ForCenS-LGM: a dataset of planktonic foraminifera species assemblage composition for the Last Glacial Maximum.

Species assemblage composition of marine microfossils offers the possibility to investigate ecological and climatological change on time scales inaccessible using conventional observations. Planktonic foraminifera - calcareous zooplankton - have an excellent fossil record and are used extensively in palaeoecology and palaeoceanography. During the Last Glacial Maximum (LGM; 19,000 - 23,000 years ago), the climate was in a radically different state. This period is therefore a key target to investigate climate and biodiversity under different conditions than today. Studying LGM climate and ecosystems indeed has a long history, yet the most recent global synthesis of planktonic foraminifera assemblage composition is now nearly two decades old. Here we present the ForCenS-LGM dataset with 2,365 species assemblage samples collected using standardised methods and with harmonised taxonomy. The data originate from marine sediments from 664 sites and present a more than 50% increase in coverage compared to previous work. The taxonomy is compatible with the most recent global core top dataset, enabling direct investigation of temporal changes in foraminifera biogeography and facilitating seawater temperature reconstructions.

Intrusion of Arabian Sea high salinity water and monsoon-associated processes modulate planktic foraminiferal abundance and carbon burial in the southwestern Bay of Bengal.

The unicellular calcareous planktic foraminifera sequester a significant portion of the carbon dioxide dissolved in the ocean, thus burying the carbon in sediments for millions of years. The global warming and associated processes are likely to affect the planktic foraminiferal abundance and diversity. Therefore, their baseline distribution has to be documented and correlated with ambient parameters to assess its fate under different climate change scenarios. Here, we report an exceptionally high abundance of planktic foraminifera and thus large carbon burial in the southwestern Bay of Bengal. The very high absolute abundance of planktic foraminifera in the Cauvery River basin is attributed to biannual productivity, warmer and saline waters. Globigerinita glutinata is the highest abundant species followed by Globigerinoides ruber and Globigerina bulloides. Globigerina bulloides is abundant on the shelf, where the upwelling is more frequent. The relative abundance of Globorotalia menardii is positively correlated with thermocline salinity and negatively correlated with thermocline temperature. Similarly, Neogloboquadrina dutertrei and Globoquadrina conglomerata are negatively correlated with mixed layer as well as thermocline temperature and mixed layer salinity. Both these species are positively correlated with thermocline salinity. Globigerina falconensis is more abundant in the southernmost transect influenced by intense winter monsoon precipitation. We report that G. ruber prefers high saline and warmer waters with the highest abundance in the southernmost transect. From the foraminiferal distribution, it is evident that the temperature and salinity of the mixed layer as well as thermocline, food availability, and monsoon-associated processes affect the planktic foraminiferal abundance and thus carbon burial in the southwestern Bay of Bengal. The changes in influx of southeastern Arabian Sea water will affect the planktic foraminiferal population and subsequent carbon burial in the southwestern Bay of Bengal.

A matter of choice: Understanding the interactions between epiphytic foraminifera and their seagrass host Halophila stipulacea.

In sub/tropical waters, benthic foraminifera are among the most abundant epiphytic organisms inhabiting seagrass meadows. This study explored the nature of the association between foraminifera and the tropical seagrass species H. stipulacea, aiming to determine whether these interactions are facilitative or random. For this, we performed a "choice" experiment, where foraminifera could colonize H. stipulacea plants or plastic "seagrasses" plants. At the end of the experiment, a microbiome analysis was performed to identify possible variances in the microbial community and diversity of the substrates. Results show that foraminifera prefer to colonize H. stipulacea, which had a higher abundance and diversity of foraminifera than plastic seagrass plants, which increased over time and with shoot age. Moreover, H. stipulacea leaves have higher epiphytic microbial community abundance and diversity. These results demonstrate that seagrass meadows are important hosts of the foraminifera community and suggest the potential facilitative effect of H. stipulacea on epiphytic foraminifera, which might be attributed to a greater diversity of the microbial community inhabiting H. stipulacea.

Impact of pesticides on marine coral reef foraminifera.

Our laboratory study looked into how pesticides affect the foraminifera species Heterostegina depressa and their obligatory algal endosymbionts. We incubated the foraminifera separately with different types of pesticides at varying concentrations (1 %, 0.01 % and 0.0001 %); we included the insecticide Confidor© (active substance: imidacloprid), the fungicide Pronto©Plus (tebuconazole), and the herbicide Roundup© (glyphosate). Our evaluation focused on the symbiont's photosynthetically active area (PA), and the uptake of dissolved inorganic carbon (DIC) and nitrogen (nitrate) to determine the vitality of the foraminifera. Our findings showed that even the lowest doses of the fungicide and herbicide caused irreparable damage to the foraminifera and their symbionts. While the insecticide only deactivated the symbionts (PA = 0) at the highest concentration (1 %), the fungicide, and herbicide caused complete deactivation even at the lowest levels provided (0.0001 %). The fungicide had the strongest toxic effect on the foraminiferal host regarding reduced isotope uptake. In conclusion, all pesticides had a negative impact on the holosymbiont, with the host showing varying degrees of sensitivity towards different types of pesticides.

New xenophyophores (Foraminifera, Monothalamea) from the eastern Clarion-Clipperton Zone (equatorial Pacific).

Xenophyophores are large, agglutinated foraminifera that dominate the benthic megafauna in some parts of the deep sea. Here, we describe an assemblage of largely fragmentary specimens from the Clarion-Clipperton Zone (CCZ), an area of the eastern abyssal Pacific hosting large, commercially significant deposits of polymetallic nodules. We recognised 18 morphospecies of which eight yielded DNA sequences. These include two new genera and three new species, Claraclippia seminuda gen. & sp. nov., Stereodiktyoma mollis gen. & sp. nov., and Aschemonella tani sp. nov., three that are assigned to known species, Abyssalia foliformis, Aschemonella monilis and Shinkaiya contorta, and two assigned to open nomenclature forms Abyssalia aff. foliformis and Stannophyllum aff. granularium. An additional ten forms are represented only by morphology. The following seven are placed in known genera, species and open-nomenclature forms: Aschemonella? sp., Homogammina sp., Psammina multiloculata, P. aff. multiloculata, P. aff. limbata form 1 sensu Gooday et al., 2018, P. aff. limbata form 2 sensu Gooday et al., 2018, and Stannophyllum spp. The other three could not be identified to genus level. This new collection brings the total of described and undescribed species and morphotypes from the CCZ to 27 and 70, respectively, reinforcing the already high diversity of xenophyophores known from this part of the Pacific.

Deciphering the impact of decabromodiphenyl ether (BDE-209) on benthic foraminiferal communities: Insights from Cell-Tracker Green staining and eDNA metabarcoding.

This study investigates the ecotoxicological effects of BDE-209, a persistent organic pollutant (POP) prevalent in Kuwait's coastal-industrial areas, on benthic foraminiferal communities. We conducted a mesocosm experiment in which we exposed benthic foraminiferal communities sampled from the coastal-industrial areas of Kuwait to a gradient of BDE-209 concentrations (0.01 to 20 mg/kg). The impact of exposure was assessed using live-staining and metabarcoding techniques. Despite the significantly different taxonomic compositions detected by the two techniques, our results show that BDE-209 significantly affects foraminiferal communities, with moderately high concentrations leading to reduced α-diversity and considerable taxonomic shifts in both molecular and morphological assemblages. At concentrations of 10 and 20 mg/kg, no living foraminifera were detected after 8 weeks, suggesting a threshold for their survival under BDE-209 exposure. The parallel responses of molecular and morphological communities confirm the reliability of both assessment methods. This study is the first to investigate the reaction of eukaryotic communities, specifically foraminifera, to POPs such as BDE-209, generating valuable insights that have the potential to enhance field studies and aid the refinement of sediment quality guidelines.

Assigning the unassigned: A signature-based classification of rDNA metabarcodes reveals new deep-sea diversity.

Environmental DNA metabarcoding reveals a vast genetic diversity of marine eukaryotes. Yet, most of the metabarcoding data remain unassigned due to the paucity of reference databases. This is particularly true for the deep-sea meiofauna and eukaryotic microbiota, whose hidden diversity is largely unexplored. Here, we tackle this issue by using unique DNA signatures to classify unknown metabarcodes assigned to deep-sea foraminifera. We analyzed metabarcoding data obtained from 311 deep-sea sediment samples collected in the Clarion-Clipperton Fracture Zone, an area of potential polymetallic nodule exploitation in the Eastern Pacific Ocean. Using the signatures designed in the 37F hypervariable region of the 18S rRNA gene, we were able to classify 802 unassigned metabarcodes into 61 novel lineages, which have been placed in 27 phylogenetic clades. The comparison of new lineages with other foraminiferal datasets shows that most novel lineages are widely distributed in the deep sea. Five lineages are also present in the shallow-water datasets; however, phylogenetic analysis of these lineages separates deep-sea and shallow-water metabarcodes except in one case. While the signature-based classification does not solve the problem of gaps in reference databases, this taxonomy-free approach provides insight into the distribution and ecology of deep-sea species represented by unassigned metabarcodes, which could be useful in future applications of metabarcoding for environmental monitoring.

Foraminifera as a model of eukaryotic genome dynamism.

In contrast to the canonical view that genomes cycle only between haploid and diploid states, many eukaryotes have dynamic genomes that change content throughout an individual's life cycle. However, the few detailed studies of microeukaryotic life cycles render our understanding of eukaryotic genome dynamism incomplete. Foraminifera (Rhizaria) are an ecologically important, yet understudied, clade of microbial eukaryotes with complex life cycles that include changes in ploidy and genome organization. Here, we apply fluorescence microscopy and image analysis techniques to over 2,800 nuclei in 110 cells to characterize the life cycle of Allogromia laticollaris strain Cold Spring Harbor (CSH), one of few cultivable foraminifera species. We show that haploidy and diploidy are brief moments in the A. laticollaris life cycle and that A. laticollaris nuclei endoreplicate up to 12,000 times the haploid genome size. We find that A. laticollaris reorganizes a highly endoreplicated nucleus into thousands of haploid genomes through a non-canonical mechanism called Zerfall, in which the nuclear envelope degrades and extrudes chromatin into the cytoplasm. Based on these findings, along with changes in nuclear architecture across the life cycle, we believe that A. laticollaris uses spatio-temporal mechanisms to delineate germline and somatic DNA within a single nucleus. The analyses here extend our understanding of the genome dynamics across the eukaryotic tree of life.IMPORTANCEIn traditional depictions of eukaryotes (i.e., cells with nuclei), life cycles alternate only between haploid and diploid phases, overlooking studies of diverse microeukaryotic lineages (e.g., amoebae, ciliates, and flagellates) that show dramatic variation in DNA content throughout their life cycles. Endoreplication of genomes enables cells to grow to large sizes and perhaps to also respond to changes in their environments. Few microeukaryotic life cycles have been studied in detail, which limits our understanding of how eukaryotes regulate and transmit their DNA across generations. Here, we use microscopy to study the life cycle of Allogromia laticollaris strain CSH, an early-diverging lineage within the Foraminifera (an ancient clade of predominantly marine amoebae). We show that DNA content changes significantly throughout their life cycle and further describe an unusual process called Zerfall, by which this species reorganizes a large nucleus with up to 12,000 genome copies into hundreds of small gametic nuclei, each with a single haploid genome. Our results are consistent with the idea that all eukaryotes demarcate germline DNA to pass on to offspring amidst more flexible somatic DNA and extend the known diversity of eukaryotic life cycles.

Benthic foraminifera in Gulf of Mexico show temporal and spatial dynamics of microplastics.

Microplastics have accumulated in the environment since plastic production began, with present-day observations that range from marine trenches to mountains. However, research on microplastics has only recently begun so it is unclear how they have changed over time in many oceanic regions. Our study addressed this gap by quantifying the temporal and spatial dynamics of microplastics in two deep-water regions of the Gulf of Mexico (GOM). We isolated agglutinated foraminifera from sediment cores and assessed microplastics that were incorporated into their tests. Our results indicated that microplastics were incorporated by agglutinated foraminifera after plastic production began. Microplastics were higher at deep-water sites and closer to the Mississippi River. This study confirms the presence of microplastic incorporation into agglutinated foraminifera tests and investigates microplastics in deep-water sediments in the GOM. Additional work is needed to fully identify the distribution of microplastics across the GOM and other oceanic basins.

Assessment of the ecological quality status of the Sepetiba Bay (SE Brazil): When metabarcoding meets morphology on foraminifera.

In recent years, the region surrounding Sepetiba Bay (SB; SE Brazil) has become a hub of intense urban expansion and economic exploitation in response to ore transport and industrial and port activities. As a result, contaminants have been introduced into the bay, leading to an overall worsening of the environmental quality. The present work applies for the first time a foraminiferal morphology-based approach (M) and eDNA-based metabarcoding sequencing (G), along with geochemical data to assess the ecological quality status (EcoQS) in the SB. Principal component analysis shows that the eDNA and morphospecies diversity as well as most of the taxa relative abundance decline in response to the environmental stress (ES) gradient related to total organic carbon (TOC) and metal pollution. Based on ecological indices, Exp(H'bc) (G), Exp(H'bc) (M), foraminifera ATZI marine biotic index (Foram-AMBI), Foram Stress Index (FSI), and geochemical indices (TOC and Potential Ecological Risk Index), the lowest values of EcoQS (i.e., bad to moderate) are inferred in the innermost part of the SB. Despite minor discrepancies among the six EcoQS indices, an agreement has been found for 63% of the stations. To improve the agreement between the ecological indices, it is necessary to fill the gap in species ecology; information on the ecology of many species is still unknown. This work reinforces the importance of molecular analysis and morphological methods in environmental impact studies and confirms the reliability of foraminiferal metabarcoding in EcoQS assessment. This is the first study evaluating the EcoQS in the South Atlantic by using combined foraminiferal eDNA metabarcoding with morphological data.

An 800-year record of benthic foraminifer images and 2D morphometrics from the Santa Barbara Basin.

The Santa Barbara Basin is an extraordinary archive of environmental and ecological change, where varved sediments preserve microfossils that provide an annual to decadal record of the dynamics of surrounding ecosystems. Of the microfossils preserved in these sediments, benthic foraminifera are the most abundant seafloor-dwelling organisms. While they have been extensively utilized for geochemical and paleoceanographic work, studies of their morphology are lacking. Here we use a high-throughput imaging method (AutoMorph) designed to extract 2D data from photographic images of fossils to produce a large image and 2D shape dataset of recent benthic foraminifera from two core records sampled from the center of the Santa Barbara Basin that span an ~800-year-long interval during the Common Era (1249-2008 CE). Information on more than 36,000 objects is included, of which more than 22,000 are complete or partially-damaged benthic foraminifera. The dataset also includes other biogenic microfossils including ostracods, pteropods, diatoms, radiolarians, fish teeth, and shark dermal denticles. We describe our sample preparation, imaging, and identification techniques, and outline potential data uses.

Influence of methane and other hydrocarbon gases on foraminifera and nematodes in the Northwestern part of the Black Sea.

This study investigates the response of foraminifera and nematodes to different types of methane and other hydrocarbon gases (alkanes, alkins) in bottom sediments of the northwestern shelf and continental slope of the Black Sea. Foraminifera are represented by 39 species from six orders, 13 families, 24 genera. Nematoda are represented by 48 species from six orders, 17 families, 27 genera. Both groups of meiobenthos are characterized by impoverished simple diversity and abundance/density as well as by much smaller sizes of foraminiferal tests compared to those from areas not affected by hydrocarbon gases, indicating stressful conditions for their life in the study areas. Among foraminifera, calcareous hyaline taxa dominate, but agglutinated forms are present as well and occasionally play a dominant role. A significant number of the identified nematode species belong to the tyobiotic type from the families Comesomatidae and Linhomoidae. They are characterized by a long, thin body and a small or completely undeveloped oral cavity, indicating their feeding on small particles of sediment or bacteria. Nine groups of foraminifera and nematodes were identified according to their tolerance and sensitivity to methane (15 species), ethane (five species), propane (eight species), butane (10 species), pentane (11 species), ethylene (20 species), propylene (two species), acetylene (three species), and isobutylene (one species). This study provides evidence for the first time that different types of methane, its homologues, and other hydrocarbon gases affect the reproduction, diversity, abundance, and growth of foraminifera, as well as density and simple diversity of nematodes.

Azoic sediments and benthic foraminifera: Environmental quality in a subtropical coastal lagoon in the gulf of California.

Marine transitional environments play an important role in human sustainability. Around these ecosystems, coastal lagoons are subject to high anthropogenic pressure from population growth. The increased demand for goods and services is associated with the elevated discharge of untreated and treated wastewater into lagoon systems. The absence of benthic organisms in lagoon environments has been linked to extreme natural conditions and severe anthropogenic impact at both spatial and temporal scales. However, the mechanisms that lead to the presence of azoic sediments in lagoon environments have yet to be studied. This study aimed to determine the vertical variability of textural groups, geochemistry, and benthic foraminiferal fauna to understand how natural and anthropogenic components generate a vertical sediment sequence with low or absent benthic foraminifera in a subtropical coastal lagoon in the southwestern end of the Gulf of California. A 41 cm-long sediment core was collected from La Paz Lagoon at a 1-m depth. The core was sectioned every centimeter, and sediment subsamples were dried and homogenized for grain size, calcium carbonate, elemental and isotopic carbon and nitrogen analyses, and benthic foraminifera quantification. Muds with fine sands towards the core's base characterized the sedimentary sequence. Organic carbon and total nitrogen increased from the base (1.4% and 0.06%, respectively) to the core-top (CT, 3.0% and 0.14%, respectively), significant from the 27 cm interval. Calcium carbonate content was very low (<0.8%). The relationship of δ13C vs. C:N ratio indicated that sedimentary organic carbon was derived from the marine and sewage source mixture. The δ15N of organic matter increased by 3.7‰, starting from the 27 cm interval towards the CT. The nitrogen sewage input source was relatively more significant than nitrogen fixation. The few individuals (<18 ind. in 10 g) and genera (Ammonia and Elphidium), as well as the absence of foraminifera in 19 of 41 intervals in the core, indicated that environmental conditions were unfavorable, even for colonization of environmentally stress-tolerant genera. The frequency of azoic sediments was higher from the 25 cm interval to the CT vs. from the base to the 25 cm interval. Moreover, the AEI revealed severe to moderate hypoxia in the study area. The limited presence of benthic foraminifera and calcium carbonate preservation corroborated that the quality of the lagoon's environment has deteriorated along with population growth, which requires strategic programs to sustain this transitional ecosystem.

Benthic foraminifera as bioindicators of marine pollution: A bibliometric approach to unravel trends, patterns and perspectives.

Benthic foraminifera, single-celled marine organisms, are known for their wide distribution, high abundance and species diversity, test (i.e., shell) preservation in the sedimentary (e.g., historical) record, and sensitivity to environmental changes. Because of these characteristics, they have been widely used as bioindicators in environmental monitoring and, more recently, as Biological Quality Elements (BQEs) in the Ecological Quality Status (EcoQS) evaluation. The global scientific literature on benthic foraminifera as bioindicators was gathered from the Scopus database (overall 966 papers from 1973 to 2022) and explored with scientometric software. The outcomes highlight that the investigation of benthic foraminiferal response to pollutants started over 50 years ago. Indeed, not only the number of published documents has recently peaked (i.e., 2021 and 2022) but there has been also a growth in the percentages of papers falling within the Decision Sciences category that deals with the application of foraminiferal indices for the EcoQS assessment.

Morphology and molecular phylogeny of two new species of Psammophaga (Rhizaria, Foraminifera) from the west coast of India.

Benthic foraminifera are unicellular eukaryotic protists that construct an organic, agglutinated, or calcareous test wall. Although single-chambered (monothalamous) foraminifera are ubiquitous in marine habitats worldwide, they are poorly known compared to their multi-chambered relatives, notably from the tropical marine environments of India. In this study, we describe two new species of marine monothalamid genus Psammophaga Arnold, 1982, from the Rajapuri Creek, coastal Maharashtra, India (Arabian Sea). Psammophaga holzmannae sp. nov. is ovoid to spherical shaped, 103-246 µm in length, single aperture, translucent to orange color cytoplasm, outer surface is composed of agglutinated fine clay particles, and ingested mineral grains are concentrated near its aperture. Psammophaga sinhai sp. nov. is oblong, elliptical, or droplet-shaped, 279-448 µm in length, single aperture, yellow olive color cytoplasm, the exterior surface formed of agglutinated fine clay particles, and the ingested mineral grains are dispersed throughout the body. Phylogenetic analyses based on partial small subunit rRNA gene sequences position new species within the Clade E of monothalamids and are genetically distinct from other Psammophaga. Elemental (SEM-EDS) analysis of engulfed mineral grains revealed preferential selection and uptake of heavy opaque titaniferous minerals from the ambient environment in the cytoplasm.

Response of Arctic benthic foraminiferal traits to past environmental changes.

The Arctic is subjected to all-encompassing disruptions in marine ecosystems caused by anthropogenic warming. To provide reliable estimates of how future changes will affect the ecosystems, knowledge of Arctic marine ecosystem responses to past environmental variability beyond the instrumental era is essential. Here, we present a novel approach on how to evaluate the state of benthic marine biotic conditions during the deglacial and Holocene period on the Northeast Greenland shelf. Benthic foraminiferal species were assigned traits (e.g., oxygen tolerance, food preferences) aiming to identify past faunal changes as a response to external forcing mechanisms. This approach was applied on sediment cores from offshore Northeast Greenland. We performed numerical rate-of-change detection to determine significant changes in the benthic foraminiferal traits. That way, the significant abrupt trait changes can be assessed across sites, providing a better understanding of the impact of climate drivers on the traits. Our results demonstrate that during the last ~ 14,000 years, bottom water oxygen is the main factor affecting the variability in the benthic foraminiferal faunas in this area. Our results show that significant changes in the traits correspond to drastic climate perturbations. Specifically, the deglacial-Holocene transition and mid-Holocene warm period exhibited significant change, with several trait turnovers.

Assessing the effect of Amphistegina lobifera invasion on infralittoral benthic foraminiferal assemblages in the Sicily Channel (Central Mediterranean).

This article documents, through a quantitative approach, the negative effect of the highly invasive species Amphistegina lobifera Larsen, 1976 on native benthic foraminiferal assemblages of coastal areas in the Sicily Channel (Central Mediterranean). A nested sampling design was applied through the comparison of benthic foraminiferal community structure across three areas that are known to be at different stages of invasion (i.e. Maltese Islands - advanced, southern Sicily - medium, and eastern Sicily - early). Results suggested that both diversity and richness of benthic foraminiferal community from the Maltese Islands were strongly modified by increased abundances of A. lobifera. In contrast, this phenomenon is less evident in southern and eastern Sicily, where the invader displayed lower abundances and the community structure was more diversified. Collected data also allowed for predicting what could happen in the near future in the whole Sicily Channel, as well as in the rest of the Mediterranean Sea.

Collection of X-ray micro computed tomography images of shells of planktic foraminifera with curated taxonomy.

Calcite shells of planktic foraminifera (Protista, Rhizaria) constitute a large portion of deep-sea sediments. The shells are constructed by sequential addition of partly overlapping chambers with diverse shapes, resulting in complex shell architectures, which are genetically fixed and diagnostic at the species level. The characterisation of the complete architecture requires three-dimensional imaging of the shell, including the partially or entirely covered juvenile chambers. Here we provide reconstructed x-ray micro computed tomography image stacks of 179 specimens of extant planktic foraminifera collected from plankton tows, sediment traps and surface sediments. The specimens have fully resolved and curated taxonomy and represent 43 of the currently recognised 48 holoplanktic species and subspecies. The image stacks form a basis for further applications, such as the characterisation of the architectural morphospace of the extant taxa, allowing studies of species functional ecology, calcification intensity and reconstructions of phylogenetic relationships.

Shared ancestry of algal symbiosis and chloroplast sequestration in foraminifera.

Foraminifera are unicellular organisms that established the most diverse algal symbioses in the marine realm. Endosymbiosis repeatedly evolved in several lineages, while some engaged in the sequestration of chloroplasts, known as kleptoplasty. So far, kleptoplasty has been documented exclusively in the rotaliid clade. Here, we report the discovery of kleptoplasty in the species Hauerina diversa that belongs to the miliolid clade. The existence of kleptoplasty in the two main clades suggests that it is more widespread than previously documented. We observed chloroplasts in clustered structures within the foraminiferal cytoplasm and confirmed their functionality. Phylogenetic analysis of 18S ribosomal RNA gene sequences showed that H. diversa branches next to symbiont-bearing Alveolinidae. This finding represents evidence of of a relationship between kleptoplastic and symbiotic foraminifera.. Analysis of ribosomal genes and metagenomics revealed that alveolinid symbionts and kleptoplasts belong to the same clade, which suggests a common ancestry.

The odd couple of protistology: Edward Heron-Allen (1861-1943) and Arthur Earland (1866-1958).

Edward Heron-Allen and Arthur Earland were among the last great amateur foraminifera researchers. Their partnership began in 1907 and ended in 1932. While close in age to one another, they shared little more than a fascination for forams and a lack of any university training. In most other aspects, the two men were completely different. Heron-Allen was a famous upper class polymath, expert not only on forams, but also on the Persian language, violins, palm reading, history, asparagus, and barnacles. He was also an accomplished novelist and poet, who frequented literary circles. In contrast to the flamboyant Heron-Allen, Earland was a discrete civil servant who admitted to working on forams as a relief from the monotony of his job. Hence, the two were improbable partners. However, together they produced 39 substantial works on forams. Their studies concerned assemblages from Southern Ocean to the North Sea and they are today credited with the original description of 186 species. Here the distinct lives of the two men are presented, and their contributions to protistology, as partners as well as individuals, are reviewed. The case is made for considering Earland's work as neglected relative to that of Heron-Allen, except perhaps by foram taxonomists.