Explaining species richness from continents to communities: the time-for-speciation effect in emydid turtles

Speciation is the process that ultimately generates species richness. However, the time required for speciation to build up diversity in a region is rarely considered as an explanation for patterns of species richness. We explored this "time-for-speciation effect" on patterns of species richness in emydid turtles. Emydids show a striking pattern of high species richness in eastern North America (especially the southeast) and low diversity in other regions. At the continental scale, species richness is positively correlated with the amount of time emydids have been present and speciating in each region, with eastern North America being the ancestral region. Within eastern North America, higher regional species richness in the southeast is associated with smaller geographic range sizes and not greater local species richness in southern communities. We suggest that these patterns of geographic range size variation and local and regional species richness in eastern North America are caused by glaciation, allopatric speciation, and the time-for-speciation effect. We propose that allopatric speciation can simultaneously decrease geographic range size and increase regional diversity without increasing local diversity and that geographic range size can determine the relationship between alpha, beta, and gamma diversity. The time-for-speciation effect may act through a variety of processes at different spatial scales to determine diverse patterns of species richness.     

Mollusk species diversity in the Southeastern Pacific: why are there more species towards the pole?

The most ubiquitous and well recognized diversity pattern at large spatial scales is the latitudinal increase in species richness near the equator and decline towards the poles. Although several exceptions to this pattern have been documented, shallow water mollusks, the most specious group of marine invertebrates, are the epitome of the monotonic decline in species diversity toward higher latitudes along the Pacific and Atlantic coasts of North America. Here we analyze the geographic diversity of 629 mollusk species along the Pacific South American shelf. Our analyses are based on the most complete database of invertebrates assembled for this region of the world, consisting of latitudinal ranges of over 95% of all described mollusks between 10° and 55°S. Along this coast, mollusk diversity did not follow the typical latitudinal trend. The number of species remained constant and relatively low at intermediate latitudes and sharply increased toward higher latitudes, south of 42°S. This trend was explained by changes in shelf area, but not by sea surface temperature, unlike the pattern documented for Northern Hemisphere mollusks. Direct sampling of soft bottom communities along the gradient suggests that regional trends in species richness are produced by increased alpha diversity, and not only by artifacts produced by the increase in sampling area. We hypothesize that increased shelf area south of 42°S, geographic isolation produced by divergence of major oceanic currents, and the existence of refugia during glaciations, enabled species diversification. Radiation could have been limited by narrow continental shelves between 10°–42°. Asymmetries in latitudinal diversity trends between hemispheres show that there is not a single general factor determining large-scale diversity patterns.     

Seaweeds, latitudinal diversity patterns, and Rapoport's Rule

We analysed the geographic distribution of 645 species of marine benthic algae along the Atlantic coast of Europe and Pacific coast of temperate South America to test for the existence of an association between geographic range size and latitude (Rapoport's Rule) and for three key components of the explanations offered for it. We found that species in high diversity areas are characterized by small geographic ranges and by low specific growth rates as compared to species with large geographic ranges, thus supporting the Rapoport-rescue hypothesis. However, the pattern is not related to species' tolerances, to abiotic conditions or to climatic variability. Further, the inverse latitudinal diversity pattern shown by the marine algal flora of temperate Pacific South America, and the opposite patterns shown by tropical and subantarctic species within this flora, stressed that small geographic ranges are linked to high diversity areas in general, and not only in relation to the pole to tropic species diversity gradient.     

Biogeographic and phylogenetic patterns in diversity of liverwort-associated endophytes

Liverworts harbor diverse fungi, including endophytes, in their healthy tissues. To address whether patterns of endophyte diversity are correlated with host phylogeny or geography, we designed a broad geographic survey with controlled phylogenetic host sampling. We collected liverworts in North Carolina, Washington, Idaho, British Columbia, Germany, and New Zealand and identified endophytes using culture-based and molecular methods. Of the major lineages of filamentous ascomycetes recovered, 53-88% belonged to the Xylariales. Endophyte accumulation curves did not saturate, and singleton sequences were dominant in each region, suggesting that liverwort endophyte communities are diverse. There was no significant difference in species richness between regional endophyte communities; however, total richness estimators indicated that North Carolina and New Zealand have richer communities than do Germany and the Pacific Northwest. This pattern reflects lower per-host endophyte density and prevalence of a common, shared sequence group in Germany and the Pacific Northwest. Although species richness was relatively low in the Pacific Northwest, the greatest phylogenetic diversity of endophytes was recovered there. Tests for regional and host specificity revealed that endophyte floras of hosts within a geographic area are more similar to one another than to those of closely related hosts. Geographic distance, not host phylogeny, best explains differences among communities.     

Spatial patterns of morphological diversity across the Indo-Pacific: analyses using strombid gastropods.

Biological diversity can be measured using various metrics, but existing knowledge of spatial patterns of diversity is largely based on species counts. There is increasing evidence that trends in species richness might not match trends in other biodiversity metrics, such as morphological diversity. Here, we use data from a large group of Indo-Pacific gastropods (family Strombidae) to show that the species richness of a region is a poor predictor of the morphological diversity present there. Areas with only a few species can harbour an impressive array of morphologies and, conversely, morphological diversity in the most species-rich regions is no higher than in regions with half their taxonomic diversity. Biological diversity in the Pacific is highly threatened by human activity and our results indicate that, in addition to species richness, morphological diversity metrics need to be incorporated into conservation decisions.     

Comparing galling insect richness among Neotropical savannas: effects of plant richness, vegetation structure and super-host presence

Plant species diversity maintains the stability of ecosystems and the diversity of consumer species such as insect herbivores. Considering that gall-inducing insects are highly specialized on their host plants and dependent on the occurrence, abundance and distribution of plants, we evaluated the diversity patterns of gall-inducing insect along Brazilian Neotropical savannas and the potential role of plant species richness, vegetation structure and super-host presence on determining these patterns. We found 1,882 individual plants that belonged to 64 different host plant species grouped in 31 families, associated to 112 galling insect species. The galling richness was positively influenced by plant species richness and the presence of the super-host genus Qualea (Vochysiaceae). Plant species richness explained 48 % of the galling richness and areas with presence of super-hosts had more than twice of galling species than areas where they were absent. On the other hand, we found no evidence that larger plants hosted more species of galling insects. We observed that for the diversity of galling insects in the Brazilian Cerrado, vegetation structure explained almost the same portion as plant richness, because structural variables did not have an effect on residuals of galling richness and plant richness regression. Our findings suggests that plant richness has a more important role on the mitigation of natural enemies and adaptive radiation of galling species, while structural aspects of the vegetation does not seem to have that effect. Furthermore, we show that the super-host taxa provide an increment in local galling richness because they present a great diversity of local number of gall morphospecies (i.e. alpha diversity) and the high turnover of morphospecies among different localities (i.e. beta diversity). Therefore we argue that the quality of resources (richness and super host presence) appears to be a most important factor for the diversity of galling insects in Neotropical systems, than the amount of resources.     

Island biogeography of Caribbean coral reef fish

Aim The goal of our study was to test fundamental predictions of biogeographical theories in tropical reef fish assemblages, in particular relationships between fish species richness and island area, isolation and oceanographic variables (temperature and productivity) in the insular Caribbean. These analyses complement an analogous and more voluminous body of work from the tropical Indo‐Pacific. The Caribbean is more limited in area with smaller inter‐island distances than the Indo‐Pacific, providing a unique context to consider fundamental processes likely to affect richness patterns of reef fish. Location Caribbean Sea. Methods We compiled a set of data describing reef‐associated fish assemblages from 24 island nations across the Caribbean Sea, representing a wide range of isolation and varying in land area from 53 to 110,860 km². Regression‐based analyses compared the univariate and combined effects of island‐specific physical predictors on fish species richness. Results We found that diversity of reef‐associated fishes increases strongly with increasing island area and with decreasing isolation. Richness also increases with increasing nearshore productivity. Analyses of various subsets of the entire data set reveal the robustness of the richness data and biogeographical patterns. Main conclusions Within the relatively small and densely packed Caribbean basin, fish species richness fits the classical species–area relationship. Richness also was related negatively to isolation, suggesting direct effects of dispersal limitation in community assembly. Because oceanic productivity was correlated with isolation, however, the related effects of system‐wide productivity on richness cannot be disentangled. These results highlight fundamental mechanisms that underlie spatial patterns of biodiversity among Caribbean coral reefs, and which are probably also are functioning in the more widespread and heterogeneous reefs of the Indo‐Pacific.     

BIODIVERSITY RESEARCH: Genetic diversity in two introduced biofouling amphipods (Ampithoe valida & Jassa marmorata) along the Pacific North American coast: investigation into molecular identification and cryptic diversity

Aim We investigated patterns of genetic diversity among invasive populations of Ampithoe valida and Jassa marmorata from the Pacific North American coast to assess the accuracy of morphological identification and determine whether or not cryptic diversity and multiple introductions contribute to the contemporary distribution of these species in the region. Location Native range: Atlantic North American coast; Invaded range: Pacific North American coast. Methods We assessed indices of genetic diversity based on DNA sequence data from the mitochondrial cytochrome c oxidase subunit I (COI) gene, determined the distribution of COI haplotypes among populations in both the invasive and putative native ranges of A. valida and J. marmorata and reconstructed phylogenetic relationships among COI haplotypes using both maximum parsimony and Bayesian approaches. Results Phylogenetic inference indicates that inaccurate species‐level identifications by morphological criteria are common among Jassa specimens. In addition, our data reveal the presence of three well supported but previously unrecognized clades of A. valida among specimens in the north‐eastern Pacific. Different species of Jassa and different genetic lineages of Ampithoe exhibit striking disparity in geographic distribution across the region as well as substantial differences in genetic diversity indices. Main conclusions Molecular genetic methods greatly improve the accuracy and resolution of identifications for invasive benthic marine amphipods at the species level and below. Our data suggest that multiple cryptic introductions of Ampithoe have occurred in the north‐eastern Pacific and highlight uncertainty regarding the origin and invasion histories of both Jassa and Ampithoe species. Additional morphological and genetic analyses are necessary to clarify the taxonomy and native biogeography of both amphipod genera.     

At least some meiofaunal species are not everywhere. Indication of geographic,ecological and geological barriers affecting the dispersion of species of Ototyphlonemertes (Nemertea,Hoplonemertea)

Most meiofaunal species are known to have a broad distribution with no apparent barriers to their dispersion. However, different morphological and/or molecular methods supported patterns of diversity and distribution that may be different among taxa while also conflicting within the same group. We accurately assessed the patterns of geographic distribution in actual genetic species of a marine meiofaunal animal model: Ototyphlonemertes. Specimens were collected from several sites around Europe, Northern and Central America, Southern America, Pacific Islands and Asia. We sequenced regions of two mitochondrial and two nuclear genes. Using single‐gene, a concatenated data set, multilocus approaches and different DNA taxonomy methods, we disentangled the actual diversity and the spatial structures of haplotypes and tested the possible correlation between genetic diversity and geographic distance. The results show (i) the importance of using several genes to uncover both diversity and highlight phylogeographic relationships among species and that (ii) independent genetic evolutionary entities have a narrower distribution than morphological species. Moreover, (iii) a Mantel test supported a positive correlation between genetic and geographical distance. By sampling from the two sides of Isthmus of Panama, we were additionally able to identify lineage divergence times that are concordant with vicariance mechanisms caused by the geological closure of the seaway across the Isthmus. We therefore propose that in addition to distance, other geological and ecological conditions are also barriers to the dispersion of and gene flow in marine meiofaunal organisms.     

Collembola,the biological species concept and the underestimation of global species richness

Despite its ancient origin, global distribution and abundance in nearly all habitats, the class Collembola is comprised of only 8000 described species and is estimated to number no more than 50 000. Many morphologically defined species have broad geographical ranges that span continents, and recent molecular work has revealed high genetic diversity within species. However, the evolutionary significance of this genetic diversity is unknown. In this study, we sample five morphological species of the globally distributed genus Lepidocyrtus from 14 Panamanian sampling sites to characterize genetic diversity and test morphospecies against the biological species concept. Mitochondrial and nuclear DNA sequence data were analysed and a total of 58 molecular lineages revealed. Deep lineage diversification was recovered, with 30 molecular lineages estimated to have established more than 10 million years ago, and the origin almost all contemporary lineages preceding the onset of the Pleistocene (~2 Mya). Thirty‐four lineages were sampled in sympatry revealing unambiguous cosegregation of mitochondrial and nuclear DNA sequence variation, consistent with biological species. Species richness within the class Collembola and the geographical structure of this diversity are substantially misrepresented components of terrestrial animal biodiversity. We speculate that global species richness of Collembola could be at least an order of magnitude greater than a previous estimate of 50 000 species.     

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.     

Zoobenthic diversity-gradients in the Baltic Sea: Continuous post-glacial succession in a stressed ecosystem

The Baltic Sea, formed after the latest glaciation, is an enclosed, low-saline, non-tidal ecosystem and has steep latitudinal and vertical gradients from sub-arctic conditions in the north to temperate in the south. The sea has undergone rapid changes since the glaciation, and the “ecological age” of the present ecosystem is only about 8000 years. Primary successional processes are still ongoing, and numerous ecological niches (e.g. large-bodied sediment bioturbators) remain available for immigration. The system is species-poor and vulnerable to the threat of exotic invasive species, and to date about 50 zoobenthic species have established populations in the Baltic Sea. The present biota is a mixture of species of different ecological and zoogeographical origin (marine to limnic; northern Arctic marine and limnic, to North Sea and Atlantic marine). The current distribution patterns of zoobenthos are illustrated, using marine, limnic and non-indigenous examples of structure and ecosystem functions. The species richness decreases from over 1600 marine benthic species in the open Skagerrak to about 500 in the western parts of the Baltic Sea, approximately 80 in the southern regions, to less than 20 in the northern regions. On the other hand, limnic species increase diversity in the inner reaches of the Gulf of Finland and the Gulf of Bothnia. Polychaetes, molluscs and echinoderms are dramatically reduced in numbers from the south to the north.     

Genetic and morphological evidence for cryptic diversity in the <Emphasis Type="Italic">Careproctus rastrinus</Emphasis> species complex (Liparidae) of the North Pacific

The molecular phylogeny of the Careproctus rastrinus species complex is presented on the basis of sequence variations in the 16S rRNA and cytochrome b genes (1,447 base pairs) of mitochondrial DNA using specimens collected from across the North Pacific and its marginal seas, including the Sea of Japan, the Pacific coast of Japan, the Sea of Okhotsk, the Bering Sea, the Gulf of Alaska, and the Arctic Ocean. Gene flow and migration between geographic regions is apparently strictly restricted in the C. rastrinus species complex, as this phylogeny revealed nine genetically diverged groups in the C. rastrinus species complex, most of which corresponded well with major geographic regions. Most of the groups were different in terms of morphological characters. Five nominal species have been considered to be members of the complex and have been variously recognized as being valid or as synonyms of C. rastrinus, but the present genetic and morphological differences suggested that the C. rastrinus species complex represents far more divergence. Such cryptic diversity of the C. rastrinus species complex defined by geographic regions may be related to their low dispersal ability, because they bear large demersal eggs. The genetic divergence suggested that colder climates from the late Pliocene and the isolation of marginal seas during the Pleistocene may have driven the divergence of the C. rastrinus species complex.     

Global pattern of phylogenetic species composition of shark and its conservation priority

The diversity of marine communities is in striking contrast with the diversity of terrestrial communities. In all oceans, species richness is low in tropical areas and high at latitudes between 20 and 40°. While species richness is a primary metric used in conservation and management strategies, it is important to take into account the complex phylogenetic patterns of species compositions within communities. We measured the phylogenetic skew and diversity of shark communities throughout the world. We found that shark communities in tropical seas were highly phylogenetically skewed, whereas temperate sea communities had phylogenetically diversified species compositions. Interestingly, although geographically distant from one another, tropical sea communities were all highly skewed toward requiem sharks (Carcharhinidae), hammerhead sharks (Sphyrnidae), and whale sharks (Rhincodon typus). Worldwide, the greatest phylogenetic evenness in terms of clades was found in the North Sea and coastal regions of countries in temperate zones, such as the United Kingdom, Ireland, southern Australia, and Chile. This study is the first to examine patterns of phylogenetic diversity of shark communities on a global scale. Our findings suggest that when establishing conservation activities, it is important to take full account of phylogenetic patterns of species composition and not solely use species richness as a target. Protecting areas of high phylogenetic diversity in sharks, which were identified in this study, could form a broader strategy for protecting other threatened marine species.     

Spatial and temporal patterns of genetic variation in the widespread antitropical deep‐sea coral Paragorgia arborea

Numerous deep‐sea species have apparent widespread and discontinuous distributions. Many of these are important foundation species, structuring hard‐bottom benthic ecosystems. Theoretically, differences in the genetic composition of their populations vary geographically and with depth. Previous studies have examined the genetic diversity of some of these taxa in a regional context, suggesting that genetic differentiation does not occur at scales of discrete features such as seamounts or canyons, but at larger scales (e.g. ocean basins). However, to date, few studies have evaluated such diversity throughout the known distribution of a putative deep‐sea species. We utilized sequences from seven mitochondrial gene regions and nuclear genetic variants of the deep‐sea coral Paragorgia arborea in a phylogeographic context to examine the global patterns of genetic variation and their possible correlation with the spatial variables of geographic position and depth. We also examined the compatibility of this morphospecies with the genealogical‐phylospecies concept by examining specimens collected worldwide. We show that the morphospecies P. arborea can be defined as a genealogical‐phylospecies, in contrast to the hypothesis that P. arborea represents a cryptic species complex. Genetic variation is correlated with geographic location at the basin‐scale level, but not with depth. Additionally, we present a phylogeographic hypothesis in which P. arborea originates from the North Pacific, followed by colonization of the Southern Hemisphere prior to migration to the North Atlantic. This hypothesis is consistent with the latest ocean circulation model for the Miocene.     

Unexpected High Genetic Diversity at the Extreme Northern Geographic Limit of Taurulus bubalis (Euphrasen, 1786)

The longspined bullhead (Taurulus bubalis, Euphrasen 1786) belongs to the family Cottidae and is a rocky shore species that inhabits the intertidal zones of the Eastern Atlantic since Iceland, southward to Portugal and also the North Sea and Baltic, northward to the Gulf of Finland, with some occurrences in the northern Mediterranean coasts eastward to the Gulf of Genoa. We analysed the phylogeographic patterns of this species using mitochondrial and nuclear markers in populations throughout most of its distributional range in west Europe. We found that T. bubalis has a relatively shallow genealogy with some differentiation between Atlantic and North Sea. Genetic diversity was homogeneous across all populations studied. The possibility of a glacial refugium near the North Sea is discussed. In many, but not all, marine temperate organisms, patterns of diversity are similar across the species range. If this phenomenon proves to be most common in cold adapted species, it may reflect the availability of glacial refugia not far from their present-day northern limits.     

The environmental basis of North American species richness patterns among Epicauta (Coleoptera: Meloidae)

Understanding regional variability in species richness is necessary for conservation efforts to succeed in the face of large-scale environmental deterioration. Several analyses of North American vertebrates have shown that climatic energy provides the best explanation of contemporary species richness patterns. The paucity of analyses of insect diversity patterns, however, remains a serious obstacle to a general hypothesis of spatial variation in diversity. We collected species distribution data on a North American beetle genus, Epicauta (Coleoptera: Meloidae) and tested several major diversity hypotheses. These beetles are generally grasshopper egg predators as larvae, and angiosperm herbivores as adults. Epicauta richness is highest in the hot, dry American southwest, and decreases north and east, consistent with the species richness-energy hypothesis. Potential evapotranspiration, which is also the best predictor of richness patterns among North American vertebrates, explains 80.2% of the variability in Epicauta species richness. Net primary productivity and variables measuring climatic heat energy only (such as PET) are not generally comparable, though they are sometimes treated as if they were equivalent. We conclude that the species richness-energy hypothesis currently provides a better overall explanation for Epicauta species richness patterns in North America than other major diversity hypotheses. The observed relationship between climatic energy and regional species richness may provide significant insight into the response of ecological communities to climate change.     

Trophic interactions and abiotic factors drive functional and phylogenetic structure of vertebrate herbivore communities across the Arctic tundra biome

Communities are assembled from species that evolve or colonise a given geographic region, and persist in the face of abiotic conditions and interactions with other species. The evolutionary and colonisation histories of communities are characterised by phylogenetic diversity, while functional diversity is indicative of abiotic and biotic conditions. The relationship between functional and phylogenetic diversity infers whether species functional traits are divergent (differing between related species) or convergent (similar among distantly related species). Biotic interactions and abiotic conditions are known to influence macroecological patterns in species richness, but how functional and phylogenetic diversity of guilds vary with biotic factors, and the relative importance of biotic drivers in relation to geographic and abiotic drivers is unknown. In this study, we test whether geographic, abiotic or biotic factors drive biome‐scale spatial patterns of functional and phylogenetic diversity and functional convergence in vertebrate herbivores across the Arctic tundra biome. We found that functional and phylogenetic diversity both peaked in the western North American Arctic, and that spatial patterns in both were best predicted by trophic interactions, namely vegetation productivity and predator diversity, as well as climatic severity. Our results show that both bottom–up and top–down trophic interactions, as well as winter temperatures, drive the functional and phylogenetic structure of Arctic vertebrate herbivore assemblages. This has implications for changing Arctic ecosystems; under future warming and northward movement of predators potential increases in phylogenetic and functional diversity in vertebrate herbivores may occur. Our study thus demonstrates that trophic interactions can determine large‐scale functional and phylogenetic diversity just as strongly as abiotic conditions.