Global biogeographic regions for ants have complex relationships with those for plants and tetrapods.

On a global scale, biodiversity is geographically structured into regions of biotic similarity. Delineating these regions has been mostly targeted for tetrapods and plants, but those for hyperdiverse groups such as insects are relatively unknown. Insects may have higher biogeographic congruence with plants than tetrapods due to their tight ecological and evolutionary links with the former, but it remains untested. Here, we develop a global regionalization for a major and widespread insect group, ants, based on the most comprehensive distributional and phylogenetic information to date, and examine its similarity to regionalizations for tetrapods and vascular plants. Our ant regionalization supports the newly proposed Madagascan and Sino-Japanese realms based on tetrapod delineations, and it recovers clusters observed in plants but not in tetrapods, such as the Holarctic and Indo-Pacific realms. Quantitative comparison suggests strong associations among different groups-plants showed a higher congruence with ants than with tetrapods. These results underscore the wide congruence of diverse distribution patterns across the tree of life and the similarities shared by insects and plants that are not captured by tetrapod groups. Our analysis highlights the importance of developing global biogeographic maps for insect groups to obtain a more comprehensive geographic picture of life on Earth.

Post-extinction recovery of the Phanerozoic oceans and biodiversity hotspots.

The fossil record of marine invertebrates has long fuelled the debate as to whether or not there are limits to global diversity in the sea1-5. Ecological theory states that, as diversity grows and ecological niches are filled, the strengthening of biological interactions imposes limits on diversity6,7. However, the extent to which biological interactions have constrained the growth of diversity over evolutionary time remains an open question1-5,8-11. Here we present a regional diversification model that reproduces the main Phanerozoic eon trends in the global diversity of marine invertebrates after imposing mass extinctions. We find that the dynamics of global diversity are best described by a diversification model that operates widely within the exponential growth regime of a logistic function. A spatially resolved analysis of the ratio of diversity to carrying capacity reveals that less than 2% of the global flooded continental area throughout the Phanerozoic exhibits diversity levels approaching ecological saturation. We attribute the overall increase in global diversity during the Late Mesozoic and Cenozoic eras to the development of diversity hotspots under prolonged conditions of Earth system stability and maximum continental fragmentation. We call this the 'diversity hotspots hypothesis', which we propose as a non-mutually exclusive alternative to the hypothesis that the Mesozoic marine revolution led this macroevolutionary trend12,13.

Global warming is causing a more pronounced dip in marine species richness around the equator.

The latitudinal gradient in species richness, with more species in the tropics and richness declining with latitude, is widely known and has been assumed to be stable over recent centuries. We analyzed data on 48,661 marine animal species since 1955, accounting for sampling variation, to assess whether the global latitudinal gradient in species richness is being impacted by climate change. We confirm recent studies that show a slight dip in species richness at the equator. Moreover, richness across latitudinal bands was sensitive to temperature, reaching a plateau or declining above a mean annual sea surface temperature of 20 °C for most taxa. In response, since the 1970s, species richness has declined at the equator relative to an increase at midlatitudes and has shifted north in the northern hemisphere, particularly among pelagic species. This pattern is consistent with the hypothesis that climate change is impacting the latitudinal gradient in marine biodiversity at a global scale. The intensification of the dip in species richness at the equator, especially for pelagic species, suggests that it is already too warm there for some species to survive.

Reply to 'Dissimilarity measures affected by richness differences yield biased delimitations of biogeographic realms'.

Recently, we classified the oceans into 30 biogeographic realms based on species' endemicity. Castro-Insua et al. criticize the choices of dissimilarity coefficients and clustering approaches used in our paper, and reanalyse the data using alternative techniques. Here, we explain how the approaches used in our original paper yield results in line with existing biogeographical knowledge and are robust to alternative methods of analysis. We also repeat the analysis using several similarity coefficients and clustering algorithms, and a neural network theory method. Although each combination of methods produces outputs differing in detail, the overall pattern of realms is similar. The coarse nature of the present boundaries of the realms reflects the limited field data but may be improved with additional data and mapping to environmental variables.

Marine biogeographic realms and species endemicity.

Marine biogeographic realms have been inferred from small groups of species in particular environments (e.g., coastal, pelagic), without a global map of realms based on statistical analysis of species across all higher taxa. Here we analyze the distribution of 65,000 species of marine animals and plants, and distinguish 30 distinct marine realms, a similar proportion per area as found for land. On average, 42% of species are unique to the realms. We reveal 18 continental-shelf and 12 offshore deep-sea realms, reflecting the wider ranges of species in the pelagic and deep-sea compared to coastal areas. The most widespread species are pelagic microscopic plankton and megafauna. Analysis of pelagic species recognizes five realms within which other realms are nested. These maps integrate the biogeography of coastal and deep-sea, pelagic and benthic environments, and show how land-barriers, salinity, depth, and environmental heterogeneity relate to the evolution of biota. The realms have applications for marine reserves, biodiversity assessments, and as an evolution relevant context for climate change studies.

Marine Biodiversity, Biogeography, Deep-Sea Gradients, and Conservation.

The oceans appear ideal for biodiversity - they have unlimited water, a large area, are well connected, have less extreme temperatures than on land, and contain more phyla and classes than land and fresh waters. Yet only 16% of all named species on Earth are marine. Species richness decreases with depth in the ocean, reflecting wider geographic ranges of deep sea than coastal species. Here, we assess how many marine species are named and estimated to exist, paying particular regard to whether discoveries of deep-sea organisms, microbes and parasites will change the proportion of terrestrial to marine species. We then review what factors have led to species diversification, and how this knowledge informs conservation priorities. The implications of this understanding for marine conservation are that the species most vulnerable to extinction will be large and endemic. Unfortunately, these species are also the most threatened by human impacts. Such threats now extend globally, and thus the only refuges for these species will be large, permanent, fully protected marine reserves.

Bimodality of Latitudinal Gradients in Marine Species Richness.

The paradigm for the latitudinal gradient in species richness is that it is unimodal with a tropical peak. For 27 published studies, and global datasets of 65 000 recent and 50 000 fossil marine species, we found that almost all datasets were significantly bimodal with a dip in species richness near the equator. The locations of mid-latitude peaks varied between taxa and were higher in the northern hemisphere where the continental shelf is greatest. Our findings support hypotheses of tropical species evolving in response to temperature variation near the edges of the tropics and available high-productivity habitat. They suggest that the equator may already be too hot for some species and that the modes may move further apart due to climate warming.