Does energy availability influence classical patterns of spatial variation in exotic species richness?

Aim At macroecological scales, exotic species richness is frequently positively correlated with human population density. Such patterns are typically thought to arise because high human densities are associated with increased introduction effort and/or habitat modification and disturbance. Exotic and native species richness are also frequently positively correlated, although the causal mechanisms remain unclear. Energy availability frequently explains much of the variation in species richness and we test whether such species–energy relationships may influence the relationships of exotic species richness with human population density and native species richness. Location Great Britain. Methods We first investigate how spatial variation in the distributions of the 10 exotic bird species is related to energy availability. We then model exotic species richness using native avian species richness, human population density and energy availability as predictors. Species richness is modelled using two sets of models: one assumes independent errors and the other takes spatial correlation into account. Results The probability of each exotic species occurring, in a 10‐km quadrat, increases with energy availability. Exotic species richness is positively correlated with energy availability, human population density and native species richness in univariate tests. When taking energy availability into account, exotic species richness is negligibly influenced by human population density, but remains positively associated with native species richness. Main conclusions We provide one of the few demonstrations that energy availability exerts a strong positive influence on exotic species richness. Within our data, the positive relationship between exotic species richness and human population density probably arises because both variables increase with energy availability, and may be independent of the influence of human density on the probability of establishment. Positive correlations between exotic and native species richness remain when controlling for the influence of energy on species richness. The relevance of such a finding to the debate on the relationship between diversity and invasibility is discussed.     

Diversity distribution of saproxylic beetles in Chilean Mediterranean forests: influence of spatiotemporal heterogeneity and perturbation

Mediterranean ecosystems have been recognized as a priority for biodiversity conservation due to their high levels of species richness and endemism. In South America, these environments are restricted to central Chile and represent a biodiversity hotspot. The study of saproxylic beetles in this area is an unexplored topic, despite the ecological role they play in these ecosystems and their potential usefulness for monitoring the degree of forest conservation. We investigated the diversity distribution of trophic guilds of saproxylic beetles in Chilean Mediterranean forests, to identify the main environmental variables that influence their distribution. We also analyzed seasonal dynamics as a key factor influencing insect communities and the effect of human disturbance on their diversity and composition. We identified characteristic species of perturbation degree as bioindicators for habitat monitoring. A total of 40 window traps were used to survey three Mediterranean forest types in the Río Clarillo National Reserve and the adjacent non-protected areas. We found that the diversity, abundance and composition of saproxylic beetles varied significantly spatiotemporally among the studied forest types and among perturbation degrees, showing different patterns depending on the trophic guild. Results indicated that conservation decisions should include the preservation of a larger range of different vegetation types and the nearby zones that have suffered low levels of disturbance or fragmentation and where actions promoting the presence of old native trees would have a significant conservation value.     

Ants and people: a test of two mechanisms potentially responsible for the large‐scale human population–biodiversity correlation for Formicidae in Europe

Aim: Recent coarse‐scale studies have shown positive relationships between the biodiversity of plants/vertebrates and the human population. Little is known about the generality of the pattern for invertebrates. Moreover, biodiversity and human population might correlate because they both covary with other factors such as energy availability and habitat heterogeneity. Here we test these two non‐mutually exclusive mechanisms with ant species‐richness data from the Fauna Europaea. Location Forty‐three European countries/regions. Methods We derived mixed models of total, native and exotic ant species richness as a function of human population size/density, controlling for country area, plant species richness (as a proxy for habitat heterogeneity), and mean annual temperature and precipitation (variables related to energy availability). Results Ant species richness increased significantly with increasing human population. This result was confirmed when controlling for variations in country area. Both for human population size/density and for ant species richness, there were positive correlations with temperature but not with precipitation. This finding is in agreement with the energy‐availability hypothesis. However, we observed a negative latitudinal gradient in ant and plant species richness, although not in human population size/density. Plant species richness was positively correlated with ant species richness but not with human population size/density. Thus, there is evidence that this type of habitat heterogeneity can play a role in the observed latitudinal gradient of ant species richness, but not in the positive correlation between ant species richness and human population. The results were confirmed for the 545 native and the 32 exotic ant species reported, and we observed a good correlation between exotic and native ant species richness. Main conclusions Ant species richness in European countries conforms to six macroecological patterns: (1) a negative latitudinal gradient; and a positive (2) species–energy relationship, (3) species–area relationship, (4) correlation with plant species richness, (5) exotic–native species richness correlation, and (6) species–people correlation. There is some evidence for the energy‐availability hypothesis, but little evidence for habitat heterogeneity as an explanation of the large‐scale human population–ant biodiversity correlation. This correlation has implications for the conservation of ant diversity in Europe.     

Negative native–exotic diversity relationship in oak savannas explained by human influence and climate

Recent research has proposed a scale-dependence to relationships between native diversity and exotic invasions. At fine spatial scales, native–exotic richness relationships should be negative as higher native richness confers resistance to invasion. At broad scales, relationships should be positive if natives and exotics respond similarly to extrinsic factors. Yet few studies have examined both native and exotic richness patterns across gradients of human influence, where impacts could affect native and exotic species differently. We examined native–exotic richness relationships and extrinsic drivers of plant species richness and distributions across an urban development gradient in remnant oak savanna patches. In sharp contrast to most reported results, we found a negative relationship at the regional scale, and no relationship at the local scale. The negative regional-scale relationship was best explained by extrinsic factors, surrounding road density and climate, affecting natives and exotics in opposite ways, rather than a direct effect of native on exotic richness, or vice versa. Models of individual species distributions also support the result that road density and climate have largely opposite effects on native and exotic species, although simple life history traits (life form, dispersal mode) do not predict which habitat characteristics are important for particular species. Roads likely influence distributions and species richness by increasing both exotic propagule pressure and disturbance to native species. Climate may partially explain the negative relationship due to differing climatic preferences within the native and exotic species pools. As gradients of human influence are increasingly common, negative broad-scale native–exotic richness relationships may be frequent in such landscapes.     

Distance to suburban/wildland border interacts with habitat type for structuring exotic plant communities in a natural area surrounding a metropolitan area in central Chile

Background: The explosive growth of urbanisation in Mediterranean ecosystems in Chile has favoured the rapid expansion of exotic plant species, yet factors driving these invasion patterns in adjacent natural areas remain poorly assessed.

Aims: To assess how distance to a suburban/wildland border, habitat type, site-scale disturbance and woody plant cover of native species influences the diversity of exotic species in a natural area surrounding the city of Santiago, Chile.

Methods: Three watersheds were chosen, and the diversity of exotic species was assessed in 36 100-m-long transects, equally distributed over two distance categories and three habitats. For each transect, we measured woody plant cover of native species and frequency of rabbit faeces as a measure of competitive exclusion and site-scale disturbance, respectively.

Results: Species diversity decreased as the distance from the suburban/wildland border increased, and it was found to be higher in north-facing habitats compared to south-facing and alluvial habitats. Neither native woody plant cover nor frequency of rabbit faeces had an effect on species diversity.

Conclusions: The current pattern of exotic plant species in this natural area is mainly influenced by the distance to suburban border and habitat type. An adequate management of conditions favouring exotic species in suburban/wildland border may prevent the spread of these into natural areas next to urban settings.     

Invasive Exotic Plants in the Tropical Pacific Islands: Patterns of Diversity

Oceanic islands are good model systems with which to explore factors affecting exotic species diversity. Islands vary in size, topography, substrate type, degree of isolation, native species diversity, history, human population characteristics, and economic development. Moreover, islands are highly vulnerable to exotic species establishment. We used AICc analyses of data on 1132 vascular plant species for 15 countries and 114 islands from the Pacific Island Ecosystems at Risk (PIER) project to examine biological, geographical, and socioeconomic correlates of exotic species richness. PIER provides data on the distribution of naturalized non-native plant species thought to pose environmental or economic risk. We hypothesized that the numbers of PIER-listed species would be positively correlated with island size, habitat diversity, and proximity to major source pools for propagules. Further, we expected numbers of PIER-listed exotic species to be similar among islands in the same country and to be greater where human populations were larger and where economic activity was high. Most species (908) were found on ≤ 10 islands. Species number was significantly correlated with island and country areas and with native plant species richness. The strongest model revealed by AICc analyses of island data included log (area) and maximum elevation as well as country membership, substrate type, and presence of an airport with paved runway (an index of economic activity). By country, AICc analyses revealed two equivalent models, both of which included log (area) and per capita gross domestic product as well as a measure of population size (either log (population size) or (population density)). Our analyses provide strong evidence of the roles of biogeographic, environmental, and socioeconomic impacts on the distribution and spread of exotic species.     

Disturbance accumulation hampers fish assemblage recovery long after the worst mining spill in the Iberian Peninsula

The influence of environmental variables on native and exotic fish species richness and diversity was analysed 8 years after one of the most environmentally harmful toxic spills worldwide. Environment‐diversity relationships were addressed on different scales, and values were also compared with those of six similar basins not affected by the spill, with the aim of determining whether this disturbance was still exerting an influence on the fish assemblage. Results showed higher native species richness in environments with low human influence, no reservoirs upstream, a large drainage area and coarse substrate reaches. For native fish, variables at both the catchment and site were equally relevant. Exotic fish were mainly favoured by site‐scale factors such as valley width downstream from the reservoir, where the alteration of the river channel and accumulated disturbances give them an advantage vs native species. Overall, 8 years after the accident, richness and diversity of the Guadiamar fish assemblage seemed more affected by anthropogenic impacts than by the long‐term influence of the toxic spill. This work highlights that the potentially synergic effects of anthropogenic factors must be taken into account when monitoring the long‐term effects of pollution events.     

Short and long term consequences of increases in exotic species richness on water filtration by marine invertebrates

Although recent research has considered the consequences of global declines in the number of species, less attention has focused on the aggregate effects of regional increases in species richness as a result of human-mediated introductions. Here we examine several potential ecosystem consequences of increasing exotic species diversity of suspension feeding marine invertebrates. First, we experimentally manipulated native and non-native suspension feeder richness and measured its effect on short-term phytoplankton clearance rates. Multispecies communities all performed similarly, regardless of whether they were dominated by natives, exotics, or an even mix of the two. Individual species varied considerably in filtration rates, but non-native species often filtered less than the most similar native. Second, we determined potential changes in integrated function over time by comparing seasonal patterns of recruitment as a proxy for the ability to quickly recover filtration capacity after a disturbance. We found that exotic species have complementary seasonal phenologies both to native species and each other. Our results suggest that the consequences of local increases in species richness due to invasions may be manifest over long (annual to interannual) time scales, even when short term changes in ecosystem function are negligible.     

Latitudinal patterns and regionalization of plant diversity along a 4270‐km gradient in continental Chile

According to the global latitudinal diversity gradient, a decrease in animal and plant species richness exists from the tropics towards higher latitudes. The aim of this study was to describe the latitudinal distribution patterns of Chilean continental flora and delineate biogeographic regions along a 4270‐km north–south gradient. We reviewed plant lists for each of the 39 parallels of continental Chile to build a database of the geographical distribution of vascular plant species comprising 184 families, 957 genera and 3787 species, which corresponded to 100%, 94.9% and 74.2% of the richness previously defined for Chile, respectively. Using this latitudinal presence–absence species matrix, we identified areas with high plant richness and endemism and performed a Cluster analysis using Jaccard index to delineate biogeographic regions. This study found that richness at family, genus and species levels follow a unimodal 4270‐km latitudinal distribution curve, with a concentration of richness in central Chile (31–42°S). The 37th parallel south (central Chile) presented the highest richness for all taxonomic levels and in specific zones the endemism (22–37°S) was especially high. This unimodal pattern contrasts the global latitudinal diversity gradient shown by other studies in the Northern hemisphere. Seven floristic regions were identified in this latitudinal gradient: tropical (18–22°S), north Mediterranean (23–28°S), central Mediterranean (29–32°S), south Mediterranean (33–37°S), north temperate (38–42°S), south temperate (43–52°S) and Austral (53–56°S). This regionalization coincides with previous bioclimatic classifications and illustrates the high heterogeneity of the biodiversity in Chile and the need for a reconsideration of governmental conservation strategies to protect this diversity throughout Chile.     

Assessing the impact of local habitat variables and landscape context on riparian birds in agricultural,urbanized, and native landscapes

Large tracts of natural habitat are being replaced by agriculture and urban sprawl in Mediterranean regions worldwide. We have limited knowledge about the effects of human activities on native species in these landscapes and which, if any, management practices might enhance the conservation of native biodiversity within them. Through a citizen volunteer bird-monitoring project, we compared bird abundance and species richness in northern Californian riparian zones surrounded by vineyards, urban areas, and natural areas. We assessed both local and landscape-level variables that may enhance native bird diversity in each land use type. We also demonstrate a new statistical approach, generalized estimating equations, to analyze highly variable data, such as that collected by volunteers. Avian abundance was highly correlated with both landscape context and local habitat variables, while avian richness was correlated with local habitat variables, specifically shrub richness, and percent of tree cover. In particular, shrub species richness has a strong positive correlation with riparian-preferring bird species. This suggests that active local management of riparian zones in human-dominated landscapes can increase our ability to retain native bird species in these areas.     

Anthropogenic impacts upon plant species richness and net primary productivity in California

We assess the importance of anthropogenic land‐use, altered productivity, and species invasions for observed productivity–richness relationships in California. To this end, we model net primary productivity (NPP) c. 1750 AD and at present (1982–1999) and map native and exotic vascular plant richness for 230 subecoregions. NPP has increased up to 105% in semi‐arid areas and decreased up to 48% in coastal urbanized areas. Exotic invasions have increased local species diversity up to 15%. Human activities have reinforced historical gradients in species richness but reduced the spatial heterogeneity of NPP. Structural equation modelling suggests that, prior to European settlement, NPP and richness were primarily controlled by precipitation and other abiotic variables, with NPP mediating richness. Abiotic variables remain the strongest predictors of present NPP and richness, but intermodel comparisons indicate a significant anthropogenic impact upon statewide distributions of NPP and richness. Exotic and native species each positively correlate to NPP after controlling for other variables, which may help explain recent reports of positively associated native and exotic richness.     

Grazing, Environmental Heterogeneity, and Alien Plant Invasions in Temperate Pampa Grasslands

Temperate humid grasslands are known to be particularly vulnerable to invasion by alien plant species when grazed by domestic livestock. The Flooding Pampa grasslands in eastern Argentina represent a well-documented case of a regional flora that has been extensively modified by anthropogenic disturbances and massive invasions over recent centuries. Here, we synthesise evidence from region-wide vegetation surveys and long-term exclosure experiments in the Flooding Pampa to examine the response of exotic and native plant richness to environmental heterogeneity, and to evaluate grazing effects on species composition and diversity at landscape and local community scales. Total plant richness showed a unimodal distribution along a composite stress/fertility gradient ranging several plant community types. On average, more exotic species occurred in intermediate fertility habitats that also contained the highest richness of resident native plants. Exotic plant richness was thus positively correlated with native species richness across a broad range of flood-prone grasslands. The notion that native plant diversity decreases invasibility was supported only for a limited range of species-rich communities in habitats where soil salinity stress and flooding were unimportant. We found that grazing promoted exotic plant invasions and generally enhanced community richness, whereas it reduced the compositional and functional heterogeneity of vegetation at the landscape scale. Hence, grazing effects on plant heterogeneity were scale-dependent. In addition, our results show that environmental fluctuations and physical disturbances such as large floods in the pampas may constrain, rather than encourage, exotic species in grazed grasslands.     

Factors Controlling Compositional Changes in a Northern Andean Páramo (La Rusia,Colombia)

In this study, we aimed to assess the processes controlling compositional change in a Northern Andean páramo highly affected by human‐induced disturbances over the last few decades (La Rusia, Colombia). Along the 3000–3800 m asl altitudinal range, we randomly sampled fifty 10 × 10 m plots. Therein, we measured altitude and variables related to soil conditions (i.e., moisture, nutrient contents, bulk density, and texture), occurrence of human‐induced disturbances (i.e., fire, vegetation clearing, potato cultivation, and cattle grazing), and land‐use history. We also recorded richness and abundance of plant species, identifying them as exotic or native. We differentiated four groups of plots according to their species composition. The groups had significant differences in altitude, soil conditions, land‐use history, and particularly, in richness of exotic species and exotic/native cover ratio. They could be ascribed to shrub‐ and grass‐páramo vegetation types based on their relative dominance of woody and herbaceous species; however, these groups were not arranged according to the hypothetical composition of altitudinal belts, but rather formed a mosaic of patches. This mosaic was determined not only by altitude but also by soil conditions and disturbance history of sites. Our results corroborate recent findings which highlight shrub‐ and grass‐páramo vegetation types as patches of contrasting species composition and structure that depend on local environmental variables, as well as human‐induced disturbances as a major determinant of compositional discontinuities in these ‘high mountain’ tropical ecosystems.     

Designing nature reserves: traditional criteria may act as misleading indicators of quality

Traditionally, numerous criteria have been used for selecting valuable areas for the establishment of nature reserves. In most cases these criteria are applied indiscriminately with the assumption that their generalized use gives them universal validity. In this study we test the value of different variables (area and shape of patches of natural vegetation, degree of internal fragmentation, and diversity of habitats at the periphery of patches) at the landscape level as indicators of plant richness and diversity, and relative abundance of native plants, in a group of natural vegetation relicts ranging from 0.004 km² to 1.027 km² and surrounded by sub-urban and industrial settings. Species richness, diversity, and number of native and exotic species increased with the area of the patches. The shape of the patch was the second most important variable to influence species richness and diversity. The number of exotic species increased with increasing numbers of native plants. Thus, patch size and plant richness should be carefully used for selecting conservation areas because it could result in choosing places threatened by the presence of exotics. Many of the other variables analysed showed no effects on biodiversity at the temporal and geographic scales considered. Ignoring these outcomes could result in choosing sub-optimal areas. We recommend the critical use of general criteria considering the selection process as an opportunity to evaluate the relevance of each criterion at the local level.     

A null model of exotic plant diversity tested with exotic and native species-area relationships

At large spatial scales, exotic and native plant diversity exhibit a strong positive relationship. This may occur because exotic and native species respond similarly to processes that influence diversity over large geographical areas. To test this hypothesis, we compared exotic and native species–area relationships within six North American ecoregions. We predicted and found that within ecoregions the ratio of exotic to native species richness remains constant with increasing area. Furthermore, we predicted that areas with more native species than predicted by the species–area relationship would have proportionally more exotics as well. We did find that these exotic and native deviations were highly correlated, but areas that were good (or bad) for native plants were even better (or worse) for exotics. Similar processes appear to influence exotic and native plant diversity but the degree of this influence may differ with site quality.     

Human impacts, energy availability and invasion across Southern Ocean Islands

Aim Ongoing biological invasions will enhance the impacts of humans on biodiversity. Nonetheless, the effects of exotic species on diversity are idiosyncratic. Increases in diversity might be a consequence of similar responses by species to available energy, or because of positive relationships between human density, energy and propagule pressure. Here we use data from the Southern Ocean island plants and insects to investigate these issues. Location The Southern Ocean Islands ranging from Tristan da Cunha to Heard Island and South Georgia. Methods Generalized linear models are used to explore the relationships between indigenous and exotic species richness for plants and insects on two different islands. Similar models are used to examine interactions between indigenous and exotic species richness, energy availability and propagule pressure at the regional scale. Results Positive relationships were found between indigenous and exotic species richness at local scales, although for plants, the relationship was partially triangular. Across the Southern Ocean Islands, there was strong positive covariation between indigenous and exotic plant species richness and insect species richness, even taking spatial autocorrelation into account. Both exotic and indigenous plant and insect species richness covaried with energy availability, as did human visitor frequency. When two islands with almost identical numbers of human visits were contrasted, it was clear that energy availability, or perhaps differences in climate‐matching, were responsible for differences in the extent of invasion. Conclusion In plants and insects, there are positive relationships between indigenous and exotic diversity at local and regional scales across the Southern Ocean islands. These relationships are apparently a consequence of similar responses by both groups and by human occupants to available energy. When visitor frequency is held constant, energy availability is the major correlate of exotic species richness, though the exact mechanistic cause of this relationship requires clarification.     

Aboveground plant community and seed bank composition along an invasion gradient

Invasive plants can reduce plant diversity and abundance in native grassland communities; however, the effect on the native seed bank is less clear. The objective of this study was to assess the effects of invasion by the exotic grass old world bluestem (OWB; Bothriochloa spp.) on native aboveground plant species composition and seed bank diversity and abundance (i.e., cover, density). In this central Great Plains grassland, OWB invasion had differential effects on native diversity and abundance of both aboveground and seed bank plant communities. Native plant species diversity and cover showed a steep decline as OWB cover increased. No change in native seed density or richness was observed in response to OWB invasion, however, OWB seed density increased with increasing invasion, thus increasing total seed density. Our results indicate that as OWB invasion increases, the native plant community decreases in diversity and abundance. Although, no effect on native seed bank diversity and density was observed in this study, as native seeds are lost through a loss of native species in the plant communities, native seed bank diversity and density is expected to decline.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.     

Indirect effects of grazing intensity on pollinators and floral visitation

1. The introduction of livestock in natural areas is a common disturbance that affects both plant and pollinator diversity and might affect their interaction. Understanding whether livestock affect a food resource for pollinators (i.e. flower abundance) and/or a pollinator assemblage (i.e. abundance and richness) has important implications for plant–pollinator interactions and still needs deeper investigation. 2. This study investigated how pollinator communities and flower abundance determined floral visitation frequency along a grazing gradient, using seven large paddocks in Patagonian Monte Desert that varied in livestock densities. Pollinator visitation frequency was measured in five of the most abundant native plant species of the region, present in all the paddocks, but that differed in reproductive strategy ranging from insect‐pollinated self‐compatible and self‐incompatible to wind‐pollinated. The influence of livestock density, insect, and flower abundance on visitation frequency was evaluated using D‐separation hierarchical path models. 3. Intermediate stocking densities showed the highest insect richness and abundance. Livestock density showed a negative quadratic relationship with insect richness; hymenopterans being the main insect group in the region. Flower density decreased with the increase in livestock density. The five plant species shared several pollinator species although each one supported a distinct assemblage. 4. The path model showed that livestock was not directly associated with pollinator visitation frequency; however, this apparent lack of association was as a result of opposite forces acting together. An increase in livestock density reduced visitation frequency through a decrease in insect abundance, yet, livestock simultaneously increased the pollinator visitation rate through decreased flower abundance. 5. This study describes how changes in the density of exotic mammals can affect pollinator and flower abundance, resulting in contrasting effects on flower visitation rates with, apparently, neutral net consequences. This illustrates the complexity of responses to plant–pollinator interactions to anthropogenic disturbances that alter the ecological context.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.