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Swarming in the storm: Collembola aggregations on sub-Antarctic Marion Island

Published online by Cambridge University Press:  06 May 2026

Daniela Marques Monsanto Open the ORCID record for Daniela Marques Monsanto [Opens in a new window] ,
Peter Convey Open the ORCID record for Peter Convey [Opens in a new window] ,
David William Hedding Open the ORCID record for David William Hedding [Opens in a new window] ,
Charlene Janion-Scheepers ,
Sandra Durand ,
Stefan Schoombie Open the ORCID record for Stefan Schoombie [Opens in a new window] ,
Monica Leitner  and
Bettine Jansen van Vuuren
Daniela Marques Monsanto*
Affiliation:
Centre for Ecological Genomics and Wildlife Conservation, Department of Zoology, University of Johannesburg , South Africa
Peter Convey
Affiliation:
Centre for Ecological Genomics and Wildlife Conservation, Department of Zoology, University of Johannesburg , South Africa Natural Environment Research Council, British Antarctic Survey , Cambridge, United Kingdom Cape Horn International Center (CHIC) , Puerto Williams, Chile Millennium Institute - Biodiversity of Antarctic and sub-Antarctic Ecosystems (BASE), Santiago, Chile School of Biosciences, University of Birmingham, Edgbaston, Birmingham, United Kingdom
David William Hedding
Affiliation:
Department of Geography, University of South Africa , Florida, South Africa
Charlene Janion-Scheepers
Affiliation:
Department of Biological Sciences, University of Cape Town , Rondebosch, South Africa Iziko South African Museum , Cape Town, South Africa
Sandra Durand
Affiliation:
Centre for Ecological Genomics and Wildlife Conservation, Department of Zoology, University of Johannesburg , South Africa
Stefan Schoombie
Affiliation:
Department of Zoology, Marine Apex Predator Research Unit (MAPRU), Institute for Coastal and Marine Research, Nelson Mandela University , Gqeberha, South Africa FitzPatrick Institute of African Ornithology, University of Cape Town , Cape Town, South Africa
Monica Leitner
Affiliation:
Department of Zoology and Entomology, University of Pretoria , Pretoria, South Africa
Bettine Jansen van Vuuren
Affiliation:
Centre for Ecological Genomics and Wildlife Conservation, Department of Zoology, University of Johannesburg , South Africa
*
Corresponding author: Daniela Marques Monsanto; Email: dmonsanto119@gmail.com
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Abstract

The intensity and frequency of extreme weather events in rapidly changing environments continue to increase, driving unusual behaviours and posing significant threats to terrestrial ecosystems. In this study, we describe the co-occurrence of 1) Collembola swarming, 2) vegetation die-off and 3) extreme weather events, particularly heavy rainfall and high temperatures, over a 5 day period on sub-Antarctic Marion Island. Taxonomic and molecular evidence confirmed that the Collembola species displaying swarming behaviour was Ceratophysella denticulata, an invasive hypogastrurid. Our observations suggest that environmental stressors may have induced vegetation die-off, which, in turn, may have directly or indirectly driven Collembola aggregation. The association of these factors highlights the potential role of Collembola as bioindicators of soil ecosystem responses to climatic extremes, and that the recognition of these interactions can be critical in the prediction and management of ecological responses to changing environments.

Keywords

Ceratophysella denticulataCollembola swarmingextreme weather eventsSouthern Oceansub-Antarctic islandsvegetation die-off

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Type
Short Note
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Antarctic Science , First View , pp. 1 - 9
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Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (https://creativecommons.org/licenses/by/4.0), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
Copyright
© The Author(s), 2026. Published by Cambridge University Press on behalf of Antarctic Science Ltd

Introduction

Collembola, commonly known as springtails, are globally distributed microarthropods that are largely terrestrial and often numerically abundant in soil ecosystems. Densities of 104–105 individuals m−2 in suitable habitats are common, sometimes reaching considerably greater numbers, with population numbers often exceeding more than half the total number of soil arthropods (Petersen & Luxton Reference Petersen and Luxton1982, Convey & Smith Reference Convey and Smith1997, Hopkin Reference Hopkin1997, Bellini et al. Reference Bellini, Weiner and Winck2023). Despite their small size (typically 0.1–3.0 mm), Collembola are important constituents of soil mesofauna, contributing to nutrient cycling and supporting complex food chains by serving as prey to numerous small predatory organisms. They, in turn, feed on microorganisms, fungal hyphae and spores, algae, plant debris, decaying animals and faeces, and, in some cases, they prey on microfaunal groups such as nematodes (Hopkin Reference Hopkin1997, Rusek Reference Rusek1998, Salmon et al. Reference Salmon, Ponge, Gachet, Deharveng, Lefebvre and Delabrosse2014). In addition, Collembola play crucial roles in soil ecosystem functioning by regulating soil microbial communities and influencing biochemical processes such as decomposition and carbon sequestration, as well as plant nutrition and growth (Petersen & Luxton Reference Petersen and Luxton1982, Hopkin Reference Hopkin1997, Potapov et al. Reference Potapov, Bellini, Chown, Deharveng, Janssens and Kováč2020, Bellini et al. Reference Bellini, Weiner and Winck2023). Their faeces and other excreted products contribute significantly to humus formation, which improves soil fertility, enhancing nutrient availability and promoting plant growth (Verhoef & Brussaard Reference Verhoef and Brussaard1990, Forey et al. Reference Forey, Coulibaly and Chauvat2015, Winck et al. Reference Winck, Chauvat, Coulibaly, Santonja, Saccol De Sá and Forey2020). The main outcomes of these processes, largely influenced by Collembola density, affect key physical soil properties such as porosity, gaseous and water exchange and stability against erosion (Rusek Reference Rusek1998, Maaß et al. Reference Maaß, Caruso and Rillig2015, Potapov et al. Reference Potapov, Bellini, Chown, Deharveng, Janssens and Kováč2020, Bellini et al. Reference Bellini, Weiner and Winck2023). Given their key role in soil ecosystems, Collembola have been the focus of studies across fields, including the effects of biological invasions (Potapov et al. Reference Potapov, Bellini, Chown, Deharveng, Janssens and Kováč2020, Chown et al. Reference Chown, Bergstrom, Houghton, Kiefer, Terauds and Leihy2022), biomass shifts due to climate change (Gruss et al. Reference Gruss, Yin, Julia, Eisenhauer and Schädler2023), abiotic parameters such as terrain ruggedness, topography and geographical distances (Monsanto et al. Reference Monsanto, Hedding, Durand, Parbhu, Adair and Emami-Khoyi2024), soil ecotoxicology and landscape stress and disturbance (Greenslade Reference Greenslade2007), responses to urbanization (Qiao et al. Reference Qiao, Wang, Yao, Li, Scheu, Zhu and Sun2022) and the influence of selective pressures on the mitochondrial genome and adaptive strategies (Monsanto et al. Reference Monsanto, Main, Janion-Scheepers, Emami-Khoyi, Deharveng and Bedos2022).

Biological invasions can pose significant threats to biodiversity, especially that of sub-Antarctic islands owing to their favourable terrestrial systems and milder temperatures compared to more extreme Antarctic landmasses (Chown et al. Reference Chown, Gremmen and Gaston1998, Greve et al. Reference Greve, Mathakutha, Steyn and Chown2017, Leihy et al. Reference Leihy, Duffy and Chown2018). Indigenous fauna are sometimes outcompeted or apparently displaced by their invasive counterparts (although this is generally an inference based on the post-invasion dominance of the invasive species in certain habitats where other native species would typically be found; e.g. Convey et al. Reference Convey, Greenslade, Arnold and Block1999), with consequential impacts on ecosystem functioning and processes (Greenslade Reference Greenslade2002, Frenot et al. Reference Frenot, Chown, Whinam, Selkirk, Convey, Skotnicki and Bergstrom2005, McGeoch et al. Reference McGeoch, Shaw, Terauds, Lee and Chown2015, Vega et al. Reference Vega, Pertierra, Benayas and Olalla-Tárraga2021). Environmental impact assessments of invasive taxa on indigenous groups have been categorized by Greve et al. (Reference Greve, Mathakutha, Steyn and Chown2017), and recent quantitative assessments of invasive organisms found that Marion Island hosts a total of 45 established non-native taxa, 25 of which are deemed invasive (Fernández Winzer et al. Reference Fernández Winzer, Greve, Le Roux, Faulkner and Wilson2025). Of these, five springtail species are considered invasive, in comparison with an overall diversity of 10 native springtails (Janion-Scheepers Reference Janion-Scheepers2025). Environmental soil variables such as resource availability, temperature and chemistry (e.g. pH, total sodium, exchangeable sodium, total nitrogen, total phosphorous, phosphate, organic carbon, moisture content) and the consequential vegetation/habitat differences strongly affect Collembola assemblages, highlighting microhabitat specificity and adaptive ability (Hopkin Reference Hopkin1997, Convey et al. Reference Convey, Greenslade, Arnold and Block1999, Gabriel et al. Reference Gabriel, Chown, Barendse, Marshall, Mercer, Pugh and Smith2001, Monsanto et al. Reference Monsanto, Main, Janion-Scheepers, Emami-Khoyi, Deharveng and Bedos2022). Notably, the group also exhibits preference for different levels of microhabitat stratification, which are categorized into ecomorphological life-forms, probably driven by factors such as food source and availability, soil type and chemistry, functional ecology and biology/morphology (Rusek Reference Rusek1998, Salmon et al. Reference Salmon, Ponge, Gachet, Deharveng, Lefebvre and Delabrosse2014, Potapov et al. Reference Potapov, Semenina, Korotkevich, Kuznetsova and Tiunov2016, Malcicka et al. Reference Malcicka, Berg and Ellers2017, Monsanto et al. Reference Monsanto, Main, Janion-Scheepers, Emami-Khoyi, Deharveng and Bedos2022). Collembola are broadly defined into six ecomorphological life-forms with major ecological, biological and morphological characteristics - or combinations thereof - that typify each life-form: atmobiotic, epiedaphic, hemiedaphic, euedaphic, myrmecophilous and hydrophilous/aquatic. Atmobiotic and epiedaphic species are generally larger in size (> 3 mm), possess ocelli, body pigmentation and dorsal patterns, long antennae, well-developed appendages (including furca) and show strong dispersal abilities. Hemiedaphic, euedaphic, myrmecophilous and hydrophilous taxa are usually smaller in size (< 3 mm) and have fewer or no ocelli, little to no body pigmentation with no dorsal patterns, shorter antennae and appendages and reduced or absent furca, and they are therefore poor dispersers (Deharveng et al. Reference Deharveng, D'Haese and Bedos2008, Potapov et al. Reference Potapov, Semenina, Korotkevich, Kuznetsova and Tiunov2016, Malcicka et al. Reference Malcicka, Berg and Ellers2017, Monsanto et al. Reference Monsanto, Main, Janion-Scheepers, Emami-Khoyi, Deharveng and Bedos2022).

Social communication among individuals is necessary to ensure the adequate functioning of a population. Social signalling between organisms can occur via various means, such as chemical, visual, acoustic, tactile or substrate-borne means (Richard & Hunt Reference Richard and Hunt2013). Among these, chemical communication through pheromones is the most common, ancient and widespread means of disseminating information (Candolin Reference Candolin2003). Chemical messaging is efficient in most conditions and can be used to convey messages pertaining to health, mate recognition and selection, aggressive behaviour and dominance, defence strategies, microhabitat selection and the search for food (Candolin Reference Candolin2003, Richard & Hunt Reference Richard and Hunt2013). Chemical communication among higher-order insects has long been known to be an adaptive feature, such as in mate attraction or enabling the development of aggregation behaviour to collectively respond to and deter predators (Messer et al. Reference Messer, Walther, Dettner and Schulz2000, Pfander & Zettel Reference Pfander and Zettel2004, Richard & Hunt Reference Richard and Hunt2013).

Aggregation or swarming behaviour has also been noted in more primitive hexapods such as springtails, and in other microarthropods such as mites (Acari). For instance, the Antarctic oribatid mite Alaskozetes antarcticus commonly aggregates in multi-instar groups around the edges where embedded rocks or carpet moss vegetation adjoin the ground surface (Block & Convey Reference Block and Convey1995), or even in abundance on the faeces of the native ducks on Bird Island, South Georgia (T. Martin, personal communication 2002). Although they probably graze on epilithic microalgae in these aggregations, aggregation also appears to be driven by movement towards favourable microhabitats in otherwise-challenging (temperature and desiccation) environments. Sometimes these aggregations form mounds or balls of tightly massed mites several to tens of centimetres across, containing tens to hundreds of thousands of individuals, which could play multiple roles including microhabitat regulation (temperature and water loss) or facilitating mating. These mites do not have native predators in their natural Antarctic habitat, so these visually obvious aggregations may not incur a ‘defence’ cost. Similarly, Collembola are frequently observed to form aggregations rafting on water and on dead foliage, decaying wood, stone structures and snow surfaces/glaciers, particularly during climatic changes, which can influence the behavioural patterns and community dynamics of populations (Turk Reference Turk1932, Krediet et al. Reference Krediet, Ellers and Berg2023, Susanti et al. Reference Susanti, Krashevska, Widyastuti, Stiegler, Gunawan, Scheu and Potapov2024, Valle et al. Reference Valle, Porco, Skarżyński, Frati, Caccianiga and Rodriguez-Prieto2024). Incidences of extremely high densities of up to millions of individuals per square metre of the Antarctic springtail Cryptopygus antarcticus have been observed in one of its preferred habitats, the nitrophilous foliose alga Prasiola crispa (P. Convey, personal observation from Signy Island in 2024). This, again, is likely to bestow a combination of advantages, including providing a stable and hydrated microhabitat within the folds and thalli of the algal carpet, thereby maximizing access to a preferred food source. These algal mats regularly and progressively dry for periods during the Antarctic summer, sometimes on a daily basis, but live springtails can be retrieved from them even when the mat texture is dry and externally crisp to the touch. Another example of aggregations observed in the Antarctic region is given by the mating swarms of the chironomid midge Parochlus steinenii from the Maritime Antarctic South Shetland Islands (Hahn & Reinhardt Reference Hahn and Reinhardt2006, Contador Mejias et al. Reference Contador Mejias, Gañan, Rendoll-Cárcamo, Maturana, Benítez and Kennedy2023). Strongly dominated by males, these swarms are found in densities of more than 5000 individuals in preferred microhabitats such as on the underside of stones, on the sheltered side of rocks and on mosses on the shores of permanent lakes and ponds, and swarms of a few hundred individuals may even be observed floating on the water surface.

Aggregations may have a protective defence function for some springtails, integrating the secretion of active compounds or chemical irritants, thereby providing an energetically more efficient defence mechanism (Hopkin Reference Hopkin1997, Bitzer et al. Reference Bitzer, Brasse, Dettner and Schulz2004, Pfander & Zettel Reference Pfander and Zettel2004). Whether this is the case in Antarctic or sub-Antarctic springtails is unknown, although, unlike the oribatid A. antarcticus, the springtail C. antarcticus is the primary prey of the mesostigmatid mite Gamasellus racovitzai in the Maritime Antarctic; however, the level of mite predation does not measurably impact the overall springtail population (Usher & Bowring Reference Usher and Bowring1984, Lister et al. Reference Lister, Block and Usher1988). Evidence of chemical communication (pheromones) has been documented for some collembolans (specifically species of the family Hypogastruridae) that frequently display aggregation/swarming behaviour often noticeable on snow, on melting snow patches or above dead foliage specifically during wet seasons (Turk Reference Turk1932, Pfander & Zettel Reference Pfander and Zettel2004, Hågvar Reference Hågvar2010, Dányi Reference Dányi2013). Indeed, in Scandinavia, coordinated swarms of springtails have been documented moving across snow surfaces in winter (commonly known as ‘snow fleas’; Hågvar Reference Hågvar2010, Valle et al. Reference Valle, Porco, Skarżyński, Frati, Caccianiga and Rodriguez-Prieto2024). However, swarming by collembolans in response to extreme abiotic (climatic) changes in environmental conditions has received relatively little attention globally. In this study, we document springtail swarming observed in two locations during a wet and hot 5 day period on sub-Antarctic Marion Island in the southern Indian Ocean, and we consider the possible causes. These observations are the first of their kind for Marion Island, and we provide a synthesis of the subject of aggregation behaviour in springtails and other invertebrates as applied to the sub-Antarctic and Antarctic regions.

Environmental setting and methodology

The islands that constitute the ‘core’ of the sub-Antarctic region (Fig. 1) are South Georgia, Marion and Prince Edward islands, the archipelagos of Crozet and Kerguelen, Heard and McDonald islands and Macquarie Island (Convey Reference Convey, Goldstein and DellaSala2020). Marion Island is characterized by hyper-maritime weather conditions, specifically high rainfall (~2200 mm/annum), high frequency of strong winds and low mean annual air temperatures (averaging at 6.3°C), and it has experienced shifts in climatic conditions on both geological and near-contemporary timescales (Rouault et al. Reference Rouault, Melice, Reason and Lutjeharms2005, Chown & Froneman Reference Chown and Froneman2008, le Roux & McGeoch Reference le Roux and McGeoch2008a, Nel et al. Reference Nel, Boelhouwers, Borg, Cotrina, Hansen and Haussmann2020, Reference Nel, Hedding and Rudolph2023, Rudolph et al. Reference Rudolph, Hedding, Fabel, Hodgson, Gheorghiu, Shanks and Nel2020). Notably, the mean annual air temperature on Marion Island has increased by 1.7°C since records began in the mid-twentieth century, the highest rate of change for any sub-Antarctic island over the same period (Nel et al. Reference Nel, Hedding and Rudolph2023). Moreover, the island experiences short-lasting ‘extreme weather events’, such as high-temperature and high-rainfall events (see Kabase Reference Kabase2024), which can have negative impacts on the island’s flora and invertebrate fauna (le Roux & McGeoch Reference le Roux and McGeoch2008b, Nyakatya & McGeoch Reference Nyakatya and McGeoch2008, Hedding & Greve Reference Hedding and Greve2018, Hugo-Coetzee & le Roux Reference Hugo-Coetzee and le Roux2018).

Figure 1.

Digital surface model map of Marion Island and its position (represented as Prince Edward Islands) within the Southern Ocean, highlighting the locations (Swartkops and Mixed Pickle) where Collembola aggregations were observed. The inset map shows the positions of sub-Antarctic islands and the Antarctic Polar Front (dashed line). m.a.s.l. = metres above sea level.

Microarthropod aggregation behaviour was observed for the first time on sub-Antarctic Marion Island (the larger of the two islands belonging to the Prince Edward Islands) in April 2019 by S. Schoombie and M. Leitner. This involved a single collembolan species at two locations on the west coast of the island, namely Swartkops (SK; 46°55’33.8"S, 37°35’38.8"E) and Mixed Pickle (MP; 46°52’38.0"S, 37°37’56.2"E). Hundreds of aggregations near SK were observed at the burrow entrances of blue petrels (Halobaena caerulea; Fig. 2a), whereas those near MP were at the wet and/or flooded burrow entrances of white-chinned petrels (Procellaria aequinoctialis; Fig. 2b,c).

Figure 2.

Aggregations of springtails of ~20 cm in size at a. the burrow entrances of blue petrels at Swartkops, b. the wet burrow entrances of white-chinned petrels of ~10 cm in diameter at Mixed Pickle and c. floating on the water surface of a flooded white-chinned petrel burrow entrance of ~50 cm in size at Mixed Pickle. d. Taxonomic and molecular techniques identified the species forming these aggregations to be the invasive hypogastrurid Ceratophysella denticulata.

Approximately 80 Collembola specimens were collected at one nest entrance from each of these two sites and stored immediately in absolute ethanol for subsequent taxonomic examination and molecular identification. Twenty individual specimens were photographed using a Zeiss Stemi 305 Stereo Microscope equipped with an AxioCam ERc 5s microscope camera. The specimens were identified using a taxonomic key for the Collembola of Marion Island (Janion-Scheepers Reference Janion-Scheepers2025). Taxonomic confirmation was made using molecular sequence data by individually placing ethanol-preserved specimens into 0.2 ml polymerase chain reaction (PCR) tubes and allowing them to air dry at room temperature to remove excess ethanol. The DNA barcoding gene (mitochondrial cytochrome c oxidase subunit I gene, or COI) was amplified using the primers described by Folmer et al. (1994). A standard PCR master mix was used containing 3 μl reaction buffer (20 mM TRIS-HCl (pH 8.0), 100 mM NaCl, 0.1 mM EDTA, 1 mM DTT, stabilizers and 50% glycerol), 3 μl 25 mM MgCl2, 3 μl 1 μM of each dNTP, 3 μl 10 μM of both forward and reverse primers, 0.3 μl Taq polymerase (5 units ml−1) and 14.7 μl ddH2O to create a reaction volume of 30 μl, which was added to each tube. The individual springtails were then homogenized using a pestle, and each PCR tube was placed into a MultiGene OptiMax thermal cycler with the following conditions: initial denaturation at 96°C for 5 min, 40 cycles of 30 s at 96°C for denaturation, 30 s at 48°C for annealing and 50 s at 72°C for extension, followed by a final extension step at 72°C for 10 min. All samples underwent post-PCR clean-up using the Macherey-Nagel NucleoSpin® Gel and PCR Clean-up Kit following the manufacturer’s recommendations. Successful amplifications were confirmed using 2% agarose gel electrophoresis with 0.5X TBE buffer and the GelRed Nucleic Acid Gel Stain (Biotium). PCR products were sequenced with the forward primer on a 3730XL DNA Analyzer (Applied Biosystems). To identify and confirm the species in this study, Basic Local Alignment Search Tools (BLAST) searches were conducted using the GenBank repository based on > 98% identity (Benson et al. Reference Benson, Cavanaugh, Clark, Karsch-Mizrachi, Lipman, Ostell and Sayers2013).

Results and discussion

Collembola aggregations were observed at the burrow entrances of blue petrels and white-chinned petrels at SK and MP, respectively. Ground observations highlighted vegetation alterations at MP with mild vegetation die-off and decay of the native flowering plant Leptinella plumosa in the areas near the aggregations (Fig. 2b), and taxonomic and molecular techniques confirmed that the Collembola species was Ceratophysella denticulata (Bagnall, 1941; Fig. 2d). At SK, extensive vegetation die-off and decay were observed affecting an area of ~200 m2 of L. plumosa (Fig. 3a,b), noticeably differing from the typical state of vegetation observed in the area (Fig. 3c,d).

Figure 3.

a. & b. Extensive decay of Leptinella plumosa of ~200 m2 at Swartkops contrasted with c. & d. when no vegetation alterations are observed in the area. The photographs for a. and b. were taken in April 2019, the photograph for c. was taken in April 2013 and the photograph for d. was taken in January 2020.

Figure 4.

Temperature and precipitation data for Marion Island for April 2019 recorded at the Marion Island research station. Intense rainfall was experienced on 20 April, 5 days before the Collembola aggregations were observed and collected on 25 April, when extremely high temperatures were recorded. Hourly weather data were obtained from the South African Weather Service.

The conspicuous Collembola aggregations observed here were found in areas comprising bare ground and poor-health/decaying vegetation in the immediate vicinity of bird nesting burrows, including within their entrances. This may suggest a possible nutrient-related cue, either from the decaying plant material or bird guano, and also raises the possibility of further zoochoric transfer of this already highly invasive springtail if carried in soil or plant material that becomes attached to the birds’ plumage. The species forming these aggregations on hyper-maritime Marion Island was confirmed to be the hypogastrurid C. denticulata (Bagnall, 1941), which has been documented to be invasive and widely distributed at lower elevations on Marion Island (Gabriel et al. Reference Gabriel, Chown, Barendse, Marshall, Mercer, Pugh and Smith2001, Frenot et al. Reference Frenot, Chown, Whinam, Selkirk, Convey, Skotnicki and Bergstrom2005, Hugo et al. Reference Hugo, Chown and McGeoch2006, Myburgh et al. Reference Myburgh, Chown, Daniels and Jansen van Vuuren2007, Slabber et al. Reference Slabber, Worland, Leinaas and Chown2007, Greve et al. Reference Greve, Mathakutha, Steyn and Chown2017). Furthermore, this species belongs to the highly invasive family Hypogastruridae (Greenslade Reference Greenslade2002), some of whose members are known to be resilient to extreme climatic conditions (Onley et al. Reference Onley, Houghton, Liu and Shaw2025), and with physiological analyses revealing C. denticulata to have a competitive advantage over its indigenous counterparts due to its higher upper lethal temperature and ability to withstand warmer temperatures (Slabber et al. Reference Slabber, Worland, Leinaas and Chown2007). Other members of the Hypogastruridae are also well-known invasive springtails with wide and even global cosmopolitan distributions. These include two representatives of Hypogastrura, Hypogastrura viatica and Hypogastrura purpurescens, which are known non-native species on sub-Antarctic South Georgia (Frenot et al. Reference Frenot, Chown, Whinam, Selkirk, Convey, Skotnicki and Bergstrom2005, Greenslade & Convey Reference Greenslade and Convey2012). Additionally, H. viatica is widespread and abundant on other sub-Antarctic islands such as Macquarie Island (alongside C. denticulata; Greenslade Reference Greenslade2018), Kerguelen Island and Possession Island (Baird et al. Reference Baird, Moon, Janion-Scheepers and Chown2019), and on Maritime Antarctic Deception Island (Greenslade et al. Reference Greenslade, Potapov, Russell and Convey2012, Greenslade Reference Greenslade2018, Hughes et al. Reference Hughes, Convey and Lee2025). Hypogastrura viatica has also been recorded at a location off the coast of the western Antarctic Peninsula, with a recent first-time report (although most probably not established) within Casey Station on the Continental Antarctic coastline (Greenslade Reference Greenslade1995, Reference Greenslade2002, Greenslade & Convey Reference Greenslade and Convey2012, Onley et al. Reference Onley, Houghton, Liu and Shaw2025, Hughes et al. Reference Hughes, Greenslade and Convey2017). These species are frequently found in dense concentrations, particularly around and under human-made debris.

Previously, Gabriel et al. (Reference Gabriel, Chown, Barendse, Marshall, Mercer, Pugh and Smith2001) concluded that native springtail richness and abundance were unaffected by non-native springtails on Marion Island. However, a recent study comparing Marion Island and Macquarie Island with the pristine Heard Island concluded that invasive springtails have the potential to cause detrimental impacts on the native taxa (Chown et al. Reference Chown, Bergstrom, Houghton, Kiefer, Terauds and Leihy2022). On the Prince Edward Islands, the current invasion risk potential of non-native springtails on native taxa remains unknown (Fernández Winzer et al. Reference Fernández Winzer, Greve, Le Roux, Faulkner and Wilson2025). Studies comparing the non-native springtails present on Marion Island and Prince Edward Island previously confirmed only one on Prince Edward Island: the already globally distributed C. denticulata, with densities on Marion Island possibly exceeding 38 000 individuals/m−2 (Greenslade Reference Greenslade2002, Hugo et al. Reference Hugo, Chown and McGeoch2006). The most recent collections from Prince Edward Island made in November 2023 revealed that a second non-native springtail, Isotomurus maculatus, is now present, a species already well-established on Marion Island (Fernández Winzer et al. Reference Fernández Winzer, Greve, Le Roux, Faulkner and Wilson2025). Genetic analyses of C. denticulata and I. maculatus revealed single haplotypes across Marion Island, consistent with single invasion events for each species, highlighting the insignificance of propagule pressure and illustrating the high invasion potential of these taxa (Myburgh et al. Reference Myburgh, Chown, Daniels and Jansen van Vuuren2007).

In this context, it is notable that both the vegetation die-off and decay and aggregation behaviour of C. denticulata were observed after extreme weather conditions, namely warm temperatures and intense rainfall, over a 5 day period between 20 and 25 April 2019 (Fig. 4). Mild vegetation die-off of the flowering plant L. plumosa around bird burrows was observed at MP (Fig. 2b), and a considerable extent of vegetation die-off and decay of ~200 m2 was observed at SK (Fig. 3a,b), standing in clear contrast to when the area experiences no such die-off and decay (Fig. 3c,d). Although causation cannot be confirmed, environmental stress and subsequent plant decay could produce chemical or physical cues that influence Collembola aggregation. Wider botanical research, albeit primarily on crop species, has shown that extreme weather conditions such as heatwaves, droughts and intense rainfall events can impose significant stresses on vegetation (Rosenzweig et al. Reference Rosenzweig, Tubiello, Goldberg, Mills and Bloomfield2002, Lesk et al. Reference Lesk, Rowhani and Ramankutty2016, Kim et al. Reference Kim, Webber, Adiku, Nóia Júnior, Deswarte, Asseng and Ewert2024). Specifically, excess precipitation and waterlogging can impose stresses as a result of submergence, lodging (i.e. weakening of stem and root system and subsequent shifting from an upright position) and the facilitating of pests and pathogens that can benefit from excess water (Thompson et al. Reference Thompson, Levin and Rodriguez-Iturbe2013, Kim et al. Reference Kim, Webber, Adiku, Nóia Júnior, Deswarte, Asseng and Ewert2024). Among higher-order taxa, strong evidence of swarming or migratory events linked to climatic episodes, such as storms, floods, atmospheric circulation changes, heatwaves and vegetation alterations, have been extensively documented, with extreme events potentially causing both population outbreaks or crashes (Harvey et al. Reference Harvey, Heinen, Gols and Thakur2020, Filazzola et al. Reference Filazzola, Matter and MacIvor2021, John et al. Reference John, Riat, Ahmad Bhat, Ganie, Endarto and Nugroho2024, Vives-Ingla et al. Reference Vives-Ingla, Capdevila, Clements, Stefanescu and Carnicer2025). Additionally, research has suggested that Collembola behaviour and community dynamics are also closely linked to extreme weather conditions, with their abundances, species compositions and activity patterns shifting in response to these changes, and with moisture and temperature being the primary drivers (Beet et al. Reference Beet, Hogg, Cary, McDonald and Sinclair2022, Zhang et al. Reference Zhang, Xie, Dou, Sun, Chang and Wu2022, Krediet et al. Reference Krediet, Ellers and Berg2023, Sanders et al. Reference Sanders, Martínez-De León, Formenti and Thakur2024, Susanti et al. Reference Susanti, Krashevska, Widyastuti, Stiegler, Gunawan, Scheu and Potapov2024, Li et al. Reference Li, Zhang, Wang, Ai, Zhang and Shao2025). During the decomposition of vegetation, volatile compounds are released that can serve as chemical cues or signals for insects to migrate towards or away from decaying material, and the subsequent secretion of aggregation pheromones in response to decaying vegetation has been reported for social insects (Kojima Reference Kojima2015, Mitaka et al. Reference Mitaka, Helms and Vargo2024). Although no current research reveals a direct link between vegetation decay and Collembola swarming, the co-occurrence of vegetation die-off and decay and aggregations on Marion Island highlights the possible association between vegetation die-off and decomposition and the swarming behaviour of soil microarthropods.

Conclusions

Our findings highlight a plausible connection between Collembola aggregation, vegetation die-off and decomposition with extreme weather events recorded on the island. The co-occurrence and interplay of these factors suggest that changes in environmental conditions, especially those brought on by climatic extremes, may significantly influence Collembola behaviour and distribution. Such responses may become more noticeable as the intensity and frequency of extreme climatic conditions continue to change. Understanding these interactions is crucial, as they reflect broader ecosystem responses to environmental stress, and they may serve as early bioindicators of soil health and vegetation change. Future studies that recognize these interactions will enhance our ability to predict and manage ecological impacts in a rapidly changing climate.

Author contributions

SS and ML collected the samples. DMM conducted the laboratory work and data analysis. DMM, PC and DWH contributed to writing the first version of the manuscript. All authors provided substantial input in editing and revising the final version.

Acknowledgements

We thank the South African Weather Service (SAWS) for providing weather data for Marion Island.

Financial support

The authors thank the National Research Foundation (NRF) South African National Antarctic Programme (SANAP) grants awarded to BJvV (NRF SANAP grant number: SANAP230501100411) and DWH (NRF SANAP grant number: SANAP230529111074). PC is supported by NERC core funding to the British Antarctic Survey’s ‘Biodiversity, Evolution and Adaptation’ Team.

Competing interests

The authors declare none.

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Figure 1. Digital surface model map of Marion Island and its position (represented as Prince Edward Islands) within the Southern Ocean, highlighting the locations (Swartkops and Mixed Pickle) where Collembola aggregations were observed. The inset map shows the positions of sub-Antarctic islands and the Antarctic Polar Front (dashed line). m.a.s.l. = metres above sea level.

Figure 1

Figure 2. Aggregations of springtails of ~20 cm in size at a. the burrow entrances of blue petrels at Swartkops, b. the wet burrow entrances of white-chinned petrels of ~10 cm in diameter at Mixed Pickle and c. floating on the water surface of a flooded white-chinned petrel burrow entrance of ~50 cm in size at Mixed Pickle. d. Taxonomic and molecular techniques identified the species forming these aggregations to be the invasive hypogastrurid Ceratophysella denticulata.

Figure 2

Figure 3. a. & b. Extensive decay of Leptinella plumosa of ~200 m2 at Swartkops contrasted with c. & d. when no vegetation alterations are observed in the area. The photographs for a. and b. were taken in April 2019, the photograph for c. was taken in April 2013 and the photograph for d. was taken in January 2020.

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Figure 4. Temperature and precipitation data for Marion Island for April 2019 recorded at the Marion Island research station. Intense rainfall was experienced on 20 April, 5 days before the Collembola aggregations were observed and collected on 25 April, when extremely high temperatures were recorded. Hourly weather data were obtained from the South African Weather Service.