Introduction

General description

Project description

Sampling methods

Sampling description: 

A set of rules and guidance was created to generate a robust standardised checklist that would meet the conditions of representation and inclusion, systematic harmonisation, proofing, transparency and dynamism. Based on these rules in order to generate the revised biodiversity checklist across groups, we constructed a generalised stepwise validation procedure tailored for eukaryotes and prokaryotes (see Fig. 1).

Figure 1.  

Biodiversity checklist compilation protocol.

List compilation. To generate an initial compendium of Antarctic biodiversity, we retrieved all the species listed in the existing major diversity compendia for the continent. We started with the Wauchope et al. (2019) list of species as a backbone and added further taxa. A list of parasitic animal species records was included from the Barbosa and Palacios (2009) compendium. In the case of Rotifera, a pre-existing checklist of species was available in the AntaBIF database to GBIF (https://www.gbif.org/dataset/b109fc97-b7b8-4432-a89a-5eaaadeee431). Additional lists of bryophytes were obtained from the Ochyra et al. (2008) (moss) and Bednarek-Ochyra et al. (2000) (liverworth) compendia. A complementary list of lichens was retrieved from Øvstedal and Lewis Smith (2001)'s compendium. A list of non-lichenised fungal species was retrieved from the BAS fungal database (Bridge et al. 2010). The SAR group redefines the previous terminology of the paraphyletic group of “protists” and includes the SAR subclades of Stramenopiles, Alveolata and Rhizaria, along with Amoeobozoa and others (Ruggiero et al. 2015). The latter are not related to SAR and we merely used them here in line with current convention. The works of Adams et al. 2006 and Thompson et al. 2019 were taken as the source compendium of SAR species. The initial diatom species list was retrieved from the recent and comprehensive biogeographical study of Verleyen et al. (2021) which includes known freshwater species; sub-Antarctic species listed in that study were excluded. The list was supplemented with information on terrestrial diatom species recorded from Antarctica in the 20^th and 21^stcenturies and taxonomic papers describing freshwater diatom taxa published after 2021. In the absence of published compendia, additional lists of species representing other SAR groups and Amoebae were retrieved directly from the relevant literature. To obtain a list of bacterial and archaeal genera found in soil and freshwater Antarctic environments, we re-analysed data from publicly available 16S rRNA gene datasets (PROKARYOTIC STUDY SOURCE LIST data set). Eukaryotic viruses and bacteriophages remain very poorly described in Antarctica. We, therefore, only identify the main lineages reported from the region in recent literature.

Lastly, for groups without an initial species list, we retrieved candidate species from review works and/or Antarctic literature searches. Firstly, participant curator co-authors screened seminal book compendia. Next, a complementary screening of literature was performed for inclusion/updating from recent discoveries. The species inventories were augmented with recently published findings (updated to October 2023). Antarctic literature searches were done in Web of Science repository using basic terms per group with the following syntax e.g. “Taxon*” AND Antarctic*”. The complete list of queries can be seen in Suppl. material 1. Participant parataxonomist co-authors were assigned subset lists of works around the taxon/taxa queried according to their main expertise. From the extracted literature, we first excluded all studies that were not specifically conducted in the continent of Antarctica. Second, we browsed through the titles and abstracts of literature to identify works that could contain previously unreported species. For instance, species inventories for some groups of flora and fauna have been recently reviewed with regards to the existence of specific type locality data (Phillips et al. 2022) providing the opportunity to grow the dataset with previous omissions and/or new species detections. Additional annotations were made with regards of the content of the manuscripts, but these were not subsequently analysed here. A particular focus was given to the publications of the last five years. The possibility exists that further new material exists, but the intention of this exercise is to be dynamically active and that further revisions will be made to the living inventories over time.

Geographic coverage

Taxonomic coverage

Traits coverage

Temporal coverage

Collection data

Usage licence

Data resources

Additional information

Results and discussion

1. Numbers of identifiable Antarctic biodiversity units and recent biodiversity findings

A total of 1870 eukaryotic species with a currently-accepted status and a total of 349,966 prokaryotic ASVs were obtained from our collations. Thirty of the 149 green algae listed as pseudo-species in Wauchope et al. (2019) were accepted by GBIF. This indicates that much of the chlorophyte taxonomy is still unconfirmed. A total of 468 non-lichenised fungi listed by Bridge et al. (2010) were also recognised in MicoBank. We report bacterial (Datasets 2-3 of the Data Package) and archaeal (Datasets 4-5 of the Data package) diversity by presenting the number of genera and ASVs found for the Antarctic soil and freshwater datasets.

ANIMALIA (METAZOA) KINGDOM 470 species total in version 1.0.

Breeding vertebrates. Twenty-six species in version 1.0. Amongst marine vertebrates utilising the continent, no new species descriptions have been proposed in more than a century. However, the taxonomy of a few species of macrofauna has been revisited as a result of advances in molecular biological methodologies that can be applied in describing their phylogenetic relationships (e.g. Techow et al. (2009), Vianna et al. (2020)).

Arthropods. A total of 189 species in version 1.0. In contrast to their marine counterparts in the Southern Ocean, arthropods are not a dominant component of Antarctic freshwater fauna, often occurring in low abundances and diversity. Our latest knowledge of the distribution of non-marine freshwater arthropods comes from Díaz et al. (2019). Only three new freshwater species have been named since the start of the 21^st century (Diacyclops kaupi, D. walkeri and D. joycei) by Karanovic et al. (2014). Arthropods are, however, prominent in soil ecosystems. The species inventories and phylogeography amongst terrestrial arthropod invertebrates, such as springtails (e.g. Carapelli et al. (2020)), midges and mites (e.g. Brunetti et al. (2021), Collins et al. (2023)) continues to be explored with sometimes highly challenging reassessments (Stevens and D'Haese 2016). In addition, new species of invertebrate ectoparasites continue to be reported (Barbosa and Palacios 2009, Montero et al. 2016).

Non-arthropod invertebrates. A total of 254 species in version 1.0. Amongst other animal phyla, there has been a recent surge in descriptions of new species of terrestrial and freshwater non-arthropod invertebrates, including both Antarctic regional or short-range endemic species, again encouraged by the application of advanced molecular phylogenetic and phylogeographic techniques. This also includes multiple instances of the identification of species-level (or greater) evolutionary divergences within species considered to date as single species and of the presence of cryptic speciation, with multiple such species yet to be formally described. Latest examples include tardigrades (e.g. Short et al. (2022)) and nematodes (e.g. Bostrom et al. (2011)), but not rotifers, for which species seem to be broadly distributed in Antarctica (Cakil et al. 2021). In addition, new species of invertebrate gastroparasites continue to be reported (Montero et al. 2016). Overall, new systematic re-classification techniques, based on molecular markers, are being explored in diverse microscopic animal groups, such as tardigrades (Vecchi et al. 2016). Inventories have recently been compiled and subsequently revised and updated for some invertebrate groups, such as rotifers (Iakovenko et al. 2015, Garlasché et al. 2019).

PLANTAE KINGDOM. A total of 306 species in version 1.0.

Embryophyte plants. A total of 154 species in version 1.0. Angiosperms and bryophytes represent one of the best known groups in Antarctica. Only two vascular plant species occur in Antarctica, with both having wider sub-Antarctic and South American distributions; therefore, no new species descriptions have been made for quite some time. Contemporary diversity research is now examining their precise evolutionary identities (Biersma et al. 2020). This is also applicable to bryophytes, which are relatively well known, though their taxonomical relationships are still being unveiled (Biersma et al. 2018, Camara et al. 2019).

Green algae. A total of 152 species in version 1.0. New species of free-living green algae are expected to be described as we continue to explore the continent. In turn, recent studies examine green algae diversity as photobionts in lichens (Ruprecht et al. 2012, Garrido‐Benavent and Pérez‐Ortega 2017) and as free-living algal forms in snow and water systems (Davey et al. 2019)

FUNGI KINGDOM. A total of 997 species in version 1.0.

Fungi. A total of 871 Ascomycota species, 95 Basidiomycota and 41 Zygomycota and others (in version 1.0). New species of lichen-forming fungi are still being described (e.g. Garrido-Benavent et al. 2016) with currently over 500 known entities identifiable in our dataset), but also current research also focuses on disentangling phylogenetic relationships (e.g. Lagostina et al. (2017)). Non-lichenised fungi represent a more cryptic group that continues to be explored, but a substantial number (> 500) of globally-distributed species have been recorded from Antarctica (Bridge et al. 2010).

AMOEBA, PROTOZOA (Ciliata and Flagellata) AND SAR/CHROMISTA KINGDOM COMPLEXES. 434 SPECIES.

SAR. A total of 418 species in version 1.0. Amongst the SAR supercomplex, biogeographical research in diatoms (Stramenophyles, Ochrophyta, Bacillariophyceae) is a growing discipline (Verleyen et al. 2021). Here, we detected 287 diatom species in total. In addition, five Xanthophyta species were also isolated from biogeographical studies (Broady 1996). However, studies on the diversity of heterotrophic protists (Alveolata, Rhizaria) are a bit more stagnant, but see pioneer efforts (Thompson et al. 2019). Their current known diversity to our knowledge involves A-R 130 species plus one Chrysophyceae.

Protozoa. Seven species in version 1.0. Flagellata and Ciliophora are also inconsistently recorded, with systematic observations in but a few regions (Adams et al. 2006). Additionally, new species continue to be described (Park et al. 2020). Only seven Antarctic protozoan species were found here, representing one of the largest Antarctic biodiversity knowledge gaps.

Oomycota (Chromista). Four species in version 1.0. A total of four Oomycota species were listed in Bridge et al. (2010).

Amoeba (Sarcodina). One accepted species in version 1.0. One single Amoeba species (Platyamoeba stenopodia) remains described (Adams et al. 2006). However, additional protozoans with incerta status are likely to be placed in the group once their classification is resolved.

BACTERIA AND ARCHAEA KINGDOMS. A total of 349,966 prokaryotic ASVs in version 1.0.

Prokaryotes. Whereas we know that microbial communities are adapted to live in diverse Antarctic challenging habitats (Dragone et al. 2021), these communities are widely unexplored. Environmental surveys from soil and freshwater Antarctic habitats routinely find unidentified microorganisms (Bowman 2018). Our re-analysis of a selection of publicly available 16S rRNA amplicon datasets resulted in only 37% and 82% of taxonomically assigned ASVs at the genus level (see PROKARYOTIC STUDY SOURCE LIST dataset). This high portion of unknown microorganisms highlights the knowledge gap in microbial communities and the consequent gaps in taxonomic reference databases (Bowman 2018). However, as both sampling and sequencing technology advance, new microbial habitats are explored, new sequencing datasets are obtained and new microorganisms are characterised (Taş et al. 2021). In addition to amplicon studies, whole shotgun metagenome is also allowing advances in microbial taxonomy explorations with the reconstruction of complete metagenome assembled genomes (MAGs), allowing the discovery of full genomes without the use of culturing techniques (Tully et al. 2018, Yang et al. 2021). Metagenomic and metatranscriptomic data are, therefore, pivotal for the exploration of microbial diversity and metabolic functions, in particular, in life-challenging environments, such as Antarctica where cell isolation and culture in laboratory settings are rarely possible (Bodor et al. 2020).

VIRAL ENTITIES. Fourteen families in version 1.0.

Viruses. We have a very coarse notion of viral diversity, which is thought to be remarkably high (Lopez-Bueno et al. 2009, Zablocki et al. 2014). Pathogenic viruses in Antarctic wildlife have received increased attention in recent years. Wildlife pathogens can be transported to Antarctic via migratory species (e.g. Antarctic fur seals, Arctic terns or skuas) arriving from areas where the pathogen already exists or through human activity (Barbosa et al. 2021). Direct or serological evidence has been detected in Antarctic wildlife for viral diseases including Infectious Bursal Disease Virus, Newcastle Disease Virus, Petrelpox Virus, Sealpox Virus, Canine Distemper Virus, Phocine Distemper Virus and Phocine Herpes virus 1 (Barbosa and Palacios 2009, Kerry and Riddle 2009, Grimaldi et al. 2014). Recent concerns regarding the risk of transmission of SARS-CoV-2 from human to Antarctic wildlife have resulted in the production of mitigation measures, although there remain no reports of COVID-19 in Antarctic wildlife to date.

2. Examination of the Antarctic Tree of Life: phylodiversity elements in the continent

Basic phylogenetic relationships between taxa were established from the Tree of Life project (http://tolweb.org/tree/). A Tree of Life for Antarctic diversity was created using the major realms, kingdoms, phyla and classes present in the region (Fig. 2). Due to the widely differing levels of diversity within each clade, we used only one representative taxon for each (once these diverged too broadly at the root of the trees to be depicted in full). This representative was typically drawn from the most recurrent higher taxa amongst each clade in our biodiversity list. Thus, only one group of viruses and phages were selected, respectively. For SAR and bacteria, only some representative supergroup lineages were taken. In turn, for fauna and flora (plant, fungi and algae), most groups were represented at the level of class.

Figure 2.  

Basic Antarctic Tree of Life. Some lesser animal phyla with less than five species total are not visually depicted (e.g. Annelida).

Overall, the Antarctic continent hosts over 400 species from at least seven animal phyla (see Fig. 2). Considering the general diversity of the region, most terrestrial phyla are present even in this “barren” continent. Amongst freshwater annelids, only a few Lumbricillus/Marionina species (five in total) are reported to be present in the continent's stream waters (Rodriguez and Rico 2008), but a non-native entrytraeid worm Christensenia blocki is also locally present in Signy Island (Dózsa-Farkas and Convey 1997, Hughes et al. 2015). No terrestrial or freshwater molluscs are known from the region but, if the intertidal zone is included, some marine species are present (Jossart et al. 2023). No non-marine bryozoans have been reported from the region. Additionally, no freshwater fish, reptiles (sensu lato) or amphibians are present. Insect diversity is remarkably low, with representatives of only one free-living order present (two species of Diptera), along with another two parasitic (Psocodea and Siphonaptera).

Plant diversity is also high in the continent, yet the full extent of Viridiplantae diversity is still uncertain. In the case of embryophytes, the most remarkable contemporary absence is that of ferns, along with the very low diversity of vascular plants (two species, one native Caryophyllaceae and another Poaceae), but significantly covering both monocots and eucots presence in the continent. No gimnosperms are present, but they were widely present at some point in the paleohistory of the continent, as observed from fossil records. A remarkable diversity of bryophytes is found, with all three major divisions present (mosses, hornworts and liverworts). Both insects and vascular plants become more diverse in the sub-Antarctic islands.

Amongst fungi, compared with global patterns, Antarctic diversity is relatively lower amongst Basidiomycota and higher in Ascomycota, but all major groups are present at the division level, including some mushroom-forming species of Basidiomycota and Ascomycota. SAR and others (formerly described as “protists”) are also diverse and include representatives of most recognised groups in the region. However, major surveying and knowledge gaps still exist and knowledge of diversity remains far from complete.

The unique combination of phylogenetic diversity in the Antarctic is threatened by the arrival and establishment of non-native species (Duffy et al. 2017). Futhermore, pre-established alien species in Antarctica present singular invasive traits, with the risk of altering the functional behaviour of the ecosytems (Pertierra et al. 2022). Fortunately, at present, these remain limited for the most part to soil arthropods (Acari, Insecta and Collembolla).

3. Challenges to constitute more accurate biodiversity compendia

Continued development of modern molecular and integrated taxonomic methodologies is paramount for further improving the assessment of species identity and representative inclusion. Detailed and extensive taxonomic revisions are required for most groups, also founded on wider mobilisation to local studies and expertise generating frequently updated species list repositories. The ongoing need for biodiversity data compilation and integration into repositories, such as GBIF and AntaBIF, is currently a major challenge for ecological research, at the same time being particularly relevant for strategic conservation planning. New primary survey data are required to generate accurate knowledge of diversity in many areas of the continent that, even today, remain unvisited by group specialists. The role of taxonomists is especially relevant for filling gaps and, therefore, the training of new taxonomists is essential. The very diverse range of poorly-understood microbiological groups that are bundled under the term ‘microbial biodiversity’, namely viruses, bacteria, archaea, microscopic fungi and algae and protozoans, represent a major research gap of the region.

Eukaryotes. No high-level faunal groups were absent from Antarctic inventories prior to the preparation of this compendium. However, the availability and application of new molecular and integrated taxonomic approaches are leading to an upsurge in new species descriptions, especially for non-arthropod micro-invertebrates and Acari and Collembola (Collins et al. 2023). Most embryophyte groups are also well studied, with robust compendia available that can be updated with recent reconsideration of a small number of species identities of mosses (Ochyra et al. 2008) and liverworths (Bednarek-Ochyra et al. 2000). In contrast, eukaryotic green algae continue to be a highly uncertain group where the known species diversity is likely to represent a fraction of the true total present. Lichen-forming fungal diversity is similarly well studied (Øvstedal and Lewis Smith 2001), with a broad spectrum of non-lichenised fungal species records in the region also available (Bridge et al. 2010). However, much fungal diversity remains unknown, with increasing evidence from eDNA and other molecular analyses supporting the presence of substantial unknown diversity within this group (Rosa et al. 2020).

Prokaryotes. Considerable advances in understanding microbial diversity in specific Antarctic regions have been made in the last decade through the application of newly-available sequencing and metagenomic technologies (Taş et al. 2021) and it is clear that the potential of these is only just beginning to be tapped. These are allowing exploration of the microbial diversity previously unexplored Antarctic regions and habitats, such as Dronning Maud Land nunataks (Staebe et al. 2019) and the Transantarctic Mountains (Dragone et al. 2021). However, while the potential of these approaches is clear, important limitations remain, in particular, relating to the large-scale incompleteness of available sequence databases against which to assign sequence identities and the lack of ability to confirm the presence or function of viable biota. Many ‘new’ ASVs are typically found in newly-analysed samples, suggesting (as with the fungi above) that Antarctic microbial diversity is far from well-characterised. Effort has also been devoted in recent years to describe microbial distribution patterns in relation to habitat conditions, bioclimatic variables and geographical distances (Chong et al. 2015, Chown et al. 2015, Almela et al. 2021). Better characterisation of these patterns, both at micro- and macro- scales, will provide valuable information on microbial diversity and how better to protect and conserve it. Again, increased sampling effort and the development of considerably more comprehensive microbial taxonomy online databases are required to facilitate improved microbial diversity characterisation. Lastly, further research is also required to enhance understanding of disease prevalence and transmission in Antarctic wildlife and the associated effects of increasing human activity and climate change in the region.

Acknowledgements

Author contributions

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