Published October 8, 2025
| Version v1
Journal article
Open
Terrestrial isopods (Crustacea, Isopoda, Oniscidea) from Mesovoid Shallow Substratum habitats of Portugal
Authors/Creators
- 1. Universidade de Lisboa, Lisbon, Portugal
- 2. Istituto di Ricerca sugli Ecosistemi Terrestri, Sesto Fiorentino (Firenze), Italy
Description
The Mesovoid Shallow Substratum (MSS) is a largely unknown subterranean habitat formed by the accumulation of rock fragments in steep slopes. In this study, we investigated the diversity, abundance and seasonal dynamics of terrestrial isopods in MSS habitats across Portugal. We identified 12 species belonging to five families, with Porcellionidae and Armadillidiidae being the most represented. The new species, Eluma cristata, endemic to the MSS of Alcobertas, is described. The terrestrial isopod communities included both surface-dwelling and subterranean-adapted species, pointing out the connectivity between these habitats and highlighting the adaptability of terrestrial isopods to diverse environments. Widespread species occurring in multiple locations, and species confined to single sites are present. Seasonality does not seem to affect the community composition. Our findings highlight the rich diversity of MSS habitats, thus their importance for conservation.
Files
SB_article_163632.pdf
Files
(5.3 MB)
| Name | Size | Download all |
|---|---|---|
|
md5:23d76d9446ad741b27ce2aa7c658e75f
|
5.3 MB | Preview Download |
System files
(197.0 kB)
| Name | Size | Download all |
|---|---|---|
|
md5:2fc78bedf3f15a4cd713c689f33ac1b7
|
197.0 kB | Download |
Linked records
Additional details
References
- Arcangeli A (1935a) Gli isopodi terrestri del Portogallo. Bollettino del Laboratorio di Zoologia generale e agraria della RealeScuola superiore d'Agricoltura in Portici 29: 1–39.
- Arcangeli A (1935b) Specie nuove di isopodi terrestri della Spagna raccolti dal Prof. F. Silvestri. Bollettino del Laboratorio di Zoologia generale e agraria del Regio Istituto superiore agrario in Portici 29: 40–46.
- Arcangeli A (1948) Schizidiinae sottofamiglia di Armadillidiidae (crostacei isopodi terrestri). Bollettino dell'Istituto e Museo di Zoologia dell'Università di Torino 1: 211–272.
- Brandt JF (1833) Conspectus Monographiae Crustaceorum Oniscodorum Latreillii. Bulletin de la Société impériale des Naturalistes de Moscou 6: 171–193.
- Brandt JF, Ratzeburg JTC (1831) Isopoda. Gleichfüßler. In: Medizinische Zoologie oder getreue Darstellung und Beschreibung der Thiere, die in der Arzneimittellehre in Betracht kommen, in systematischer Folge herausgegeben. Trowitzsch und Sohn, Berlin 2: 70–84. [pls 12–13]
- Budde-Lund G (1885) Crustacea Isopoda terrestria per familias et genera et species descripta. Nielsen and Lydiche, Copenhagen. https://doi.org/10.5962/bhl.title.109769
- Cifuentes J (2019) Contribution à la connaissance des Trichoniscoides Sars, 1899 ibériques, avec la description de deux nouvelles espèces: Trichoniscoides marinae n. sp. et Tr. enoli n. sp (Crustacea, Isopoda, Trichoniscidae). Boletín de la Asociatión española de Entomologia 43(1–2): 27–54.
- Cifuentes J, Da Silva LP (2023) The genus Eluma in the Iberian Peninsula, with the description of a new terrestrial isopod: Eluma matae sp. nov. (Oniscidea, Armadillidiidae, Eluminae). European Journal of Taxonomy 860: 27–41. https://doi.org/10.5852/ejt.2023.860.2047
- Cifuentes J, Da Silva LP (2024) Revision of the genus Porcellionides Miers, 1877 (Isopoda: Porcellionidae) in the Ibero-Balearic region. European Journal of Taxonomy 939: 1–51. https://doi.org/10.5852/ejt.2024.939.2573
- Cifuentes J, Martínez SL, Pardo JLM, Barranco P (2021) Isópodos (Crustacea, Oniscidea) del Medio Subterráneo Superficial (MSS) en el Sureste Ibérico (Almería, España). Boletín de la Asociatión española de Entomologia 45(3–4): 261–276.
- Clarke KR (1993) Non‐parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18(1): 117–143. https://doi.org/10.1111/j.1442-9993.1993.tb00438.x
- Cruz A (1991) Especies nuevas o poco conocidas de isópodos terrestres de la Peninsula Ibérica. II. Isópodos epigeos de España y Portugal (Crustacea, Oniscidea). Bulletin de la Société d'Histoire naturelle de Toulouse 127: 71–75.
- Culver DC, Pipan T (2009) The biology of caves and other subterranean habitats. Oxford University Press, New York. https://doi.org/10.1093/acprof:oso/9780199646173.001.0001
- Culver DC, Pipan T (2014) Shallow subterranean habitats: ecology, evolution, and conservation. First edition. Oxford University Press, Oxford; New York.
- Dixie B, White H, Hassall M (2015) Effects of microclimate on behavioural and life history traits of terrestrial isopods: Implications for responses to climate change. ZooKeys 515: 145–157. https://doi.org/10.3897/zookeys.515.9399
- Dollfus A (1889a) Liste préliminaire des isopodes extramarins recueillis aux Açores pendant les campagnes de l'Hirondelle (1887–1888) par M. Jules de Guerne. Bulletin de la Société zoologique de France 14: 125–132.
- Dollfus A (1889b) Isopodes terrestres recueillis aux Açores en 1887,1888 et 1889 par MM. Dr. Th. Barrois et le Lieutenant Chaves. Revue biologique du Nord de la France 1: 306–392.
- Dollfus A (1892) Catalogue raisonné des isopodes terrestres de l'Espagne. Anales de la Sociedad española de Historia natural 21: 161–190.
- Dollfus A (1896a) Sur la distribution géographique des Armadilliens en Europe. Compte-rendu des Séances du Troisième Congrès international de Zoologie Leyden, 1895, 356–358.
- Dollfus A (1896b) Les isopodes terrestres du nord de l'Afrique, du Cap Blanc à Tripoli (Maroc, Algérie, Tunisie, Tripolitaine). Mémoires de la Société zoologique de France 9: 523–553.
- Dollfus A (1897) Notes de géographie zoologique. Les Crustacés Isopodes terrestres a grande dispersion. La Feuille des Jeunes Naturalistes 27: 205–212.
- Duméril A (1816) Armadille. Dictionnaire des Sciences naturelles 3: 115–117.
- Eaton AE (1882) Note on Rhacodes inscriptus, Koch, and Armadillo officinalis, Duméril, terrestrial Isopoda. The Annals and Magazine of Natural History 10: 360–362. https://doi.org/10.1080/00222938209459726
- Eusébio RP, Enghoff H, Solodovnikov A, Michelsen A, Barranco P, Salgado JM, Sendra A, Reboleira ASPS (2021) Temporal and spatial dynamics of arthropod groups in terrestrial subsurface habitats in central Portugal. Zoology 147: 125931. https://doi.org/10.1016/j.zool.2021.125931
- Gregory SJ, Lee P, Read HJ, Richards P (2012) Woodlice (Isopoda: Oniscidea) collected from northwest Spain and northern Portugal in 2004 by the British Myriapod and Isopod Group. Bulletin of the British Myriapod and Isopod Group 26: 6–23.
- Hornung E (2011) Evolutionary adaptation of oniscidean isopods to terrestrial life: Structure, physiology and behavior. Terrestrial Arthropod Reviews 4: 95–130. https://doi.org/10.1163/187498311X576262
- ICNB (2000) Plano Sectorial da Rede Natura 2000: Habitats Naturais (8130). https://www.icnf.pt/biodiversidade/natura2000/planosetorial
- ICNF (2023) Livro Vermelho dos Invertebrados de Portugal Continental. https://lvinvertebrados.uac.pt/docs/Livro-Vermelho-dos-Invertebrados-de-Portugal-Continental.pdf
- IUCN (2025) The IUCN Red List of Threatened Species. https://www.iucnredlist.org/en
- Jackson HG (1926) Woodlice from Spain and Portugal, with an Account of Benthana, a Sub-genus of Philoscia–Crustacea. Proceedings of the Zoological Society of London 96(1): 183–201. https://doi.org/10.1111/j.1096-3642.1926.tb01541.x
- Jiménez-Valverde A, Gilgado ID, Sendra A, Pérez-Suárez G, Herrero-Borgoñón JJ, Ortuño VM (2015) Exceptional invertebrate diversity in a scree slope in Eastern Spain. Journal of Insect Conservation 19: 713–728. https://doi.org/10.1007/s10841-015-9794-1
- Juberthie C, Delay D, Bouillon M (1980) Extension du milieu souterrain en zone non calcaire: description d'un nouveau milieu et de son peuplement par les Coléoptères troglobies. Mémoires de Biospéologie 8: 77–93.
- Kinahan J (1857) Analysis of certain allied genera of terrestrial isopods; with description of a new genus, and a detailed list of the British species of Ligia, Philougria, Philoscia, Porcellio, Oniscus and Armadillium [sic]. Natural History Review 4: 258–282.
- Koch L (1856) Crustacea. In: Rosenhauer WG. Die Thiere Andalusiens nach dem Resultate einer Reise zusammengestellt, nebst den Beschreibungen von 249 neuen oder bis jetzt noch unbeschriebenen Gattungen und Arten. Erlangen: Theodor Blaesing, 418–423.
- Latreille PA (1804) Histoire naturelle, générale et particulière, des crustacés et des insectes. Cloportides 7: 25–49.
- Mammola S, Amorim IR, Bichuette ME, Borges PAV, Cheeptham N, Cooper SJB, Culver DC, Deharveng L, Eme D, Ferreira RL, Fišer C, Fišer Ž, Fong DW, Griebler C, Jeffery WR, Jugovic J, Kowalko JE, Lilley TM, Malard F, Manenti R, Martínez A, Meierhofer MB, Niemiller ML, Northup DE, Pellegrini TG, Pipan T, Protas M, Reboleira ASPS, Venarsky MP, Wynne JJ, Zagmajster M, Cardoso P (2020) Fundamental research questions in subterranean biology. Biological Reviews 95(6): 1855–1872. https://doi.org/10.1111/brv.12642
- Mammola S, Cardoso P, Culver DC, Deharveng L, Ferreira RL, Fišer C, Galassi DMP, Griebler C, Halse S, Humphreys WF, Isaia M, Malard F, Martinez A, Moldovan OT, Niemiller ML, Pavlek M, Reboleira ASPS, Souza-Silva M, Teeling EC, Wynne JJ, Zagmajster M (2019) Scientists' Warning on the Conservation of Subterranean Ecosystems. BioScience 69(8): 641–650. https://doi.org/10.1093/biosci/biz064
- Mammola S, Giachino PM, Piano E, Jones A, Barberis M, Badino G, Isaia M (2016) Ecology and sampling techniques of an understudied subterranean habitat: the Milieu Souterrain Superficiel (MSS). The Science of Nature 103(11): 88. https://doi.org/10.1007/s00114-016-1413-9
- Mammola S, Piano E, Giachino PM, Isaia M (2017) An ecological survey of the invertebrate community at the epigean/hypogean interface. Subterranean Biology 24: 27–52. https://doi.org/10.3897/subtbiol.24.21585
- Miers EJ (1877) On a collection of Crustacea, Decapoda and Isopoda, chiefly from South America, with descriptions of new genera and species. Proceedings of the Zoological Society of London 1877: 653–679.
- Nitzu E, Nae A, Giurginca A, Popa I (2010) Invertebrate communities from the mesovoid shallow substratum of the Carpatho-Euxinic area: eco-faunistic and zoogeographic analysis. Travaux de l'Institut Spéol. "Émile Racovitza" 49: 41–79.
- Nitzu E, Popa I, Giurginca A (2011) Invertebrate fauna (Coleoptera, Collembola, Diplopoda, Isopoda) collected in the karst areas of the Aninei-Locvei Mountains. Travaux de l'Institute de Spéologie "Émile Racovitza" 50: 15–35.
- Nitzu E, Nae A, Băncilă R, Popa I, Giurginca A, Plăiaşu R (2014) Scree habitats: ecological function, species conservation and spatial-temporal variation in the arthropod community. Systematics and Biodiversity 12(1): 65–75. https://doi.org/10.1080/14772000.2013.878766
- Norman A (1899) The land Isopoda of Madeira. Annals and Magazine of natural History (Series 7) 3: 66–70. [pl. 6] https://doi.org/10.1080/00222939908678078
- Oksanen J, Simpson GL, Blanchet FG, Kindt R, Legendre P, Minchin PR, O'Hara RB, Solymos P, Stevens MHH, Szoecs E, Wagner H, Barbour M, Bedward M, Bolker B, Borcard D, Carvalho G, Chirico M, Caceres MD, Durand S, Evangelista HBA, FitzJohn R, Friendly M, Furneaux B, Hannigan G, Hill MO, Lahti L, McGlinn D, Ouellette MH, Cunha ER, Smith T, Stier A, Braak CJFT, Weedon J, Borman T (2025) vegan: Community Ecology Package.
- Preudhomme de Borre A (1886) Crustacés Isopodes recueillis par feu Camille Van Volxem, pendant son voyage en Portugal, en 1871. Comptes-Rendus des Séances de la Société entomologique de Belgique (Serie III) 72: 112–113.
- R Core Team (2024) R: A language and environment for statistical computing (Version 4.4.2). R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/
- Reboleira ASPS, Eusébio RP (2021) Cave-adapted beetles from continental Portugal. Biodiversity Data Journal 9: e67426. https://doi.org/10.3897/BDJ.9.e67426
- Reboleira ASPS, Eusébio RP (2023) Cave-adapted millipedes from Portugal: species conservation profiles. Biodiversity Data Journal 11: e110382. https://doi.org/10.3897/BDJ.11.e110382
- Reboleira ASPS, Eusébio RP, Taiti S (2022) Species conservation profiles of cave-adapted terrestrial isopods from Portugal. Biodiversity Data Journal 10: e78796. https://doi.org/10.3897/BDJ.10.e78796
- Reboleira ASPS, Goncalves F, Oromí P, Taiti S (2015) The cavernicolous Oniscidea (Crustacea: Isopoda) of Portugal. European Journal of Taxonomy 161: 1–61. https://doi.org/10.5852/ejt.2015.161
- Recuero E, Rodríguez-Flores PC, García-París M (2022) Homoplasy and morphological stasis revealed through multilocus phylogeny of new myrmecophilous species in Armadillidiidae (Isopoda: Oniscidea). Zoological Journal of the Linnean Society 194(4): 1312–1340. https://doi.org/10.1093/zoolinnean/zlab066
- Schmalfuss H (2003) World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde (Serie A) 654: 1–341.
- Schmölzer K (1955) Landasseln aus Spanien, gesammelt von Prof. Dr. Ing. H. Franz. Ein Beitrag zur Kenntnis der Spanischen Isopodenfauna. Eos 31: 311–321.
- Schmölzer K (1965) Ordnung Isopoda (Landasseln). Bestimmungsbücher zur Bodenfauna Europas, Issues 4 and 5. Academie Verlag, Berlin.
- Schmölzer K (1971) Die Landisopoden der Iberischen Halbinsel. Monografías de Ciencia Moderna 80: 1–161.
- Taiti S, Rossano C (2015) Terrestrial isopods from the Oued Laou basin, north-eastern Morocco (Crustacea: Oniscidea), with descriptions of two new genera and seven new species. Journal of Natural History 49(33–34): 2067–2138. https://doi.org/10.1080/00222933.2015.1009512
- Vandel A (1946) Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anais da Faculdade de Ciências do Porto 30: 135–427.
- Vandel A (1947) L'énigme de la reproduction des arthropodes cavernicoles et plus particulièrement des isopodes terrestres. Bulletin de la Société d'Histoire naturelle de Toulouse 82: 263–275.
- Vandel A (1948) La faune isopodique française (Oniscoïdes ou Isopodes terrestres). Sa répartition, ses origines et son histoire. Revue française d'Entomologie 15: 101–139.
- Vandel A (1952) Les trichoniscides (Crustacés – Isopodes) de l'hémisphère austral. Mémoires du Muséum national d'Histoire naturelle (Série A) 6: 1–116.
- Verhoeff KW (1901) Über paläarktische Isopoden (7. Aufsatz). Zoologischer Anzeiger 24: 403–408, 417–421.
- Verhoeff KW (1907) Über Isopoden, 10. Aufsatz. Zur Kenntnis der Porcellioniden. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin 8: 229–281.
- Verhoeff KW (1908) Über Isopoden. 15. Aufsatz. Archiv für Biontologie 2: 335–387. [pls 29 –31]
- Wickham H (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. https://doi.org/10.1007/978-3-319-24277-4_9