European Journal of Taxonomy 161: 1-61

http://dx.doi.org/10.5852/ejt.2015.161

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2015 • Reboleira A.S.P.S. et al.

Monograph

urn:lsid:zoobank.org:pub:2297E4A3-D279-4D0A-923C-D5E0D5DCB3C0

The cavernicolous Oniscidea (Crustacea: Isopoda) of Portugal

Ana Sofia P.S. REBOLEIRA 1 *, Fernando GONCAI.VKS 2 .

Pedro OROMI 3 & Stefano TAITI 4

lj2 Departamento de Biologia & CESAM, Universidade de Aveiro,

Campus Universitario de Santiago, 3810-193 Aveiro, Portugal.

13 Departamento de Biologia Animal, Facultad de Biologia, Universidad de La Laguna,

38206 La Laguna, Tenerife, Islas Canarias, Spain.

1 Zoological Museum, Natural History Museum of Denmark, University of Copenhagen,

Universitetsparken 15, 2100 Copenhagen 0, Denmark.

4 Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche,

Via Madonna del Piano 10, 50019 Sesto Fiorentino (Firenze), Italy.

* Corresponding author: [email protected]

2 Email: [email protected]

3 Email: [email protected]

4 Email: [email protected]

1 urn:lsid:zoobank.org:author:338DE845-4839-4EF5-B684-587C021F076C

2 urn:lsid:zoobank.org:author:4B8A2E04-3543-465F-9949-B7E1745DA931

3 urn:lsid:zoobank.org:author:00F2ACC4-308B-4BEE-BBA4-DE6AC8905449

4 urn:lsid:zoobank.org:author:62E97059-6AE5-4984-9ABB-7FB6F7358BD6

Abstract. The study of subterranean Oniscidea in Portugal has been neglected for nearly 70 years,

but recent investigations have revealed high diversity. All the terrestrial isopods known from caves

of mainland Portugal, including data from the literature and new material, are listed, revealing new

biodiversity patterns. Twenty-seven species, belonging to 15 genera and six families, are known, of

which 16 species are considered to be exclusively from subterranean ecosystems, i.e., troglobionts.

Trichoniscidae is the most diverse family. Seven species in the family Trichoniscidae and one species in the

family Styloniscidae are described as new (all with Reboleira & Taiti as authors): Trichoniscoides bellesi

sp. nov. from the Montejunto Massif; T. sicoensis sp. nov. from the Sico Massif; Metatrichoniscoides

salirensis sp. nov. from the typhonic valley of Caldas da Rainha; Troglonethes olissipoensis sp. nov. from

the Lisbon Peninsula; T. arrabidaensis sp. nov. from the Arrabida Massif; Miktoniscus longispina sp.

nov. from the Sico Massif and Cesaredas Plateau; Moserius inexpectatiis sp. nov. from the Estremenho

Massif; and Cordioniscus lusitanicus sp. nov. from Alentejo and Algarve, the southernmost provinces

of Portugal. The subgenus Trogleluma Vandel, 1946 (Armadillidiidae) is raised to genus level. In this

geographic region terrestrial isopods are the richest group of cave-adapted animals.

Key words. Isopoda, new species, troglobiont, subterranean ecosystems, Iberian Peninsula.

Reboleira A.S.P.S., Gongalves R, Oromi P. & Taiti S. 2015. The cavernicolous Oniscidea (Crustacea: Isopoda) of

Portugal. European Journal of Taxonomy 161: 1-61. http://dx.doi.org/10.58552/eit. 2015,161

European Journal of Taxonomy 161: 1-61 (2015)

Table of contents

Introduction .3

Material and methods.3

Results .6

Family Trichoniscidae Sars, 1899 .6

Genus Trichoniscoides Sars, 1899 .6

T. broteroi Vandel, 1946 .6

T. machadoi Vandel, 1946 .6

T. ouremensis Vandel, 1946 .7

T. serrai Cruz, 1993 .7

T. subterraneus Vandel, 1946 .8

T. meridionalis Vandel, 1946 .8

T. bellesi Reboleira & Taiti sp. nov.9

T. sicoensis Reboleira & Taiti sp. nov..13

Genus Metatrichoniscoides Vandel, 1943 .17

M. salirensis Reboleira & Taiti sp. nov.17

Genus Troglonethes Cruz, 1989 .21

T. olissipoensis Reboleira & Taiti sp. nov.21

T. arrabidensis Reboleira & Taiti sp. nov.25

Genus Miktoniscus Kesselyak, 1930 .28

M. longispina Reboleira & Taiti sp. nov.28

Genus Haplophthalmus Schobl, 1860 .32

EL. siculus Vandel, 1946 .32

Genus Moserius Strouhal, 1940 .32

M. inexpectatus Reboleira & Taiti sp. nov.32

Family Styloniscidae Vandel, 1952 .35

Genus Cordioniscus Grave, 1914.35

C. lusitanicus Reboleira & Taiti sp. nov..35

Family Philosciidae Kinahan, 1947 .39

Genus Anaphiloscia Racovitza, 1907 .39

A. sicula Arcangeli, 1934 .39

Genus Ctenoscia Verhoeff, 1938 .40

C. minima (Dollfus, 1892).40

Family Platyarthridae Verhoeff, 1949 .40

Genus Trichorhina Budde-Tund, 1908 .40

T. anophthalma Arcangeli, 1935 .40

Family Porcellionidae Brandt, 1831 .42

Genus Porcellionides Miers, 1877 .42

P. pruinosus (Brandt, 1833) .42

P. sexfasciatus (Budde-Tund, 1885) .43

P. cingendus (Kinahan, 1857) .43

Genus Porcellio Tatreille, 1804 .43

P. cavernicolus Vandel, 1946 .43

P. dilatatus dilatatus Brandt, 1831 .46

P. dispar Verhoeff, 1901 .47

P. echinatus Tucas, 1849 .47

Family Armadillidiidae Brandt, 1833 .47

Genus Eluma Budde-Tund, 1885 .47

E. caelata (Miers, 1877).47

E. tuberculata Cruz, 1991 .49

REBOLEIRAA.S.P.S. et al., Oniscidea from caves of Portugal

Genus Paraschidium Verhoeff, 1918 .50

Paraschizidium sp.50

Genus Trogleluma Vandel, 1946 .50

T. machadoi (Vandel, 1946) .51

Discussion .54

Acknowledgements .57

References .57

Introduction

Terrestrial isopods (suborder Oniscidea) play a very important role in the decomposition of organic

matter in land ecosystems (Hornung 2011). In subterranean ecosystems they occupy a key-ecological

position, recycling organic detritus and being frequent prey of cave predators (Rezac et al. 2008).

Terrestrial isopods are successful land colonizers, but a great majority of species have a high dependence

on hygrophilous habitats (Hornung 2011). In fact, they are quite abundant in subterranean ecosystems.

Several species of oniscideans have adapted to the aphotic environment, where they can be found in the

terrestrial and in the aquatic compartment, including species with amphibian behaviour (Taiti & Xue

2012; Broly et al. 2013; Campos-Filho et al. 2014).

Oniscidea represent the most diverse group among the Portuguese cave-obligate fauna (Reboleira et al.

2011a, 2013b). Taxonomic studies on the Oniscidea from mainland Portugal are scarce and included

mainly in two publications recording epigean and subterranean species by Arcangeli (1935a) and Vandel

(1946), based on material collected by A. Barros Machado (for a complete list of Portuguese terrestrial

isopods see Schmalfuss 2003).

Prior to this study, only seven species of Oniscidea had been recorded as cave-adapted from karst massifs

in mainland Portugal (Reboleira et al. 2011a, 2013b). Five species in the family Trichoniscidae were

known: one from the north of the country ( Trichoniscoides serrai Cruz, 1993) and four from central

Portugal, i.e., Trichoniscoides broteroi Vandel, 1946 from the Sico karst massif, and T. subterraneus

Vandel, 1946, T. meridionalis Vandel, 1946 and T. ouremensis Vandel, 1946 from the Estremenho karst

massif. The family Porcellionidae was represented by the species Porcellio cavernicolus Vandel, 1946,

known from Sico, and the family Armadillidiidae by Troglarmadillium ( Trogleluma) machadoi Vandel,

1946, found in caves of the Algarve in the south of the country (Vandel 1946; Cruz 1993).

Recent investigations in caves of karst areas from Portugal revealed the presence of several new species

of terrestrial isopods. The oniscidean fauna from the subterranean environment of mainland Portugal is

here discussed based on literature data and the new material examined.

Material and methods

Field work was performed between 2007 and 2014 in many karst caves from mainland Portugal and

terrestrial isopods have been collected in 29 of these caves (Table 1, Fig. 1). The original Portuguese

names for caves are maintained throughout the text, i.e., “Grata” for a horizontal cave, “Algar” for a

vertical cave and “Lapa” for a small cave.

Each cave was monitored in two different zones: just beyond the twilight zone and in its deeper parts.

Specimens were collected using baited pitfall traps and active search in each cave, and most were

preserved in 70% ethanol, while some specimens were preserved in 100% ethanol for future molecular

analyses. Temperature was measured with an Aspiration Psychrometer 761 (Lambrecht, Gottingen).

Identifications are based on morphological characters and the pertinent literature. The terminology used

in species descriptions is mainly based on Vandel (1960a, 1962). Specimens were studied under Wild

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Table 1 . Studied caves in Portugal. Coordinates: UTM WGS; E = entrance altitude in metres; T = mean

temperature at soil level in °C.

Cave

Karst area

Coordinates

Grata de Santo Adriao

Vimioso

41°32’53.9” N, 6°26’40.1” W

580

12.0

Grata d’el Rey

Cantanhede

40°17’38.8” N, 8 0 32’49.3” W

13.2

Grata da Arrifana

Sico

40°05’22.53” N, 8°30 , 55.76” W

300

16.7

Grata do Algarinho

Sico

39°53 , 04.5 ,, N, 8°23 , 05.0 ,, W

208

14.9

Grata do Soprador do Carvalho

Sico

39°59 , 10.1 ,, N, 8°22 , 57.7 ,, W

15.1

Grata da Senhora da Estrela

Sico

39°55 , 41.2 ,, N, 8°32 , 59.5 ,, W

380

13.8

Grata da Ceramica

Sico

39°55’36.6” N, 8°31 , 03.6 ,, W

355

15.0

Algar da Ervilha

Sico

39°56’23.54” N, 8°32’24.39” W

335

15.1

Grata de Sao Simao

Sico

39°57 , 47.31”N, 8°28’13.62” W

335

14.6

Grata Abrigo Tomar I

Estremenho

39°39 , 34.7” N, 8°25’03.6”W

14.6

Grata do Papagaio

Estremenho

39°36’ 12.17” N, 8°41 ’ 11.26” W

419

15.6

Grata do Burro

Estremenho

39°36’48.43” N, 8°4E21.10” W

390

15.4

Algar do Ladoeiro

Estremenho

39°3E33.01”N, 8°45’15.58”W

485

15.5

Grata dos Moinhos Velhos

Estremenho

39°32’25.5” N, 8°42’15.5”W

307

18.0

Grata do Almonda

Estremenho

39°30 , 17.10” N, 8°36’54.40” W

18.0

Algar do Ze de Braga

Estremenho

39°29’ 16.02” N, 8°42 , 56.21” W

255

17.0

Algar do Vale da Pena

Estremenho

39°26 , 48.0” N, 8°55 , 56.8”W

294

14.3

Grata de Salir

Caldas da Rainha

39°29’ 11.64” N, 9°09 , 56.39” W

17.3

Grata dos Bolhos

Cesaredas

39°18 , 52.25” N, 9°16 , 61.37” W

145

16.3

Algar do Javali

Montejunto

39°1E96.5”N, 9°01’7l.8”W

380

14.4

Grata da Assafora

Lisbon Peninsula

38°54’24.80” N, 9°25’ 17.31” W

16.7

Grata da Alvide

Lisbon Peninsula

38°42 9 35.7”N, 9°25 , 32.8”W

18.3

Grata do Frade

Arrabida

38°25’46.56” N, 9°08’10.22” W

21.5

Lapa da Furada

Arrabida

38°25 , 45.8”N, 9°10 , 28.3”W

159

18.1

Algar de Santo Antonio

Alandroal

34°42 , 14.2” N, 7 0 23’59.3”W

370

17.6

Grata de Ibne Ammar

Algarve

37°09 , 24.9”N, 8 0 29’59.4” W

18.9

Algarao do Remexido

Algarve

37°14 , 29.5” N, 8°16 , 35.6”W

131

18.1

Grata do Vale Telheiro

Algarve

37°10 , 13.1”N, 8°02 , 05.3” W

239

17.0

Grata da Senhora

Algarve

37°06 , 20.5”N, 7°46 , 35.2”W

17.8

M5 and M20 microscopes and figures drawn with the aid of a camera lucida. Parts of specimens were

mounted in micropreparations with Hoyer’s liquid (Anderson 1954).

The ecological classification of species was made according to their degree of dependence on the

subterranean environment using the Schiner-Racovitza system and following the same criteria as

in Reboleira et al. (2011a): trogloxene species are those whose occurrence in caves is accidental;

troglophiles, species with affinities to the subterranean environment and frequently found in it, but

lacking troglomorphic traits, i.e., classical adaptations to subterranean life-style; and troglobionts,

species that exhibit a combination of depigmentation and lack or reduction of eyes, and are found

exclusively in subterranean ecosystems.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

All material was collected by A.S.P.S. Reboleira and has been deposited in the following collections:

MZUF = Museo di Storia Naturale dell’Universita, Sezione di Zoologia “La Specola”, Florence, Italy

SR = Sofia Reboleira collection, Universidade de Aveiro, Portugal

ZMUC = Zoological Museum, Natural History Museum of Denmark, University of Copenhagen

Fig. 1 . Location of the main karst areas of Portugal. 1. Vimioso. 2. Cantenhede-Outil. 3. Sico.

4. Estremenho. 5. Cesaredas. 6. Montejunto. 7. Lisbon Peninsula. 8. Arrabida. 9. Estremoz-Cano.

10. Algarve.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Results

Class Malacostraca Latreille, 1802

Order Isopoda Latreille, 1817

Suborder Oniscidea Latreille, 1802

Family Trichoniscidae Sars, 1899

Genus Trichoniscoides Sars, 1899

Trichoniscoides broteroi Vandel, 1946

Trichoniscoides broteroi andel, 1946: 178, figs 22-26.

Trichoniscoides broteroi - Vandel 1948: 129, fig. 14; 1952a: 264, figs 2, 12. — Schmolzer 1965: 43,

figs 120-122; 1971: 5, 78, 140, map 4. — Schmalfuss 2003: 302. —Reboleira et al. 2011a: 7, table 1.

Records

Grata dos Alqueves, Coimbra, Sico Massif (Vandel 1946).

Remarks

No specimens of this species were collected during our research. The type locality is now closed by the

municipality and the whole surrounding area is urbanized.

Distribution

Presently known only from the type locality.

Trichoniscoides machadoi Vandel, 1946

Fig. 2A-C

Trichoniscoides machadoi Vandel, 1946: 168, figs 8-12.

Trichoniscoides machadoi -Vandel 1947: 270; 1948: 129, fig. 14; 1952a: 262, figs 2, 9. — Schmolzer

1965: 37, figs 79-82; 1971: 4, 75, 139, map 4. — Schmalfuss 2003: 303.

Material examined

PORTUGAL: 3 $$, 4 § §, Grata d’el Rey, Cantanhede-Outil Massif, 8 Oct. 2009 (MZUF); 2 same

locality, 19 May 2009 (SR); 1 S, 1 $, Grata da Senhora, Algarve Massif, 29 Dec. 2009 (MZUF); 1 S,

1 5, Grata Abrigo Tomar I, Sico Massif, 26 Sep. 2011 (ZMUC).

Previous records

Serra do Geres, Terras de Bouro, Braga; Portunhos; Pampilhosa, Mealhada; Algar da Serra, Alvaiazere;

Algar da Rocha, Souto, Sico; Serra de Minde, Porto de Mos; Algarve da Terra da Rolha, Montejunto;

Buracos Mineiros, Montejunto; Algarve n°l da Senhora das Neves, Montejunto; Serra do Montejunto;

Cruz Quebrada, Oeiras; Grata do Algar, Mexilhoeirinha, Lagos (Vandel 1946).

Remarks

The diagnostic features of this species, i.e., the male pereopod 7 and male pleopods 1 and 2, are illustrated

in Fig. 2 from specimens collected in Grata d’el Rey.

Distribution

Species endemic to Portugal where it is widely distributed, from Serra do Geres in the north to the

Algarve in the south.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Ecological notes

This species is both cavemicolous and endogean.

Trichoniscoides ouremensis Vandel, 1946

Trichoniscoidespseudomixtus ouremensis Vandel, 1946: 177, figs 20-21.

Trichoniscoides ouremensis -Vandel 1952a: 263, figs 2, 10. — Schmolzer 1965: 38, figs 91-92; 1971: 5,

76, 139, map 4. — Schmalfuss 2003: 304. — Reboleira et al. 2011a: 7, table 1.

Records

Lapa da Salgada, Fatima, Vila Nova de Ourem (Vandel 1946).

Distribution

Species endemic to Portugal, presently only known from the type locality.

Trichoniscoides serrai Cruz, 1993

Trichoniscoides serrai Cruz, 1993: 18, fig. 2a-h.

Trichoniscoides serrai - Schmalfuss 2003: 304.

Fig. 2. Trichoniscoides machadoi Vandel, 1946 from Gruta d'el Rey, Cantanhede-Outil, $. A. Pereo-

pod 7. B. Pleopod E C. Pleopod 2.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Records

Gruta de Santo Adriao, Tras-os-Montes, Miranda do Douro (Cruz 1993).

Distribution

Species endemic to northern Portugal, known only from the type locality.

Trickoniscoides subterraneus Vandel, 1946

Trichoniscoides machadoi subterraneus V andel, 1946: 173, figs 13-15.

Trichoniscoides subterraneus - Vandel 1952a: 264, figs 2, 11. — Schmolzer 1965: 42, figs 116-117;

1971: 5, 77, 140, map 4. — Schmalfuss 2003: 304. —Reboleira et al. 2011a: 7, table 1.

Records

Gruta Alta do Cabetpo dos Mosqueiros, Aljubarrota, Alcobatpa (Vandel 1946).

Distribution

Species endemic to central Portugal, known only from the type locality.

Trichoniscoides meridionalis Vandel, 1946

Fig. 3A-C

Trichoniscoidespseudomixtus meridionalis V andel, 1946: 175, figs 16-19.

Fig. 3. Trichoniscoides meridionalis Vandel, 1946 from Gruta dos Moinhos, Velhos, Estremenho, f.

A. Pereopod 7. B. Pleopod 1. C. Pleopod 2.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Trichoniscoidesmeridionalis -Vandel 1952a: 266, figs 2,13. — Schmolzer 1965: 37, figs 83-84; 1971: 5,

75, 139, map 4. —Reboleira et al. 2011a: 7, table 1.

Material examined

PORTUGAL: 4 $$, 2 Gruta dos Moinhos Velhos, Estremenho Massif, 24 Mar. 2007 (MZUF);

3 2 $ 5, Algar do Vale da Pena, Estremenho Massif, 26 Jul. 2011 (SR); 1 $, 3 $ $, 1 juv., Algar

do Burro, Estremenho Massif, 5 May 2014 (MZUF); 1 $, Algar do Ze de Braga, Estremenho Massif,

16 Feb. 2013 (SR); 7 4 55, Algar do Ladoeiro, Estremenho Massif, 3 Feb. 2007 (MZUF); 3

1 5, same locality, 6 Jan. 2007 (SR); 1 $, 1 5,3 juvs, Gruta do Almonda, Estremenho Massif, 24 Aug. 2013

(MZUF); 1 5, same locality, 17 Feb. 2013 (SR); 3 1 5» same locality, 25 Aug. 2013 (ZMUC); 1 $,

Gruta do Papagaio, Estremenho Massif, 5 Apr. 2014 (MZUF).

Previous records

Lapa da Cha de Cima, Porto de Mos, Leiria; Gruta das Alcobertas, Rio Maior, Santarem (Vandel 1946).

Lapa dos Ladoeiros, Porto de Mos; Lapa da Ovelha, Serra de Minde, Santarem (Vandel 1952a).

Distribution

This species is restricted to the Estremenho karst massif, in central-western Portugal.

Remarks

This form was described by Vandel (1946) as a subspecies of Trichoniscoidespseudomixtus ( Arcangeli,

1935), a species present in some caves of northeastern Spain. Vandel (1952a) elevated this form to

species level. The male pereopod 7 and male pleopods 1 and 2 are illustrated in Fig. 3 from specimens

collected in Gruta dos Moinhos Velhos.

Ecological notes

This species can be found from the entrances to the deepest parts of the caves, normally associated with

decomposing wood. It is distributed along the three main subunits of the Estremenho Massif, being

the only troglobiotic oniscidean so far known from this large karst area. Contrary to other troglobiotic

arthropods from the same massif, it does not present closely related species along the three subunits

(Reboleira 2007; Reboleira et al. 2009). The biocoenosis of this karst area includes other troglobionts,

i.e., the spider Nesticus lusitanicus Fage, 1931, the campodeid Podocampa cf. fragiloides Silvestri,

1932, and the ground beetles Trechus gamae Reboleira & Serrano, 2009, T. lunai Reboleira & Serrano,

2009 and T. machadoi Jeannel, 1941 (Reboleira & Ortuno 2011; Reboleira et al. 2010b).

Trichoniscoides bellesi Reboleira & Taiti sp. nov.

urn:lsid:zoobank.org:act:FE37F766-5E77-430F-87C2-C5CB28A78D5D

Figs 4A-G, 5A-E, 6A-D

Diagnosis

A blind and colourless Trichoniscoides characterised by the male pleopod 1 exopod having a broadly

rounded outer margin and two equal distal lobes, and the male pleopod 2 endopod having a distal article

thickset for 2 Z of its length, ending with a narrow point.

Etymology

The new species is named after Prof. Xavier Belles for his invaluable contribution to the synthesis of

knowledge on the subterranean fauna from the Iberian Peninsula.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Fig. 4. Trichoniscoides bellesi Reboleira & Taiti sp. nov. from Algar do Javali, Montejunto. —Paratype,

§. A. Adult specimen in dorsal view. B. Dorsal scale-seta. C. Cephalon in frontal view. D. Cephalon in

dorsal view. — Paratype, $. E. Pleonites 4, 5, telson and right uropod. F. Antemiula. G. Antenna.

REBOLEIRA A.S.P.S. et al ., Oniscidea from caves of Portugal

Material examined

Holotype

PORTUGAL: S, Algar do Javali, Montejunto Massif, 26 Dec. 2012 (MZUF).

Paratypes

PORTUGAL: 1 1 same data as holotype (MZUF).

Description

Maximum size: $ 1.7 x 0.8 mm ; § 1.8 x 0.8 mm. Body colourless and slightly convex (Fig. 4A). Cephalon

and pereon with granulated dorsal surface; granules on pereonites arranged in double rows; each granule

bearing a triangular scale-seta (Fig. 4B) on top. Cephalon (Fig. 4C-D) with quadrangular frontal lateral

lobes not protruding frontwards. Eyes absent. Pereonites 1—4 (Fig. 4A) with rounded posterior corners;

pereonites 5-7 (Fig. 4A) with epimera pointing backwards. Pleon (Fig. 4A, E) narrower than pereon;

pleonites 3-5 with small epimera and short posterior points. Telson (Fig. 4E) more than twice as wide as

Fig. 5. Trichoniscoides bellesi Reboleira & Taiti sp. nov. from Algar do Javali, Montejunto, paratype, $.

A. Right mandible. B. Left mandible. C. Maxillula. D. Maxilla. E. Maxilliped.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

long, distal part trapezoidal with concave sides and slightly convex apex. Antennula (Fig. 4F) with three

articles; third article with distinct spine and three long aesthetascs at apex. Antenna (Fig. 4G) with fifth

article of peduncle shorter than flagellum; flagellum with three articles, with three long aesthetascs on

second article. Mandibles (Fig. 5A-B) with two penicils on the right and three on the left; molar process

without penicils. Maxillula (Fig. 5C) with inner branch bearing three penicils at apex, proximal one

distinctly longer than other two; outer branch with 11 teeth and two thin stems, one among outer group

and one among inner group of teeth. Maxilla (Fig. 5D) apically bilobed, with outer lobe smaller than

inner one; inner lobe with several long, stout setae. Maxilliped (Fig. 5E) endite triangular, with stout

penicil at apex; palp distally rounded, with long setae, basal article with two short, compound setae.

Fig. 6. Trichoniscoides bellesi Reboleira & Taiti sp. nov. from Algar do Javali, Montejunto, paratype, S-

A. Pereopod 1. B. Pereopod 7. C. Pleopod 1. D. Pleopod 2 endopod.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Uropod (Fig. 4E) with exopod distinctly longer than endopod; exopod with several long, pointed setae

and endopod with a long and a short seta at apex.

Male. Pereopod 1 (Fig. 6A) and pereopod 7 (Fig. 6B) with no particular modifications. Pleopod 1 (Fig.

6C) exopod almost as long as wide, with broadly rounded outer margin and two equal distal lobes each

bearing a short seta at apex; endopod with triangular and flagelliform basal article and setose distal

article. Pleopod 2 exopod missing in the specimen examined; endopod (Fig. 6D) biarticulated, distal

article about twice as long as basal one, thickset for first 2 /a and ending with narrow point.

Remarks

In the shape of the male pleopod 2 exopod, with a thickset distal article ending in a narrow point, the

new species is similar to Trichoniscoides lusitanus Vandel, 1946, T. broteroi and T. heroldi Vandel,

1952. It differs from T. lusitanus , an epigean species from north-western Spain and northern Portugal,

in lacking eyes and pigment, in the male pleopod 1 endopod having a triangular basal article, and in

the male pleopod 2 endopod having a stouter distal article ending in a shorter point; from T. broteroi in

lacking the hook on the male pereopod 7 merus, and in the triangular shape of the basal article of the

male pleopod 1 endopod; from T. heroldi , an epigean species from eastern France, in the lack of eyes

and the shape of the male pleopod 1.

Ecological notes

Trichoniscoides bellesi sp. nov. is typically a troglobiotic species, lacking eyes and body pigment. It was

discovered in the deepest and most thermally insulated parts of Algar do Javali, around 10 m deep. Javali

is currently the richest cave of the Montejunto karst massif. Four other caves in the same massif were

also sampled (Algar do Escorpiao, Ralo das Fontainhas, Algar da Terra da Rolha and Buracos Mineiros)

without recovering any specimens of T. bellesi. The subterranean community of Algar do Javali includes

other troglobiotic species, such as the campodeid Podocampa cf. fragiloides , a species of the oniscidean

genus Paraschizidium (see below), the ground beetle Trechus tatai Reboleira & Ortuno, 2010, a new

pselaphid beetle, and the pseudoscorpions Chthonius cardosoi Zaragoza, 2012 and Roncocreagris

occidentals Zaragoza & Reboleira, 2013 (Reboleira et al. 2010a, 2013c).

Trichoniscoides sicoensis Reboleira & Taiti sp. nov.

urn:lsid:zoobank.org:act:FCBE373B-42CE-4305-A932-28D88355FFB8

Figs 7A-G, 8A-E, 9A-E

Diagnosis

A blind and colourless Trichoniscoides characterised by the male pereopod 7 merus having a lobe on the

mid-sternal margin, the male pleopod 1 exopod having a broadly rounded outer margin and two unequal

setae, the endopod having a fusiform distal article with a distinct circular suture in the middle, and the

male pleopod 2 endopod having thickset distal article bearing two short triangular lobes and two setae

at the apex.

Etymology

The new species is named after the Sico karst area, where the species occurs.

Material examined

Holotype

PORTUGAL: Grata da Ceramica, Sico Massif, 21 Mar. 2010 (MZUF).

Paratypes

PORTUGAL: 10 $$, 12 $$, Grata da Ceramica, Sico, 21 Mar. 2010 (MZUF); 7 11 $$, same

locality, 28 Nov. 2009 (SR); 4 SS, 4 same locality, 10 Jun. 2011 (ZMUC); 4 5 same

European Journal of Taxonomy 161 : 1-61 ( 2015 )

locality, 2 Mar. 2013 (MZUF); 12 33 , 8 $ Gruta da Senhora da Estrela, Sico, 20 Nov. 2009 (MZUF);

5 33 , 12 5 5, same locality, 29 Aug. 2009 (SR); 3 33 , same locality, 11 Jun. 2009 (ZMUC); 1 3 , Gruta

do Soprador do Carvalho, Sico, 12 Jun. 2011 (SR); 2 33 , 3 55, same locality, 8 Jun. 2013 (MZUF);

3 33 , 2 ??, Gruta do Algarinho, Sico, 9 Jun. 2013 (SR); 3 33 , 1 5 » Gruta da Arrifana, Sico, 12 Jan.

2013 (ZMUC); 1 3 , 2 5 5, Gruta de Sao Simao, Sico, 30 Nov. 2013 (MZUF).

Fig. 7. Trichoniscoides sicoensis Reboleira & Taiti sp. nov. from Gruta da Ceramica, Sico. — Paratype,

5- A. Specimen in dorsal view. — Paratype, 3■ B. Dorsal scale-seta. C. Cephalon in frontal view.

D. Cephalon in dorsal view. E. Pleonites 4, 5, telson and right uropod. F. Antennula. G. Antenna.

REBOLEIRAA.S.P.S. et al., Oniscidea from caves of Portugal

Description

Maximum size: S 3.5 x 1.2 mm; $ 3.9 x 1.2 mm Body colourless and slightly convex (Fig. 7A). Dorsum

smooth, with scattered triangular scale-setae (Fig. 7B). Cephalon (Fig. 7C-D) with rounded frontal

lateral lobes not protruding frontwards. Eyes absent. Pereonites 1-4 (Fig. 7A) with rounded posterior

corners; pereonites 5-7 (Fig. 7A) with epimera pointing backwards. Pleon (Fig. 7A, E) narrower than

pereon; pleonites 3-5 with small epimera and short posterior points. Telson (Fig. 7E) about three times

as wide as long, with distal part trapezoidal with concave sides. Antennula (Fig. 7F) with three articles;

third article with distinct spine and three long aesthetascs at apex. Antenna (Fig. 7G) with fifth article

of peduncle shorter than flagellum; flagellum with five articles with two long aesthetascs on second

article. Mandibles (Fig. 8A-B) with two penicils on the right and three on the left; molar process without

penicils. Maxillula (Fig. 8C) with inn er branch bearing three penicils at apex, inner one distinctly longer

than other two; outer branch with 11 teeth and two thin stems. Maxilla (Fig. 8D) apically bilobed and

setose, with outer lobe smaller than inner one; inner lobe with several long stout setae. Maxilliped (Fig.

8E) endite triangular, with stout triangular penicil at apex; palp distally rounded, long setae, and basal

Fig. 8. Trichoniscoides sicoensis Reboleira & Taiti sp. nov. from Grata da Ceramica, Sico, paratype, S-

A. Right mandible. B. Left mandible. C. Maxillula. D. Maxilla. E. Maxilliped.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Fig. 9. Trichoniscoides sicoensis Reboleira & Taiti sp. nov. from Gruta da Ceramica, Sico, paratype, <$■

A. Pereopod 1. B. Pereopod 7. C. Genital papilla. D. Pleopod 1. E. Pleopod 2.

REBOLEIRAA.S.P.S. et al., Oniscidea from caves of Portugal

article with two short compound setae. Uropod (Fig. 7E) with exopod distinctly longer than endopod

and more distally inserted; exopod with several long, pointed setae and endopod with a long and a short

seta at apex.

Male. Pereopods 1 (Fig. 9A) to 3 with an area of short quadrangular scales on sternal margin of carpus and

merus. Pereopod 7 (Fig. 9B) merus with distinct cylindrical lobe, with short seta at apex on mid-sternal

margin. Genital papilla (Fig. 9C) fusiform. Pleopod 1 (Fig. 9D) exopod as long as wide, with broadly

rounded outer margin and two unequal setae on distal lobe; endopod with basal article quadrangular

with convex inner margin and concave outer margin, distal article fusiform with distinct suture in middle

and setose apex. Pleopod 2 (Fig. 9E) exopod trapezoidal with large seta at apex; endopod biarticulated,

distal article about 1.5 times as long as basal one, thickset, with two short triangular lobes and setae at

apex.

Remarks

This new species is tentatively included in the genus Trichoniscoides since it has all the characters of the

genus, except for the distal article of the male pleopod 1 endopod, which shows a distinct circular suture

in the middle, and the thickset distal part of the male pleopod 2 endopod. These characters are present

also in an epigean species described by Gregory et al. (2012: 7, fig. 4) as “? Trichoniscoides species C”

from Avion, valley of Rio Valdeiras, Orense, north-western Spain, and Castanheira, Viana do Castelo,

northern Portugal. The new species differs from the one recorded by Gregory et al. (2012) in the shape

of the basal article of the male pleopod 1 endopod and of the distal part of the male pleopod 2 endopod.

Trichoniscoides sicoensis Reboleira & Taiti sp. nov., together with the species recorded by Gregory et al.

(2012), might belong to a distinct genus, but this can only be confirmed when other species with similar

characters are found. For the presence of a lobe on the mid-sternal margin of the male pereopod 7 merus,

the new species also resembles T. broteroi , from which it is distinguishable by the shape of the male

pleopods 1 and 2. A hook-like lobe on the sternal margin of the male pereopod 7 merus is present also in

the “aquitano-languedocien” group of species ( sensu Vandel 1960a), but in the species of this group the

lobe is near the base instead of being on the mid-part of the sternal margin.

Ecological notes

This species seems to be endemic to caves of the Sico karst area. In Grata do Soprador do Carvalho one

specimen was collected under a stone completely submerged in the cave stream. Despite the amphibian

behaviour and submersion tolerance, this species is mostly found in the superficial parts of caves,

where roots are abundant. Several troglobiotic species are known to share this habitat with T. sicoensis

Reboleira &Taiti sp. nov.: the pseudoscorpions Roncocreagris blothroides (Beier, 1962), R. borgesi

Zaragoza & Reboleira, 2013 and R. cavernicola Vachon, 1946; the millipede Scutogona minor Enghoff

& Reboleira, 2013; the woodlice T. broteroi, Miktoniscns longispina Reboleira & Taiti sp. nov. and

Porcellio cavernicolus ; and the rove beetle Domene lusitanica Reboleira & Oromi, 2011 (Reboleira et

al. 2011b, 2013b; Enghoff & Reboleira 2013a).

Genus Metatrichoniscoides Vandel, 1942

Metatrichoniscoides salirensis Reboleira & Taiti sp. nov.

urndsid: zoobank.org: act: 7E09D1C5 -19F 8-44DF-A 1AE-B2D741C411AE

Figs 10A-G, 11A-E, 12A-E

Diagnosis

A blind and colourless Metatrichoniscoides characterised by the male pleopod 1 exopod having two

long distal setae subequal in length, and the male pleopod 2 endopod having a thickset distal article,

ending in a thinner sinuous part with a beak-like small lobe medially directed.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Etymology

The new species is named after Gruta de Salir, the cave where the specimens were collected, in the

municipality of Caldas da Rainha.

Fig. 10. Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. from Gruta de Salir, Caldas da Rainha

typhonic valley. — Paratype, §. A. Adult specimen in dorsal view. B. Dorsal scale-seta. C. Cephalon

in frontal view. D. Cephalon in dorsal view. — Paratype, f. E. Pleonites 4, 5, telson and right uropod.

F. Antennula. G. Ante nn a

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Material examined

Holotype

PORTUGAL: S, Grata de Salir, Salir do Porto, Caldas da Rainha, 26 Dec. 2012 (MZUF).

Paratypes

PORTUGAL: 1 <$,2 $ §, same data as holotype (MZUF); 1 S,2 $5, same data as holotype (SR).

Fig. 11. Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. from Grata de Salir, Caldas da Rainha,

paratype, $. A. Right mandible. B. Left mandible. C. Maxillula. D. Maxilla. E. Maxilliped.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Description

Maximum size: S and $ 2.2 x 0.8 mm. Body colourless, elongated and slightly convex (Fig. 10A).

Cephalon and pereon with granulated dorsal surface; granules on pereonites arranged in double rows;

each granule bearing a triangular scale-seta (Fig. 10B). Cephalon (Fig. 10C-D) with no frontal lateral

lobes. Eyes absent. Pereonites 1M (Fig. 10A) with rounded posterior comers; pereonites 5-7 (Fig. 10A)

with epimera directed backwards. Pleon (Fig. 10A,E) narrower than pereon; pleonites 3-5 with small

epimera and short posterior points. Telson (Fig. 10E) twice as wide as long, with distal part trapezoidal

with concave sides. Antennula (Fig. 10F) with three articles; third article distinctly longer than first

Fig. 12. Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. from Grata de Salir, Caldas da Rainha,

paratype, S. A. Pereopod 1. B. Pereopod 7. C. Genital papilla. D. Pleopod 1. E. Pleopod 2.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

and second and bearing four long aesthetascs at apex. Antenna (Fig. 10G) with fifth article of peduncle

slightly shorter than flagellum; flagellum with three articles, with two aesthetascs on second article.

Mandibles (Fig. 11A-B) with two penicils on the right and three on the left; molar process without

penicils. Maxillula (Fig. 11C) with inner branch bearing three short penicils at apex; outer branch with

11 teeth and two thin stems. Maxilla (Fig. 11D) apically bilobed, with outer lobe shorter than inner

one; inner lobe with some long, stout setae near medial margin. Maxilliped (Fig. 11E) endite triangular,

ending with triangular point and subapical penicil; palp distally rounded, with long setae, and basal

article with two short compound setae. Uropod (Fig. 10E) with exopod slightly longer than endopod;

exopod with several long, pointed setae and endopod with long seta at apex.

Male. Pereopod 1 (Fig. 12A) and pereopod 7 (Fig. 12B) with no peculiar modifications. Genital papilla

(Fig. 12C) fusiform, with rounded apex. Pleopod 1 (Fig. 12D) exopod as long as wide, with broadly

rounded apical margin bearing two long setae of equal length; endopod with basal article triangular, baton¬

like distal article inserted on medial margin of basal article. Pleopod 2 (Fig. 12E) exopod trapezoidal,

with short seta at apex; endopod biarticulated, distal article thickset, ending in thinner sinuous part with

beak-like small lobe medially directed.

Remarks

Until now the genus Metatrichoniscoides included only four species (Schmalfuss 2003): M. ley digit

(Weber, 1880) from western France, Belgium, the Netherlands, western Germany and greenhouses in

the Czech Republic, Sweden and Finland; M. nemausiensis Vandel, 1942 and M. fouresi Vandel, 1950

from France; and M. celticus Oliver & Trew, 1981 from Wales (Vandel 1942; Oliver & Trew 1981).

Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. is readily distinguished from all the other

species in the genus by the male pleopod 1 exopod bearing two long setae and the peculiar shape of

the male pleopod 2 endopod. This represents the first record of the genus for Portugal and the Iberian

Peninsula.

Ecological notes

Grata de Salir is a peculiar, beautiful cave located near the sea, with a genesis related to previous sea

levels, evidenced by the presence of marine sand in the lowest parts. This is the first troglobiotic species

known from this cave and this karst area.

Genus Troglonethes Cruz, 1989

Troglonethes olissipoensis Reboleira & Taiti sp. nov.

urn:lsid:zoobank.org:act:7E09DlC5-19F8-44DF-AlAE-B2D741C411AE

Figs 13A-G, 14A-E, 15A-D

Diagnosis

A species of Troglonethes characterised by the antenna having five flagellar articles, the male pleopod 1

exopod triangular, as wide as long, and the male pleopod 2 endopod with the distal article bearing a basal

and a distal hook-like process.

Etymology

The new species is named after Olissipo, the old name of Lisbon, on the peninsula of which the Grata

de Alvide is located.

Material examined

Holotype

PORTUGAL: Grata da Alvide, Cascais, Lisbon Peninsula, 26 Dec. 2009 (MZUF).

LUUI l

European Journal of Taxonomy 161: 1-61 (2015)

Paratypes

PORTUGAL: 5 SS, 2 5 ?, 3 juvs, same data as holotype (MZUF); 5 SS, 2 $same data as holotype

(SR); 5 SS, 2$?, same data as holotype (ZMUC).

Fig. 13. Troglonethes olissipoensis Reboleira & Taiti sp. nov. from Gruta de Alvide, Lisbon Peninsula.

— Paratype, $. A. Adult specimen in dorsal view. — Paratype, f. B. Dorsal scale-seta. C. Cephalon

in frontal view. D. Cephalon in dorsal view. E. Pleonites 4, 5, telson and left uropod. F. Antennula.

G. Ante nn a.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Description

Ma xim um size: S 3.5 x 0.8 mm; 5 4.2 x 1.2 mm Body colourless and elongated (Fig. 13A). Dorsum of

cephalon and pereon granulated; each granule equipped with a triangular scale-seta (Fig. 13B); pleon

and telson smooth. Cephalon (Fig. 13C-D) with quadrangular frontal lateral lobes slightly protruding

outwards. Eyes absent. Pereonites 1—4 (Fig. 13A) with rounded posterior corners; pereonites 5-7 (Fig.

13A) with epimera pointing backwards. Pleon (Fig. 13A, E) narrower than pereon; pleonites 3-5 with

small epimera and very short posterior points. Telson (Fig. 13E) about twice as wide as long, with distal

part trapezoidal with concave sides. Antennula (Fig. 13F) with three articles; third article with spine and

three long aesthetascs at apex. Antenna (Fig. 13G) with third, fourth and fifth article of peduncle with

distinct tubercles bearing scale-setae; fifth article as long as flagellum; flagellum with five articles, with

two long aesthetascs on second and third article. Mandibles (Fig. 14A-B) with one penicil on the right

and three on the left. Maxillula (Fig. 14C) with inner branch bearing three penicils at apex, inner one

distinctly longer than other two; outer branch with 12 teeth and thin, setose stem among outer group of

teeth. Maxilla (Fig. 14D) apically bilobed and setose, with outer and inner lobes subequal; inner lobe

with several long stout setae along margin. Maxilliped (Fig. 14E) endite triangular, with stout triangular

Fig. 14. Troglonethes olissipoensis Reboleira & Taiti sp. nov. from Grata de Alvide, Lisbon Peninsula,

paratype, S- A. Right mandible. B. Left mandible. C. Maxillula. D. Maxilla. E. Maxilliped.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

penicil at apex; palp distally rounded, with long setae, basal article with two short, compound setae.

Uropod (Fig. 13E) with exopod slightly longer than endopod and more distally inserted; exopod with

several long pointed setae and endopod with a long and a short seta at apex.

Fig. 15. Troglonethes olissipoensis Reboleira & Taiti sp. nov. from Gruta de Alvide, Lisbon Peninsula,

paratype, f. A. Pereopod 1. B. Pereopod 7. C. Genital papilla and pleopod 1. D. Pleopod 2.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Male. Pereopod 1 (Fig. 15A) with no particular modifications. Pereopod 7 (Fig. 15B) ischium with

straight sternal margin, carpus with row of scales on distal margin, and propodus with long, thin setae

on distal half of tergal margin. Genital papilla (Fig. 15C) fusiform, with rounded apex, much longer than

pleopod 1. Pleopod 1 (Fig. 15C) exopod triangular, as wide as long, with broadly rounded apex; endopod

with basal article elongated, longer than exopod, with sinuous outer margin, distal article flagelliform

with setose apex. Pleopod 2 (Fig. 15D) exopod triangular, with broadly rounded apex with few short

setae; endopod biarticulated, basal article about three times as long as distal one, with parallel sides,

distal article with outer margin bearing hook-like process at base and one at apex.

Remarks

The genus Troglonethes was erected by Cruz (1989) for the new species T. aurowci from a cave north of

Valencia, Spain. Tabacaru (1993) included the genus in the tribe Speleonethini. The new species from

Portugal is included in the genus since it has all the characters listed in the diagnosis. Troglonethes

olissipoensis Reboleira & Taiti sp. nov. differs from T. aurowci in having an antennal flagellum with

five, instead of four, articles, the male pleopod 1 exopod shorter than the basal article of the endopod,

and the male pleopod 2 endopod with a basal hook-like process on the outer margin of the distal article.

Ecological notes

This species was only found in Grata de Alvide, located in an overarbanized area, with part of the cave

ceiling used as the base of a residential building. The specimens were collected in the deepest parts of

the cave, the so-called third level. It is the second troglobiotic species from karst caves in the Lisbon

Peninsula, after the record of the bristletail Coletinia sp. in Grata de Colaride (Reboleira et al. 2012a).

Unidentified springtails and blaniulid millipedes were also collected.

Troglonethes arrabidaensis Reboleira & Taiti sp. nov.

um:lsid:zoobank.org:act:78A12550-F7CA-4892-9E90-7F85446423D3

Figs 16A-G, 17A-D

Diagnosis

A species of Troglonethes characterised by the antenna having three flagellar articles, the male pereo¬

pod 7 carpus enlarged in the basal part, the pleopod 1 exopod triangular, as wide as long, and the male

pleopod 2 endopod with the distal article bearing an apical hook-like process.

Etymology

The new species is named after the type locality, the Arrabida karst massif.

Material examined

Holotype

PORTUGAL: $, Grata do Frade, Arrabida Massif, 16 Aug. 2013 (MZUF).

Paratypes

PORTUGAL: 3 13 $ ?, same data as holotype (MZUF, SR and ZMUC).

Description

Maximum size: $ and $ 2.7 x 0.8 mm. Body colourless and elongated (Fig. 16A). Dorsum of cephalon

and pereonites with two rows and one row of large granules, respectively; each granule with a scale-seta

(Fig. 16B); pleon and telson smooth. Cephalon (Fig. 16C-D) with short, quadrangular frontal lateral

lobes. Eyes absent. Pereonites 1-3 with rounded posterior corners; pereonites 4-7 with epimera pointing

backwards (Fig. 16A). Pleon (Fig. 16A, E) slightly narrower than pereon; pleonites 3-5 with small

European Journal of Taxonomy 161: 1-61 (2015)

epimera and very short posterior points. Telson (Fig. 16E) about twice as wide as long, with distal part

trapezoidal, with concave sides and broadly rounded apex. Antennula (Fig. 16F) with three articles; third

article with three long aesthetascs at apex. Antenna (Fig. 16G) with fourth and fifth article of peduncle

with distinct tubercles bearing scale-setae; fifth article slightly shorter than flagellum; flagellum with

Fig. 16. Troglonethes arrabidaensis Reboleira & Taiti sp. nov. from Gruta do Frade, Arrabida. —

Paratype, §. A. Adult specimen in dorsal view. B. Dorsal scale-seta. C. Cephalon in frontal view.

D. Cephalon in dorsal view. E. Pleonites 4, 5, telson and left uropod. — Paratype, S. F. Antennula.

G. Ante nn a.

REBOLEIRA A.S.P.S. et al, Oniscidea from caves of Portugal

three articles, with two long aesthetascs on second article. Buccal pieces as in previous species. Uropod

(Fig. 16E) with exopod slightly longer than endopod, exopod and endopod inserted at same level.

Male. Pereopod 1 (Fig. 17A) with no particular modifications. Pereopod 7 (Fig. 17B) ischium with

straight sternal margin, carpus enlarged in basal part. Genital papilla (Fig. 17C) fusiform, with rounded

apex, much longer than pleopod 1. Pleopod 1 (Fig. 17C) exopod triangular, as wide as long, with broadly

rounded apex; endopod with basal article elongated, shorter than exopod, with sinuous outer margin,

distal article flagelliform. Pleopod 2 (Fig. 17D) exopod triangular, with broadly rounded apex with few

short setae; endopod biarticulated, basal article more than twice as long as distal one, with parallel sides,

distal article with hook-like process at apex.

Fig. 17. Troglonethes arrabidaensis Reboleira & Taiti sp. nov. from Grata do Frade, Arrabida, para-

type, S- A. Pereopod E B. Pereopod 7. C. Genital papilla and pleopod 1. D. Pleopod 2.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Remarks

The new species is very similar to Troglonethes aurouxi in the dorsal granulation and shape of the male

pleopod 2. It differs in having the antennal flagellum with three, instead of four, articles, and a wider

male pleopod 1 exopod. It is readily distinguishable from T. olissipoensis Reboleira & Taiti sp. nov. in

the larger and less numerous dorsal granules, the enlarged basal part of the male pereopod 7 carpus, the

male pleopod 1 endopod with the basal article shorter than the exopod, and the distal article of the male

pleopod 2 endopod lacking the hook-like process at the base.

Ecological notes

This species was only found in the Gruta do Frade in the Arrabida Massif. The cave entrance is by the

seashore and there are several anchialine lakes inside, influenced by the sea tides. It is the first record of

troglobiotic terrestrial isopods in Arrabida.

Genus Miktoniscus Kesselyak, 1930

Miktoniscus longispina Reboleira & Taiti sp. nov.

urn:lsid:zoobank.org:act:6B9A5E8B-53F8-408D-B3BA-8E10BCE20717

Figs 18A-F, 19A-E, 20A-D

Diagnosis

A species of Miktoniscus characterised by a depigmented body, the male pereopod 7 having a long and

stout seta on the distal corner of the ischium, and a triangular male pleopod 1 exopod.

Etymology

From Latin longus = long + spina = spine. The name of the new species refers to the long, thick seta on

the distal part of the male pereopod 7.

Material examined

Holotype

PORTUGAL: S, Gruta dos Bolhos, Cesaredas Plateau, 26 Dec. 2010 (MZUF).

Paratype

PORTUGAL: 1 $, same locality as holotype, Feb. 2009 (MZUF).

Additional material

PORTUGAL: 1 2 §§, Gruta da Ceramica, Sico Massif, 10 Jun. 2011 (MZUF); 2 3 §§, same

locality, 2 Mar. 2013 (SR); 1 c?, 1 $, Algar da Ervilha, Sico, 30 Mar. 2014 (ZMUC).

Description

Maximum size: $ and $ 3.5 x 1.0 mm. Body depigmented, elongated (Fig. 18A). Dorsum of cephalon

and pereon distinctly granulated, each granule bearing a scale-seta on top (Fig. 18B). Cephalon (Fig.

18C) with no frontal lobes. Eye consisting of single black ocellus (lacking in specimens from Gruta da

Ceramica and Algar da Ervilha). Pereonites 1—4 with rounded posterior corners; pereonites 5-7 with

epimera pointing backwards (Fig. 18A). Pleon (Fig. 18A, D) slightly narrower than pereon; pleonites

3-5 with reduced epimera and very short posterior points. Telson (Fig. 18D) about twice as wide as long,

with rectangular basal part and trapezoidal distal part, with almost straight sides. Antennula (Fig. 18E)

with three articles; third article with short spine and four long aesthetascs at apex (five in specimens

from Gruta da Ceramica and Algar da Ervilha). Antenna (Fig. 18F) with fifth article of peduncle slightly

longer than flagellum; flagellum with four articles, with one aesthetasc on second article and two

REBOLEIRA A.S.P.S. et al, Oniscidea from caves of Portugal

aesthetascs on third article (two and four, respectively, in specimens from Grata da Ceramica and Algar

da Ervilha). Mandibles (Fig. 19A-B) with one penicil on the right and two on the left; molar process

without penicils. Maxillula (Fig. 19C) with inner branch bearing three penicils at apex, diminishing in

size from proximal to distal; outer branch with nine teeth and thin stem. Maxilla (Fig. 19D) apically

bilobed, outer lobe smaller and shorter than inner one; inner lobe with line of several long, stout setae

near apical and inner margins. Maxilliped (Fig. 19E) endite triangular, with stout triangular penicil at

Fig. 18. Miktoniscus longispina Reboleira & Taiti sp. nov. from Grata dos Bolhos, Cesaredas Plateau.

— Paratype, $. A. Specimen in dorsal view. —Paratype, S- B. Dorsal scale-seta. C. Cephalon in dorsal

view. D. Pleonite 5, telson and left uropod. E. Antennula. F. Antenna.

European Journal of Taxonomy 161: 1-61 (2015)

apex; palp elongated, with long setae on margin of apical part, and basal article with two short, simple

setae. Uropod (Fig. 18E) with exopod distinctly longer than endopod and more distally inserted.

Fig. 19. Miktoniscus longispina Reboleira & Taiti sp. nov. from Gruta dos Bolhos, Cesaredas Plateau,

paratype, $. A. Right mandible. B. Left mandible. C. Maxillula. D. Maxilla. E. Maxilliped.

REBOLEIRA A.S.P.S. et al ., Oniscidea from caves of Portugal

Male. Pereopod 1 (Fig. 20A) with no distinct sexual modifications. Pereopod 7 (Fig. 20B) ischium

triangular, with strong seta, as long as merus, at corner between distal and sternal margin; sternal margin

of merus sinuous, with one recurved seta and one pointed seta in the middle; carpus with distal part

enlarged, with some strong setae on sternal margin. Pleopod 1 (Fig. 20C) exopod triangular, about

hS"N

. 0 $,

VJ\ 'u

V "S V v' \

' vV V

' V

N ^ ^ V

\VV" S v

i 1

\ '

r\ \

/ *

i ) \

i / j

j i/y/. 1 ' ji>>

D V -

1 ;V tfj.j JH )

/ ^

7 ' ■ — — _ i

Fig. 20. Miktoniscus longispina Reboleira & Taiti sp. nov. from Gruta dos Bolhos, Cesaredas Plateau,

paratype, S- A. Pereopod 1. B. Pereopod 7. C. Pleopod E D. Pleopod 2.

European Journal of Taxonomy 161: 1-61 (2015)

twice as long as wide; endopod with basal article distinctly shorter than distal article, which is slightly

enlarged subapicaly, with triangular and crenulated apical part. Pleopod 2 (Fig. 20D) exopod triangular,

with distinctly concave outer margin; endopod biarticulated, distal article fusiform, elongated, about

four times as long as first article.

Remarks

At present, the genus Miktoniscus includes 15 species distributed in the Americas and Western Europe,

including the Atlantic islands (Schmalfuss 2003). The new species is readily distinguishable from all

species in the genus by the presence of a strong and long seta at the distal corner of the male pereopod 7

ischium. In continental Portugal and northwestern Spain only one species was previously known,

Miktoniscis bisetosus Vandel, 1946, from which M. longispina Reboleira & Taiti sp. nov. also differs in

lacking a recurved seta on the male pereopod 7 carpus and in having the male pleopod 1 exopod with

an acute, rather than broadly rounded, apex. The new species occurs in two distinct massifs of central

Portugal, on the west border of Cesaredas Plateau near the Atlantic coast, and in Grata da Ceramica

and Algar da Ervilha in the centre of Sico Massif. Some differences are present in the specimens from

Sico compared with the type specimens from Cesaredas, i.e., the lack of visible eyes and the different

number of aesthetascs on the antennula and antennal flagellum. For this reason, we have not designated

the additional material as paratypes. However, since the male characters are the same in all these

populations, we consider them as belonging to the same species.

Genus Haplophthalmus Schobl, 1860

Haplophthalmus siculus Dollfus, 1896

Haplophthalmus siculus Dollfus, 1896: 5, fig. 3.

Haplophthalmus siculus -Vandel 1946: 192, figs 36M1.

Records

Cova da Adi£a, Moura; Grata do Algar, Mexilhoeirinha, Lagoa (Vandel 1946).

Distribution

Portugal, Spain, southern Italy and Sicily.

Genus Moserius Strouhal, 1940

Moserius inexpectatus Reboleira & Taiti sp. nov.

um:lsid:zoobank.org:act:18B5B016-F7A6-4047-A2CB-28B4394797CF

Figs 21A-E, 22A-C

Diagnosis

A species of Moserius characterised by the male pereopod 7 carpus having a distal lobe on the sternal

margin, and the pleopod 1 exopod having a truncate and sinuous distal point.

Etymology

From Latin inexpectatus = unexpected. The name refers to the location, which is far from the previously

known distribution range of the genus.

Material examined

Holotype

PORTUGAL: Grata do Almonda (= Grata do Olho da Fonte), Estremenho Massif, 24 Aug. 2013

(MZUF).

REBOLEIRA A.S.P.S. et al, Oniscidea from caves of Portugal

Description

Size: 1.5 x 0.6 mm. Body colourless, ovoidal, outline not interrupted between pereon and pleon; pereon

tergites very convex, with epimera obliquely directed outwards (Fig. 21 A). Dorsum bearing ribs and

tubercles (Fig. 21 A): cephalon with large central tubercle with 3 + 3 rounded tips, and 3+3 tubercles near

rear margin, the two external ones fused at base; pereonites 2-6 with 3 + 3 longitudinal ribs as long as

tergites; pereonite 7 with 2+2 ribs and small tubercle per side between two ribs; pleonite 3 with 2 large

paramedian tubercles; pleonites 4 and 5 andtelson with 2 small paramedian tubercles. Cephalon (Fig. 2 IB)

with short, quadrangular frontal lateral lobes, directed outwards and not protruding compared with median

lobe. Eyes absent. Pereonites with rounded posterior corners; epimera 4-7 slightly pointing backwards

Fig. 21. Moserius inexpectatus Reboleira & Taiti sp. nov. from Grata do Almonda, Estremenho,

paratype, S- A. Specimen in dorsal view. B. Cephalon in dorsal view. C. Pleonite 5, telson and uropod.

D. Ante nn a E. Pereopod 1.

European Journal of Taxonomy 161: 1-61 (2015)

(Fig. 21 A). Pleonites 3-5 (Fig. 21 A, C) with large epimera directed backwards. Telson (Fig. 21C)

about twice as wide as long, with distal part trapezoidal, with concave sides and truncate apex. Antenna

(Fig. 2 ID) with thickset articles of peduncle; flagellum with three articles, with j-A long aesthetascs

on second article. Uropod (Fig. 21C) with flattened protopod; exopod slightly longer than endopod,

with tuft of four apical setae; endopod with single apical seta. Pereopod 1 (Fig. 2 IE) with no particular

modifications. Pereopod 7 (Fig. 22A-B) ischium with slightly convex sternal margin, carpus with large

distal lobe on sternal margin. Pleopod 1 (Fig. 22C) exopod quadrangular, with truncate and sinuous

distal point, sinuous outer margin and straight medial margin.

Fig. 22. Moserius inexpectatus Reboleira & Taiti sp. nov. from Grata do Almonda, Estremenho, Estremoz-

Cano Massif, paratype, <f. A. Pereopod 7, caudal view. B. Pereopod 7, rostral view. C. Pleopod 1.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Remarks

This new blind species belongs to the subfamily Haplophthalminae and is included in Moserius since

it shows the tergal ornamentation typical of the genus, i.e., 3 + 3 ribs on the pereonites and two large

tubercles on pleonite 3. The genus Moserius was previously known only for two species: M. percoi

Strouhal, 1940, originally described from the cave Belinca Jama, Slovenia, and later recorded from

Ligurian and Tuscan caves (Strouhal 1940; Brian 1963; Taiti & Ferrara 1995), and M. elbanus Taiti &

Ferrara, 1995, from a small cave on Elba Island, Tuscany, Italy. The new species is readily distinguished

from the other two members of the genus by the peculiar shape of the male pleopod 1 exopod, with a

truncate and sinuous, rather than triangular, distal point. The location of M. inexpectatus (Portugal) is

very far from those of the other two species (northern Italy and Slovenia), but it is quite probable that

other species are present in the countries encompassing the northwestern Mediterranean.

Ecological notes

This species occurs in the Grata do Almonda, the largest Portuguese cave, with at least 10 km of mapped

subterranean galleries. It shares habitat with other troglobiotic species, namely the spider Nesticus

lusitanicus Fage, 1931, the woodlouse Trichoniscoides meridionalis , and the beetle Trechus lunai

Reboleira & Serrano, 2009 (Reboleira et al. 2009).

Family Styloniscidae Vandel, 1952

Genus Cordioniscus Grave, 1914

Cordioniscus lusitanicus Reboleira & Taiti sp. nov.

urn:lsid:zoobank.org:act:BlF2D756-D90F-43BF-9F42-4F8EAE6C79F0

Figs 23A-G, 24A-E, 25A-E

Diagnosis

A blind species of Cordioniscus characterised by a colourless body, the male pereopod 7 ischium having

a rounded hyaline basal lobe, the triangular male pleopod 1 exopod, as long as the endopod, and the

complex apical part of the male pleopod 2 endopod.

Etymology

From Latin lusitanicus = Portuguese. The name refers to the country where the specimens were collected.

Material examined

Holotype

PORTUGAL: $, Algar de Santo Antonio, Estremoz-Cano Massif, 30 Mar. 2009 (MZUF).

Paratypes

PORTUGAL: 1 $, 7 $$, same data as holotype, 30 Mar. 2009 (MZUF); 2 $$, 1 §, Grata de Ibne

Ammar, Algarve Massif, 23 May 2009 (MZUF); 3 same locality and date (SR); 3 $ $, Algarao do

Remexido, Algarve Massif, 5 Sep. 2009 (SR); 1 $, same locality, 30 Jan. 2009 (MZUF); 2 same

locality, 29 Dec. 2009 (SR); 1 $ juv., 2 §§, Grata da Senhora, Algarve Massif, 3 Jul. 2011 (ZMUC);

1 $, same locality, 18 May 2013 (SR).

Description

Maximum size: $ 5.0 x 1.0 mm ; § 3.0 x 0.8 mm. Body colourless and elongated (Fig. 23A). Dorsum of

cephalon and pereon finely granulated and equipped with triangular scale-setae (Fig. 23B); pleon and

telson smooth. Cephalon (Fig. 23C-D) with short, rounded frontal lateral lobes. Eyes absent. Pereonites

European Journal of Taxonomy 161: 1-61 (2015)

1-3 with rounded posterior comers; pereonites 4-7 with epimera pointing backwards (Fig. 23A).

Pleon (Fig. 23A, E) narrower than pereon; pleonites 3-5 with epimera reduced and no posterior points.

Fig. 23. Cordioniscus lusitanicus Reboleira & Taiti sp. nov. from Algar de Santo Antonio, Estremoz-

Cano. — Paratype, §. A. Adult specimen in dorsal view. — Paratype, f. B. Dorsal scale-seta.

C. Cephalon in frontal view. D. Cephalon in dorsal view. E. Pleonites 4, 5, telson and uropods. F. Anten-

nula. G. Ante nn a.

REBOLEIRA A.S.P.S. et al, Oniscidea from caves of Portugal

Telson (Fig. 23E) about twice as wide as long, distal part with concave sides and broadly rounded

apex. Antennula (Fig. 23F) with third article longer than first and second articles, with small triangular

posterior point, and with one subapical and four apical, long aesthetascs. Antenna (Fig. 23G) with

articles of peduncle bearing scale-setae; fifth article distinctly longer than flagellum; flagellum with four

articles, with very long apical organ. Right mandible (Fig. 24A) with one penicil between lacinia mobilis

and molar process, which bears long penicil; left mandible (Fig. 24A) with two penicils. Maxillula

(Fig. 24C) inner branch bearing two thickset penicils at apex and long subapical penicil; outer branch

Fig. 24. Cordioniscus lusitanicus Reboleira & Taiti sp. nov. from Algar de Santo Antonio, Estremoz-

Cano, paratype, $. A. Right mandible. B. Feft mandible. C. Maxillula. D. Maxilla. E. Maxilliped.

European Journal of Taxonomy 161: 1-61 (2015)

with 10 teeth and thin, setose stem among outer group of teeth. Maxilla (Fig. 24D) apically bilobed

and setose, with outer and inner lobes subequal; inner lobe with several long, stout setae along margin.

Maxilliped (Fig. 24E) basis enlarged in middle; endite triangular, with stout triangular penicil at apex;

palp distally rounded, with long setae at apex, and basal article with two short compound setae. Uropod

(Fig. 23E) with exopod distinctly longer than endopod, endopod inserted proximally to exopod.

Male. Pereopod 1 (Fig. 25A) with no particular modifications. Pereopod 7 (Fig. 25B) ischium with

slightly convex sternal margin and bearing rounded hyaline lobe at base. Genital papilla (Fig. 25C)

lanceolate with pointed apex, shorter than pleopod 1. Pleopod 1 (Fig. 25D) exopod triangular, as long as

endopod; endopod with short triangular basal article and flagelliform distal article. Pleopod 2 (Fig. 25E)

Fig. 25. Cordioniscus Insitanicus Reboleira & Taiti sp. nov. from Algar de Santo Antonio, Estremoz-

Cano, paratype, f. A. Pereopod 1. B. Pereopod 7. C. Genital papilla. D. Pleopod 1. E. Pleopod 2.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

exopod small, ovoidal; endopod biarticulated, distal article thickset, about three times as long as basal

article, with parallel sides and complex apical part.

Remarks

At present the genus Cordioniscus includes with certainty 14 species (Andreev 2002; Schmalfuss 2003),

mainly distributed in caves of Mediterranean countries. For a definition of the genus see Schmalfuss &

Erhard (1998). Species of Cordioniscus are mainly distinguishable by the shape of the male pleopod 2

endopod. In the Iberian Peninsula, only Cordioniscus stebbingi (Patience, 1907) was previously known

from some caves in Valencia and Castellon provinces, Spain (Vandel 1952b). The new species differs

from C. stebbingi in the lower number of aesthetascs on the antennula (5 vs 8), the presence of a hyaline

lobe on the male pereopod 7 ischium, and the thickset and complex, rather than tapering, distal part

of the male pleopod 2 endopod. In the presence of a lobe at the base of the male pereopod 7 ischium,

C. lusitanus Reboleira & Taiti sp. nov. is similar to C. bulgaricus Andreev, 1986 from Bulgaria and

C. andreevi Schmalfuss & Erhard, 1998, C. beroni Vandel, 1968, C. graecus Vandel, 1959 and

C. kithnosi Andreev, 1986 from Greece (Vandel 1959, 1968; Andreev 1986a, 1986b; Schmalfuss &

Erhard 1998). It differs from all these species in the different structure of the male pleopod 2 endopod.

Ecological notes

This species was collected in two isolated karst areas, Alentejo and Algarve. These two areas are more

than 200 km apart and are separated by the extended dry, flat areas of Alentejo Province. In the Algarve

Massif this species is frequent in caves. In the northern locality, Algar de Santo Antonio in Alentejo, the

specimens were collected in deep layers of soil at the bottom of the first pit (-20 m), where it cohabits

with the troglobiotic millipede Sireuma nobile Reboleira & Enghoff, 2014 (Reboleira & Enghoff 2014b).

Family Philosciidae Kinahan, 1857

Genus Anaphiloscia Racovitza, 1907

Anaphiloscia sicula Arcangeli, 1934

Anaphiloscia sicula Arcangeli, 1934: 152, figs 1-19.

Anaphiloscia sicula -Vandel 1946: 202, figs 50-59.

Records

Algueirao, Pechao; Algarao Menor do Paulino, Louie, Algarve Province (Vandel 1946).

Distribution

Portugal, Sicily, Ustica Island, Pantelleria Island and Malta.

Remarks

According to Schmalfuss (2003), this blind and depigmented species might be a junior synonym of

Anaphiloscia simoni Racovitza, 1907, described on the basis of a female specimen from a cave on

Mallorca, Balearic Islands (Racovitza 1907; Arcangeli 1934) and later recorded from western Spain,

Morocco, Algeria and Libya (Cyrenaica). Vandel (1972) seems to have considered the two species as

synonymous, without formally discussing the issue (Schmalfuss 2003). Since no illustrations of the male

characters of A. simoni exist, it is impossible to clarify this taxonomic problem without a re-examination

of the type material. No specimens of this genus have been collected during our investigations.

Ecological notes

Epigean species.

European Journal of Taxonomy 161: 1-61 (2015)

Genus Ctenoscia Verhoeff, 1928

Ctenoscia minima (Dollfus, 1892)

Philoscia minima Dollfus, 1892: 187, 3 figs.

Ctenoscia minima - Vandel 1946: 196, figs 42^19.

Material examined

PORTUGAL: 1 §, Gruta da Assafora, Sintra, 4 Apr. 2011 (SR).

Previous records

Algarao da Ribeira de Alte, Paderne; Algarao do Barrocal do Esguicho, Louie, Algarve Province (Vandel

1946).

Distribution

Portugal, Spain (including Canary and Balearic Islands), Corsica, Sardinia, northwestern Italy, Sicily

and surrounding islands, Malta, Tunisia and Morocco (Rif).

Remarks

This epigean species was recently discussed and fully illustrated by Taiti & Rossano (2015), who

considered it to be a senior synonym of Ctenoscia dorsalis (Verhoeff, 1928).

Ecological notes

Epigean species.

Family Platyarthridae Verhoeff, 1949

Genus Trichorhina Budde-Lund, 1908

Trichorhina anophthalma Arcangeli, 1935

Figs 26A-G, 27A-C

Trichorhina anophthalma Arcangeli, 1935b: 23, figs 19-24.

Trichorhina anophthalma - Vandel 1946: 216, figs 62-63. — Schmolzer 1965: 281, fig. 1104; 1971: 28,

89, 146. — Schmalfuss 2003: 315.

Material examined

PORTUGAL: 1 3 , 1 ?, Algar de Santo Antonio, Estremoz-Cano Massif, 30 Dec. 2009 (MZUE); 9 $

4 1 juv., same locality, 30 Mar. 2009 (MZUE); 5 33 , 3 same locality, 22 May 2009 (SR);

4 33 , 4 55 , same locality, 4 Sep. 2009 (ZMUC); 1 3 , Gruta do Vale Telheiro, Algarve Massif, 30 Jun.

2011 (MZUF); 2 Gruta da Senhora, Algarve Massif, 3 Jul. 2011 (SR).

Previous records

Serpa, Alentejo (Arcangeli 1935b). Palmela, Serra da Arrabida, Setubal (Vandel 1946).

Distribution

Known with certainty from southern Portugal and southern Spain (Schmolzer 1971).

REBOLEIRA A.S.P.S. et al, Oniscidea from caves of Portugal

Remarks

Trichorhina anophthalma is fully illustrated here (Figs 26-27) to facilitate its identification. Hoese

(1984) recorded this species from the Canary Islands (Fuerteventura and Tenerife) and considered

T. hoestlandti Vandel, 1960 from Madeira (Vandel 1960b) to be a junior synonym of that species.

Contrary to this, Schmalfuss (2003) regarded both species as valid. A re-examination of specimens

of Trichorhina hoestlandti from Madeira is necessary in order to define the taxonomic status of this

species. The specimens of Trichorhina from the Canary Islands recorded by Hoese (1984) also need to

be examined for a correct identification.

Ecological notes

This species is probably endogean and troglophile rather than a true troglobiont.

Fig. 26. Trichorhina anophthalma Ancangeli, 1935 from Algar de Santo Antonio, Estremoz-Cano, <$■

A. Adult specimen in dorsal view. B. Scale-seta. C. Cephalon in frontal view. D. Pleonite 5, telson and

right uropod. E. Antennula. F. Ante nn a G. Pereopod 1.

European Journal of Taxonomy 161: 1-61 (2015)

Family Porcellionidae Brandt, 1831

Genus Porcellionides Miers, 1877

Porcellionicles pruinosus (Brandt, 1833)

Porcellio pruinosus Brandt, 1833: 481.

Metoponorthus ( Metoponorthus ) pruinosus - Van del 1946: 272, figs 87-90.

Fig. 27. Trichorhina anophthalma Ancangeli, 1935 from Algar de Santo Antonio, Estremoz-Cano, $.

A. Pereopod 7. B. Pleopod 1. C. Pleopod 2.

REBOLEIRA A.S.P.S. et al. , Oniscidea from caves of Portugal

Material examined

PORTUGAL: 1 §, Algar de Santo Antonio, Estremoz-Cano Massif, 22 May 2009 (SR).

Previous records

Gratas dos Moinhos da Rocha, Tavira, Faro (Vandel 1946).

Distribution

Cosmopolitan species of Mediterranean origin.

Ecological notes

Epigean species, occasionally found at cave entrances.

Porcellionides sexfasciatus (Budde-Lund, 1885)

Metoponorthus sexfasciatus Budde-Lund, 1885: 167.

Metoponorthus ( Polytretus ) sexfasciatus lusitanus - Vandel 1946: 269, figs 80b-d.

Records

Grata do Algar, Mexilhoeirinha, Lagoa; Algarao, Pechao, Algarve Province (Vandel 1946).

Distribution

Western Mediterranean region, Atlantic coast of Europe and northern Africa, including Atlantic islands.

It has also been introduced to many other parts of the world.

Ecological notes

Epigean species, trogloxene.

Porcellionides cingendus (Kinahan, 1857)

Porcellio cingendus Kinahan, 1857: 279, pi. 19, figs 1-2, 4, 6, 8-9.

Metoponorthus ( Lusitanoniscus ) cingendus - Vandel 1946: 274, figs 91-97.

Records

Grata Baixa do Cabe£o dos Mosqueteiros, Aljubarrota; Cova dos Mouros de Vale Ribeiro, Lagoa do

Cao, Alcobaga (Vandel 1946).

Distribution

Southern British Isles, Atlantic coasts of France, Portugal and Spain.

Ecological notes

Epigean species, trogloxene.

Genus Porcellio Latreille, 1804

Porcellio cavernicolus Vandel, 1946

Figs 28A-G, 29A-F

Porcellio cavernicolus Vandel, 1946: 330, figs 137-143.

Porcellio cavernicolus - Schmolzer 1965: 208, figs 745-746; 1971: 37, 95, 149. — Schmalfuss 2003:

220. —Reboleira et al. 2011a: 27, fig. 3, table 1.

European Journal of Taxonomy 161: 1-61 (2015)

Material examined

PORTUGAL: 8 33 , 5 Grata d’el Rey, Cantanhede-Outil Massif, 19 May 2009 (MZUF); 4 33 ,

5 $ 5, 6 juvs, same locality, 8 Oct. 2009 (SR); 5 33 , 4 55> Grata da Senhora da Estrela, Sico Massif,

11 Jun. 2009 (MZUF); 3 33 , 2 5?, same locality, 20 Nov. 2009 (SR); 1 3 , 1 5, same locality, 29

Aug. 2009 (ZMUC); 1 3 , Grata do Soprador do Carvalho, Sico Massif, 30 Jul. 2007 (MZUF); 2 33 ,

same locality, 17 Jan. 2009 (SR); 1 3, 2 55> same locality, 30 Jul. 2009 (ZMUC); 1 juv., same locality,

21 Mar. 2009 (SR); many 33 and $ $, Grata do Algarinho, Sico Massif, 21 Mar. 2012 (MZUF); many

33 and 55, same locality, 10 Jun. 2012 (SR); 1 5,2 juvs. Grata da Ceramica, Sico Massif, 28 Nov.

in dorsal view. B. Cephalon in dorsal view. C. Epimeron of pereonite 1. D. Pleonites 3-5, telson and

uropods. E. Antennula. F. Antenna. — 3 from Grata d’el Rey. G. Pleonite 4, 5, telson and uropods.

REBOLEIRA A.S.P.S. et al ., Oniscidea from caves of Portugal

Fig. 29. Porcellio cavernicolus Vandel, 1946. — S from Gruta da Ceramica, Sico. A. Pereopod 1.

B. Pereopod 7. C. Pleopod 1. D. Pleopod 2. — $ from Gruta d’el Rey, Cantanhede-Outil Massif

E. Pereopod E F. Pereopod 7.

European Journal of Taxonomy 161: 1-61 (2015)

2009 (SR); many $$ and $ same locality, 28 Nov. 2009 (MZUF); many $$ and 5 same locality,

21 Mar. 2010 (SR); 1 same locality, 29 Aug. 2009 (SR); many $$ and $$, same locality, 27 Dec.

2010 (ZMUC); 1 $, 3 juvs, Gruta Abrigo Tomar I, Sico Massif, 26 Sep. 2011 (SR).

Previous records

Algar da Lapa, Ansiao; Gruta dos Alqueves, Sao Martinho do Bispo, Coimbra (Vandel 1946). 3

2^?, Gruta Maior da Furjaca, Pampilhosa, Coimbra, 26 Feb. 1987, A. Serra leg. (Cruz 1990).

Distribution

Species endemic to caves in central Portugal, from Cantanhede-Outil to the southernmost border of the

Sico Massif, but so far absent from the Estremenho karst massif.

Remarks

This troglobiotic species is fully illustrated here (Figs 28-29) to facilitate its recognition. The finding

of this species in Gruta d’el Rey, Cantanhede-Outil Massif, another isolated karst area far from the Sico

localities from where it was described, considerably enlarges the distribution of this species. Populations

from the two distinct karst areas show some small differences which, however, do not seem to justify

a split of the species. The specimens from Gruta d’el Rey have the apex of the telson more rounded

(compare Fig. 28D with 28G), the male pereopod 1 has less numerous setae on the sternal margin of the

merus (compare Fig. 29 A with 29E), and the male pereopod 7 has a more concave sternal margin of the

ischium (compare Fig. 29B with 29F) than specimens from the type locality.

Ecological notes

This species inhabits the most superficial parts of the caves. It can frequently be found on roots hanging

from the ceiling, from which specimens can easily be distinguished by their whitish coloration.

Porcellio dilatatus dilatatus Brandt, 1831

Porcellio dilatatus Brandt in Brandt & Ratzeburg, 1831: 78, pi. 12, fig. 6c-d.

Porcellio {Porcellio) dilatatus - Arcangeli 1935b: 12.

Porcellio dilatatus -Vandel 1946: 316, fig. 135.

Material examined

PORTUGAL: many fyfy and $ $, Gruta d’el Rey, Cantanhede-Outil Massif, 8 Oct. 2009 (MZUF); many

SS and $$, Gruta dos Moinhos Velhos, Estremenho Massif, 26 Dec. 2006 (ZMUC); 1 fy, 1 $, Gruta

dos Bolhos, Cesaredas Plateau, 18 Sep. 2009 (SR); 1 5, Gruta de Salir, Caldas da Rainha typhonic

valley, 23 May 2013 (SR); many SS and 55, Lapa da Furada, Arrabida Massif, 14 Jun. 2009 (SR).

Previous records

Beja (Arcangeli 1935b). Gruta de Ferreiros, Santo Adriao, Miranda do Douro, Braganga; Ermesinde,

Valongo, Porto; Gruta de Sabadal, Portunhos, Cantanhede, Coimbra; Gruta dos Alqueves, S. Martinho

do Bispo, Coimbra; Gruta de Legacao, Rabagal, Penela, Coimbra; Algar N. 1 das Corujeiras, Pombal,

Leiria; Algar da Lapa, Anciao, Leiria; l a Cova da Moura da Ribeira do Sirol, Leiria; Loca dos Vales,

Cortes, Leiria; Gruta Alta do Cabego dos Mosqueiros, Aljubarrota, Leiria; Gruta Baixa do Cabego dos

Mosqueiros, Aljubarrota, Leiria; Algar de Joao Ramos, Turquel, Leiria; Cova da Moura do Cabego de

Turquel, Alcobaga, Leiria; Algar n° 2 de Pinheiro, Benedita, Leiria; Casa da Moura, Cesareda, Peniche,

Leiria; Gruta do Almonda, Torres Novas, Santarem; Gruta da Senhora da Luz, Rio Maior, Santarem;

Gruta das Alcobertas, Rio Maior, Santarem; Algarve da Terra da Rolha, Cadaval, Lisboa; Gruta N. do

Furadoro, Cadaval, Lisboa; Cova da Moura, Cereal, Cadaval, Lisboa; Lapa dos Morcegos, Sesimbra,

Setubal; Lapa do Medico, Portinho da Arrabida, Palmela, Setubal; Cova da Adiga, Moura, Beja; Gruta

REBOLEIRA A.S.P.S. et al. , Oniscidea from caves of Portugal

da Seiceira, Aljezur, Faro; Algarao menor do Paulino, Cerro da Cabe 9 a Gorda, Louie, Faro; Solestreira

Pequena, Querela; Igrejinha de Soidos, Alte, Louie, Faro; Algueirao dos Mouros, Pechao, Olhao,

Faro; Abismo Novo, Moncarapacho, Olhao, Faro; Algarao do Garrafao, Cerro da Cabe 9 a, Olhao, Faro;

Ladroeira Pequena, Cerro da Cabe 9 a, Olhao, Faro (Vandel 1946).

Distribution

Species widely distributed in Europe and introduced to many other parts of the world.

Ecological notes

This species is a common inhabitant of caves and other subterranean habitats. It has no morphological

adaptations to cave life and can be considered as a troglophile.

Porcellio dispar Verhoeff, 1901

Porcellio dispar Verhoeff 1901: 407.

Porcellio dispar andel 1946: 289, figs 110-111.

Records

Algar n° 2 do Pinheiro, Benedita (Vandel 1946).

Distribution

Portugal.

Ecological notes

Epigean species, trogloxene.

Porcellio echinatus Lucas, 1849

Porcellio echinatus Lucas, 1849: 69, pi. 7, fig. 1.

Porcellio echinatus - Vandel 1946: 304, figs 125-130.

Records

Algar da Serra, Alvaiazere (Vandel 1946).

Distribution

Southern Portugal, southern Spain, northern Morocco, and north-western Algeria.

Ecological notes

Epigean species, trogloxene.

Family Armadillidiidae Brandt, 1833

Genus Eluma Budde-Lund, 1885

Eluma caelata (Miers, 1877)

Armadillidium caelatum Miers, 1877: 665, pi. 7, fig. 3.

Elumapurpurascens -V andel 1946: 357, fig. 158 (not Budde-Lund, 1885).

Records

Algarve das Gralhas, Cadaval, Montejunto Massif (Vandel 1946).

European Journal of Taxonomy 161: 1-61 (2015)

Distribution

Southern British Isles, western France, Portugal, Azores, Madeira, Canary Islands, mainland Spain and

north-western Africa. It has been introduced to Tasmania and French Guyana.

Fig. 30. Eluma tuberculata Cruz, 1991 from Grata do Soprador do Carvalho, Sico. — $. A. Specimen

in lateral view. B. Cephalon in frontal view. C. Pleonites 4, 5, telson and uropods. — $. D. Antenna.

E. Pereopod 1. F. Pereopod 7.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Remarks

The genus Eluma includes three epigean species: E. caelata , E. tuberculata Cruz, 1991 from Portugal

(see below), and E. praticola Taiti & Rossano, 2015 from the Rif region in Morocco. Eluma caelata has

been fully illustrated by Taiti & Rossano (2015: figs 32-33).

Ecological notes

Epigean species, trogloxene.

Eluma tuberculata Cruz, 1991

Figs 30A-F, 31A-B

Eluma tuberculata Cruz, 1991: 73, fig. la-f.

Eluma tuberculata - Schmalfuss 2003: 107.

Material examined

PORTUGAL: many SS and $ §, mesovoid shallow substratum (MSS) near Gruta das Alcobertas, Estre-

menho Massif (MZUF); 1 Gruta do Soprador do Carvalho, Sico Massif, 30 Aug. 2007 (SR); 4 SS,

4 $ 2 juvs, same locality, 30 Aug. 2009 (SR); 5 SS, 6 $ §, same locality, 20 Nov. 2009 (ZMUC).

Fig. 31. Eluma tuberculata Cruz, 1991 from Gruta do Soprador do Carvalho, Sico, <$. A. Pleopod 1.

B. Pleopod 2.

European Journal of Taxonomy 161: 1-61 (2015)

Previous records

Sao Mamede, Fatima (Cruz 1991).

Distribution

This species seems to be endemic to central Portugal.

Remarks

Eluma tuberculata was described by Cruz (1991) based on two females from Sao Mamede, Fatima.

The numerous specimens collected during our research allow us to illustrate the species with its most

important features, i.e., the tuberculated body and the schisma on pereonite 1 (Fig. 30A), the cephalon

(Fig. SOB), the telson and uropods (Fig. 30C), and the antenna (Fig. 30D). This species has a very plastic

morphology concerning the dorsal tubercles, which may be more or less developed. The presence of

several adult males permits the description of the male characters which were previously unknown:

Pereopods 1 (Fig. 30E) to 6 with brushes of setae on sternal margins of merus and carpus. Pereopod 7

(Fig. 3OF) ischium narrow, with straight sternal margin and a ridge on distal part of caudal surface.

Pleopod 1 (Fig. 31 A) exopod with long, rounded median lobe directed outwards, endopod with distal

half having parallel sides and small recurved and pointed apical part not protruding outwards. Pleopod 2

(Fig. 3 IB) exopod triangular and slightly shorter than endopod.

Ecological notes

This troglophilic species was collected with traps in the MSS (mesovoid shallow substratum, or ‘milieu

souterrain superficieF sensu Juberthie et al. 1980) in the Estremenho Massif as well as in a cave (Grata

Soprador do Carvalho) in Sico.

Genus Paraschizidium Verhoeff, 1918

Paraschizidium sp.

Fig. 32A-E

Material examined

PORTUGAL: 4 Algar do Javali, Montejunto Massif, 19 Sep. 2009 (MZUF); 6 $$, same locality,

24 Dec. 2009 (SR).

Remarks

At present, the genus Paraschizidium includes with certainty only three species: P. coeculum (Silvestri,

1897), distributed from Spain to the Balkans, P. hispanum Arcangeli, 1935 from southern Spain, and

P. roubali Frankenberger, 1940 from Prague, Czech Republic (Frankenberger 1940), which is probably

a junior synonym of P. coeculum (see Manicastri & Taiti 1994). Other species from Greece originally

included in the genus Paraschizidium by Schmalfuss (1981) and Sfenthourakis (1992, 1995) were

transferred to the genus Schizidium Verhoeff, 1901 by Schmalfuss (2008). Our specimens certainly

belong to the genus Paraschizidium'. they are depigmented and blind, have no schisma at the postero¬

lateral corner of the pereonite 1 (Fig. 32 A), the cephalon has oblique antennal lobes (Fig. 32B-C), the

telson is short and triangular (Fig. 32D) and the antennula has two articles (Fig. 32E). Unfortunately, the

absence of males in our material does not permit the identification of these specimens to species level.

Genus Trogleluma Vandel, 1946

Remarks

Trogleluma was erected by Vandel (1946) as a subgenus of Troglarmadillidium Verhoeff, 1900 to

include the new species Troglarmadillidium ( Trogleluma ) machadoi from two caves in South Portugal

(see below). At present, the genus Troglarmadillidium only includes the monospecific subgenera

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Troglarmadillidium and Trogleluma, with the species T. ( Troglarmadillidiam ) stygium (Verhoeff, 1900)

from Herzegovina, and T. ( Trogleluma ) machadoi. Both these species have a depigmented body, are

blind, have no schisma at the postero-lateral corner of pereonite 1, a triangular telson, and uropods

with the exopodite flattened, longer than wide. The two subgenera show quite distinct morphological

traits in the cephalic structure: in Troglarmadillidium the triangular frontal scutellum of the cephalon,

frontal line and antennal lobes are missing, while in Trogleluma these structures are present. These

characters have been confirmed after the examination of a fremale specimen of T. ( Troglarmadillidium )

stygium from Montenegro (Crna Gora, Danilovgrad, Tunjevo, Milojevica vrela, 26 Apr. 1997, leg. and

det. I. Karaman). On the basis of these characters we consider Trogleluma as a distinct genus from

Troglarmadillidium. Trogleluma includes the type species, Trogleluma machadoi , and a few undescribed

new species from Sardinia and Tuscany in Italy (Taiti 2007, unpublished results).

Trogleluma machadoi (Vandel, 1946)

Figs 33A-J, 34A-E

Troglarmadillidium ( Trogleluma ) machadoi V andel, 1946: 361, figs 148-157.

Troglarmadillidium ( Trogleluma ) machadoi - Schmolzer 1965: 307, figs 1232-1233; 1971: 62.

Troglarmadillidium machadoi - Schmolzer 1971: 106, 155. — Schmalfuss 2003: 319. —Reboleira et

al. 2011a: 7, table 1.

Fig. 32. Paraschizidium sp. from Algar do Javali, Montejunto, $. A. Specimen in lateral view.

B. Cephalon in frontal view. C. Cephalon in dorsal view. D. Pleonite 5, telson and uropods. E. Antennula.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Fig. 33. Trogleluma machadoi (Vandel, 1946) from Algarao do Remexido, Algarve. — $. A. Specimen

in lateral view. B. Scale-seta. — S.C. Cephalon in frontal view. D. Cephalon in dorsal view. E. Cephalon

in lateral view. F. Epimeron of pereonite 1 in dorsal view. G. Epimeron of pereonite 2 in ventral view.

H. Pleonite 5, telson and right uropod. I. Antennula. J. Antenna.

REBOLEIRA A.S.P.S. et al., Oniscidea from caves of Portugal

Fig. 34. Trogleluma machadoi (Vandel, 1946) from Algarao do Remexido, Algarve, S- A. Uropod.

B. Pereopod 1. C. Pereopod 7. D. Pleopod 1. E. Pleopod 2.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

Material examined

PORTUGAL: many 33 and §§, Grata de Ibne Ammar, 23 May 2009 (MZUF); many 33 and $

Algarao do Remexido, 5 Sep. 2009 (MZUF); many S3 and §§, same locality, 29 Dec. 2009 (SR);

5 S3, 7 55 , same locality, 23 May 2009 (ZMUC); 6 S3, 8 same locality, 15 Mar. 2009 (SR);

1 juv., same locality, 5 Sep. 2009 (SR); many 33 and §§, Grata da Senhora; all caves in the Algarve

Massif (MZUF).

Previous records

Algarao Menor do Paulino, Serro da Cabe£a Gorda, Louie, Faro; Abismo Novo, Moncarapacho, Olhao,

Faro, Algarve Province (Vandel 1946).

Distribution

Endemic to caves of the Algarve, the southernmost province of Portugal.

Remarks

The species is here fully illustrated (Figs 33-34) to facilitate its recognition.

Ecological notes

Troglobiotic species. The specimens were mostly found inside the clay, which often covers their

integument, an observation already pointed out by Vandel (1946). Some individuals were also found

walking on cave walls and completely clean of clay. In the caves of the Algarve, this species occurs

together with other remarkable troglobiotic species: the pseudoscorpions Titanobochica magna Zaragoza

6 Reboleira, 2010 and Lusoblothrus aenigmaticus Zaragoza & Reboleira, 2012; the spiders Harpactea

stalitoides Ribera, 1993 and Teloleptoneta synthetica Machado, 1951; the millipedes Boreviulisoma

barrocalense Reboleira & Enghoff, 2013, Acipes machadoi Enghoff & Reboleira, 2013, and A. bifillum

Enghoff & Reboleira, 2013; the campodeid Litocampa mendesi Sendra & Reboleira, 2010; and the giant

thysanuran Squamatinia algharbica Mendes & Reboleira, 2012 (Reboleira et al. 2010c, 2012a, 2012b;

Reboleira & Enghoff 2013; Enghoff & Reboleira 2013b).

Discussion

Terrestrial isopods are the most diverse taxonomic group of troglobionts (Reboleira et al. 2013b) in the

caves of Portugal, with the number of known species doubled with the present work. The Portuguese

oniscidean fauna is now composed of 16 troglobiotic species in mainland Portugal, 5 troglophiles and

6 trogloxenes (Table 2). Two additional species are known from the volcanic caves of the Portuguese

Atlantic islands: Trichoniscus bassoti Vandel, 1960 from Madeira (Vandel 1960b; Reboleira et al.

2011a), and a new, still undescribed, trichoniscid from the Azores (Borges et al. 2012). For comparison,

27 species of troglobiotic Oniscidea are known from Spain and 64 from Italy (including Sardinia and

Sicily), most of which belonging to the family Trichoniscidae (25 and 57 species, respectively).

Trichoniscidae is also the richest family in Portugal, with 14 species frequently found in caves of mainland

Portugal, 12 of which are considered to be true troglobionts. Trichoniscoides is the most diverse genus.

Trichoniscoides machadoi is an endogean and cavernicolous species occurring in different regions of

Portugal, while the remaining species are restricted to single karst areas: T. serrai to the palaeokarst of

Vimioso in northeastern Portugal; T. broteroi to the northern part of the Sico Massif; T. sicoensis to the

central area of Sico; and three species to the Estremenho Massif, the largest karst area of Portugal, i.e.,

T. meridionalis distributed in caves along its three main central subunits (Serra dos Candeeiros, Santo

Antonio Plateau and Serra de Aire/Sao Mamede Plateau), T. subterraneus in the Aljubarrota Plateau at

REBOLEIRAA.S.P.S. et al., Oniscidea from caves of Portugal

Table 2. Terrestrial isopod species recorded in subterranean habitats of karst areas from Portugal, with

ecological classification. Tb = troglobiont; Tph = troglophile; Tx = trogloxene.

Family

Species

Ecology

Karst area

Trichoniscidae

Trichoniscoides broteroi

Sico

Trichoniscoides machadoi

Tph

widespread

Trichon isco ides o uremensis

Estremenho

Trichoniscoides serrai

Vimioso

Trichoniscoides subterraneus

Estremenho

Trichoniscoides meridionalis

Estremenho

Trichoniscoides bellesi

Sico

Trichoniscoides sicoensis

Sico

Metatrichoniscoides salirensis

Caldas da Rainha

Troglonethes olissipoensis

Lisbon Peninsula

Troglonethes arrabidaensis

Arrabida

Miktoniscus longispina

Cesaredas and Sico

Haplophthalmus siculus

Lisbon Peninsula

Moserius inexpectatus

Estremenho

Styloniscidae

Cordioniscus lusitanicus

Alentejo

Philosciidae

Anaphiloscia sicula

Tph

Algarve

Ctenoscia minima

Algarve and Lisbon Peninsula

Platyarthridae

Trichorhina anophthalma

Tph

Alentejo

Porcellionidae

Porcellionides pruinosus

widespread

Porcellionides sexfasciatus

Algarve

Porcellionides cingendus

Estremenho

Porcellio cavernicolus

Sico and Catanhede

Porcellio dilatatus

Tph

widespread

Armadillidiidae

Eluma caelata

Montejunto

Elurna tuberculata

Tph

Sico and Estremenho

Paraschizidium sp.

Montejunto

Trogleluma machadoi

Algarve

the western part of the massif, and T. ouremensis in the Fatima Plateau, the eastern part of Estremenho.

Despite the sampling effort at their type localities, the troglobiotic species Trichoniscoides broteroi ,

T. serrai , T. ouremensis and T. subterraneus were not found during our research. The Iberian genus

Troglonethes now also includes two species from central Portugal: T. olissipoensis Reboleira & Taiti

sp. nov. from a cave of the Lisbon Peninsula, and T. arrabidaensis Reboleira & Taiti sp. nov. from a

cave in the Arrabida Massif Miktoniscus longispina Reboleira & Taiti sp. nov. was collected in two

different karst areas: Sico and the Cesaredas Plateau, the specimens from these localities presenting

slight morphological differences from each other.

The styloniscid Cordioniscus lusitanicus Reboleira & Taiti sp. nov. is described from caves of the

Alentejo and Algarve karst areas, more than 200 km apart in the south of Portugal.

European Journal of Taxonomy 161 : 1-61 ( 2015 )

The family Platyarthridae is represented by the species Trichorhina anophthalma , described from

endogean habitats in Palmela, the Arrabida karst massif and Serpa (Baixo Alentejo), but recently also

found in karst caves of Alandroal in central Alentejo and in the Algarve.

The family Porcellionidae has one troglobiotic species, Porcellio cavernicolus , described from the

Sico Massif, but now found in caves of the Outil-Cantanhede and Mealhada karst areas, north of the

Mondego River, and also inhabiting caves in the Tomar region, significantly increasing its former area of

distribution. Slight morphological differences can easily be observed between the extremes of its area of

distribution, and molecular studies may clarify the taxonomy of these isolated populations. This species

is particularly abundant in shallow caves with roots pending from the ceiling. The troglophilic Porcellio

dilatatus dilatatus is very abundant in the most superficial parts of caves from all of Portugal, a feature

already pointed out by Vandel (1946).

The family Armadillidiidae is represented by four species belonging to the genera Eluma, Paraschizidium

and Trogleluma. The genus Eluma includes the epigean E. caelata and E. tuberculata, originally

described from specimens collected under stones in the eastern subunit of the Estremenho Massif (Cruz

1991) and now found in the MSS on the western border of the same massif and also in one cave of

Sico. It appears to be distributed along the MSS, being found inside caves only in their most superficial

areas. One cave-adapted species of Paraschizidium was found in a cave of the Montejunto Massif and

represents the first record of this genus in Portugal. Trogleluma machadoi is widely distributed in caves

along the Barrocal area in the Algarve karst massif.

From a biospeleological point of view, mainland Portugal is traditionally divided into two main areas,

the Lusitanian district in central Portugal, and the Baetic district in the south, including the Algarve and

extending to Andalucia in Spain (Belles 1987; Reboleira et al. 2011a). The pattern of troglobiont richness

for terrestrial isopods reinforces this separation, with the Tusitanic district clearly rich in troglobiotic

species of the genus Trichoniscoides and the Baetic district characterised by the presence of the genera

Trogleluma and Cordioniscus.

The recent rise in the number of known subterranean species is a result of the extensive use of a standard

sampling methodology in a large number of karst caves, which has also provided new taxa of other cave-

adapted arthropod groups (Reboleira 2012; Reboleira et al. 2009, 2010a, 2010b, 2010c, 2011b, 2012a,

2012b, 2013b; Reboleira & Enghoff 2014a). The richness of terrestrial isopods in the subterranean

ecosystems of Portugal, and by extension of the Iberian Peninsula, requires a consistent sampling effort

to reach a complete level of evaluation.

Terrestrial isopods have been used as bioindicators in land ecosystems, since they are extremely

sensitive to contaminants (Paoletti & Hassall 1999; Paoletti et al. 2007). In the same way, the presence

of troglobiotic Oniscidea can be used as indicators of subterranean ecosystem health. A strong argument

in support of this is their basal trophic position in cave communities. Decomposing detritus and their

strong dependence on water makes them more vulnerable to contamination. The presence of troglobiotic

oniscidean species is generally related to deep, stable and undisturbed parts of caves, while in the most

disturbed areas only trogloxenes or, at most, troglophile isopods are present, suggesting that they can

be used as indicators of the quality of the subterranean ecosystems that they occupy. An example of this

is provided by the Gruta dos Moinhos Velhos, with a 300 m long show cave branch, where only the

troglophile Porcellio dilatatus dilatatus is abundant, while the troglobiont Trichoniscoides meridionalis

occurs only in the undisturbed galleries, far from human impact. All troglobiotic species of Oniscidea

in Portugal lack formal protection (Reboleira et al. 2013a) and their biodiversity patterns should be

considered for sustainable management of karst areas in such an important world biodiversity hotspot.

REBOLEIRAA.S.P.S. et al., Oniscidea from caves of Portugal

Acknowledgements

We express our gratitude to all the Portuguese caving groups: NEUA, GPS, CEAE-LPN, Geonauta,

CEEAA, NEL, AES, ECLER, NALGA, Desnivel, Alto relevo, DIR-SPE, CIES, SAGA and NEC for

their human and material support in the field work and for providing us with speleological information.

For the same reasons we are also grateful to F. Regala, R. Francisco, R. Cerqueira, O. Mateus, P Robalo

and S. Medeiros. We are indebted to Parque Natural das Serras de Aire e Candeeiros (PNSAC/ICNF),

Grutas de Mira d’Aire, S.A., Camara Municipal de Penela-CISED, and Camara Municipal do Alandroal

for logistic support during the field work. We thank Dr. Ivo Karaman for kindly sending material of

Trogloarmadillidium stygium. The Portuguese Foundation for Science and Technology (FCT, Portugal)

provided Ana Sofia Reboleira with a Postdoc grant (SFRH/BPD/92435/2013) and support for a stay of

one month at the Istituto per lo Studio degli Ecosistemi, CNR, Florence. All specimens were collected

under permits of the Instituto de Conserva£ao da Natureza e das Florestas.

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Manuscript received: 26 September 2015

Manuscript accepted: 3 November 2015

Published on: 11 December 2015

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, De nm ark