Planktonic foraminiferal species identification is central to many paleoceanographic studies, from selecting species for geochemical research to elucidating the biotic dynamics of microfossil communities relevant to physical oceanographic processes and interconnected phenomena such as climate change. However, few resources exist to train students in the difficult task of discerning amongst closely related species, resulting in diverging taxonomic schools that differ in species concepts and boundaries. This problem is exacerbated by the limited number of taxonomic experts. Here we document our initial progress toward removing these confounding and/or rate-limiting factors by generating the first extensive image library of modern planktonic foraminifera, providing digital taxonomic training tools and resources, and automating species-level taxonomic identification of planktonic foraminifera via machine learning using convolution neural networks. Experts identified 34,640 images of modern (extant) planktonic foraminifera to the species level. These images are served as species exemplars through the online portal Endless Forams (endlessforams.org) and a taxonomic training portal hosted on the citizen science platform Zooniverse (zooniverse.org/projects/ahsiang/ endless-forams/). A supervised machine learning classifier was then trained with~27,000 images of these identified planktonic foraminifera. The best-performing model provided the correct species name for an image in the validation set 87.4% of the time and included the correct name in its top three guesses 97.7% of the time. Together, these resources provide a rigorous set of training tools in modern planktonic foraminiferal taxonomy and a means of rapidly generating assemblage data via machine learning in future studies for applications such as paleotemperature reconstruction.

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The size structure of plankton communities is an important determinant of their functions in marine ecosystems. However, few studies have quantified how organism size varies within species across biogeographical scales. Here, we investigate how planktonic foraminifera, a ubiquitous zooplankton group, vary in size across the tropical and subtropical oceans of the world. Using a recently digitized museum collection, we measured shell area of 3,799 individuals of nine extant species in 53 seafloor sediments. We first analyzed potential size biases in the collection. Then, for each site, we obtained corresponding local values of mean annual sea‐surface temperature (SST), net primary productivity (NPP), and relative abundance of each species. Given former studies, we expected species to reach largest shell sizes under optimal environmental conditions. In contrast, we observe that species differ in how much their size variation is explained by SST, NPP, and/or relative abundance. While some species have predictable size variation given these variables ( Trilobatus sacculifer, Globigerinoides conglobatus, Globigerinella siphonifera, Pulleniatina obliquiloculata, Globorotalia truncatulinoides ), other species show no relationships between size and the studied covariates ( Globigerinoides ruber , Neogloboquadrina dutertrei , Globorotalia menardii, Globoconella inflata ). By incorporating intraspecific variation and sampling broader geographical ranges compared to previous studies, we conclude that shell size variation in planktonic foraminifera species cannot be consistently predicted by the environment. Our results caution against the general use of size as a proxy for planktonic foraminifera environmental optima. More generally, our work highlights the utility of natural history collections and the importance of studying intraspecific variation when interpreting macroecological patterns.

Biodiversity is expected to change in response to future global warming. However, it is difficult to predict how species will track the ongoing climate change. Here we use the fossil record of planktonic foraminifera to assess how biodiversity responded to climate change with a magnitude comparable to future anthropogenic warming. We compiled time series of planktonic foraminifera assemblages, covering the time from the last ice age across the deglaciation to the current warm period. Planktonic foraminifera assemblages shifted immediately when temperature began to rise at the end of the last ice age and continued to change until approximately 5,000 years ago, even though global temperature remained relatively stable during the last 11,000 years. The biotic response was largest in the mid latitudes and dominated by range expansion, which resulted in the emergence of new assemblages without analogues in the glacial ocean. Our results indicate that the plankton response to global warming was spatially heterogeneous and did not track temperature change uniformly over the past 24,000 years. Climate change led to the establishment of new assemblages and possibly new ecological interactions, which suggests that current anthropogenic warming may lead to new, different plankton community composition.

Anthropogenic climate change is altering global biogeographical patterns. However, it remains difficult to quantify how bioregions are changing because pre‐industrial records of species distributions are rare. Marine microfossils, such as planktonic foraminifera, are preserved in seafloor sediments and allow the quantification of bioregions in the past. Using a recently compiled data set of pre‐industrial species composition of planktonic foraminifera in 3802 worldwide seafloor sediments, we employed multivariate and statistical model‐based approaches to study spatial turnover in order to 1) quantify planktonic foraminifera bioregions and 2) understand the environmental drivers of species turnover. Four latitudinally banded bioregions emerge from the global assemblage data. The polar and temperate bioregions are bi‐hemispheric, supporting the idea that planktonic foraminifera species are not limited by dispersal. The equatorial bioregion shows complex longitudinal patterns and overlaps in sea surface temperature (SST) range with the tropical bioregion. Compositional‐turnover models (Bayesian bootstrap generalised dissimilarity models) identify SST as the strongest driver of species turnover. The turnover rate is constant across most of the SST gradient, showing no SST threshold values with rapid shifts in species composition, but decelerates above 25°C, suggesting SST is less predictive of species composition in warmer waters. Other environmental predictors affect species turnover non‐linearly, and their importance differs across regions. In the Pacific ocean, net primary productivity below 500 mgC m−2 day−1 drives fast compositional change. Water depth values below 3000 m (which affect calcareous microfossil preservation) increasingly drive changes in species composition among death assemblages in the Pacific and Indian oceans. Together, our results suggest that the dynamics of planktonic foraminifera bioregions are expected to be highly responsive to climate change; however, at lower latitudes, environmental drivers other than SST may affect these dynamics.

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The Henry Buckley Collection of Planktonic Foraminifera at the Natural History Museum in London (NHMUK) consists of 1665 single-taxon slides housing 23 897 individuals from 203 sites in all the major ocean basins, as well as a vast research library of Scanning Electron Microscope (SEM) photomicrographs. Buckley picked the material from the NHMUK Ocean-Bottom Deposit Collection and also from fresh tow samples. However, his collection remains largely unused as he was discouraged by his managers in the Mineralogy Department from working on or publicizing the collection. Nevertheless, Buckley published pioneering papers on isotopic interpretation of oceanographic and climatic change and was one of the first workers to investigate foraminiferal wall structure using the SEM technique. Details of the collection and images of each slide are available via the NHMUK Data Portal (http://dx.doi.org/10.5519/0035055). The Buckley Collection and its associated Ocean-Bottom Deposit Collection have great potential for taxon-specific studies as well as geochemical work, and both collections are available on request