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A new epigean species of Trichoniscoides (Isopoda: Oniscidea: Trichoniscidae) from the northwest of the Iberian Peninsula

Abstract

The genus Trichoniscoides Sars, 1899 is composed mostly of cave-dwelling species. In the Iberian Peninsula, 32 species, and three additional species of uncertain status (partially described but not named), are currently known to occur. We were able to study one of these unnamed species, clarify its status, and name it. Most of the Iberian species of this genus are troglobiotic (22) and the ten remaining are epigean or troglophilic. The description of the new species from the surface can help in understanding the variability observed in the troglobitic species. The new, formally described, species has morphological characteristics that fit in several currently proposed groups for the genus, therefore, it is not possible to attribute Trichoniscoides gallaecus n. sp. to any of them.

Keywords:
New species; Portugal; Spain; taxonomy; terrestrial isopod

INTRODUCTION

The genus TrichoniscoidesSars, 1899Sars GO 1899. An account of the Crustacea of Norway, with short descriptions and figures of all the species. Vol. II. Isopoda. Bergen, Bergen Museum, 270p. https://www.biodiversitylibrary.org/item/100338-page/9/mode/1up
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is part of the family Trichoniscidae Sars, 1899 that is mostly comprised of cave-dwelling species. So far, the genus comprises 52 described species, of which 32 have been recorded in the Iberian Peninsula with 22 being further troglobiotic (Cifuentes et al., 2021Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
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; Cifuentes and Prieto, 2022Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
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). The genus has a known Atlantic distribution, being mostly present in Western Europe, with very few species reported from outside this area. Only Trichoniscoides sarsiPatience, 1908Patience, A. 1908. On Trichoniscoides albidus (Budde-Lund) and T. sarsi, Patience (nom. nov.). Annals and Magazine of Natural History (Ser. 8), 2: 84−88. https://doi.org/10.1080/00222930808692454
https://doi.org/10.1080/0022293080869245...
was found in greenhouses in Finland and introduced into North America (Schmallfus, 2003Schmalfuss H 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde, Serie A, 654: 1−341. https://www.biodiversitylibrary.org/page/60882444
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). The genus was recently revised in the Iberian Peninsula and an identification key is available (Cifuentes, 2019aCifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
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), although a few more species have been described since then (e.g., Cifuentes and Prieto, 2021Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
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; 2022Cifuentes J and Prieto CE 2022. Revisitando las antiguas localidades del Sistema Ibérico de Trichoniscoides cavernicola (Budde-Lund, 1885) (Crustacea: Isopoda: Trichoniscidae), con la descripción de tres especies nuevas (España). Boletín de la Sociedad Entomológica Aragonesa, 70: 335−343.). Nevertheless, more species remain undescribed in the region. For example, there are three potential species based on morphology with an uncertain status, since they were not formally described nor named, but simply mentioned as Trichoniscoides species A, B, and C (Gregory et al., 2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
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). More work is needed to better know the diversity of this genus and have a better understanding of its radiation and evolution.

A survey carried out on Portuguese isopods allowed collection of several specimens that match the short description made by Gregory et al. (2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
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) of Trichoniscoides species A. Considering that there has been a decade since this publication, the description of these authors is not exhaustive and does not provide important information on this species morphology, and that it was not formally described, we believe it is convenient to name it, provide a more detailed description, and discuss the morphological characters that allow it to be associated with, but also differentiated from, the rest of the species in the genus.

MATERIAL AND METHODS

Oniscidea specimens were captured in the northwest of Portugal, in the west of the Minho region (Fig. 1), by direct search underneath rocks, litter, dead wood, and tree bark. The region is composed of a heterogenous landscape with forest, agricultural, and urban areas. Forest areas are dominated by Eucalyptus globulus plantations, but there are other tree species plantations, namely that of Pinus pinaster. Some areas, mostly along watercourses or the edge of fields and plantations, have native tree cover usually dominated by Quercus robur or with invasive species such as Acacia melanoxylon, Acacia dealbata and Acacia longifolia. Agricultural areas are diverse, mainly with subsistence farming or corn plantations. Urbanized areas range from small rural villages with a large proportion of agricultural and forest areas, to cities with a few tens of thousands of inhabitants. The region is mainly composed of metamorphic and granitic rocks from the Palaeozoic (Direcção Geral de Minas e Serviços Geológicos, 1952Direcção Geral de Minas e Serviços Geológicos 1952. Carta geológica de Portugal. Direcção Geral de Minas e Serviços Geológicos, Serviços Geológicos, Portugal.). The region has annual mean temperatures between 11 and 13 °C and between 2,200 and 2,500 mm of precipitation, being one of the rainiest regions in Portugal (IPMA, 2019IPMA (Instituto Português do Mar e da Atmosfera) 2019. Portal do Clima. Instituto Português do Mar e da Atmosfera. Available at Available at http://portaldoclima.pt/pt/ . Accessed on 28 June 2022.
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).

Figure 1.
Location of the Iberian Peninsula (Spain and Portugal) in western Europe and distribution of Trichoniscoides gallaecus n. sp. (bibliographical references in yellow circles; current material studied in red circles).

The holotype and one paratype are deposited in the Museo Nacional de Ciencias Naturales de Madrid (MNCN), 4 paratypes are in the invertebrate collection of InBIO Barcoding Initiative (IBI, reference start with INV), from Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO), Campus de Vairão, University of Oporto, and the remaining two are in the collection of the first author (JC).

Distribution map (Fig. 1) was created using QGIS Desktop 3.22.2, using the coordinates obtained in the field during the capture of the analyzed Portuguese specimens and the coordinates of the bibliographic locations available in Gregory et al. (2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
https://www.bmig.org.uk/sites/www.bmig.o...
). To facilitate species identification, a figure of the studied species was included (Fig. 2). The drawings (Fig. 3) were prepared with the graphic editor InKscape (https://inkscape.org/es/).

Figure 2.
Trichoniscoides gallaecus n. sp. A, Habitus; B, cephalon and anterior pereonites, lateral view; C, male pereopod 1, carpus hyaline scales; D, male pereopod 7, merus curved hook; E, male pleopod I exopod; F, male pleopod I exopod, protuberances; G, male pleopod II endopod and exopod; H, male pleopod II endopod, tip.

Figure 3.
Trichoniscoides gallaecus n. sp. A, Tegument granules and scale-setae; B, antennule; C, antenna; D, male pereopod 1; E, male pereopod 7; F, male pleopod I endopod; G, male pleopod I exopod; H, male pleopod II endopod; I, male pleopod II exopod.

SYSTEMATICS

Family Trichoniscidae Sars, 1899Sars GO 1899. An account of the Crustacea of Norway, with short descriptions and figures of all the species. Vol. II. Isopoda. Bergen, Bergen Museum, 270p. https://www.biodiversitylibrary.org/item/100338-page/9/mode/1up
https://www.biodiversitylibrary.org/item...

Genus Trichoniscoides Sars, 1899Sars GO 1899. An account of the Crustacea of Norway, with short descriptions and figures of all the species. Vol. II. Isopoda. Bergen, Bergen Museum, 270p. https://www.biodiversitylibrary.org/item/100338-page/9/mode/1up
https://www.biodiversitylibrary.org/item...

Trichoniscoides gallaecus n. sp.

(Figs. 2, 3; Gregory et al., 2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
https://www.bmig.org.uk/sites/www.bmig.o...
: 12, fig. 2)

Zoobank: urn:lsid:zoobank.org:act:3D8DCDF5-B8E0-4980-A492-802D7A223D3B

Type material. Holotype: male (MNCN 20.04/20602), 27 May 2022, Fradelos, Braga, Portugal, (41°21'16"N 8°36'22"W), coll. L.P. da Silva. Paratypes: 2 females (INV18000, INV18001), 27 March 2022, same data as holotype; 1 male (JC667), same data as holotype; 4 females (INV18002, INV18003, JC668, MNCN 20.04/20568), same data as holotype.

Diagnosis. Dark red coloration. Eye apparatus composed of single large ocellus. Integument with granulations. Antennae with scaly tubercles and 3 flagellar segments. Male pereopods 1 and 7 with sexual dimorphism. Male pleopods: endopod I with large triangular distal lobe; exopod I trapezoidal, with 2 very different hairy protuberances; endopod II very wide, narrowing sharply near tip; exopod II trapezoidal, lacking inner tip.

Description. Maximum observed length: female 2.5 mm, male 2.1 mm. Coloration intense dark red (Fig. 2A, B), stable in ethanol for at least 6 months, interrupted by light color from muscle attachments. Tegument covered by granulations (Fig. 2A, B) with large triangular scale-setae with crenated edge surrounded by accessory scales (Fig. 3A). Cephalon with 5 granulation rows, including 1 row on posterior edge. Pereon with 3 rows on first pereonite and 2 rows on remaining pereonites, plus another row on posterior edge of each tergite. These granulations fading towards posterior region. Pleon with row of small granulations on posterior edge of all pleonites. Eye apparatus formed by large ocellus (Fig. 2B) in black pigmented spot. Antennules with 3 segments, last one with group of 3 long aesthetascs (Fig. 3B). Antennae short, and not reaching posterior edge of 2nd pereon tergite and presenting hairy tubercles (Fig. 3C). Antennal flagellum composed of 3 segments, first the shortest, second with group of long aesthetascs inserted in middle and extending behind insertion of 3rd segment. Pereopod 1 of both sexes bearing fan-shaped group of setae in carpus tip (Fig. 3D).

Male sexual characters: Pereopod 1 merus and carpus with hyaline scales (Figs. 2C, 3D), but in merus only present in distal half. Pereopod 7 merus with strong curved hook (Figs. 2D, 3E) inserted in middle and with small seta at hook tip. Pleopod I: endopod triangular, with large distal lobe (Fig. 3F); exopod trapezoidal, with 2 hairy protuberances of very different sizes, internal one very small, external one very long and thick, getting progressively thinner until ending in sharp tip (Figs. 2E, F, 3G). Pleopod II: endopod surrounded by hyaline membrane (Figs. 2G, 3H), with distal segment 3 times longer than basal segment, sharply narrowing at two-thirds of its length, ending in fine but flattened tip (Figs. 2H, 3H); exopod trapezoidal, lacking distal tip and bearing small hairy protuberance surrounded by many fan-shaped setae; these setae also on inner and outer edges (Figs. 2G, 3I).

Etymology. The specific epithet is a reference to gallaecus, a Latin word used by the Romans to identify the indigenous peoples who lived north of the Douro River in Portugal and a large part of Galicia (Spain).

Ecology. Epigean species found below (or within) deep moist litter (usually > 10cm), mostly composed of dead leaves of Q. robur. It was found together with Porcellio herminiensisVandel, 1946Vandel A 1946b. Especes francaises, nouvelles ou peu connues, de Trichoniscidae (Isopodes terrestres). Bulletin de la Societe Zoologique de France, 71: 53−72.a, Porcellio debueniDollfus, 1892Dollfus A 1892. Catalogue raisonné des isopodes terrestres de l’Espagne (espèces signalées jusqu’à ce jour et description d’espèces nouvelles). Anales de la Sociedad Española de Historia Natural, 21: 161-190. https://www.biodiversitylibrary.org/item/235119 - page/175/mode/1up
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, and Trichoniscus pusillus s.l. Brandt, 1833Brandt I 1833. Conspectus Monographiae Crustaceorum Oniscodorum Latreillii. Byulleten moskovskogo Obshchestva Ispŷtateleĭ Prirodŷ, 6: 171-193.. Gregory et al. (2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
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) mentioned that it can also be found beneath dead wood and stones among leaf-litter.

Distribution. The species is so far known from the northwest of Portugal, in the Minho region, and in the southwest of Galicia, Spain (Fig. 1).

Remarks. Gregory et al. (2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
https://www.bmig.org.uk/sites/www.bmig.o...
) found females of 3 mm length and males of 2.25 mm. They also mentioned that all pigmentation is rapidly lost after preservation in alcohol.

DISCUSSION

Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
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) classified the species in the genus Trichoniscoides into four groups based on the structure of the endopod of the male pleopod II. These groups are an Atlantic group, with a long and fine tip; a Pyrenean group, with a short, thin, sharp, and recurved tip; an Aquitanian-Languedocian group, with a very short, broad, and recurved tip; and an Alpine group, with the tip straight or recurved, but less long than in the Atlantic group. In the Alpine group, Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
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) included five species: Trichoniscoides helveticus (Carl, 1908Carl J 1908. Monographie der Schweizerischen Isopoden. Denkschriften Schweizerischen Naturforschenden Gesellschaft, 42: 107−242.) known from Germany, Belgium, France, the Netherlands, and the Czech Republic (Schmalfuss, 2003Schmalfuss H 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde, Serie A, 654: 1−341. https://www.biodiversitylibrary.org/page/60882444
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); Trichoniscoides heroldiVandel, 1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362., and Trichoniscoides mixtusRacovitza, 1908Racovitza E-G 1908. Isopodes terrestres (Second Serie). Biospéologica. IX. Isopodes terrestres (2 serie). Archives de Zoologie Experimentale et Generale (IVe Série), 9: 239−415. https://www.biodiversitylibrary.org/item/28438 - page/338/mode/1up
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, that are only known from France (Schmalfuss, 2003Schmalfuss H 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde, Serie A, 654: 1−341. https://www.biodiversitylibrary.org/page/60882444
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); Trichoniscoides pulchellusLegrand, 1950Legrand J 1950. Les isopodes terrestres des Alpes de Savoie. Bulletin de la Société zoologique de France, 75: 45−62. cited from France and Switzerland (Schmalfuss, 2003Schmalfuss H 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde, Serie A, 654: 1−341. https://www.biodiversitylibrary.org/page/60882444
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) and Trichoniscoides remyiBonnefoy, 1945Bonnefoy F 1945. Note sur quelques Trichoniscidae (Isopodes terrestres) des Alpes francaises. Bulletin Societe Zoologique de France, 70: 3−7. also only known from France (Schmalfuss, 2003Schmalfuss H 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beiträge zur Naturkunde, Serie A, 654: 1−341. https://www.biodiversitylibrary.org/page/60882444
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). Among these species the endopod differs in structure. In T. mixtus and T. pulchellus, the endopod tip has a wide hyaline membrane; in T. helveticus, the endopod tip is complex, with two appendages; and in T. heroldi and T. remyi, the endopod tip is short and flattened, similar to that present in T. gallaecus n. sp. Considering this character alone, the new species should belong to the Alpine group, despite its known distribution being very different from any of the previous species.

In this genus there is an important derived character, sexual dimorphism in the pereopods, that can be in the first pair, the first two or three pairs, or in the seventh pair, but it is possible to find different combinations of these, in different species. For example, in Trichoniscoides cantabricusCifuentes and Prieto, 2021Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
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, the dimorphism is only present in the anterior pereopods; in Trichoniscoides picturarumVandel, 1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362., it is only in the seventh pair; and, finally, in some species the dimorphism occurs in both the anterior pereopod pairs and the seventh pair, as in T. gallaecus n. sp., that has sexual dimorphism in the first and seventh pairs. In the anterior pairs, the dimorphism is characterized by the presence or absence? of hyaline scales on the inner side of the merus and carpus. The differences in the seventh pair are more variable and can be found in species belonging to different groups, as for example in the Atlantic group, Trichoniscoides broteroiVandel, 1946aVandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
https://www.fc.up.pt/fa/index.php?p=nav&...
, Trichoniscoides irregularisSchmölzer, 1955Schmölzer K 1955. Isopoda terrarum mediterranearum. 1. Mitteilung: Über neue und bekannte Landasseln der Pyrenaenhalbinsel. Eos (Madrid), 31: 155-215. https://digital.csic.es/bitstream/10261/154616/1/1955-31-1_2-006.pdf
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, and Trichoniscoides subterraneusVandel, 1946aVandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
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; in the Pyrenean group, Trichoniscoides calcarisCruz and Dalens, 1989Cruz A and Dalens H 1989. Especies nuevas o poco conocidas de isopodos terrestres de la Peninsula Iberica. 1. Isopodos cavernicolas de la España oriental (Crustacea; Oniscidea). Bulletin de la Societe d'Histoire Naturelle de Toulouse, 125: 91−98. https://gallica.bnf.fr/ark:/12148/bpt6k6556448v/f93.image.r=Oniscedea
https://gallica.bnf.fr/ark:/12148/bpt6k6...
, Trichoniscoides picturarum and Trichoniscoides vandeliDalens, 1966Dalens H 1966. Une nouvelle espece endogene de Trichoniscoides, T. vandeli n. sp. provenant des Asturies: T. cassagnaui n. sp. et T. asturiensis n. sp. (crustaces, isopodes terrestres). Annales de Speleologie, 21: 651−653.; in the Aquitanian-Languedocian group, Trichoniscoides bonnetiVandel, 1946bVandel A 1946b. Especes francaises, nouvelles ou peu connues, de Trichoniscidae (Isopodes terrestres). Bulletin de la Societe Zoologique de France, 71: 53−72., Trichoniscoides cadurcensisVandel, 1934Vandel A 1934. Description d'une nouvelle espece d'Isopode terrestre, Trichoniscoides cadurcensis, n. sp., suivie de remarques sur la distribution géographique des especes du genre Trichoniscoides. Bulletin de la Societe d'Histoire Naturelle de Toulouse, 70: 193−196. https://gallica.bnf.fr/ark:/12148/bpt6k6555582x/f201.item
https://gallica.bnf.fr/ark:/12148/bpt6k6...
, Trichoniscoides consoranensisVandel, 1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362., Trichoniscoides drescoiVandel, 1954 Vandel A 1954. Une nouvelle espèce cavernicole de Trichoniscoides, T. drescoi n. sp., provenant des Asturies (crustacés; isopodes terrestres). Notes biospéologiques, 9: 67−69.and Trichoniscoides sarsi or even Trichoniscoides sicoensisReboleira and Taiti, 2015Reboleira AS; Gonçalves F; Oromí P and 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
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which is not clear to which of the groups established by Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
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) it belongs, as indicated by Reboleira and Taiti (2015Reboleira AS; Gonçalves F; Oromí P and 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
https://doi.org/10.5852/ejt.2015.161...
). Nevertheless, none of the species that Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
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) included in the Alpine group has sexual dimorphism in the seventh pair of pereopods. A hook with a short seta on the merus is one of the different forms of dimorphism indicated by Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
https://faunedefrance.org/bibliotheque/d...
) important to define the Aquitanian-Languedocian group. This character is present in the five species previously mentioned as belonging to this group, but is also present in T. gallaecus n. sp. The structure of this hook in T. gallaecus n. sp. allows it to resemble more closely T. broteroi, T. cadurcensis, T. consoranensis, T. sarsi, and T. sicoensis. In T. consoranensis and T. sarsi, the hook is located at the merus base and the endopod II does not end in a narrow, flattened tip as in T. gallaecus n. sp. Trichoniscoides cadurcensis has the male exopod of the pleopod II very similar to that of T. gallaecus n. sp., but in T. cadurcensis, in pereopod 7, the position of the hook is at the base of the merus and the tip of the pleopod II is different from T. gallaecus n. sp. Trionychoids broteroi and T. sicoensis also have the hook in the middle of the merus?, but in T. broteroi the male exopod I has small, equally hairy, protuberances, which are subequal in T. sicoensis. The male endopod II of both species is very different from T. gallaecus n. sp., where in the exopod I the outer protuberance is hypertrophied, the inner protuberance is very small, and the endopod II tip is narrow and flattened. Summarizing, the structure of the endopod of the male pleopod II would place the newly described species in the Alpine group of Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
https://faunedefrance.org/bibliotheque/d...
), a group that for this author is very close to the Atlantic group, with which the new species shares its distribution, being known from Portugal and Spain (Fig. 1), but from which it is separated by the sexual dimorphism of the male pereopod 7. It also has some characteristics similar to the species in the Aquitanian-Languedocian group, but it cannot be included in this group if we stick to the original divisions of Vandel (1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
https://faunedefrance.org/bibliotheque/d...
). It would be useful to make new criteria for the division of the groups of this genus, especially after having more knowledge of other European species, outside the Iberian Peninsula, allowing more effective grouping of all Trichoniscoides species, which currently cannot be confidently placed in any of the groups referred to above.

As already mentioned, the genus Trichoniscoides is composed mostly of cave-dwelling species, which have accessed the underground from the surface, likely through a secondary colonization in the face of unfavorable environmental conditions. In the Iberian Peninsula there are 32 valid species of the 52 known species of the genus (Cifuentes et al., 2021Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
https://doi.org/10.3989/graellsia.2021.v...
; Cifuentes and Prieto, 2022Cifuentes J and Prieto CE 2022. Revisitando las antiguas localidades del Sistema Ibérico de Trichoniscoides cavernicola (Budde-Lund, 1885) (Crustacea: Isopoda: Trichoniscidae), con la descripción de tres especies nuevas (España). Boletín de la Sociedad Entomológica Aragonesa, 70: 335−343.). Of these 32 species, 22 are troglobiotic, indicating the importance of the subterranean environment as a shelter and area for species diversification. Two of the 10 remaining Iberian species are troglophiles, Trichoniscoides jeanneliVandel, 1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362. known from the Spanish provinces of Huesca and Navarra (Vandel, 1952Vandel A 1952. Biospeologica LXXIII. Isopodes terrestres (troisième série). Archives de Zoologie expérimentale et générale, 88: 2313−2362.; 1960Vandel A 1960. Faune de France, vol. 64. Isopodes terrestres (première partie). Paris, Lechevalier, 416p. https://faunedefrance.org/bibliotheque/docs/VANDEL(FdeFr64)IsopodeTerrestre.pdf
https://faunedefrance.org/bibliotheque/d...
; Schmölzer, 1971Schmölzer K 1971. Die Landisopoden der Iberischen Halbinsel. Madrid, Consejo Superior de Investigaciones Científicas, 161p.; Cifuentes, 2019aCifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
; Cifuentes et al., 2021Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
https://doi.org/10.3989/graellsia.2021.v...
) and Trichoniscoides machadoiVandel, 1946aVandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
https://www.fc.up.pt/fa/index.php?p=nav&...
, known from the Portuguese districts of Aveiro, Braga, Coimbra, Faro, Leiria, and Lisbon (Vandel, 1946aVandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
https://www.fc.up.pt/fa/index.php?p=nav&...
; Reboleira et al., 2015Reboleira AS; Gonçalves F; Oromí P and 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
https://doi.org/10.5852/ejt.2015.161...
). Finally, the eight remaining species are epigean: Trichoniscoides ancaresiSchmölzer, 1955Schmölzer K 1955. Isopoda terrarum mediterranearum. 1. Mitteilung: Über neue und bekannte Landasseln der Pyrenaenhalbinsel. Eos (Madrid), 31: 155-215. https://digital.csic.es/bitstream/10261/154616/1/1955-31-1_2-006.pdf
https://digital.csic.es/bitstream/10261/...
, known in Spain from Asturias, Cantabria, Lugo, and Pontevedra (Schmölzer, 1955Schmölzer K 1955. Isopoda terrarum mediterranearum. 1. Mitteilung: Über neue und bekannte Landasseln der Pyrenaenhalbinsel. Eos (Madrid), 31: 155-215. https://digital.csic.es/bitstream/10261/154616/1/1955-31-1_2-006.pdf
https://digital.csic.es/bitstream/10261/...
; 1971Schmölzer K 1971. Die Landisopoden der Iberischen Halbinsel. Madrid, Consejo Superior de Investigaciones Científicas, 161p.; Cifuentes, 2019bCifuentes J 2019b. Los isópodos terrestres de Galicia, España (Crustacea: Isopoda, Oniscidea). Graellsia, 75(2): e098. https://doi.org/10.3989/graellsia.2019.v75.243.
https://doi.org/10.3989/graellsia.2019.v...
); the Asturian species Trichoniscoides asturiensisDalens, 1972Dalens H 1972. Deux nouvelles espèces de Trichoniscoides provenant des Asturies: T. cassagnani n. sp. et T. asturiensis n. sp. (crustacés; isopodes terrestres). Bulletin de la Société d’Histoire naturelle de Toulouse, 108: 426−432. https://gallica.bnf.fr/ark:/12148/bpt6k6556269b/f94.image.r=Dalens
https://gallica.bnf.fr/ark:/12148/bpt6k6...
and Trichoniscoides cassagnaniDalens, 1972 Dalens H 1972. Deux nouvelles espèces de Trichoniscoides provenant des Asturies: T. cassagnani n. sp. et T. asturiensis n. sp. (crustacés; isopodes terrestres). Bulletin de la Société d’Histoire naturelle de Toulouse, 108: 426−432. https://gallica.bnf.fr/ark:/12148/bpt6k6556269b/f94.image.r=Dalens
https://gallica.bnf.fr/ark:/12148/bpt6k6...
cited by Dalens (1972Dalens H 1972. Deux nouvelles espèces de Trichoniscoides provenant des Asturies: T. cassagnani n. sp. et T. asturiensis n. sp. (crustacés; isopodes terrestres). Bulletin de la Société d’Histoire naturelle de Toulouse, 108: 426−432. https://gallica.bnf.fr/ark:/12148/bpt6k6556269b/f94.image.r=Dalens
https://gallica.bnf.fr/ark:/12148/bpt6k6...
), Trichoniscoides enoliCifuentes, 2019 Cifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
by Cifuentes (2019aCifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
) and Trichoniscoides irregularis by Schmölzer (1955Schmölzer K 1955. Isopoda terrarum mediterranearum. 1. Mitteilung: Über neue und bekannte Landasseln der Pyrenaenhalbinsel. Eos (Madrid), 31: 155-215. https://digital.csic.es/bitstream/10261/154616/1/1955-31-1_2-006.pdf
https://digital.csic.es/bitstream/10261/...
) and Cifuentes (2019aCifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
). Trichoniscoides lusitanusVandel, 1946a Vandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
https://www.fc.up.pt/fa/index.php?p=nav&...
is the only epigean species with a large known distribution, since it has been cited from the Portuguese districts of Braga, Porto, and Viana do Castelo (Vandel, 1946aVandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
https://www.fc.up.pt/fa/index.php?p=nav&...
; Gregory et al., 2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
https://www.bmig.org.uk/sites/www.bmig.o...
) and from the Spanish provinces of A Coruña, Cantabria, Orense, and Pontevedra (Schmölzer, 1955Schmölzer K 1955. Isopoda terrarum mediterranearum. 1. Mitteilung: Über neue und bekannte Landasseln der Pyrenaenhalbinsel. Eos (Madrid), 31: 155-215. https://digital.csic.es/bitstream/10261/154616/1/1955-31-1_2-006.pdf
https://digital.csic.es/bitstream/10261/...
; 1971Schmölzer K 1971. Die Landisopoden der Iberischen Halbinsel. Madrid, Consejo Superior de Investigaciones Científicas, 161p.; Gregory et al., 2012Gregory S; Lee P; Read HJ and 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 & Isopod Group, 26: 6−23. https://www.bmig.org.uk/sites/www.bmig.org.uk/files/bulletin/BullBMIG26(2012)p6-23_Gregory-etal_NWSpain.pdf
https://www.bmig.org.uk/sites/www.bmig.o...
; Cifuentes, 2019aCifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
; 2019bCifuentes J 2019b. Los isópodos terrestres de Galicia, España (Crustacea: Isopoda, Oniscidea). Graellsia, 75(2): e098. https://doi.org/10.3989/graellsia.2019.v75.243.
https://doi.org/10.3989/graellsia.2019.v...
). Trichoniscoides pitarquensisCruz, 1993Cruz A 1993. Especies nuevas o poco conocidas de isópodos terrestres de la Peninsula Ibérica. III. Trichoniscoides pitarquensis sp. n. y T. serrai sp. n. (Crustacea, Oniscidea, Trichoniscidae). Bulletin de la Société d’Histoire naturelle de Toulouse, 127: 15−21. is only known from the Spanish province of Teruel (Cruz, 1993Cruz A 1993. Especies nuevas o poco conocidas de isópodos terrestres de la Peninsula Ibérica. III. Trichoniscoides pitarquensis sp. n. y T. serrai sp. n. (Crustacea, Oniscidea, Trichoniscidae). Bulletin de la Société d’Histoire naturelle de Toulouse, 127: 15−21.) and Trichoniscoides saeroeensisLohmander, 1924Lohmander H 1924. Trichoniscoides saeroensis n. sp. Entomologisk Tidskrift, 45: 80-82. is only known from the Iberian Peninsula, in the province of Cantabria (Erhard, 1997Erhard F 1997. Das pleonale Skelet-Muskel-System von Titanethes albus (Synocheta) und weiterer Taxa der Oniscidea (Isopoda), mit Schlußfolgerungen zur Phylogenie der Landasseln. Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie), 550: 1−70. https://www.zobodat.at/pdf/Stuttgarter-Beitraege-Naturkunde_550_A_0001-0070.pdf
https://www.zobodat.at/pdf/Stuttgarter-B...
; Cifuentes et al., 2021Cifuentes J; Beruete E and Prieto CE 2021. Isópodos oniscídeos en cuevas de la región Vasco-Cantábrica, España (Crustacea: Isopoda). Boletín de la Sociedad Entomológica Aragonesa, 68: 270−288.). Summing up, the epigean species, with the exception of T. pitarquensis, are located, to the best of our current knowledge, in a narrow fringe with a climatic Atlantic influence in the north of the Iberian Peninsula, between Porto (Portugal) and Cantabria (Spain), just like T. gallaecus n. sp. The troglophile species have a larger known distribution, reaching the south of Portugal in the Faro district in the case of T. machadoi, but in this district it is only known to occur in caves (Vandel, 1946aVandel A 1946a. Crustacés isopodes terrestres (Oniscoïdea) épigés et cavernicoles du Portugal. Anaïs da Faculdade de Ciências do Porto, 30: 135−427. https://www.fc.up.pt/fa/index.php?p=nav&f=books.0066.0
https://www.fc.up.pt/fa/index.php?p=nav&...
; Reboleira et al., 2015Reboleira AS; Gonçalves F; Oromí P and 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
https://doi.org/10.5852/ejt.2015.161...
), and reaching the Pyrenean region, in the case of T. jeanneli, in the Spanish provinces of Huesca and Navara. It is important to note that there is no interruption between the Cantabrian and the Pyrenees Mountains, that are in contact in Navarra. The troglobiont species of Trichoniscoides from the Iberian Peninsula are not only more diverse, but some species are found far from the previously mentioned areas, as for example Trichoniscoides arlanzaCifuentes and Prieto, 2022Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
https://doi.org/10.3989/graellsia.2021.v...
in Cueva Negra (Hortigüela, Burgos); Trichoniscoides galianaCifuentes and Prieto, 2022Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
https://doi.org/10.3989/graellsia.2021.v...
in the Cueva de La Galiana Alta (Ucero, Soria); Trichoniscoides jonfernandeziCifuentes and Prieto, 2022 Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
https://doi.org/10.3989/graellsia.2021.v...
in the Cueva de la Mora (Arcos de Jalón, Soria) (see Cifuentes and Prieto, 2022Cifuentes J and Prieto CE 2021. Descripción de dos nuevas especies cavernícolas de Trichoniscinae Verhoeff, 1908 de la Cordillera Cantábrica (Crustacea: Isopoda: Trichoniscidae). Graellsia, 77(1): e124. https://doi.org/10.3989/graellsia.2021.v77.275
https://doi.org/10.3989/graellsia.2021.v...
); and Trichoniscoides marinaeCifuentes, 2019 Cifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
in Cueva del Chorrillo (Tamajón, Guadalajara) according to Cifuentes (2019aCifuentes J 2019a. 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 Asociación española de Entomología, 43(1−2): 27−54. http://www.entomologica.es/publicaciones-boletin/art1773
http://www.entomologica.es/publicaciones...
). These suggest an early colonization of these areas by endogean species, whose distribution likely retracted in the past and is now considerably smaller. The discovery and description of new species inhabiting the surface can help to understand the variability and diversity observed in troglobiont species.

ACKNOWLEDGMENTS

We would also like to thank Dra. Begoña Sánchez from the National Museum of Natural Sciences of Madrid, for her kindness in facilitating the access to the specimens deposited in the collections; and Dra. Vanessa Mata and two anonymous reviewers for reviewing the manuscript.

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ADDITIONAL INFORMATION AND DECLARATIONS

  • Consent for publication

    Both authors declare that they have reviewed the content of the manuscript and gave their consent to submit the document.
  • Data availability

    All study data are included in the article.
  • Funding and grant disclosures

    LPS benefits from an individual research contract (CEECIND/02064/2017) funded through Fundação para a Ciência e a Tecnologia (FCT/MEC). This work was co-funded by the project NORTE-01-0246-FEDER-000063, supported by Norte Portugal Regional Operational Programme (NORTE2020), under the PORTUGAL 2020 Partnership Agreement, through the European Regional Development Fund (ERDF).
  • Study association

    Not applicable.
  • Study permits

    Not applicable.

Edited by

Associate Editor:

Spyros Sfenthourakis

Data availability

All study data are included in the article.

Publication Dates

  • Publication in this collection
    17 July 2023
  • Date of issue
    2023

History

  • Received
    24 Oct 2022
  • Accepted
    18 Jan 2023
Sociedade Brasileira de Carcinologia Instituto de Biociências, UNESP, Campus Botucatu, Rua Professor Doutor Antônio Celso Wagner Zanin, 250 , Botucatu, SP, 18618-689 - Botucatu - SP - Brazil
E-mail: editor.nauplius@gmail.com