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Literature Survey, Bibliographic Analysis and a Taxonomic Catalogue of Subterranean Fauna from Portugal

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Literature Survey, Bibliographic Analysis and a Taxonomic Catalogue of Subterranean Fauna from Portugal Subterranean Biology 10: 51-60, 2012 (2013) Hypogean taxa from Portugal 51 doi: 10.3897/subtbiol.10.4025 Literature survey, bibliographic analysis and a taxonomic catalogue of subterranean fauna from Portugal Ana Sofia P. S. REBOLEIRA (1,2,*), Fernando GONÇALVES (1), Pedro OROMÍ (2) (1) Departamento de Biologia & Centro de Estudos do Ambiente e do Mar, Universidade de Aveiro, Campus Universitário de San- tiago, 3810-193 Aveiro. Portugal, e-mail: [email protected] (2) Departamento de Biología Animal, Facultad de Biología, Universidad de La Laguna. 38206 La Laguna. Tenerife. Islas Canarias, Spain * corresponding author ABSTRACT A bibliographic analysis of the hypogean biological studies in Portugal is made, compiling 138 publications related to the subter- ranean invertebrate fauna, since its begining in 1870 until November 2012. A catalogue of hypogean endemic taxa is provided, listing 27 troglobionts and 63 stygobionts, described to be obligate hypogean and endemic from mainland Portugal (Macaronesian archipelagos excluded). The first impetus on troglobiont studies was provided by the prospections of Barros Machado during 1940’s and by an expedition of Lindberg in the spring of 1961; and the major information about stygobiont species was provided by the former Instituto de Zoologia “Dr. Augusto Nobre” from Porto University. Key words: Bibliography, checklist, troglobionts, stygobionts, Portugal INTRODUCTION A literature survey allows a critical assessment of the The great momentum in the history of Portuguese state of subterranean biology, and is a crucial labor to put subterranean biology was given by António de Barros order in the knowledge of the hypogean fauna, and to Machado, an eminent arachnologist that began methodical avoid repetition of research due to the difficulties of get- exploration of caves in Portugal in the late 1930’s. He per- ting information in old bibliographical sources (Fan 2006). formed an intense sampling effort on terrestrial hypogean The subterranean habitat harbours an extremely special- fauna along the karstic caves of Portugal, and co-authored ized fauna, being a true laboratory for the study of ecology, by his brother Bernardino de Barros Machado described natural selection and evolution, central subjects in biology more than 300 caves in the “Inventário das cavernas cal- (Culver & Pipan 2009). The study of the subterranean fau- cárias de Portugal”. In the same decade, Fernando Frade na began limited to the perspective of ‘human size’ caves as from Lisbon University described Proasellus lusitanicus the domain of the cave-dwelling fauna (Vandel 1964), but (Frade, 1938) from the Alviela spring in the Estremenho the evolution of knowledge in subterranean biology leads karst massif, paving the way for the study of stygobionts in to a broad view of the subterranean habitats as a large vari- the following decades by researchers of the former Instituto ety of places with aphotic conditions, extending their limits de Zoologia “Dr. Augusto Nobre” from Porto University. far beyond the caves (Pipan & Culver 2012). Most of the biological hypogean studies in Portugal The first published register of subterranean fauna have been done in caves and dark wells. A large cryp- from Portugal (Chiroptera excluded) was in the mon- tic biodiversity still remains to unravel, especially in the ography “Trechorum oculatorum” where specimens of mesocavernous shallow substratum (MSS as described a ground beetle collected by Schaufuss in portuguese by Juberthie et al 1980) and in the anchialine waters of caves were recorded by Putzeys (1870: 18). The speci- coastal karsts. mens were assigned to Trechus fulvus Dejean, 1831 in More recently several efforts by Reboleira and col- spite of exhibiting a larger and more rounded pronotum leagues, among others, to access subterranean biodiversi- than is proper to this species, and neither the caves nor ty in Portugal in karst areas are providing new interesting the geographic area of Portugal were specified. information (Reboleira et al 2011). The first hypogean taxon described from Portugal was the troglobiont spider Nesticus lusitanicus Fage, 1931, based on specimens collected during a cave survey and METHODS exploration by the archaeologist Henri Breuil and the ge- ologist Ernest Fleury, in the beginings of the last century The entire bibliography on Portuguese subterranean (Gama & Afonso 1994). biology has been revised (excluding that on Macarone- Received 23 September; Accepted 5 December 2012 52 A.S.P.S. Reboleira et al. sian archipelagos and references to Chiroptera) since the Chronologically, the proportion of higher number of first publication in 1870 until November 2012. publications is not necessarily the result of higher num- The term hypogean is used for the species that live ber of new described taxa (Fig. 1). exclusively in subterranean habitats beneath the soil The decade of 1960 was the most productive in num- layers, displaying clear adaptations to the subterranean ber of such publications, but the largest number of new environment, expressed morphologically as convergent hypogean taxa were described in the decades of 1940 and evolution, such as integument depigmentation, absence especially 1980 (Fig. 1). or reduced ocular structures, elongated appendages, and The knowledge of the Portuguese stygofauna in- wing reduction in insects. The endogean fauna occurring creased noticeably from the 40’s until the 60’s, but the inside the soil is excluded of the aim of this catalogue. greatest expansion was in the 80’s with the description of 19 new species (Fig. 1). The researchers of the former Instituto de Investigação “Dr. Augusto Nobre” from Porto Results University gave the greatest contribution to the descrip- tion of new species, being only produced out of this in- As much as 138 scientific publications have been stitute 11% of this knowledge. José Maria Braga (25 spe- published between 1870 and November 2012, dealing on cies) and Odette Afonso (21 species) contributed to 74% the hypogean fauna of mainland Portugal, from where 27 of the Portuguese stygobiont species (Fig. 2). troglobiont and 63 stygobiont species have been described The 1940’s were the most productive decade of taxo- so far (see Table 1 and 2 for taxonomic information). nomical studies on the terrestrial hypogean fauna as a re- Fig. 1 - Number of publications and described hypogean taxa per decade until August 2012. Fig. 2 - Top contributors first authors with five or more publications on hypogean fauna in Portugal and their number of described species until November 2012. Hypogean taxa from Portugal 53 Fig. 3 - Number of hypogean species per higher taxonomic groups, until November 2012. A) Troglobionts and B) Stygobionts. sult of the great sampling efforts of Barros Machado (Fig. Juberthie, C., D. Delay, M. Bouillon. 1980. Extension 2), but the activity almost stopped during the 70´s and the du milieu souterrain en zone non-calcaire: description 80’s. The most important contribution in the past, in num- d’un nouveau milieu et de son peuplement par les Colé- ber of new troglobiontic taxa was afforded by Vandel, optéres troglobies. Mémoires de Biospéologie 7: 19-52. who described 6 species of terrestrial isopods (Vandel Pipan, T., D.C. Culver. 2012. Convergence and diver- 1946) (Fig. 3). Isopods are the most diverse group (Fig. gence in the subterranean realm: a reassessment. Biologi- 3) in the Portuguese subterranean fauna, characterized cal Journal of the Linnean Society 107: 1-14. by the absence of hypogean species belonging to groups Reboleira A.S.P.S., P.A.V. Borges, F. Gonçalves, A. normally with troglobionts in neighbouring Spain, such Serrano, P. Oromí. 2011. The subterranean fauna of a biodi- as Gastropods, Palpigrads, Diplopods, Leptodirine bee- versity hotspot region - Portugal: an overview and its con- tles, and other Arthropod groups. servation. International Journal of Speleology 40, 1: 23-37. Vandel, A. 1964. Biospéologie: La Biologie des Ani- maux cavernicoles. Paris: Ed. Gauthier-Villars. ACKNOwLEDGEMENTS we would like to express our gratitude to F. Tatá BIBLIOGRAPHIC catalogue OF SUBTERRA- Regala, V.M. Ortuño and Luís Mendes for kindly pro- NEAN fauna FROM Portugal viding some bibliography. The Portuguese Foundation for Science and Technology (FCT, Portugal) financed A Ana Sofia Reboleira by means of a Ph.D. grant (SFRH/ Afonso, O. 1978a. Une nouvelle espèce d’Isopode BD/45744/2008) and within the framework of KarstRisk troglobie du Portugal. Publicações do Instituto de Zoo- project (PTDC/AAC-AMB/114781/2009). logia «Dr. Augusto Nobre», 61: 11-23. Afonso, O. 1978b. Amphipoda hypogés du Portugal. Publicações do Instituto de Zoologia “Dr. Augusto No- REFERENCES bre”, 142: 11-26. Afonso, O. 1979. Un Proasellus (Crust. Isopoda) Culver, D.C., T. Pipan. 2009. Biology of caves and oth- nouveau du Portugal. Anais da Faculdade de Ciências er subterranean habitats. Oxford: Oxford University Press. Universidade do Porto, 61(1-4): 91-104. Fan, Q.H. 2006. Bibliographic analysis of acarolog- Afonso, O. 1982a. Description d’une nouvelle espèce ical papers published in Zootaxa from 2001 to 2005, de Proasellus (Crust. Isopoda) du Portugal et des consi- with a catalogue of described new taxa. Zootaxa 1385: dérations sur la phylogénie de son genre. Publicações do 53-66. Instituto de Zoologia «Dr. Augusto Nobre», 163: 1-15. 54 A.S.P.S. Reboleira et al. Afonso, O. 1982b. Sur un Proasellus (Crustacea, Isopo- Bellés, X. 1987. Fauna cavernícola i intersticial de la da, Asellidae) cavernicole nouveau
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