate diets when habitats are relatively intact. Our results suggest that species with similar resource requirements typically coexist in relatively intact natural settings. By contrast, exclusion becomes more common (though not dominant) when habitats are altered. Altered habitats thus fail to support the coexistence of diverse competitive interactions, reversing patterns observed in the wild