Taxonomic revision of the subfamily Toradjiinae (Crustacea: Oniscidea: Scleropactidae)

Zoological Journal of the Linnean Society (1995), 113: 351–459. With 73 figures

Taxonomic revision of the subfamily Toradjiinae (Crustacea: Oniscidea: Scleropactidae) F. FERRARA, C. MELI and S. TAITI Centro di Studio per la Faunistica ed Ecologia Tropicali del Consiglio Nazionale delle Ricerche, Via Romana 17, 50125 Firenze, Italy Received June 1994, accepted for publication December 1994

In the present revision four genera and 32 species are recognized. One genus, Paratoradjia, and 17 species, namely Adinda malaccensis, A. sumatrana, A. palniensis, A. carli, A. nilgiriensis, A. lobata, A. triangulifera, A. riedeli, A. lamellata, Protoradjia paeninsulae, P. insularis, P. montana, P. pilosa, Paratoradjia beroni, P. vietnamensis, P. sulcata and Toradjia hirsuta, are described as new. The neotype of Toradjia cephalica and lectotypes of Adinda weberi, Protoradjia jacobsoni, Toradjia gorgona and T. celebensis are established. Adinda calegarii is considered to be a junior synonym of Adinda weberi, and A. conglobator, a species inquirenda. Toradjia dollfusi is transferred to the genus Adinda, and T. indosinensis to Paratoradjia. The characters of the group and its taxonomic position within the Oniscidea are discussed: it is transferred from the Eubelidae to the Scleropactidae, of which it constitutes a separate subfamily, the Toradjiinae. A map and comments on the distribution of genera and species are included. ADDITIONAL KEY WORDS:—Isopoda – taxonomy – new genus – new species – distribution – Oriental Region.

CONTENTS Introduction . . . . . . History . . . . . . . Material . . . . . . . Abbreviations . . . . . . Morphological characters . . . Definition and systematic position . Evolution . . . . . . . Biogeography . . . . . . Taxonomy . . . . . . . Genus Adinda Budde-Lund, 1904 . Weberi group . . . . . Adinda weberi (Dollfus, 1898) . . Adinda malaccensis sp. nov. . Adinda sumatrana sp. nov. . Adinda conglobator (Budde-Lund, 1902) Stebbingi group . . . . . Adinda stebbingi (Collinge, 1914) . Adinda travancorensis (Stebbing, 1911) Adinda gigas (Collinge, 1915) . . Adinda palniensis sp. nov. . Adinda carli sp. nov. . . . Adinda nilgiriensis sp. nov. . Adinda lobata sp. nov. . . 0024–4082/95/040351+109 $08.00/0

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F. FERRARA ET AL. Adinda triangulifera sp. nov. . . . . . Adinda scabra (Collinge, 1916) . . . . . . Adinda pulchra (Collinge, 1916). . . . . . Riedeli group . . . . . . . . . Adinda riedeli sp. nov. . . . . . . Adinda platyperaeon (Schultz, 1982) . . . . . Adinda lamellata sp. nov. . . . . . Dollfusi group . . . . . . . . . Adinda dollfusi (Richardson Searle, 1922) comb. nov. Genus Protoradjia Arcangeli, 1955 . . . . . Protoradjia jacobsoni Arcangeli, 1955 . . . . Protoradjia paeninsulae sp. nov. . . . . Protoradjia insularis sp. nov. . . . . Protoradjia montana sp. nov. . . . . Protoradjia pilosa sp. nov. . . . . . Genus Paratoradjia nov. . . . . . . Paratoradjia beroni sp. nov. . . . . . Paratoradjia indosinensis (Arcangeli, 1948) comb. nov. Paratoradjia vietnamensis sp. nov. . . . Paratoradjia sulcata sp. nov. . . . . . Genus Toradjia Dollfus, 1898 . . . . . . Toradjia gorgona Dollfus, 1898 . . . . . . Toradjia celebensis Dollfus, 1898 . . . . . Toradjia cephalica Dollfus, 1898 . . . . . Toradjia hirsuta sp. nov. . . . . . . Key to genera and species . . . . . . . Acknowledgements . . . . . . . . . References . . . . . . . . . . .

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INTRODUCTION

Taiti, Ferrara & Schmalfuss (1991) revised the family Eubelidae giving a new definition and listing all the genera belonging to that family. They also stated that three genera from the Oriental region (Toradjia Dollfus, 1898, Adinda Budde-Lund, 1904, of which Paraperiscyphis Stebbing, 1911 is a junior synonym, and Protoradjia Arcangeli, 1955), previously included in the Eubelidae, belong instead to the Scleropactidae. In this contribution we discuss this statement in detail and give a revision of these three genera together with the description of a related new genus. This study represents the first comprehensive review of this enigmatic and, up to now, poorly known group, and once more points out how scanty is the knowledge of the Oniscidea in general and from tropical areas in particular. HISTORY

The first record of this group was made by Dollfus (1898), who described three species of the new genus Toradjia (T. celebensis, T. gorgona and T. cephalica) and one of the genus Periscyphus (a mis-spelling of Periscyphis) (P. weberi). BuddeLund (1902) described a new species of Toradjia from Malaysia (T. conglobator) and transferred to this genus also Periscyphis weberi. Two years later, BuddeLund (1904: 37) erected the new genus Adinda for P. weberi without giving any definition or discussion. Stebbing (1911) established the genus Paraperiscyphis for a species from south-western India (P. travancorensis), stating that it was probable that Periscyphis weberi would also have to be ascribed to this genus, obviously ignoring the existence of Adinda. Collinge (1914) described a new species of Paraperiscyphis (P. stebbingi) from southern India and a year later (Collinge, 1915)

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yet another, again from southern India, which he included in the genus Periscyphis (P. gigas), without any justification for this ascription. Collinge (1916) described two more new species of Paraperiscyphis from Sri Lanka (P. pulcher and P. scabrus ) and corrected the mistake made the year before by transferring Periscyphis gigas to the genus Paraperiscyphis. Later, Richardson Searle (1922) described a new species of Toradjia from Java (T. dollfusi), and Omer-Cooper (1926), in his revision of the genus Periscyphis, first recognized Paraperiscphis Stebbing, 1911 as a junior synonym of Adinda Budde-Lund, 1904. Arcangeli (1927), who apparently was not aware of Omer-Cooper’s paper, described a new species of Paraperiscyphis from Sumatra (P. calegarii) and later Jackson (1936) recorded this species in Malaysia and, tentatively, Adinda scabra in northern Borneo. Arcangeli (1948, 1952) is the first author to discuss this group, establishing the new subfamily Toradjiinae (family Eubelidae) for the genera Toradjia and Adinda, mainly characterized by the presence of Trachelipus-type pleopodal lungs in pleopods 1 and 2. Arcangeli (1955) defined a new genus, Protoradjia, for the new species P. jacobsoni, and better characterized this group highlighting, besides the respiratory structures of the pleopods, the exoantennate ability to roll up into a ‘semicomplete’ ball and the peculiar structure of telson and uropods, similar to that of the ‘Circoniscinae’, at that time also included in the Eubelidae (now instead in the Scleropactidae). According to Arcangeli (1955) the Toradjiinae included the following species: (1) (2) and (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13)

Toradjia celebensis Dollfus, 1898 from Sulawesi; T. gorgona Dollfus, 1898 from an unknown locality in ‘East Dutch India’ Java; T. cephalica Dollfus, 1898 from Java; T. dollfusi Richardson Searle, 1922 from Java; Adinda weberi (Dollfus, 1898) from Sumatra; A. conglobator (Budde-Lund, 1902) from Malaysia; A. travancorensis (Stebbing, 1911) from southern India; A. stebbingi (Collinge, 1914) from southern India; A. gigas (Collinge, 1915) from southern India; A. pulchra (Collinge, 1916) from Sri Lanka; A. scabra (Collinge, 1916) from Sri Lanka; A. calegarii (Arcangeli, 1927) from Sumatra; Protoradjia jacobsoni Arcangeli, 1955 from Sumatra.

Two more species must be added to this list, i.e. Toradjia indosinensis Arcangeli, 1948 from Vietnam (obviously forgotten by Arcangeli) and Paraperiscyphis platyperaeon from Borneo, described by Schultz (1982). MATERIAL

Specimens of the species described in this paper were collected in Malaysia during field trips organized by the Centro di Studio per la Faunistica ed Ecologia Tropicali, Florence (these have been deposited in the Museo di Storia Naturale, Sezione di Zoologia ‘La Specola’, Florence), and in southern India, Sri Lanka, Vietnam, Indonesia, Brunei, Philippines and New Guinea by colleagues from various institutions.

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The type specimens of Adinda weberi, Paraperiscyphis platyperaeon, Toradjia dollfusi, Protoradjia jacobsoni, T. gorgona and T. celebensis were re-examined. However, we could not re-examine those described by Stebbing (1911) (A. travancorensis) and Collinge (1914, 1915, 1916) (A. stebbingi, A. gigas, A. pulchra and A. scabra) which were originally deposited in the collections of the Zoological Survey of India, Calcutta. Since our requests for the loan of material were not granted, we do not know whether the specimens are still present in the collections of that institution or have been lost. The type specimens of Toradjia indosinensis Arcangeli, 1948 and T. cephalica Dollfus, 1898, which should be in the Museo Regionale di Scienze Naturali, Turin, and the Zoo¨logisch Museum, Amsterdam, respectively, are no longer present in the collections of those museums and must be considered as lost (E. Gavetti and D. Platvoet, personal communication). Identifications are based on morphological characters. Normally, species are very well characterized by the male sexual dimorphisms, which in several cases are truly spectacular. Abbreviations BM  Natural History Museum, London HNHM  Hungarian Natural History Museum, Budapest MHNG  Muse´um d’Histoire Naturelle, Geneva MNHN  Muse´um National d’Histoire Naturelle, Paris MZUF  Museo di Storia Naturale, Sezione di Zoologia ‘La Specola’ dell’Universita`, Florence NHMB  Naturhistorisches Museum, Basel NNHMS  National Natural History Museum, Sofia NNM  Nationaal Natuurhistorisch Museum, Leiden SMF  Senckenberg-Museum, Frankfurt SMNS  Staatliches Museum fu¨r Naturkunde, Stuttgart USNM  National Museum of Natural History, Smithsonian Institution, Washington, D.C. ZIUL  Zoological Institute, University of Lund ZMA  Zoo¨logisch Museum, Amsterdam ZMC  Zoologisk Museum, Copenhagen ZMH  Zoologisches Museum, Hamburg. MORPHOLOGICAL CHARACTERS

In the taxa considered here, the length of the body ranges from 3 mm in Toradjiahirsuta to 27 mm in Adinda platyperaeon. Usually the dorsal surface bears small scale-spines, which are triangular or petaliform, but sometimes piliform. The noduli laterales (one per side) are small and inserted on the posterior margin of the pereonites. In most species the dorsal surface is smooth, in some it is rough or distinctly granulated. The structure of the cephalon shows great differences within the group, according to the degree of specialization for rolling up and for antennal protection. In the less specialized forms (Fig. 1A) it is antero-posteriorly compressed, the profrons is flattened with no special structures to hold the antennae when the animal rolls up into a ball, and the upper margin of the

REVISION OF TORADJIINAE

Figure 1. A, Adinda malaccensis sp. nov., cephalon, dorsal view; B, Adinda stebbingi, cephalon, frontal view; C, Paratoradjia beroni sp. nov., cephalon, dorsal view. White bar  1 mm.

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frontal shield protrudes above the vertex from which it is separated by a deep groove. This groove has transverse stripes on both anterior and posterior walls and often a row of lamellar scales on its posterior margin; it is continuous in Adinda and Protoradjia, but interrupted in the middle in Paratoradjia and Toradjia. In more specialized forms the profrons shows a median longitudinal carina and two paramedian depressions which hold the antennae (as in species of Adinda of the stebbingi, riedeli and dollfusi groups) (Fig. 1B). The frontal margin may be bent over the vertex and fused with it in the middle as in Paratoradjia (Fig. 1C) or even interrupted as in Toradjia (see Fig. 65A–E); in the latter genus two deep paramedian antennal grooves run across the whole vertex. Eyes are generally large and the number of ommatidia varies (11–28) according to the size of the species. Pereonite 1 does not bear any particular structure connected to the rolling up ability in Adinda, with the exception of A. dollfusi where a rounded lobe is present at the sides of the posterior margin. In the other genera a schisma is present: in Protoradjia this cuts the posterior corner and is very shallow, with the outer lobe distinctly larger and protruding backwards in relation to the inner one; in Paratoradjia and Toradjia it is far from the posterior corner and separates two lobes normally protruding backwards to the same degree. In species of Toradjia, pereonite 1 has two variably-developed transverse grooves on the anterior margin, which are continuous with those present on the cephalon and hold the distal segment of the peduncle and flagellum of the antenna. In the forms possessing a schisma, pereonite 2 (and sometimes 3), bears a transverse ridge or a rounded lobe on the ventral surface, close to the anterior margin of the epimeron; only in Toradjia celebensis is a ventral tooth present on pereonite 2. Pleonites 3–5, with rectangular epimera, continue the body outline. The telson is semicircular, except in Adinda triangulifera and Paratoradjia beroni, where it is triangular. In all the species it is shorter than the posterior margin of the uropodal protopod and endopod. The antennule consists of three articles; the distal one is the longest, is conical and has one or more superimposed rows of aesthetascs on the medial margin. In the forms less specialized for rolling up, the antennae are long and frail (e.g. Adinda platyperaeon), while in the most specialized forms they are short and stout (Toradjia spp.). The antennal flagellum consists of two articles, the second one with some aesthetascs, and it ends with a long apical organ (half the total length of the flagellum in Toradjia and Paratoradjia). The molar penicil of the mandible may be simple, semidichotomized or dichotomized and also the number of the free penicils can vary in the different species. The outer branch of the maxillule has 10 main teeth, the inner ones often apically cleft; among the outer group of teeth a small accessory tooth and a thin stalk are present (Fig. 2A). The inner branch has two penicils, normally long and thin (e.g. Adinda weberi), rarely short and stout (e.g. Adinda stebbingi). The maxillipedal endite is setose with a short penicil on the medial corner. All pereopods bear a conspicuous, apically plumose dactylar seta (Fig. 2B). The basis of pereopod 7 has a longitudinal groove bordered by one or two rows of scales on the rostral surface (Fig. 2C). This structure, connected with

REVISION OF TORADJIINAE

Figure 2. A, Adinda malaccensis sp. nov., particular of the outer branch of maxillule with stalk and accessory tooth; B, Adinda scabra, dactylar seta; C, Adinda malaccensis sp. nov., {, pereopod 7; D, Adinda malaccensis sp. nov., water conducting system on ventral epimeron of pereonite 7. White bar  0.1 mm (A, B); 1 mm (C, D).

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the ventral grooves of the water conducting system (Fig. 2D), is similar to that present in Ligia and other Oniscidea and shows that also in this group the system is open. This is very unusual among Crinocheta, in which only the Scyphacoidea (sensu Schmalfuss, 1989) possess an open water conducting system, while all the others having a closed one (Hoese, 1981, 1982). The uropodal protopod is flattened with conspicuous exopod and endopod, both medially inserted, that fill the gap between telson and protopod, completing the body outline. The rolling up ability is exoantennate in all the species, i.e. antennae rest on the dorsum and remain outside the ball. As in other ‘rollers’ (for instance Armadillidiidae, Armadillidae and Eubelidae) there are great differences concerning the specialization of the rolling up ability: there are forms with incomplete ability, in which the animal simply bends in half like a pocketknife (e.g. Adinda platyperaeon), and others with a very convex body, in which the ball becomes completely spherical (e.g. Toradjia spp.). Only pleopods 1 and 2 exopod possess, in a dorsal position, open lungs (i.e. a respiratory area without spiracles), with two different degrees of complexity within the group. In the more simple type (Fig. 3A) the respiratory surface is only waved and completely exposed. This type, similar to that in species of the European genus Oniscus and the tropical genus Nagurus, is present in species of the genera Paratoradjia and Toradjia, as well as in Adinda dollfusi. In the more complex type (Fig. 3B) the respiratory surface is composed of irregular anastomosed folds; moreover, this surface is partly covered by the dorsal wall of the exopod. Compared to the simpler type the respiratory surface is greatly increased and the partial invagination of the respiratory folds leads to greater protection from desiccation. This type, similar to that described in the Palaearctic genus Trachelipus, is present in most species of Adinda and in Protoradjia. DEFINITION AND SYSTEMATIC POSITION

All the species considered certainly constitute a homogeneous group defined by: (1) exoantennate rolling up ability (which is more or less developed); (2) a cephalon that, though with remarkable modifications (carina, grooves to hold the antennae) within the group due to the adaptations connected with antennal protection, shows a common basic pattern: a frontal shield with upper margin protruding above the vertex and separated from it by a deep groove with transverse stripes on its walls; (3) a convex telson, normally semicircular, distinctly shorter than the tips of pleonite 5 and uropodal protopod; (4) antennal flagellum with two articles and a long apical organ; (5) maxillule inner branch with two penicils, outer one with 4 + 6 main teeth and a long stalk; (6) maxillipedal endite setose and bearing a penicil on the medial corner; (7) pereopods with long, conspicuous dactylar seta, setose in the distal part; basis of pereopod 7 rostrally with a longitudinal groove; (8) open lungs more or less developed on pleopods 1 and 2 exopod; (9) uropodal protopod flattened with both exopod and endopod medially inserted, distinctly surpassing posterior margin of telson and filling the space between protopod and telson; and (10) an open water conducting system. No character is exclusive to this group but the combination of all the characters easily distinguishes and well characterizes it.

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Figure 3. A, Paratoradjia beroni sp. nov. |, open lung of pleopod 1 exopod; B, Adinda malaccensis sp. nov. {, open lung of pleopod 1 exopod. White bar  0.1 mm.

As already proposed by Taiti et al. (1991) the Toradjia-group certainly does not belong to the Eubelidae on account of very important characters such as: absence of sulcus arcuatus on pereonite 1, telson without separation between basal and distal parts and always distinctly shorter than uropodal protopods and endopods, uropods with the exopod inserted on the medial, instead of posterior, margin of the protopod, and endopod extending beyond the tip of the telson. The structure of the telson and uropods, length of the antennal apical organ, presence of a stalk on the outer branch of the maxillule (identical to that present in Scleropactes concinnus Budde-Lund, 1885, Circoniscus gaigei Pearse, 1917 and Colomboscia cordillierae Vandel, 1972) prove that this group belongs to the family Scleropactidae. However, none of the members of this family examined by us (Circoniscus gaigei, Scleropactes tristani Arcangeli, 1930, S. concinnus, Spherarmadillo cavernicola Mulaik,

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1960, Chileoniscus marmoratus Taiti, Ferrara & Schmalfuss, 1986, Neosanfilippia zoiai Manicastri, 1991, Colomboscia bituberculata Taiti, Allspach & Ferrara, in press, and C. cordillierae) possesses a groove on the basis of pereopod 7 and thus an open water conducting system as found in the Toradjia-group, nor do other genera certainly related to the Scleropactidae, such as Hekelus Barnard, 1932 and Exzaes Barnard, 1932, both from Africa. We consider this character important enough to define a separate subfamily within the Scleropactidae, the Toradjiinae, the members of which also show a homogeneous distribution. Arcangeli (1948, 1955) first established this subfamily within the Eubelidae, pointing out the similarity with the other subfamily Cyrconiscinae (now Scleropactidae). Curiously, though, none of the defining characters of the Toradjiinae listed by that author, i.e. ‘semicomplete’ exoantennate ability to roll up, shape of the telson, inner ramus of the maxillule with two penicils, pleopod 1 and 2 exopods with Trachelipus-type lungs, is exclusive to the group, since they are present together also in the typical Scleropactidae (for instance in Scleropactes concinnus). EVOLUTION

As already pointed out by Vandel (1968) for the other Scleropactidae, in the Toradjiinae also evolution has proceeded towards a progressive specialization of the rolling up ability and of structures for protection of the antennae. In less specialized forms this ability is still imperfect and the animal simply bends with the anterior part of the body leaning on the posterior (as in species of Adinda). In these forms the body is moderately convex in cross-section and the epimera are more or less laterally directed with no particular modifications related to the rolling up ability. More specialized forms roll into perfect balls; the epimera slide one on the other and the animal, when completely rolled up, resembles a sphere (as in species of Toradjia). In these forms the body cross-section is strongly convex and the epimera are vertical and have specialization, such as a schisma on the posterior corner of pereonite 1 and teeth or transverse lobes or ridges on the ventral surfaces of the epimera of pereonite 2, which make the sphere stronger and more stable. While the anterior part of the body shows a high degree of morphological variability, the posterior part is very conservative. In almost all the species, the telson is semicircular (only in two species is it triangular) and always distinctly shorter than the uropodal protopod. Uropods show a monotonous structure, with a flattened protopod that fills a large part of the space between pleonite 5 and the telson; the stout exopod is inserted medially and endopod distinctly protrudes backwards in relation to the telson and fills the space between telson and protopod. It is interesting to observe that the evolution of the rolling up ability and the evolution of the structures for antennal protection are not parallel: in fact some species of the genus Adinda, which is the least specialized for rolling up, have the anterior part of the cephalon modified to hold the basal part of the antennae, i.e. a central carina and two large concavities at its sides (Adinda of the stebbingi, riedeli and dollfusi groups). In contrast, the species of Protoradjia, which are more specialized for rolling up (vertical epimera, schisma and ventral ridges on the epimera of pereonite 2) do not have structures on the cephalon for antennal protection, as in the species of the Adinda weberi

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group. While species of Adinda show such plasticity for antennal protection, the modifications in the species of the other genera are very stable. BIOGEOGRAPHY

The opinion of Vandel (1968), that the Scleropactidae are endemic to the Neotropical Region, with a distribution between the Tropic of Cancer and the Tropic of Capricorn, was commonly accepted until a few years ago. The French author pointed out, however, the exception of the genus Sphaerobathytropa Verhoeff, 1908 with two species, S. antarctica Vandel, 1963 from Patagonia and S. ribauti Verhoeff, 1908 from the Pyrenees (according to Schmalfuss (1980), this genus most probably belongs to the Armadillidae). Yet a form of Scleropactidae (Kefalloniscus hauseri Schmalfuss, 1986) has been described from Cephalonia in Greece and there are two genera from South Africa, Hekelus and Exzaes, which probably belong to that family (Ferrara & Schmalfuss, 1976; Schmalfuss, 1986). Furthermore, the Toradjia-group (Taiti et al., 1991) has been tentatively related to the Scleropactidae, suggesting a much wider distribution of this family. Our conclusions fully confirm this suggestion: the Toradjiinae in fact occur in a large part of the Oriental Region and in the Austro-malayan Subregion (Fig. 4). They would appear to be absent throughout much of India, the Indochinese peninsula, Philippines, etc., but this is certainly due to the lack of investigations in those regions. While the species of Protoradjia are concentrated in northern Sumatra, in the southern part of West Malaysia and Singapore, and those of Toradjia are limited

Figure 4. Distribution of the subfamily Toradjiinae: 1, Adinda weberi; 2, A. malaccensis; 3, A. sumatrana; 4, A. conglobator; 5, A. stebbingi; 6, A. travancorensis; 7, A. gigas; 8, A. palniensis; 9, A. carli; 10, A. nilgiriensis; 11, A. lobata; 12, A. triangulifera; 13, A. scabra; 14, A. pulchra; 15, A. riedeli; 16, A. platyperaeon; 17, A. lamellata; 18, A. dollfusi; 19, Protoradjia jacobsoni; 20, P. paeninsulae; 21, P. insularis; 22, P. montana; 23, P. pilosa; 24, Paratoradjia beroni; 25, P. indosinensis; 26, P. vietnamensis; 27, P. sulcata; 28, Toradjia gorgona; 29, T. celebensis; 30, T. cephalica; 31, T. hirsuta.

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to Java, Sarawak and Sulawesi, the species of Adinda and Paratoradjia are more widely distributed. Species of Adinda populate a vast area between southern India and the Philippines and those of Paratoradjia are distributed, apparently in a discontinuous manner, in a still larger area from Sri Lanka to New Ireland. Even if very little is known about the ecology of the single species, the Toradjiinae seem to populate exclusively montane and lowland tropical rain forests. TAXONOMY

Genericlevel In all the species examined many characters, such as the telson and uropods, antenna, mandible, maxilliped, dactylar seta, water conducting system and pleopodal lungs, are very similar. Yet the species can be grouped according to morphological specializations such as the ability to roll up into a ball and structures for antennal protection. Considering the large number of species examined and the stability of these specializations, they can be considered to be good characters with which genera can be defined. Four genera are recognized: Adinda, the richest in number of species (18) with no distinct structures connected with the ability to roll up into a ball; Protoradjia (5 species) characterized by a shallow schisma at the posterior corner of the pereonite 1; Paratoradjia (4 species) with frontal shield bent over vertex in the middle and partially fused with it; and Toradjia (4 species), with deep grooves for antennal protection on the cephalon and pereonite 1. Specificlevel Species are well characterized by the male sexual modifications that can involve all pereopods, pleopods and, in some species of Adinda, even sternites of pereonites 2–5. In some cases the shape of the cephalon and telson and the colour pattern are distinct enough to allow safe specific identifications, whereas dorsal granulation must be considered carefully since in some species (e.g. Adinda palniensis) juveniles are strongly granulated while adults are almost smooth. Genus Adinda Budde-Lund, 1904 Type species: Mesarmadillo [Budde-Lund’s mistake for Periscyphus] weberi Dollfus, 1898 by original designation and monotypy. Synonym: Paraperiscyphis Stebbing, 1911. Type species: Paraperiscyphis travancorensis Stebbing, 1911 by original designation and monotypy. Diagnosis Animals with incomplete exoantennate ability to roll up into a ball. Cephalon: profrons flattened or with a longitudinal carina in the middle, more or less protruding; large frontal lateral lobes, often distinctly protruding above vertex and directed forwards; frontal lamina separated from vertex by a deep groove.

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Pereonite 1 with posterior margin usually concave, sometimes straight or even deeply incised at base of epimera; posterior corner without schisma; no lobes or teeth on ventral surface. Epimera of pereonite 2 ventrally with no transverse ridge. Telson normally semicircular, rarely triangular, always shorter than uropodal protopods. Antennule of three articles. Antenna with flagellum of two articles and long apical organ. Mandible with molar penicil simple (i.e. consisting of a single plumose seta) or semidichotomized (i.e. consisting of some plumose setae arising from a common stem). Outer branch of maxillule with 4 + 6 teeth, an accessory small tooth and a slender stalk with small nodes in the distal part; inner branch with two penicils. Maxilliped with setose endite bearing a penicil on medial corner. Pereopods with long dactylar seta, bifid and setose at apex. Basis of pereopod 7 with a longitudinal deep groove bordered with scales on rostral surface. Pleopodal exopods 1–2 with open lungs similar to those in Trachelipus. Uropods with protopod flattened, posteriorly broadly rounded or subtruncated; exopod inserted in a notch of the medial margin of protopod; endopod distinctly surpassing posterior margin of telson. Remarks Budde-Lund (1904: 37), in his key to the Spherilloninae, established the genus Adinda for the species Mesarmadillo weberi Dollfus, 1898, a combination never published by the French author; obviously Budde-Lund meant Periscyphus weberi Dollfus, 1898. According to Omer-Cooper (1926: 351), Paraperiscyphis Stebbing, 1911 is a junior synonym of Adinda. The species present some differences in the structure of the cephalon and buccal pieces. In A. weberi the frontal shield is almost flattened, with no distinct carina in the middle, and with small lateral lobes not protruding forwards, while in A. platyperaeon a sharp medial carina is present and the lateral lobes are broad and protrude forwards. These differences must be considered as within genus variability since, between these two extreme conditions, intermediate states are also present. As regards the buccal pieces, the differences concern mandibles and maxillules. In some species (e.g. A. riedeli), the molar penicil of the mandible is clearly simple, while in others (e.g. A. weberi) it is semidichotomized. The maxillule also varies: the teeth of the outer branch may be entire (A. stebbingi) or some apically cleft (A. weberi); the two penicils of the inner branch may be long and thin (A. weberi) or short and stout (A. stebbingi). In the systematics of the Oniscidea the structure of buccal pieces is considered to be of high taxonomic value in some groups (e.g. Philosciidae) but not in others (e.g. Armadillidae). For the genus Adinda, we cannot exclude that these differences may be important enough to characterize distinct taxa (such as subgenera). However, considering the uniformity of all the other characters, the still fragmentary knowledge of this group (certainly several other species will be discovered), that some species have not been properly investigated (A. conglobator, A. travancorensis and A. gigas) and that also in species of Toradjia, certainly a homogenous group, there are some differences in the buccal appendages (see below), we prefer for the time being to delineate only species-groups. Although they have no taxonomic status, they serve as a useful means of grouping species that have combinations of characters in common. Four groups are recognized: the weberi group, the stebbingi group, the riedeli group and the dollfusi group. Their characters are summarized in Table 1.

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TABLE 1. Synopsis of the diagnostic characters of the four species-groups of Adinda. ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––– — Cephalic Mandibular Maxillular Maxillular GROUP carina molar penicil inner teeth penicils ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––– — weberi absent semidichotomized cleft long and thin (4. spp.) stebbingi present semidichotomized simple short and stout (10 spp.) riedeli present simple cleft long and thin (3 spp.) dollfusi present semidichotomized simple long and thin (1 sp.) ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––– —

It is interesting to note that each group is distributed in a homogeneous area that does not overlap with that of the others: the stebbingi group is the western one (southern India and Sri Lanka); the weberi group the central one (peninsular Malaysia and Sumatra); the riedeli group the eastern one (Sarawak and Philippines); and the dollfusi group, in fact only one species, the southern one (Java). Weberi group Adinda weberi (Dollfus, 1898) (Figs 5–7) PeriscyphusWeberi Dollfus, 1898: 371, fig. 16. Toradjiaweberi ; Budde-Lund, 1902: 380. AdindaWeberi ; Budde-Lund, 1904: 37; Arcangeli, 1948: 34; Arcangeli, 1955: 13. Paraperiscyphis Calegarii Arcangeli, 1927: 79, fig. 1 (syn. nov.). AdindaCalegarii ; Arcangeli, 1948: 34. ? Adinda calegarii; Jackson, 1936: 81. Typematerial examined INDONESIA: 1 { (Lectotype), 2 {{, 1 | (Paralectotypes), Sumatra, Kajoe Tanam, leg. M. Weber, 1888, ZMA; 1 | (Paralectotype), same data, MNHN; 2 {{, 2 || (Paralectotypes), Sumatra, Manindjiau, leg. M. Weber, 1888, ZMA; 2 {{, 1 | (Paralectotypes), same data, MNHN; 1 {, 2 || (Paralectotypes), Sumatra, Padang, Apenberg, leg. M. Weber, 1888, ZMA. Material examined INDONESIA: 2 {{, 8 ||, Sumatra, Lembah Anai, 500 m, 1926, leg. E. Jacobson, ZMA; 1 {, 2 ||, same data, MZUF. Previousrecords Indonesia: Sumatra: Kajoe Tanam; Manindjiau; Padang, Apenberg (Dollfus, 1898); Padang (Arcangeli, 1927); Airbangis, Pad.-Boveulanden (Arcangeli, 1955). ? Malaysia: Perak: near river Yum and Kuala Legap, Plus Valley (Jackson, 1936).

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Figure 5. Adinda weberi. A, Adult male, lateral view; B, cephalon, frontal view; C, cephalon, dorsal view; D, cephalon, posterior view; E, pleonite 5, telson and uropods; F, antennule; G, antenna.

Description Maximum length: {, 13 mm; |, 11 mm. Colour (in alcohol) grey-brownish with the usual pale muscle spots; one round pale spot at base of pereon epimera; pereon and pleon epimera lighter than tergites; antenna with distal part of fifth article of peduncle and flagellum pale. Dorsum smooth, with no

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Figure 6. Adinda weberi. A, Left mandible; B, right mandible; C, maxillule; D, maxilliped; E, uropod, ventral view.

scale-spines. Eye with 24–26 ommatidia, arranged in four rows. Cephalon with a flattened frontal shield clearly protruding above vertex and separated from this by a narrow and deep groove; upper margin of frontal shield convex and lateral lobes subright in frontal view. Pereonite 1 with posterior corners rounded and directed backwards, posterior margin regularly concave at sides; pereonites 2–7 with quadrangular epimera. Telson broadly rounded, shorter than epimera of pleonite 5. Antennule with second article shorter than the others and third article conical, very elongated, with some rows of aesthetascs in the distal part. Antenna with fifth article of peduncle reaching posterior margin of pereonite 1; flagellum with second article twice as long as the first. Mandible with molar penicil semidichotomized; left mandible with 2 + 2 penicils; right mandible with 3 + 1 penicils. Maxillule outer branch with five out of 10 teeth cleft; inner branch with two long penicils. Maxilliped with setose endite bearing a stout

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Figure 7. Adinda weberi, {. A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

penicil on medial corner. Uropodal protopod subquadrangular, with posterior margin protruding in relation to epimera of pleonite 5; endopod about twice as long as exopod; exopod and endopod slightly surpassing tip of protopod.

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Male. Pereopods 1–3 carpus and, to a lesser extent, merus with lines of long trifid spines; sternal margin of carpus of pereopods 1–3 and merus of pereopods 1–4 with a brush of small scales. Pereopod 7 ischium with concave sternal margin, a setose area at the base and one on the distal part of the rostral surface; merus with a setose, clearly protruding lobe. Pleopod 1 exopod with a long triangular posterior point, outer margin with several spines; endopod straight, with a hyaline lobe at apex. Pleopod 2 endopod longer than exopod. Pleopods 3–5 exopod as in Fig. 7E–G. Distribution Indonesia (Sumatra) and Malaysia (?). Remarks This species is characterized by the cephalon with the frontal shield flattened and protruding over the vertex, the male pereopod 7 ischium with concave sternal margin and merus with a setose basal lobe, the male pleopod 1 exopod with a long triangular posterior point. According to its original description (Arcangeli, 1927), Paraperiscyphis calegarii presents no differences from A. weberi, and the locality where the two species were collected is the same (Sumatra, Padang). In our opinion P. calegarii must be considered as a junior synonym of A. weberi. Jackson’s record (1936) from Malaysia is doubtful and re-examination of this material is necessary. Most probably these specimens belong to one of the other species recorded from Malaysia (see below). Adinda malaccensis sp. nov. (Figs 1A, 2A, C, D, 3B, 8, 9) Material examined MALAYSIA: 1 { (Holotype), 12 {{, 11 ||, 3 juvs (Paratypes), Pahang, Genting Highlands, 1725 m, edge of forest, leg. S. Taiti & L. Bartolozzi, 18.xi.1987, MZUF; 1 {, 1 | (Paratypes), same data, SMNS; 2 || (Paratypes), same locality, 1600 m, edge of forest, leg. S. Taiti & L. Bartolozzi, 20.xi.1987, MZUF; 1 {, 2 || (Paratypes), Pahang, along the road to Genting Highlands, 1320 m, leg. S. Taiti, 7.xi.1985, MZUF; 4 {{, 3 || (Paratypes), same locality, 1650 m, leg. S. Taiti & L. Bartolozzi, 13.xi.1987, MZUF; 3 ||, 1 juv. (Paratypes), Pahang, Genting Tea Estate, about 40 km NE of Kuala Lumpur, 610 m, leg. S. Taiti, 2.xi.1985, MZUF; 1 { (Paratype), same locality, leg. S. Taiti & L. Bartolozzi, 16.xi.1987, MZUF; 2 || (Paratypes), Pahang, Bukit Rengit, near Lanchang, 120 m, lowland forest, near stream, leg. S. Taiti & L. Bartolozzi, 30.xi.1987, MZUF; 5 {{, 2 ||, 11 juvs (Paratypes), same locality, leg. S. Taiti, 15.xi.1987, MZUF; 1 juv. (Paratype), Selangor, Ulu Gombak, 24 km NE of Kuala Lumpur, 230 m, lowland forest, leg. S. Taiti, 20.x.1985, MZUF; 1 juv. (Paratype), same locality, leg. S. Taiti, 22.x.1985, MZUF. Description Maximum length: {, 19 mm; |, 24 mm. Colour (in alcohol) grey-brownish with the usual pale muscle spots; one pale spot at the base of pereon and

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Figure 8. Adinda malaccensis sp. nov. A, Adult male, lateral view; B, cephalon, frontal view; C, cephalon, dorsal view; D, cephalon, posterior view; E, pleonite 5, telson and uropods; F, antennule; G, antenna.

pleon epimera; antenna with distal half of fifth article of peduncle and flagellum colourless (red in vivo?). Dorsum smooth with no visible scale-spines. Eye with about 26 ommatidia arranged in four rows. Cephalon similar to that of A.

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Figure 9. Adinda malaccensis sp. nov., {. A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

weberi, but with frontal shield having upper margin slightly sinuous and lateral lobes more rounded. Pereon, pleon, telson, antennule, antenna, buccal pieces and uropod as in A. weberi. Male. Pereopods 1–4 with lines of trifid spines on carpus and, more sparsely,

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on merus; a brush of small scales on sternal margin of carpus and merus. Pereopod 7 ischium narrower and longer than in A. weberi, with straight sternal margin; merus with basal lobe little protruding and bearing long setae. Pleopod 1 exopod with a short triangular posterior point with some spines; straight endopod with no particular modifications at apex. Pleopods 2–5 similar to those of A. weberi. Etymology The name refers to the Malay peninsula (Malacca) where these specimens were collected. Distribution Malaysia. Remarks Adinda malaccensis is morphologically very similar to A. weberi, from which it is distinguished by the larger maximum dimensions, the frontal shield with the upper margin sinuous instead of regularly convex, the male pereopod 7 which has a narrower ischium with straight, instead of concave, sternal margin, merus with the basal lobe less protruding, and the male pleopod 1 exopod with shorter posterior point.

Adinda sumatrana sp. nov. (Figs 10–12) Material examined INDONESIA: 1 { (Holotype), 1 | (Paratype), Sumatra, west coast, Tandjunggadang, 1000 m, leg. E. Jacobson, 1926, ZMA; 1 | (Paratype), same locality, leg. E. Jacobson, 1925, MZUF; 2 ||, 1 juv. (Paratypes), Sumatra, Bukittingi, Gunung Merapi, 2000–2200 m, leg. A. Riedel, 18.x.1990, SMNS; 1 { (Paratype), same data, MZUF; 1 {, 1 |, 1 juv. (Paratypes), Sumatra, Bukittingi, Gunung Singgalang, leg. A. Riedel, 16.x.1990, SMNS; 1 | (Paratype), same locality, 1800 m, leg. E. Jacobson, 1925, ZMA. Description Maximum length: {, 19 mm; |, 22 mm. Colour (in alcohol), as in A. weberi. Dorsum with some small triangular scale-spines. Cephalon with frontal shield similar to A. malaccensis. Pereonites 1–3 with posterior corners subacute. Pleon, telson, antennule, antenna and buccal pieces as in A. weberi. Uropodal protopod narrower than in the previous two species. Male. Pereopods 1–4 carpus and, to a lesser extent, merus with lines of long trifid spines and a brush of small scales on sternal margin. Pereopods 3–4 ischium with some conspicuous tubercles on sternal margin. Pereopod 7 ischium with sternal margin sinuous in the distal part; merus with basal lobe similar to that of A. malaccensis, but with shorter setae. Pleopod 1 exopod with a stout

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Figure 10. Adinda sumatrana sp. nov. A, Adult female, lateral view, B, dorsal scale-spine; C, cephalon frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and left uropod; G, antennule; H, antenna.

triangular posterior point; endopod with distal part slightly directed backwards bearing a hyaline lobe at apex. Pleopods 2–5 similar to those of A. weberi. Etymology The name refers to the island of Sumatra, where these specimens were collected.

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Figure 11. Adinda sumatrana sp. nov., {. A, Pereopod 1; B, pereopod 4; C, pereopod 7.

Distribution Indonesia (Sumatra). Remarks This species is very similar to A. weberi and A. malaccensis and is distinguished from both essentially by the narrower uropodal protopod, the tubercles on the male pereopods 3–4 ischium, the male pereopod 7 ischium with sternal margin sinuous, and the male pelopod 1 exopod with a larger posterior point. Species inquirenda Adinda conglobator (Budde-Lund, 1902) Toradjiaconglobator Budde-Lund, 1902: 380. Adindaconglobator ; Arcangeli, 1948: 34.

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Figure 12. Adinda sumatrana sp. nov., {. A, Pleopod 1; B, pleopod 2; C, pleopod 3 exopod; D, pleopod 4 exopod; E, pleopod 5 exopod.

Previousrecords Malaysia: Ajenz (?Aring), Kelantan (Budde-Lund, 1902). Distribution Malaysia. Remarks Budde-Lund (1902) did not properly describe or illustrate this species. He only stated that it was ‘‘most nearly allied’’ to A. weberi from which it differed in having ‘‘shorter antennae, with the flagellum short, white, and basal joint very short’’. Without a re-examination of the type material, which appears not to be deposited in any of the institutions contacted by us, nothing can be said about this species, except that it certainly belongs to Adinda.

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Stebbingi group Adinda stebbingi (Collinge, 1914) (Figs 1B, 13–16) Paraperiscyphisstebbingi Collinge, 1914: 207, pl. XXIV figs 1–10; Collinge, 1916: 115. AdindaStebbingi ; Arcangeli, 1948: 34. Material examined INDIA: 11 {{, 14 ||, 9 juvs, Anamalai Hills, Valparai, under rotting logs, leg. J. Carl, 4.iii.1927, MHNG; 2 {{, 1 |, same data, MZUF; 1 {, 1 |, same data, SMNS. Previousrecords India: Anamalai Hills, Madras Pres. (Collinge, 1914); Kavalai, Cochin State (Collinge, 1916). Description Maximum length: {, 23 mm; |, 20 mm. Colour (in alcohol) pale brown, probably due to the long conservation (dark brown according to Collinge, 1914). Dorsum strongly granulated with larger granules on tergites and on posterior margin of pereonites; several petaliform scale-spines surrounded by small scales. Eye with 28 ommatidia disposed in four rows. Cephalon with a distinct carina on profrons; frontal shield protruding above vertex and separated from this by a deep groove bordered by a row of scales; frontal shield with medial lobe triangularly raised; lateral lobes directed forwards, more protruding than carina. Pereonite 1 with posterior margin clearly concave at the base of epimera; posterior corner acute. Pereonites 2–4 with epimera directed backwards and with posterior corners progressively less acute. Pereonites 5–7 with posterior margin slightly sinuous at sides. Telson wider than long, with broadly rounded apex. Antennule with second article slightly shorter than first; third article conical with several rows of aesthetascs on medial margin. Antenna with fifth article of peduncle reaching the posterior margin of pereonite 2; second article of flagellum about 1.5 times as long as first. Mandibles with molar penicil semidichotomized; left mandible with 6 + 2 penicils; right mandible with 4 + 1 penicils. Maxillule outer branch with all teeth apically entire; inner branch with two stout subequal penicils. Uropodal protopod with distal margin rounded; endopod and exopod slightly protruding in relation to the protopod tip. Male. Sternites of pereonites 2–5 with two medial points protruding downwards. Pereopods 1–4 carpus and merus with lines of long trifid spines and a large area of short setae on the rostral surface. Pereopods 2–4 merus with sternal margin clearly sinuous, and ischium with some large tubercles disposed in two lines on sternal margin. Pereopods 5–7 ischium, merus and carpus with lines of long spines on sternal margin. Pleopod 1 exopod with a long triangular posterior point bearing some spines on outer and medial margin; endopod with no particular modifications. Pleopod 2 endopod slightly longer than exopod. Pleopods 3–5 exopod as in Fig. 16D–F.

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Figure 13. Adinda stebbingi. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pereonite 4, posterior view; G, antennule; H, pleonite 5, telson and uropods.

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Figure 14. Adinda stebbingi. A, Left mandible; B, right mandible; C, maxillule; D, maxilliped.

Distribution Southern India. Remarks The specimens examined correspond quite well to Collinge’s description of A. stebbingi and the locality of their collection is the same as the type locality (Anamalai Hills). A. stebbingi is characterized by a strongly granulated body, the cephalon with a distinct frontal carina and lateral lobes protruding forwards, the pereonite 1 with posterior margin concave at the sides, and by the male characters, such as the triangular points on the sternites of the pereonites 2–5 and the modifications in all pereopods.

Adinda travancorensis (Stebbing, 1911) (Fig. 17) Paraperiscyphistravancorensis Stebbing, 1911: 184, pl. XI. Adindatravancorensis ; Arcangeli, 1948: 34.

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Figure 15. Adinda stebbingi, {. A, Pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E, pereopod 5; F, pereopod 6.

REVISION OF TORADJIINAE

Figure 16. Adinda stebbingi, {. A, Pereopod 7; B, pleopod 1; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

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Figure 17. Adinda travancorensis. A, Adult specimen, dorsal view; B, adult specimen, lateral view; C, cephalon and pereonite 1, dorsal view; D, antenna [Adapted from Stebbing, 1911].

Previousrecords India: Travancore, Maddathorai (Stebbing, 1911). Distribution Southern India. Remarks According to Stebbing (1911), this species is characterized by the posterior margin of pereonite 1 which is ‘‘angularly produced backwards in an exceptional manner’’. None of the specimens examined by us from southern India shows this characteristic and can be ascribed to this species. The original illustrations that show the peculiar shape of pereonite 1 are included here. This species and the following one (A. gigas) are only tentatively included in the stebbingi group due to the lack of detailed information on the buccal pieces.

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Adinda gigas (Collinge, 1915) Periscyphisgigas Collinge, 1915: 148, pl. IX figs 1–10. Paraperiscyphisgigas ; Collinge, 1916: 116. Adindagigas ; Arcangeli, 1948: 34. Previousrecords India: Ponmudi, Travancore [about 35 km NE of Trivandrum, Kerala State] (Collinge, 1915). Remarks The original description and illustrations of A. gigas are not useful to fully characterize this species. Nevertheless, none of the specimens examined by us from southern India shows the peculiar colour pattern described by Collinge, 1915 (‘‘horny-brown with the lateral plates of the 1st, 5th and 6th mesosomatic and the 3rd and 4th metasomatic segments yellow’’) and can be ascribed to this species. Adinda palniensis sp. nov. (Figs 18–20) Material examined INDIA: 1 { (Holotype), 2 {{, 3 ||, 6 juvs (Paratypes), Upper Palni Hills, Vandaravu, Sholas, about 2300 m, leg. J. Carl, 7.iv.1927, MHNG; 5 ||, 1 juv. (Paratypes), Upper Palni Hills, Kukkal-Shola, about 1900 m, under rotting logs, leg. J. Carl, 1.iv.1927, MHNG; 1 { (Paratype), same data, MZUF; 1 {, 7 || (Paratypes), same data, MHNG; 3 {{, 5 ||, 3 juvs (Paratypes), same locality and collector, 3.iv.1927, MHNG; 2 {{, 3 || (Paratypes), Upper Palni Hills, Bombay-Shola, near Kodaikanal, 2200 m, under logs and stones, leg. J. Carl, 21.iii.1927, MHNG; 1 {, 1 | (Paratypes), same data, MZUF; 1 { (Paratype), same data, SMNS; 1 |, 2 juvs (Paratypes), Lower Palni Hills, Shola near Shembaganur (Tiger-Shola?), about 1600 m, leg. J. Carl, 17.iv.1927, MHNG; 1 {, 1 |, 1 juv. (Paratypes), Upper Palni Hills, Mariyan-Shola, 2400 m, forest, leg. J. Carl, 11–14.iv.1927, MHNG; 1 {, 5 juvs (Paratypes), Travancore, Vattavadai Valley (between Palni and Anamalai Hills), primary forest, 1800–1850 m, leg. J. Carl, 10.iv.1927, MHNG. Description Maximum length: {, 17 mm; |, 19 mm. Colour (in alcohol) brownish with pale muscle spots and a round pale spot at the base of pereonal epimera. Dorsum almost smooth in adults, distinctly granulated in young specimens, with several small petaliform scale-spines. Eye with about 25 ommatidia. Cephalon with a very narrow frontal carina protruding forwards; lateral lobes at right angle, directed forwards, clearly separated from vertex by a deep groove. Pereonite 1 with posterior margin slightly sinuous, posterior corners acute with rounded apex. Telson broadly rounded, about twice as long as wide. Antennule similar to that of A. stebbingi but with articles proportionally shorter. Antenna with fifth article reaching the posterior margin of pereonite 1 and with first article of flagellum about half the length of the second. Buccal pieces as in A.

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Figure 18. Adinda palniensis sp. nov. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna; I, uropods, ventral view.

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Figure 19. Adinda palniensis sp. nov., {. A, Pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E, pereopod 5; F, pereopod 6.

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Figure 20. Adinda palniensis sp. nov., {. A, Pereopod 7; B, pleopod 1 exopod; C, pleopod 1 endopod; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

stebbingi. Uropodal protopod short, compared to that of the other species, quadrangular, ventrally clearly depressed in the medial part; endopod slightly more protruding than exopod. Male. Sternite of pereonite 4 with two medial points protruding downwards.

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Pereopods 1–2 carpus with a setose area on the rostral surface close to the sternal margin. Pereopods 1–4 carpus and distal part of merus with a brush of short scales on sternal margin. Pereopods 2–4 ischium with distinct tubercles on sternal margin. Pereopods 5–7 carpus and merus with lines of long spines on sternal margin. Pereopod 7 ischium with lines of spines on the sternal margin of the distal part and with a triangular lobe at the base, directed proximally. Pleopods 1–5 similar to those of A. stebbingi. Etymology The name of the new species refers to the site of collection of the specimens: the Palni Hills. Distribution Southern India. Remarks Adinda palniensis is characterized by the dorsal surface of the body with no granulations, the frontal carina distinctly protruding forwards, short and quadrangular uropodal protopods, the presence of two medial points on the sternite of the male pereonite 4, and the modifications of the male pereopods. In the almost smooth dorsum it corresponds to A. gigas from which it is readily distinguished by the colour pattern. It is also similar to A. stebbingi but is distinct from that species essentially by the absence of dorsal granulations, telson relatively larger, uropods larger and shorter, and the different male characters, mainly the presence of medial points only on the sternite of the pereonite 4, the absence of distinct modifications on pereopods 1–4, and the presence of a spur-like lobe directed proximally at the base of the pereopod 7 ischium. Adinda carli sp. nov. (Figs 21, 22) Material examined INDIA: 1 { (Holotype), 1 {, 5 ||, 3 juvs (Paratypes), Lower Palni Hills, Tandikudi, Cardamon Estate, about 1500 m, under logs, leg. J. Carl, 23.iv.1927, MHNG; 1 {, 1 | (Paratypes), same data, MZUF; 1 { (Paratype), same data, SMNS; 1 {, 2 ||, 2 juvs (Paratypes), Lower Palni Hills, Maryland, Shola, 1600 m, leg. J. Carl, 20.iv.1927, MHNG. Description Maximum length: {, 14 mm; |, 17 mm. Colour (in alcohol) brownish with several pale spots. Dorsum strongly granulated with more developed granules in the central part and on the posterior margin of tergites. Eye with 22–23 ommatidia. Cephalon with frontal shield bearing a narrow central carina, less protruding forwards than lateral lobes; frontal shield separated from vertex by a deep groove bordered by same scales; a supra-antennal line interrupted in the middle is present. Pereonite 1 with posterior margin slightly sinuous at base of epimera; posterior corner acute. Telson about twice as wide as long

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Figure 21. Adinda carli sp. nov. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna.

with apex broadly rounded. Antennule similar to previous species. Antenna with fifth article of peduncle reaching half of pereonite 2; second article of flagellum about twice as long as first. Buccal pieces as in A. stebbingi. Uropodal protopod subquadrangular, slightly longer than wide, ventrally depressed in the medial part; exopod less and endopod more protruding than protopod apex. Male. Sternites of pereonites 4 and, to a lesser extent, 5 with two medial points protruding downwards. Pereopod 1 carpus with a setose area close to

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Figure 22. Adinda carli sp. nov., {. A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

the sternal margin. Pereopods 1–4 carpus and merus with a brush of short scales on sternal margin. Pereopods 5–7 carpus and merus with lines of long spines on sternal margin. Pereopod 7 ischium with a small basal lobe directed

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proximally; merus with a rounded setose lobe on rostral surface. Pleopods 1–5 similar to those of A. palniensis. Etymology The new species is named after Prof. J. Carl, Geneva, who collected these specimens. Distribution Southern India. Remarks This species is very similar to A. stebbingi and A. palniensis. It is distinguished from the former by the different shape of the uropod, modifications on the ischium and merus of the male anterior pereopods, and the presence of a small lobe at the base of the ischium and rounded lobe on the merus of the male pereopod 7. It is distinguished from the latter by the presence of dorsal granulations, longer uropodal protopod, absence of tubercles on the ischium of the male pereopods 2–4, and presence of a rounded lobe on the merus of the male pereopod 7. Adinda nilgiriensis sp. nov. (Figs 23–25) Material examined INDIA: 1 { (Holotype), 6 {{, 8 ||, 7 juvs (Paratypes), Nilgiri Hills, Avalanche, about 2000 m, forest, under relatively dry logs, leg. J. Carl, 18.i.1927, MHNG; 1 {, 1 | (Paratypes), same data, MZUF; 1 { (Paratype), same data, SMNS; 5 {{, 4 || (Paratypes), Nilgiri Hills, jungle below Coonoor, 1600 m, under dead leaves and stones on deep soil, leg. J. Carl, 24.xii.1926, MHNG; 1 { (Paratype), same data, MZUF; 1 juv. (Paratype), Nilgiri Hills, Coonoor, Lady Cumings-Seat, about 1700 m, under very damp dead leaves, leg. J. Carl, 29.xii.1926, MHNG; 1 { (Paratype), Nilgiri Hills, Dodabetta, reserved forest, 2400 m, Ootacamund Region, leg. J. Carl, 11.i.1927, MHNG; 2 {{, 2 ||, 1 juv. (Paratypes), Nilgiri Hills, Old Street of Nilgiri, 1–4 mi below Coonoor, 1600 m, under dead leaves and ground, leg. J. Carl, 4.i.1927, MHNG; 1 | (Paratype), Nilgiri Hills, S of Coonoor, forest, 1600 m, under dead leaves, leg. J. Carl, 21–24.xii.1936, MHNG; 1 | (Paratype), Nilgiri Hills, Coonoor, forest, under leaves, 1800 m, leg. J. Carl, 17.xii.1926, MHNG. Description Maximum length: {, 18 mm; |, 19 mm. Colour (in alcohol) brownish with several small pale spots; antennae with distal part of the fifth article of peduncle and first article of flagellum pale. Dorsum slightly granulated, with several small petaliform scale-spines. Eye with about 21 ommatidia. Cephalon with frontal shield bearing a narrow carina protruding forwards and upwards; lateral lobes triangular, directed forwards, separated from vertex by a deep groove with a row of scales; a supra-antennal line interrupted in the middle is present. Pereonite 1 with posterior margin slightly sinuous at base of epimera, posterior

REVISION OF TORADJIINAE

Figure 23. Adinda nilgiriensis sp. nov. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, telson and uropods; G, antennule; H, antenna.

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Figure 24. Adinda nilgiriensis sp. nov., {. A, Pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E, pereopod 5; F, pereopod 6.

REVISION OF TORADJIINAE

Figure 25. Adinda nilgiriensis sp. nov., {. A, Pereopod 7; B, pleopod 1; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

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corner acute with rounded apex. Telson twice as wide as long, broadly rounded. Antennule as in the previous species. Antenna with fifth article of peduncle reaching the posterior margin of pereonite 1; second article of flagellum about twice as long as first. Buccal pieces as in A. stebbingi. Uropodal protopod wide and ventrally depressed in the medial part; exopod slightly less and endopod more protruding in relation to protopod apex. Male. Sternites of pereonites 2–4 with two small points and pereonite 5 with two well-developed ones, directed downwards. Pereopods very similar to those of A. palniensis, except pereopod 7 which presents ischium with basal lobe perpendicular to sternal margin and merus with a protruding rounded lobe on tergal margin. Pleopods 1–5 as in A. palniensis. Etymology The name refers to the Nilgiri Hills, where the specimens were collected. Distribution Southern India. Remarks This species, because of the presence of a spur-like lobe on the proximal part of the ischium of the male pereopod 7, is similar to A. palniensis and A. carli. It is distinguished from the former in having a slight dorsal granulation, frontal carina less developed, sternal spines on the male pereonites 2–5 instead of only on pereonite 4, absence of a setose area on the male pereopod 2 carpus, and the different orientation of the basal lobe of the pereopod 7 ischium. It is distinguished from the latter species by the posterior corner of pereopod 1 protruding further backwards, shorter uropodal protopod, presence of tubercles on the male pereopods 2–4 ischium and the larger and differently oriented basal lobe of the male pereopod 7 ischium. Adinda lobata sp. nov. (Figs 26–28) Material examined INDIA: 1 { (Holotype), 1 | (Paratype), Anamalai Hills, Valparai, Naduar Estate, about 1200 m, leg. J. Carl, iii.1927, MHNG. Description Maximum length: { and |, 8 mm. Colour (in alcohol) grey-brownish with posterior part of vertex and epimera lighter; antenna with distal part of the fifth article of peduncle and first article of flagellum pale. Dorsum rough but not granulated, with tiny petaliform scale-spines. Eye with 17–19 ommatidia. Frontal shield of cephalon with central carina obtuse, slightly protruding forwards and upwards; lateral lobes rounded, directed slightly forwards and separated from vertex by a deep groove, provided with a row of scales; supra-antennal line continuous, almost straight. Pereonite 1 with posterior margin slightly sinuous at the base of epimera, posterior corner rounded. Telson about twice as wide as long, broadly rounded. Antennule with third article bearing two

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Figure 26. Adinda lobata sp. nov. A, Adult female, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna.

rows of aesthetascs on medial margin and two aesthetascs at apex. Antenna with fifth article of peduncle reaching posterior margin of pereonite 1; second article of flagellum about 1.5 times as long as first. Buccal pieces as in A. stebbingi, but the penicils of mandibles are 2 + 1 in the right one and 2 + 2 in the left one. Uropodal protopod with the distal part narrower than in all the previous species; stout exopod and endopod, distinctly protruding in relation to protopod tips. Male. Pereopods 1–3 carpus, 1–6 merus and 2–6 ischium with a brush of

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Figure 27. Adinda lobata sp. nov., {. A, Pereopod 1; B, pereopod 2; C, pereopod 6; D, pereopod 7.

REVISION OF TORADJIINAE

Figure 28. Adinda lobata sp. nov., {. A, Pleopod 1 exopod; B, pleopod 1 endopod; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

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small scales on sternal margin. Pereopod 7 ischium distally enlarged to form a rounded lobe, with 2 strong spines, sternal margin straight; merus slightly swollen on sternal margin. Pleopod 1 exopod wider than long, with short and rounded posterior point; endopod straight with rounded apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 28C–F. Etymology The name refers to the male pereopod 7 ischium which is distally enlarged in the form of a round lobe. Distribution Southern India. Remarks Adinda lobata is readily distinguished from the other species by the uropodal protopods distally narrower and exopods stouter, the male pereopod 7 ischium clearly enlarged, and the male pleopod 1 exopod with a short rounded posterior point.

Adinda triangulifera sp. nov. (Fig 29) Material examined INDIA: 1 | (Holotype), 2 || (Paratypes), Anamalai Hills, Attakatti, Shola, Ibex-Hill, 1200 m, leg. J. Carl, 26.ii.1927, MHNG; 1 | (Paratype), same data, MZUF. Description Maximum length: 18 mm. Colour (in alcohol) yellow-brownish; articles 1–3 and distal part of fifth article of antennal peduncle pale. Dorsum strongly granulated and covered by several small petaliform scale-spines. Eye with about 20 ommatidia. Cephalon with frontal shield bearing an acute central carina protruding forwards and upwards; lateral lobes triangular, directed forwards, separated from vertex by a deep groove bordered with scales; supra-antennal line interrupted in the middle. Pereonite 1 with posterior margin slightly sinuous, posterior corner with rounded apex. Telson about twice as wide as long, triangular, with slightly concave sides and rounded apex. Antennule with third article provided with four rows of aesthetascs on medial margin and two aesthetascs at apex. Antenna with fifth article of peduncle reaching posterior margin of pereonite 1; second article of flagellum twice as long as the first. Buccal pieces as in A. stebbingi, except mandibles which have 3+1 penicils on the right and 3 + 2 on the left. Uropodal protopod subquadrangular; exopod and endopod at the same level, not protruding in relation to the posterior margin of protopod.

REVISION OF TORADJIINAE

Figure 29. Adinda triangulifera sp. nov. A, Adult female, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleon and telson; G, pleonite 5, telson and uropods.

Etymology The name refers to the triangular shape of the telson. Distribution Southern India.

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Remarks Even if no male specimens have been examined, this species deserves to be described due to its peculiar telson shape (triangular with concave sides) which immediately distinguishes it from all the other species in the group. In the marked dorsal granulations, A. triangulifera resembles A. stebbingi, A. travancorensis, A. carli and A. scabra; however, it is readily distinguished form those species, besides its telson shape, also by the frontal carina more acute and protruding, and the uropodal protopod subquadrangular. Adinda scabra (Collinge 1916) (Figs 2B, 30–32) Paraperiscyphisscabrus Collinge, 1916: 117, pl. IX figs 6–10. Adindascabrus ; Arcangeli, 1948: 34. necAdinda scabrus ; Jackson, 1936: 84. Material examined SRI LANKA: 2 ||, Kandy, Udawattakele Sanctuary, 2100 ft, D. R. Davis & W. H. Rowe, 10–23.i.1970, USNM; 1 {, 1 |, Kandy, leg. C. Be´suchet & I. Lo¨bl, 14.ii.1970, MHNG; 1 {, same locality and collectors, 22.i.1970, MHNG; 1 {, 2 ||, same locality, forest, under stones, leg. O. Lo¨w-Beer, 21.ii.1912, SMF; 1 {, Haputale, leg. C. Be´suchet & I. Lo¨bl, 23.vii.1970, MHNG; 1 {, Central Prov., Horton Plains, 11 mi SSE of Nuwara-Eliya, 6700 ft, indigenous forest slope, leg. Lund University Ceylon Expedition, 19– 20.iii.1962, ZIUL; 1 |, 1 juv., same data, ZIUL; 1 |, same locality, leg. C. Be´suchet & I. Lo¨bl, 15.ii.1970, MHNG; 1 {, Sabaragamuwa Prov., Deerwood, Kuruwita, 6 mi NNW of Ratnapura, leg. Lund University Ceylon Expedition, 18–21.ii.1962, ZIUL; 1 |, 1 juv., Kuruwita, leg. P. Beron, 1.xii.1984, NNHMS; 1 |, Hatton, leg. C. Be´suchet & I. Lo¨bl, 9.xi.1970, MHNG; 1 {, Western Prov., Yongammulla, 3 mi E Yakkala, 18 mi NE of Colombo, leg. Lund University Ceylon Expedition, 19.i.1962, ZIUL; 3 {{, 1 |, same data, MZUF; 1 {, 2 ||, 5 juvs, same locality, leg. Lund University Ceylon Expedition, 24.i– 6.iii.1962, ZIUL; 1 |, 2 juvs, Peradeniya, leg. C. Be´suchet & I. Lo¨bl, 19.i.1970, MHNG. Previousrecords Sri Lanka: Peradeniya (Collinge, 1916). Description Maximum length: {, 12 mm; |, 18 mm. Colour (in alcohol) grey-brownish with yellowish spots disposed as follows: pereonites with two paramedian spots close to the anterior margin and a median one, an oblique one at the base of epimera; pleonites 3–5 with one spot at the base of epimera; anterior and posterior corners of pereonites and pleonites pale, telson with two paramedian spots; antenna with distal part of articles 3–5 of peduncle pale. Dorsum distinctly granulated, each granulation bearing a petaliform scale-spine. Eye with about 23 ommatidia. Cephalon with frontal shield provided with an obtuse median carina slightly protruding forwards and upwards, and separated from

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Figure 30. Adinda scabra. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna.

vertex by a deep groove bordered by a row of scales; lateral lobes rounded, directed forwards; supra-antennal line interrupted in the middle. Pereonite 1 with posterior margin distinctly sinuous at the base of epimera; posterior corner with rounded apex. Telson about twice as wide as long, broadly rounded. Antennule with third article elongated, bearing some rows of aesthetascs on medial margin and two aesthetascs at apex. Antenna with fifth article of peduncle reaching the posterior margin of pereonite 1; second article of

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Figure 31. Adinda scabra, {. A, Pereopod 1; B, pereopod 5; C, pereopod 6; D, pereopod 7.

flagellum slightly longer than the first. Buccal pieces as in A. stebbingi, except mandibles which bear 4 + 1 penicils on the right and 5 + 2 on the left. Uropodal protopod with narrow distal part, outer margin rounded; exopod and endopod reaching about the same level and protruding in relation to tip of protopod. Male. Pereopods 1–4 carpus and merus with a brush of short scales and some long trifid spines on sternal margin. Pereopods 5–6 basis with a distal lobe on sternal margin; pereopod 6 ischium with a proximal distinct tubercle on sternal margin. Pereopod 7 ischium with a brush of short setae on distal half of sternal margin. Pleopod 1 exopod slightly longer than wide with a posterior triangular point, apically rounded; endopod straight, without particular structures at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 32C–F.

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Figure 32. Adinda scabra, {. A, Pleopod 1 exopod; B, pleopod 1 endopod; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

Distribution Sri Lanka. Remarks This species is essentially characterized by the dorsal surface distinctly granulated, uropodal protopod with a narrow distal part, presence of a distal process on the male pereopods 5 and 6 basis and a tubercle on pereopod 6 merus, presence of a brush of setae in the distal half of the sternal margin of the male pereopod 7 ischium. Jackson (1936) tentatively ascribed some specimens from Borneo (Sabah, from

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Kiau to Tenompak) to A. scabra, but this record certainly refers to another species. Adinda pulchra (Collinge, 1916) (Figs 33, 34) Paraperiscyphispulcher Collinge, 1916: 116, pl. IX figs 1–5. Adindapulcher ; Arcangeli, 1948; 34.

Figure 33. Adinda pulchra. A, Adult female, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna.

REVISION OF TORADJIINAE

Figure 34. Adinda pulchra, {. A, Pereopod 1; B, pereopod 7; C, pleopod 1 exopod; D, pleopod 1 endopod; E, pleopod 2; F, pleopod 3 exopod; G, pleopod 4 exopod; H, pleopod 5 exopod.

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Material examined SRI LANKA: 1 |, Kuruwita, leg. P. Beron, 1.xii.1984, NNHMS; 2 juvs, Central Prov., Ramboda, 7 mi NW of Nuwara-Eliya, leg. Lund University Ceylon Expedition, 4.iii.1962, ZIUL; 1 {, same data, MZUF; 1 {, 1 juv., Kegalla, leg. C. Be´suchet & I. Lo¨bl, 14.i.1970, MHNG. Previousrecords Sri Lanka: Peradeniya (Collinge, 1916). Description Maximum length: {, 12 mm; |, 17 mm. Colour (in alcohol) brownish with several yellowish spots; antenna as in A. scabra. Dorsum rough not granulated, with several small triangular scale-spines. Eye with about 22 ommatidia. Cephalon with frontal shield bent over vertex in the median part, bearing a medial carina slightly protruding forwards; lateral lobes rounded and slightly directed forwards, separated from vertex by a deep groove provided with a row of scales; supra-antennal line visible only at sides. Pereonite 1 with posterior margin distinctly sinuous at the base of epimera; posterior corner acute with rounded apex. Telson broadly rounded, about twice as wide as long. Antennule with third article provided with four rows of aesthetascs on medial margin and two aesthetascs at apex. Antenna with fifth article of peduncle reaching posterior margin of pereonite 1; second article of flagellum about 1.5 times as long as first. Buccal pieces as in A. stebbingi, except mandibles with 4 + 1 penicils on the right and 3 or 4 + 2 on the left. Uropodal protopod with rounded posterior margin; stout exopod and endopod, distinctly protruding in relation to protopod tip. Male. Pereopods 1–3 carpus, merus, ischium and pereopods 4–6 ischium with a brush of small scales on sternal margin. Pereopod 7 ischium with sternal margin almost straight, bearing a brush of long setae in the distal half, merus with a brush of setae on sternal margin. Pleopod 1 exopod slightly wider than long, posterior point rounded; endopod with no particular modifications. Pleopod 2 and pleopods 3–5 exopod as in Fig. 34E–H. Distribution Sri Lanka. Remarks Adinda pulchra is very similar to A. scabra from which it is distinguished by the absence of dorsal granulations, frontal carina less protruding forwards, and absence of distinct sexually dimorphic modifications on the male pereopods 5–6. Riedeli group Adinda riedeli sp. nov. (Figs 35–38) Material examined MALAYSIA: 1 { (Holotype), 2 || (Paratypes), Sarawak, Belaga District, Long Linau, leg. A. Riedel, 17–21.iii.1990, SMNS; 1 |, 3 juvs (Paratypes),

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Figure 35. Adinda riedeli sp. nov. A, Adult female, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule.

Sarawak, Belaga, leg. A Riedel, 14–16.iii.1990, SMNS; 1 { (Paratype), same data, MZUF; 1 | (Paratype), Sarawak, Serian District, Penrissen Road 12 mi from Kuching, Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forestry Department, 50 m, litter at base of big trees, leg. B. Hauser, 8.xii.1987, MHNG; 1 | (Paratype), Sarawak, road Kuching-Matang, Gunung Serapi, 320 m, forest litter along road to the TV station, leg. B. Hauser, 9.xii.1987, MHNG. BRUNEI: 1 {, 1 |, 5 juvs (Paratypes), Brunei-Muara District, road from Tutong, 33 km from Bandar Seri Begawan, near bridge on Sungai Lubang Barus, leg. B. Hauser, 16.xi.1988, MHNG; 1 | (Paratype), Belait District, Andulau Forest Reserve, 3.5 km S Sungai Liang, 39.5 km from Labi, primary forest, leg. B. Hauser, 19.xi.1988, MHNG.

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Figure 36. Adinda riedeli sp. nov. A, Right mandible; B, left mandible; C, maxillule; D, maxilliped.

Description Maximum length: {, 9 mm; |, 11 mm. Very convex body with almost vertical epimera. Colour (in alcohol) grey-brownish with some irregular yellowish spots; antenna with distal part of the fifth article of peduncle and flagellum pale. Dorsum rough and pruinose due to the presence of many long triangular scale-spines. Eye with about 25 ommatidia. Cephalon with frontal shield protruding over vertex, median carina distinctly protruding with a depression in the upper part; lateral lobes triangular, directed forwards; frontal shield separated from vertex by a deep groove, wider at sides. Pereonite 1 with posterior margin slightly sinuous at the base of epimera; posterior corner

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Figure 37. Adinda riedeli sp. nov., {. A, Antenna; B, pereopod 1; C, pereopod 7.

rounded. Telson with posterior margin subtruncated, twice as wide as long. Antennule with third article bearing five rows of aesthetascs on medial margin and two aesthetascs at apex. Stout antenna reaching posterior margin of pereonite 2; second article of flagellum about twice as long as first. Mandibles with molar penicil simple, 1 + 1 penicils on the right and 1 + 2 on the left. Maxillule outer branch with five out of 10 teeth cleft; inner branch with two long penicils. Maxilliped endite setose with penicil. Uropodal protopod subquadrangular; exopod and endopod equally protruding in relation to protopod tip. Male. Anterior pereopods with no particular modifications, only pereopods 1–4 carpus with small scales on sternal margin. Pereopod 7 ischium with straight sternal margin and a relief covered with setae in the distal part of the rostral surface; merus with a distinct conical lobe covered with short setae. Pleopod 1 exopod longer than wide, with a long acute posterior point; endopod stout, with tuft of setae at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 38B–E.

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Figure 38. Adinda riedeli sp. nov., {. A, Pleopod 1; B, pleopod 2; C, pleopod 3 exopod; D, pleopod 4 exopod; E, pleopod 5 exopod.

Etymology The new species is named after A. Riedel, Stuttgart, who collected some of the specimens. Distribution Malaysia (Sarawak). Remarks Adinda riedeli is distinguished from the other species in the group by the presence of long triangular scale-spines on the dorsum, the shape of the telson

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and uropods, and by the presence of a distinct conical lobe on the male pereopod 7 ischium. Adinda platyperaeon (Schultz, 1982) (Figs 39–42) Paraperiscyphisplatyperaeon

Schultz, 1982: 104, figs 3–5.

Typematerial examined MALAYSIA: 1 { (Holotype), Sarawak, Gunung Mulu National Park, Gua Air Jernih (Clearwater Cave), BM 1981 : 332 : 1; 1 | (Allotype), same data, BM 1981 : 333 : 1; 3 {{, 1 | (Paratypes), same data, BM 1981 : 334 : 15. Material examined MALAYSIA: 1 {, 2 ||, Sarawak, Gunung Mulu National Park, Gua Ajaib (Wonder Cave), BM 1981 : 335 : 3; 3 {{, 1 |, Sarawak, Gunung Mulu National Park, Lubang Angin, Lubang Hijau, Lubang Ular (Cave of the Winds, Green Cave, Snake Cave), BM 1981 : 336 : 7; 2 {{, Sarawak, Gunung Mulu National Park, Windsor Cave, Clearwater River Cave, BM 1981:337:2 (specimens identified by Schultz, 1982). Previousrecords Malaysia: Sarawak, Gunung Mulu National Park: Gua Air Jernih, Lubang Angin, Lubang Hijau, Lubang Ular, Windsor Cave, Clearwater River Cave, Gua Ajaib (Schultz, 1982). Description Maximum length: {, 21.5 mm; |, 27 mm. Body enlarged and flattened, convex only in the median part, epimera directed obliquely outwards. Antennae and pereopods frail and long. Colour brown-yellowish, antenna with distal part of the fifth article of peduncle and flagellum pale. Dorsum smooth with small setae. Eye with about 12 ommatidia, about 20 in the specimens from Gua Ajaib. Cephalon with frontal shield bearing a median carina distinctly protruding forwards and upwards; lateral lobes triangular, directed forwards and separated from vertex by a deep groove. Pereonite 1 with posterior margin straight and posterior corner at right angle. Telson about 1.5 times as wide as long, broadly rounded. Antennule with third article bearing five superimposed rows of aesthetascs and two aesthetascs at apex. Antenna reaching the posterior margin of pereonite 5; fifth article of peduncle about twice as long as flagellum; flagellar articles subequal. Buccal pieces as in A. riedeli. Uropodal protopod with outer margin rounded; exopod and endopod slightly protruding in relation to protopod tip. Male. Pereopods 1–4 carpus and merus with lines of long trifid spines and a brush of short scales on sternal margin. Pereopod 7 ischium with sternal margin concave and provided with a distal setose lobe on rostral surface; merus with a brush of setae on basal part of sternal margin and a small setose lobe on rostral surface. Pleopod 1 exopod with a long triangular posterior point

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Figure 39. Adinda platyperaeon, specimens from Gua Air Jernih, Sarawak. A, Adult male, dorsal view; B, adult male, lateral view; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antenna.

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Figure 40. Adinda platyperaeon, { from Gua Air Jernih, Sarawak. A, Antennule; B, pereopod 1; C, pereopod 7; D, pleopod 1 exopod; E, pleopod 1 endopod.

bent outwards; endopod straight without particular modifications at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 41. Distribution Malaysia (Sarawak). Remarks Specimens from Gua Ajaib (Fig. 42) have larger eyes, frontal shield with smaller lateral lobes and carina, shorter uropodal protopod so that exopod and

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Figure 41. Adinda platyperaeon, { from Gua Air Jernih, Sarawak. A, Pleopod 2; B, pleopod 3 exopod; C, pleopod 4 exopod; D, pleopod 5 exopod.

Figure 42. Adinda platyperaeon, specimens from Gua Ajaib, Sarawak. A, Cephalon, frontal view; B, cephalon, dorsal view; C, cephalon, posterior view; D, pleonite 5, telson and uropods.

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endopod distinctly protrude backwards. For all the other characters, including the male modifications, these specimens correspond to the types. Though it cannot be excluded that they belong to a distinct taxon, we agree with Schultz (1982) that these differences are too small to justify specific separation. Adinda platyperaeon is readily distinguished from the other species in the group by the enlarged and flat body, the elongated antennae and pereopods, and the structure of the male pereopod 7. Adinda lamellata sp. nov. (Figs 43, 44) Material examined BRUNEI: 1 { (Holotype), 1 {, 1 { juv., 4 ||, 35 juvs (Paratypes), BruneiMuara District, Bandar Seri Begawan, leg. B. Hauser, 15.xi.1988, MHNG; 1 {, 1 | (Paratypes), same data, MZUF; 2 {{, 2 {{ juvs, 2 || juvs (Paratypes), Brunei-Muara District, road from Tutong, 33 km from Bander Seri Begawan, near bridge on Sungai Lubang Barus, leg. C. Lienhard, 16.xi.1988, MHNG; 1 {, 3 || juvs (Paratypes), Brunei-Maura District, road from Tutong, 33 km from Bandar Seri Begawan, near bridge on Sungai Lubang Barus, leg. B. Hauser, 16.xi.1988, MHNG; 1 {, 2 juvs (Paratypes), Brunei-Muara District, Berakas Forest Reserve, N Bandar Seri Begawan, 19.5 km from Muara, tropical heath forest, leg. C. Lienhard, 20.xi.1988, MHNG. MALAYSIA: 1 { juv., Sarawak, Bau, environs Gua Puang near Kampong Pelaman Sekiang, escarpment of Gunung Jambusan, 10–30 m, leg. C. Lienhard, 4.xii.1987, MHNG. PHILIPPINES: 1 |, 2 juvs, Palawan, S. Lucia, Puerto Princesa, leg. G. Osella, 5.ii.1981, MZUF; 1 |, Palawan, Mantalingajan, Tagerubung, 1150 m, leg. Noona Dan Expedition, 19.ix.1961, ZMC; 1 |, Balabac, Dalawan Bay, in Malaise-traps, leg. Noona Dan Expedition, 6.x.1961, ZMC. Description Maximum length: {, 9 mm; |, 11 mm. Colour (in alcohol) yellowish with many small brown spots; antenna with articles 1–3 of peduncle pale. Dorsum slightly rough with many small triangular scale-spines. Eye with about 24 ommatidia. Cephalon with shield bearing an obtuse median carina protruding upwards but protruding much less forwards in relation to the lateral lobes; frontal shield separated from vertex by a very narrow groove, more apparent at the sides; supra-antennal line interrupted in the middle. Pereonite 1 with posterior margin distinctly sinuous at the base of epimera; posterior corner rounded. Pereonite 2 with a transverse depression on anterior margin of epimera. Telson semicircular, twice as wide as long. Antennule with third article apically pointed, bearing five superimposed rows of aesthetascs. Short antenna reaching rear margin of pereonite 2; fifth article of peduncle about 1.5 times longer than flagellum; second flagellar article about twice as long as first. Buccal pieces as in A. riedeli. Uropod with quadrangular protopod; exopod and endopod protruding in relation to protopod tip. Male. Pereopods 1–4 carpus and merus with lines of long trifid spines and a brush of short scales. Pereopod 7 ischium with sternal margin distinctly

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Figure 43. Adinda lamellata sp. nov. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna.

REVISION OF TORADJIINAE

Figure 44. Adinda lamellata sp. nov., {. A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

415

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concave, distally with a small triangular protrusion, rostral surface distally with a setose process; merus with a large lamellar semicircular protrusion on sternal margin, rostral surface with a large setose lobe. Pleopod 1 exopod with triangular posterior point, bent outwards; endopod with a small hyaline lobe at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 44D–G. Etymology The name refers to the large lamellar protrusion on the sternal margin of the male pereopod 7 merus. Distribution Brunei, Malaysia (Sarawak) and Philippines. Remarks The specimens from Malaysia (Sarawak) and the Philippines correspond quite well in general morphology to the type series from Brunei and they are ascribed to this species. However, since no adult males are present in the material from these two countries, we prefer not to designate those specimens as paratypes. Adinda lamellata is easily distinguished from all the other species in the genus by the structure of the male pereopod 7 and pleopod 1. Dollfusi group Adinda dollfusi (Richardson Searle, 1922) comb. nov. (Figs 45, 46) Toradjia dollfusi Richardson Searle, 1922 : 3, pl. I fig. 2. Typematerial examined INDONESIA: 1 { (‘Type’  Holotype), Java, Nongkodjadjar, 1200 m, USNM; 1 { (‘Cotype’  Paratype), same data, NNM. Previousrecords Indonesia: Java, Nongkodjadjar (Richardson Searle, 1922). Description Maximum length: 9 mm. Body enlarged, moderately convex. Colour (according to Richardson Searle, 1922) brown with yellow spots. Dorsum smooth, pruinose due to the presence of numerous small scale-spines. Eye with about 20 ommatidia. Cephalon with frontal shield bearing an obtuse carina protruding upwards; lateral lobes right-angled, directed forwards; frontal shield separated from vertex by a narrow and shallow groove, provided with a row of short scales. Pereonite 1 with epimera directed outwards; posterior margin straight with a rounded lobe at sides which slides under the anterior margin of pereonite 2. Telson semicircular, slightly wider than long. Antennule with third article long and conical, provided with aesthetascs, Antennae absent in the specimens examined. Mandibles with molar penicils semidichotomized. Maxillule outer branch with all teeth simple; inner branch with two long penicils. Maxilliped with a setose endite and a penicil on medial corner. Pleopods 1–2 exopod

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Figure 45. Adinda dollfusi. A, Holotype, dorsal view; B, cephalon, frontal view; C, cephalon, dorsal view; D, cephalon, posterior view; E, pereonite 1, right side, dorsal view; F, pleonite 5, telson and uropods.

provided with open lungs with a large respiratory surface similar to those in Oniscus. Uropodal protopod subquadrangular; exopod and endopod not protruding in relation to the posterior margin of protopod. Male. Pereopod 7 ischium with sternal margin almost straight with a distal

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Figure 46. Adinda dollfusi, {. A, Pereopod 7, caudal surface; B, pereopod 7, rostral surface; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

setose rounded process. Pleopod 1 exopod with a short rounded posterior point; endopod with no particular modifications. Pleopod 2 exopod with very long triangular posterior point. Pleopods 3–5 exopod as in Fig. 46E–G.

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Distribution Indonesia (Java). Remarks Adinda dollfusi is the only species in the genus that has a rounded lobe at the sides of the posterior margin of the pereonite 1 and Oniscus-type respiratory structures. This species is also readily recognizable by the distal process of the male pereopod 7 ischium. Genus Protoradjia Arcangeli, 1955 Type species: Protoradjia jacobsoni Arcangeli, 1955 by monotypy. Diagnosis Animals with exoantennate ability to roll up into a ball. Cephalon with frontal shield slightly bulbous in the middle and distinctly protruding above vertex, separated from this by a deep groove. Pereonite 1 with a very shallow notch (schisma) at posterior corner, that separates two lobes, of which the outer one protrudes distinctly backwards in relation to the inner one. Pereonite 2 with transverse ridge on ventral surface of epimera. Telson semicircular, distinctly shorter than tips of pleonite 5 and uropodal protopods. Antennule of three articles. Antennal flagellum of two articles with a long apical organ. Mandible with molar penicil simple or semidichotomized; right mandible with 1 + 1 penicils and left one with 2 + 1 penicils. Maxillule outer branch with 4 + 6 (5 cleft) teeth, a small accessory tooth and a stalk; inner branch with two long penicils. Maxilliped with setose endite bearing a penicil. Pereopods with long and plumose bifid or trifid dactylar seta. Pereopod 7 basis with a deep groove bordered with scales on rostral surface. Pleopods 1–2 exopod with open lungs similar to those in Trachelipus. Uropodal protopod flattened; exopod inserted in a notch of the medial margin; endopod distinctly surpassing posterior margin of telson. Remarks Protoradjia is morphologically similar to Adinda, particularly to the species of the weberi group, in having the frontal shield with no carina. It is distinguished from that genus by the presence of a schisma on the pereonite 1 and a transverse ridge on the ventral surface of the epimera of the pereonite 2. The very shallow schisma at the posterior corner of the pereonite 1, with the outer lobe distinctly protruding backwards in relation to the inner one, immediately characterizes this genus. Protoradjia jacobsoni Arcangeli, 1955 (Figs 47, 48) ProtoradjiaJacobsoni

Arcangeli, 1955: 10, pl. I figs 1–4.

Typematerial examined INDONESIA: 1 { (Lectotype), 1 | (Paralectotype), Sumatra, Sinabang, leg. E. Jacobson, iii.1913, NNM.

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Figure 47. Protoradjia jacobsoni. A, Adult female, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, cephalon and pereonites 1–3, lateral view; G, pleonite 5, telson and uropods.

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Figure 48. Protoradjia jacobsoni. A, Antennule; B, antenna; C, pereopod 7, lectotype; D, pleopod 1 exopod, lectotype; E, pleopod 1 endopod, lectotype; F, pleopod 2, lectotype.

Material examined INDONESIA: 1 |, Sumatra, Medan, Bukit Lawang, leg. A. Riedel, x.1990, SMNS. Previousrecords Indonesia: Sumatra: Somang near Nalung; Nalung, Talatang; Sinabang (Arcangeli, 1955). Description Maximum length: |, 16 mm. Very convex body with vertical epimera. Colour grey-brownish with a yellow spot at the base of pereon and pleon epimera; antenna with distal part of fifth article of peduncle and flagellum

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pale. Dorsum smooth with small triangular scale-spines. Eye with about 25 ommatidia. Cephalon with frontal shield bulbous in the median part, slightly bent over vertex, from which it is separated by a deep groove bordered by scales; supra-antennal line straight. Pereonite 1 dorsally with a thin groove in the anterior part, close and parallel to the lateral margin; posterior margin slightly sinuous; outer lobe of schisma protruding in relation to inner one. Pereonites 2–7 with rectangular epimera. Pereonite 2 ventrally with a thin transverse ridge. Telson semicircular, about twice as wide as long. Antennule with third article long, conical, with five rows of superimposed aesthetascs and two subapical aesthetascs. Antenna reaching posterior margin of pereonite 2; second article of flagellum about twice as long as first. Mandibles with molar penicil semidichotomized. Uropod with protopod subquadrangular; endopod slightly protruding in relation to exopod and protopod tip. Male. Pereopod 7 ischium with sternal margin slightly concave. Pleopod 1 exopod with large rounded posterior lobe; endopod with no particular modifications. Pleopod 2 as in Fig. 48F. Distribution Indonesia (Sumatra). Remarks Protoradjia jacobsoni, type species of the genus, is characterized by the presence of a thin but distinct groove near the lateral margin of the pereonite 1 and the male pleopod 1 exopod with a large rounded posterior lobe. Protoradjia paeninsulae sp. nov. (Figs 49–51) Material examined MALAYSIA: 1 { (Holotype), 1 {, 2 ||, 4 juvs (Paratypes), Pahang, Cameron Highlands, Gunung Jasar, forest, 1500–1600 m, leg. S. Taiti & L. Bartolozzi, 29.xi.1987, MZUF; 1 |, 2 juvs (Paratypes), Pahang, Cameron Highlands, leg. A. Riedel, 1–8.iv.1990, SMNS; 3 ||, 1 juv. (Paratypes), Pahang, Cameron Highlands, Gunung Beremban, edge of forest, 1500 m, leg. S. Taiti & L. Bartolozzi, 28.xi.1987, MZUF; 1 | (Paratype), same data, SMNS; 1 |, 1 juv. (Paratypes), same locality, 1700 m, 28.xi.1987, MZUF; 3 {{, 2 ||, 1 juv. (Paratypes), Pahang, Genting Highlands, forest, 1500 m, leg. S. Taiti & L. Bartolozzi, 17.ix.1987, MZUF; 6 {{, 5 ||, 17 juvs (Paratypes), Pahang, Fraser Hill, edge of forest, along the path to waterfall, leg. S. Taiti, 8.xi.1985, MZUF. Description Maximum length: {, 17 mm; |, 13 mm. Colour (in alcohol) grey-brown, with a median yellow spot on pereonites 1–7, two paramedian ones on pereonites 1–6 and pleonites, and one per side at the base of pereon and pleon epimera. Dorsum slightly rough with many small petaliform scale-spines. Eye with about 22 ommatidia. Cephalon with frontal shield bulbous in the middle, protruding above vertex, from which it is separated by a narrow groove; supra-antennal line not visible. Pereonite 1 with posterior margin almost

REVISION OF TORADJIINAE

Figure 49. Protoradjia paeninsulae sp. nov. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pereonites 1, 2, ventral view; G, pleonite 5, telson and uropods; H, antennule; I, antenna.

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Figure 50. Protoradjia paeninsulae sp. nov., {. A, Left mandible; B, right mandible; C, maxillule; D, maxilliped; E, pereopod 1; F, pereopod 7.

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Figure 51. Protoradjia paeninsulae sp. nov., {. A, Pleopod 1 exopod; B, pleopod 1 endopod; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

straight, schisma with outer lobe rounded, protruding backwards in relation to inner one. Pereonite 2 ventrally with a small transverse ridge on epimeron. Telson semicircular, wider than long. Antennule with third article long, conical, bearing four rows of aesthetascs on medial margin and two aesthetascs at apex. Antenna reaching the posterior margin of the pereonite 2; second article of flagellum twice as long as first. Mandibles with molar penicil semidichotomized. Uropodal protopod subquadrangular; exopod and endopod not protruding in relation to posterior margin of protopod. Male. Pereopods 1–4 carpus and merus with a brush of short scales on sternal margin, carpus with lines of long trifid spines. Pereopod 7 ischium with sternal margin slightly concave, with a ridge covered with setae on distal part of rostral surface; merus with setose sternal margin and a setose lobe on rostral surface. Pleopod 1 exopod with a long narrow posterior point bent outwards; endopod with two small lobes at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 51C–F.

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Etymology The name refers to the Malay peninsula where the specimens were collected. Distribution Malaysia. Remarks This species is distinguished from P. jacobsoni by the absence of the groove near the lateral margin of the pereonite 1, the presence of lobes on the pereopod 7 ischium and merus, and the male pleopod 1 exopod with a long narrow posterior point. Protoradjia insularis sp. nov. (Figs 52, 53) Material examined INDONESIA: 1 { (Holotype), 2 juvs (Paratypes), Sumatra, Doerian ( Durian), Pulau Durian, Riau Archipelago, leg. K. W. Dammerman, xi.1923, NNM; 1 | (Paratype), same data, MZUF. SINGAPORE: 2 ||, 1 juv. (Paratypes), Island Country Club, between Lower Peirce Reservoir and Windsor Park Estate, remains of primary forest, leg. B. Hauser, 12.xi.1988, MHNG. Description Maximum length: { and |, 8 mm. Colour (in alcohol) brown-yellowish with antennae, cephalon and epimera pale. Dorsum smooth with tiny scale-spines. Eye with about 24 ommatidia. Cephalon with frontal shield slightly bulbous in the middle, bent over vertex, from which it is separated by a groove visible only at the sides; lateral lobes rounded, not protruding forwards. Pereonite 1 with posterior margin straight; outer lobe of schisma broadly rounded. Pereonites 2 ventrally with a transverse ridge on epimera. Pereonites 2–4 with epimera triangular, apically rounded. Telson semicircular, about twice as wide as long. Antennule with third article long, conical, bearing some rows of aesthetascs on medial margin. Antenna with second flagellar article about twice as long as first. Mandible with molar penicil semidichotomized. Uropod as in previous species. Male. Pereopods 1–4 merus and carpus with a brush of short scales; carpus with lines of long trifid spines. Pereopod 7 ischium with sternal margin concave, with an area of long setae in the basal and distal part; merus with sternal margin covered with long setae in the basal part. Pleopod 1 exopod with posterior point triangular; endopod straight with a small rounded lobe at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 53E–H. Etymology The name refers to the Riau and Singapore islands, where these specimens were collected. Distribution Indonesia (Sumatra) and Singapore. Remarks This species is distinguished from P. jacobsoni by the frontal shield clearly bent over the vertex, the absence of the groove on the lateral margin of the

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Figure 52. Protoradjia insularis sp. nov. A, Adult male, lateral view; B, cephalon, frontal view; C, cephalon, dorsal view; D, cephalon, posterior view; E, pleonite 5, telson and uropods; F, antennule; G, antenna.

pereonite 1, the pereonites 2–4 with triangular instead of quadrangular epimera, the presence of setose areas on the male pereopod 7 ischium and merus, and the male pleopod 1 exopod with the posterior point triangular instead of rounded. It is distinguished from P. paeninsulae by the frontal shield bent over the vertex, the absence of lobes and ridge on the male pereopod 7 ischium and merus, the male pleopod 1 exopod with a posterior point much larger and straight instead of bent outwards.

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Figure 53. Protoradjia insularis sp. nov., {. A, Pereopod 1; B, pereopod 7; C, pleopod 1 exopod; D, pleopod 1 endopod; E, pleopod 2; F, pleopod 3 exopod; G, pleopod 4 exopod; H, pleopod 5 exopod.

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Protoradjia montana sp. nov. (Figs 54, 55) Material examined INDONESIA: 1 { (Holotype), 1 {, 4 ||, 4 juvs (Paratypes), North Sumatra, Kotacane, Gunung Sinabung, 2000 m, leg. A. Riedel, 7–8.x.1990, SMNS; 1 {, 1 | (Paratypes), same data, MZUF. Description Maximum length: {, 8 mm; |, 7 mm. Colour (in alcohol) brownish, lighter in females, with pale spots disposed as follows: a median one on posterior margin of cephalon, pereonites and pleonites 1–4; two paramedian ones and two at the base of epimera of pereonites; two on telson. Dorsum smooth with many small petaliform scale-spines. Eye with about 20 ommatidia. Cephalon with frontal shield bulbous in the middle, separated from vertex by a narrow groove; lateral lobes rounded, not protruding forwards. Pereonite 1 with posterior margin slightly sinuous at sides; outer lobe of schisma rounded. Pereonite 2 with epimera rectangular with rounded corners, ventrally with a thin ridge. Telson semicircular. Antennule with third article long, conical, with four rows of aesthetascs on medial margin. Antenna reaching about middle of pereonite 2; second flagellar article more than twice as long as first. Mandible with molar penicil simple. Uropodal protopod subtriangular with rounded outer margin; exopod and endopod protruding backwards in relation to posterior margin of protopod. Male. Pereopods 1–3 carpus and merus with a brush of short scales and some long trifid spines on sternal margin. Pereopod 7 ischium with sternal margin slightly concave, covered with setae; merus with sternal margin convex and an area of long setae on rostral surface. Pleopod 1 exopod with posterior point short, triangular, broadly rounded apically; endopod with no particular modifications. Pleopod 2 and pleopods 3–5 exopod as in Fig. 55D–G. Etymology The name refers to the site of collection of the specimens, Mt Sinabung, at a height of about 2000 m. Distribution Indonesia (Sumatra). Remarks This species is distinguished from P. jacobsoni by the absence of a groove near the lateral margin of the pereonite 1, the male pereopod 7 ischium and merus with setose areas, and male pleopod 1 exopod with a shorter posterior point. It is distinguished from P. paeninsulae by the absence of lobes on the male pereopod 7 ischium and merus, and male pleopod 1 exopod with posterior point wider. P. montana is morphologically close to P. insularis from which it is recognisable by the frontal shield of cephalon not bent over the vertex, wider male pereopod 7 merus with a larger setose area, and the male pleopod 1 exopod with posterior point shorter and rounded apically.

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Figure 54. Protoradjia montana sp. nov. A, Adult male, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods; G, antennule; H, antenna.

Protoradjia pilosa sp. nov. (Fig. 56) Material examined INDONESIA: 1 | (Holotype), Sumatra, Tandjunggadang, west coast, 1000 m, leg. E. Jacobson, 1926, ZMA. Description Length: 10 mm. Colour (in alcohol) grey-brownish with a pale spot in the middle of pereonites, two paramedian spots and two at the base of pereon

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Figure 55. Protoradjia montana sp. nov., {. A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

and pleon epimera. Dorsum smooth with many long piliform and some triangular or lanceolate scale-spines. Eye with about 20 ommatidia. Cephalon with frontal shield bulbous in the middle, slightly bent over vertex in the median part, separated by a deep and narrow groove; lateral lobes rounded, slightly directed forwards. Pereonite 1 with posterior margin slightly sinuous;

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Figure 56. Protoradjia pilosa sp. nov. A, Holotype, lateral view; B-D, dorsal scale-spines; E, cephalon, frontal view; F, cephalon, dorsal view; G, cephalon, posterior view; H, pleonite 5, telson and uropods; I, antennule.

outer lobe of schisma broadly rounded. Pereonite 2 ventrally with a thin but distinct transverse ridge on epimeron. Telson twice as wide as long, very broadly rounded. Antennule with third article long, conical, bearing four rows of aesthetascs on medial margin. Antenna reaching the posterior margin of pereonite 2; second flagellar article about twice as long as first. Mandibles with

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molar penicil semidichotomized. Uropod with subquadrangular protopod; exopod and endopod slightly protruding in relation to protopod tip. Etymology The name refers to the presence of long piliform scale-spines on the dorsum. Distribution Indonesia (Sumatra). Remarks Even if this species is known only from one female specimen, it is worth a description because it is readily distinguished from all the other species in the genus by the presence of piliform scale-spines on the dorsal body surface. Genus Paratoradjia nov. Typespecies:

Paratoradjia beroni sp. nov.

Diagnosis Animals with exoantennate ability to roll up. Cephalon with frontal shield distinctly protruding above vertex at the sides, separated from this by a deep groove bordered with scales, and bent over vertex and partially fused with it in the median part; a median carina along the whole frontal shield is present. Pereonite 1 with a notch (schisma) on posterior margin, far from the posterior corner; inner and outer lobes of schisma rounded, equally protruding backwards. Pereonite 2 with a transverse ridge on ventral surface of epimera. Telson semicircular or triangular, shorter than tips of pleonite 5 and uropodal protopod. Antennule of three articles. Antennal flagellum of two articles with long apical organ. Pereopods with long dactylar seta, distally plumose. Pereopod 7 basis with groove bordered by a row of scales. Mandibles with molar penicil simple. Maxillule outer branch with 4 + 6 teeth, a small accessory tooth and a stalk; inner branch with two long penicils. Maxilliped with setose endite and penicil. Pleopods 1–2 exopod with open lung similar to that in Oniscus. Uropods with flattened protopod distinctly surpassing apex of telson; exopod inserted in a notch of the medial margin of protopod. Remarks The four species described here, even if clearly differentiated by the male modifications, constitute a very homogeneous group characterized mainly by the cephalic structure. Moreover, Paratoradjia is easily distinguished from Adinda by the presence of a schisma on the pereonite 1 and a ventral lobe on the pereonite 2, and from Protoradjia by the deeper schisma, far from the posterior corner. The new genus is morphologically similar to Toradjia, from which it is distinguished essentially by the different structure of the cephalon: in Paratoradjia the frontal shield is bent over the vertex and partially fused with it in the middle, the frontal margin is continuous and, when the animal rolls up, the antennae simply rest on the vertex; in Toradjia the frontal shield is interrupted in the middle by two deep antennal grooves which continue on the vertex.

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The peculiar cephalic structure immediately distinguishes Paratoradjia from all the other genera in the subfamily. Paratoradjia beroni sp. nov. (Figs 1C, 3A, 57–59) Material examined SRI LANKA: 1 { (Holotype), 3 || (Paratypes), Kuruwita, leg. P. Beron, 1.xii.1984, NNHMS; 1 | (Paratype), same data, MZUF; 1 | (Paratype), Sabaragamuwa Prov., Deerwood, Kuruwita, 6 mi NNW of Ratnapura, under logs, leg. Lund University Ceylon Expedition, 18–21.ii.1962, ZIUL; 1 { (Paratype), same data, ZIUL; 1 { (Paratype), same data, MZUF; 1 | (Paratype), same locality, little gorge, leg. Lund University Ceylon Expedition, 18.ii.1962, ZIUL; 2 ||, 1 juv. (Paratypes), Southern Prov., Haycock Mountain, 21 mi NNE of Galle, 300–450 m, under stones, leg. Lund University Ceylon Expedition, 29.i.1962, ZIUL. Description Maximum length: {, 7 mm; |, 6 mm. Colour (in alcohol) grey-brownish; antennae pale. Dorsum with distinct prominent granulations; dorsal surface with numerous circular fossettae and small petaliform scale-spines. Eye with about 19 ommatidia. Cephalon with frontal shield bent over vertex in the middle, separated from this by a ridge; lateral lobes obliquely directed forwards, separated from vertex by a groove bordered with scales; a protruding carina on profrons, which reaches posterior margin of frontal shield, separates two longitudinal concavities against which the peduncles of antennae lean when the animal rolls up; profile of carina sinuous; supra-antennal line interrupted in the middle. Pereonite 1 with posterior margin almost straight. Telson triangular with slightly concave sides and rounded apex. Antennule with third article stout and short, bearing a tuft of aesthetascs subapically and two aesthetascs apically. Short antenna almost reaching posterior margin of pereonite 1; second flagellar article about twice as long as first, with long apical organ. Right mandible with 1 + 1 penicils; left mandible with 1 + 2 penicils. Maxillule outer branch with some cleft teeth. Uropodal protopod with outer margin regularly rounded; exopod and endopod slightly protruding in relation to protopod tip. Male. Pereopod 7 ischium with sternal margin straight and setose in the basal part, a small lobe covered with short setae distally on rostral surface; merus with a distinct setose triangular lobe on rostral surface. Pleopod 1 exopod cordiform with some spines on outer margin; endopod with enlarged apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 59D–G. Etymology The new species is named after Dr P. Beron, Sofia, who collected some of the specimens. Distribution Sri Lanka. Remarks Paratoradjia beroni is characterized by the distinct dorsal granulation, the telson with triangular apex, the modifications of the male pereopod 7 and the shape of the male pleopod 1.

REVISION OF TORADJIINAE

Figure 57. Paratoradjia beroni sp. nov. A, Adult female, lateral view; B, dorsal scale-spine; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, cephalon, lateral view; G, pereonites 1, 2, dorsal view; H, pereonites 1, 2, ventral view; I, pleonite 5, telson and uropods.

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Figure 58. Paratoradjia beroni sp. nov. A, Antennule; B, antenna; C, left mandible; D, right mandible; E, maxillule; F, maxilliped.

Paratoradjia indosinensis (Arcangeli, 1948) comb. nov. (Fig. 60) Toradjiaindosinensis

Arcangeli, 1948: 30, pls I–IV figs 1–14.

Previousrecords Vietnam: Nha-Trang; Da-Lat; Banov (Arcangeli, 1948). Distribution Vietnam.

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Figure 59. Paratoradjia beroni sp. nov., {: A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

Remarks As already pointed out, the type specimens of this species apparently are no longer present in the collections of the Museo Regionale di Scienze Naturali,

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Figure 60. Paratoradjia indosinensis. A, Cephalon, dorsal view; B, pereonite 1, right side, dorsal view; C, pleonite 5, telson and uropods; D, antennal flagellum; E, left uropod; F, pereopod 7, {; G, pleopod 1 exopod, {; H, pleopod 2 exopod, { [Adapted from Arcangeli, 1948].

Turin. However, the description and illustrations provided by Arcangeli (1948) show that Toradjia indosinensis possesses all the characters of Paratoradjia, particularly the structure of the cephalon and of the pereonite 1, and thus it must be transferred to this genus. P. indosinesis has the largest body size in the genus

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(up to 12 mm long) and is characterized by the slightly tuberculated dorsum, the male modifications of pereopod 7 with the ischium bearing a longitudinal lamellar process on the sternal margin, and the shape of the male pleopod 1 exopod with outer margin deeply incised and a short posterior point apically rounded. The main characters of P. indosinensis, redrawn from Arcangeli’s original illustrations, are shown in Fig. 60. Paratoradjia vietnamensis sp. nov. (Figs 61–63) Material examined VIETNAM: 1 { (Holotype), Cuc Phuong Prov., Ninh-Binh, extracted from soil of Asplenium nidusavis, leg. Topal, 6.v.1966, HNHM. Description Length: 7 mm. Colour (in alcohol) brown-reddish; antennae lighter. Dorsum with distinct granulations on anterior half with many small scale-spines. Eye with 16 ommatidia. Cephalon similar to that of P. beroni, from which it is distinguished by the profile of carina not sinuous in the central part, and the frontal shield which in the median part is bent over the vertex but is not distinctly separated from it. Pereonite 1 with epimeron slightly bent outwards, posterior margin almost straight. Telson semicircular with two dorsal paramedian longitudinal reliefs. Antenna reaching pereonite 2; flagellum with second article about twice as long as first. Buccal pieces not observed due to their poor preservation. Uropodal protopod subquadrangular, with rounded posterior margin; exopod and endopod protruding in relation to posterior margin of protopod. Pereopods 1–5 merus with lines of small triangular spines on sternal margin. Pereopod 6 ischium strongly enlarged in the distal part, sternal margin concave covered with long setae in the distal half, tergal margin with a distinct process bearing a strong spine apically; merus with a subrectangular process and two stout spines on tergal margin. Pereopod 7 ischium with sternal margin almost straight and a distal depression with small triangular spines on rostral surface; merus with a rounded setose lobe on rostral surface. Pleopod 1 exopod with a short triangular posterior point apically rounded; endopod with no particular modifications. Pleopod 2 and pleopods 3–5 exopod as in Fig. 63C–F. Etymology The name refers to Vietnam where the specimen was collected. Distribution Vietnam. Remarks This species is readily distinguished from all the others in the genus by the distinct granulations on the anterior part of the body, the two paramedian longitudinal reliefs on the telson, and the peculiar features of the male pereopod 6.

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Figure 61. Paratoradjia vietnamensis sp. nov. A, Holotype, lateral view; B, cephalon, lateral view; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pereonites 1, 2, dorsal view; G, pereonites 1, 2, ventral view; H, pleonite 5, telson and uropods; I, antenna.

REVISION OF TORADJIINAE

Figure 62. Paratoradjia vietnamensis sp. nov., {. A, Pereopod 1; B, pereopod 2; C, pereopod 6; D, pereopod 7.

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Figure 63. Paratoradjia vietnamensis sp. nov., {. A, Pleopod 1 exopod; b, pleopod 1 endopod; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

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Paratoradjia sulcata sp. nov. (Fig. 64) Material examined PAPUA NEW GUINEA: 1 | (Holotype), Bismarck Islands, New Ireland, Lelet Plateau, soil, leg. R. Enery, viii.1979, MHNG. Description Length: 7 mm. Colour (in alcohol) light brown. Dorsum smooth, pruinose due to the presence of small scale-spines. Eye with about 20 ommatidia. Cephalon with frontal shield bent over vertex in the median part, separated from this by a distinct ridge; lateral lobes obliquely directed forwards, separated from vertex by a groove bordered with short scales; median carina obtuse, slightly protruding. Pereonite 1 with a narrow depression parallel to the lateral margin; posterior margin straight. Telson semicircular, about twice as wide as long. Antennule as in P. beroni. Antenna reaching posterior margin of pereonite 1; second flagellar article twice as long as first. Buccal pieces, dactylar seta of the pereopods and respiratory structures as in P. beroni. Uropodal protopod with posterior margin oblique; exopod and endopod protruding in relation to protopod tip. Etymology The name refers to the depression parallel to the lateral margin of the pereonite 1. Distribution New Guinea. Remarks Even if only a female specimen has been examined, this species is worthy of description because it is clearly distinguishable from all the other species and considerably enlarges the distributional range of the genus. The new species is easily distinguished by the smooth dorsum, less developed frontal carina and mainly by the presence of a distinct depression close the lateral margin of the pereonite 1. Genus Toradjia Dollfus, 1898 Type species: Toradjia celebensis Dollfus, 1898, by original designation. Diagnosis Animals with exoantennate and complete ability to roll up into a ball. Cephalon with frontal shield protruding above vertex at the sides, separated from this by a deep groove; frontal shield interrupted in the middle by two deep antennal grooves, separated by a narrow carina; these grooves, parallel in the frontal region, continue obliquely on the vertex and transversely on the anterior part of pereonite 1; when the animals roll up, the antenna fits into these grooves. Pereonite 1, in addition to the above-mentioned grooves, dorsally with a transverse ridge (not visible in T. hirsuta); a schisma on posterior margin

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Figure 64. Paratoradjia sulcata sp. nov. A, Holotype, lateral view; B, cephalon, lateral view; C, cephalon, frontal view; D, cephalon, dorsal view; E, cephalon, posterior view; F, pleonite 5, telson and uropods.

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similar to that present in Paratoradjia, i.e. far from the posterior corner. Pereonite 2 ventrally with triangular tooth or a transverse lobe more or less developed. Telson semicircular. Antennule of three articles. Antenna with flagellum of two articles and long apical organ. Mandibles with molar penicil simple or dichotomized. Maxillule outer branch with 4 + 6 teeth, a small additional tooth and a stalk; inner branch with two long penicils. Maxilliped with setose endite and a penicil on medial corner. Pereopods with long dactylar seta, bifid and setose in the distal part. Pereopod 7 basis with a groove bordered with scales. Pleopods 1 and 2 exopod with open lungs as in Paratoradjia (Oniscus-like). Uropodal protopod with posterior margin rounded, exopod inserted medially in a notch of the medial margin of protopod. Remarks Toradjia is characterized by the presence of antennal grooves on the frontal region, vertex and pereonite 1. Within the Toradjiinae, this genus is the most specialized for rolling up and for antennal protection. Toradjia gorgona Dollfus, 1898 (Figs 65–67) ToradjiaGorgona 1955: 13.

Dollfus, 1898: 366, fig. 11a–c, tab. XIV fig. 11; Arcangeli,

Typematerial examined INDONESIA: 1 { (Lectotype), uncertain locality, MNHN; 1 | (Paralectotype), uncertain locality, leg. M. Weber, 1889, ZMA. Material examined INDONESIA: 1 |, Java, ‘Tjibodas’ ( Cibodas), leg. Kraepelin (specimen identified by Budde-Lund), ZMH; 1 |, Java, Cibodas, Lithocarpus-Castanopsis forest above Botanical Garden, tourist path to the waterfall, 1300–1380 m, leg. C. Lienhard, 26.xi.1987, MHNG. Previousrecords Indonesia: locality ? (Dollfus, 1898); Java (Arcangeli, 1955). Description Maximum length: |, 7 mm. Very convex body with vertical epimera. Colour grey-brownish. Dorsum of the anterior part of the body rough with large granules. Eye with about 11 ommatidia. Cephalon with thick lateral lobes slightly protruding above vertex and separated from it by a deep groove; profrons with a narrow lamelliform carina that continues on vertex. Pereonite 1 with epimeron slightly directed outwards and posterior margin straight; inner and outer lobes of schisma rounded, equally protruding backwards. Pereonite 2 ventrally with a transverse rounded lobe on epimeron. Pereonites 3 and 4 ventrally with a transverse ridge on epimeron. Mandibles with molar penicil simple; right mandible with 1 + 1 penicils; left mandible with 1 + 2 penicils.

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Figure 65. Toradjia gorgona. A, Adult female, lateral view; B, cephalon, frontal view; C, cephalon and pereonite 1, fronto-dorsal view; D, cephalon and pereonite 1, dorsal view; E, cephalon and pereonites 1, 2, lateral view.

Maxillule outer branch with all teeth simple. Uropodal protopod subquadrangular with rounded posterior margin; exopod slightly protruding backwards. Male. Pereopods 5 and 6 with carpus and dactylus swollen. Pereopod 7 ischium with sternal margin straight and distally with a triangular setose process on caudal surface; carpus and dactylus distinctly swollen. Pleopod 1 exopod with a short triangular posterior point, narrowly rounded at apex; endopod with thin distal part, slightly enlarged at apex. Pleopod 2 and pleopods 3–5 exopod as in Fig. 67D–G.

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Figure 66. Toradjia gorgona. A, Pereonites 1–3, ventral view; B, pleonite 5, telson and uropods; C, right mandible; D, left mandible; E, maxillule; F, maxilliped.

Distribution Indonesia (Java). Remarks Dollfus described this species on the basis of two specimens from an unknown locality without indicating the sex or designating the holotype. The male specimen deposited in MNHN is designated here as lectotype of T. gorgona. Arcangeli (1955: 13) stated that he examined material of this species from Java, with no information about the number, sex or collection sites of the specimens,

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Figure 67. Toradjia gorgona, {. A, Pereopod 1; B, pereopod 7; C, pleopod 1; D, pleopod 2; E, pleopod 3 exopod; F, pleopod 4 exopod; G, pleopod 5 exopod.

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and that probably also the specimens identified by Dollfus were collected on that island. Our records confirm this assumption.

Toradjia celebensis Dollfus, 1898 (Fig. 68) Toradjiacelebensis

Dollfus, 1898: 365, fig. 10a–c, pl. XIV fig. 10.

Typematerial examined INDONESIA: 1 | (Lectotype), Celebes (Sulawesi), Soputan, 1200 m, leg. F. and P. Sarasin, MNHN; 1 | (Paralectotype), same data, humus, NHMB. Previousrecords Indonesia: Sulawesi, Soputan (Dollfus, 1898). Description Maximum length: 5 mm. Very convex body with vertical epimera. Colour (according to Dollfus, 1898) grey with small pale stripes; telson and uropods red. Dorsum smooth except pereonite 1 which has some granulations. Eye with about 20 ommatidia. Cephalon and pereonite 1 similar to those of T. gorgona, except the lateral margin of pereonite 1 that is distinctly directed outwards. Pereonite 2 ventrally with an oblique triangular tooth on epimeron. Telson semicircular, about twice as wide as long. Buccal pieces as in T. gorgona, except molar penicil of mandible that is dichotomized. Uropodal protopod with rounded posterior margin; exopod and endopod equally protruding backwards in relation to protopod tip. Remarks Toradjia celebensis is easily distinguished from all the other species by the lateral margin of the pereonite 1 directed outwards and the presence of a ventral tooth on the pereonite 2. The specimen deposited in the MNHN is designated here as lectotype of this species.

Toradjia cephalica Dollfus, 1898 (Figs 69–71) Toradjia cephalica Dollfus, 1898: 367, fig. 12a, pl. XIV fig. 12. Material examined INDONESIA: 1 { (Neotype), 1 {, 1 juv., Java, Cibodas, Lithocarpus-Castanopsis forest above botanical Garden, along a gorge, 1350–1480 m, leg. B. Hauser, 26.xi.1987, MHNG; 1 {, 1 |, same data, MZUF.

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Figure 68. Toradjia celebensis. A, Cephalon and pereonite 1, dorsal view; B, cephalon and pereonite 1, frontal view; C, cephalon and pereonites 1, 2, lateral view; D, pereonites 1–3, ventral view; E, telson and uropods; F, right mandible; G, left mandible.

Previousrecords Indonesia: Java, Tjibodas (Cibodas) (Dollfus, 1898). Description Maximum length: {, 5.2 mm. Very convex body with vertical epimera. Grey-brownish, some irregular pale spots on pereonites 1–3; telson and uropods

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Figure 69. Toradjia cephalica. A, Adult male, lateral view; B, cephalon and pereonite 1, frontal view; C, cephalon and pereonite 1, fronto-dorsal view; D, cephalon and pereonite 1, dorsal view; E, cephalon and pereonites 1, 2, lateral view.

pale (red in vivo ?). Dorsum slightly rough, equipped with small scale-spines. Eye with 15 ommatidia. Cephalon similar to the preceding species. Pereonite 1 dorsally with a transverse arched ridge and anteriorly with distinct antennal grooves; lateral part of epimeron slightly bent outwards; inner and outer lobes of schisma rounded, equally protruding backwards. Pereonite 2 ventrally with a reduced rounded lobe. Telson semicircular. Antennule with third article bearing a subapical row of five aesthetascs and two aesthetascs at apex. Antenna with second flagellar article almost three times as long as first; apical organ

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Figure 70. Toradjia cephalica, {. A, Pereonites 1–3, ventral view; B, pleonite 5, telson and uropods; C, antennule; D, antenna; E, pereopod 1; F, pereopod 7.

shorter than second article of flagellum. Buccal pieces as in T. gorgona. Uropodal protopod quadrangular; endopod distinctly surpassing exopod and protopod tip. Male. Pereopod 7 ischium with sternal margin straight and distally with a triangular setose process on caudal surface. Pleopod 1 exopod with very short

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Figure 71. Toradjia cephalica, {. A, Pleopod 1 exopod; B, pleopod 1 endopod; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

rounded posterior point; endopod with straight narrow distal part. Pleopod 2 and pleopods 3–5 exopod as in Fig. 71C–F. Distribution Indonesia (Java). Remarks The specimens examined correspond well to Dollfus’ description of T. cephalica except for the pereonite 2 which has a distinct, even if reduced, ventral lobe on the epimeron. According to Dollfus the pereonite 2 ‘‘paraissant de´pourvu de duplicature coxale’’ but the French author probably overlooked this character. Moreover also the collection locality (Cibodas) of the material examined is the same as that of the single type specimen of T. cephalica, which, as stated in

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the Introduction, is no longer present in the collections of ZMA and must be considered lost. All the species in Toradjia are morphologically very similar to each other and the description of T. cephalica does not permit a safe recognition of the species. Thus in order to ensure taxonomic stability and material for future comparisons, we designate one male specimen deposited in the MHNG as the neotype of Toradjia cephalica, since all the conditions for designation of a neotype specified in Art. 75 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1985) are satisfied. This species is morphologically very similar to T. gorgona, from which it differs in the absence of dorsal granules, the posterior margin of uropodal protopod subtruncated instead of rounded, and the male pleopod 1 exopod with shorter posterior point. Moreover the position of the triangular process on the male pereopod 7 ischium is different in the two species: in T. gorgona it is close to the tergal margin while in T. cephalica it is close to the sternal one.

Toradjia hirsuta sp. nov. (Figs 72, 73) Material examined MALAYSIA: 1 | (Holotype), Sarawak, Serian District, Penrissen Road, 12 mi from Kuching, Semongok Wildlife Rehabilitation Centre, Nursery Centre of the Forestry Department, 20–50 m, leg. B. Hauser, 8.xii.1987, MHNG. Description Length: 3 mm. Colour grey-brownish. Dorsum with long piliform scale-spines. Eye with 12 ommatidia. Cephalon similar to that of the other species in the genus. Pereonite 1 with no transverse ridge and distinct antennal grooves on anterior part, only a wide depression parallel to the anterior margin is present; posterior margin straight; lateral margin slightly directed outwards; inner lobe of schisma rounded, protruding backwards in relation to outer one. Pereonite 2 ventrally with a rounded lobe on epimeron. Telson semicircular. Antenna with second article of flagellum twice as long as first; apical organ as long as second flagellar article. Buccal pieces as in Toradjia gorgona except outer lobe of maxillule which has five out of 10 teeth apically cleft. Uropodal protopod with posterior margin rounded; endopod and exopod slightly surpassing protopod tip. Etymology The name refers to the characteristic piliform scale-spines on the dorsum. Remarks Even if only a female specimen has been studied, it deserves to be described because it represents a distinct and well characterized species. T. hirsuta differs from all the other species by the presence of piliform scale-spines, absence of a distinct antennal groove on the pereonite 1, reduction of the outer lobe of the schisma, and outer lobe of the maxillule with some apically cleft teeth.

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Figure 72. Toradjia hirsuta sp. nov. A, Holotype, lateral view; B, cephalon and pereonite 1, frontal view; C, cephalon and pereonite 1, fronto-dorsal view; D, cephalon and pereonite 1, dorsal view; E, cephalon and pereonites 1, 2, lateral view.

KEY TO GENERA AND SPECIES1

1. — 2. — 3. 1

Posterior corner of pereonite 1 entire (Fig. 5A) Adinda 2 Posterior corner of pereonite 1 notched (Figs 47F, 58G, 66E) 17 Cephalon with no frontal carina (Fig. 5B,C) 3 Cephalon with frontal carina (Figs 13C, D, 35C, D, 45B, C) 5 Male pereopods 3, 4 ischium with conspicuous tubercles on sternal margin (Fig. 11B) A. sumatrana sp. nov.

Adinda conglobator and A. gigas are not included in this key due to their poor descriptions.

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Figure 73. Toradjia hirsuta sp. nov. A, Pereonites 1–3, ventral view; B, pleonite 5, telson and uropods; C, antenna; D, maxillule.

— Male pereopods 3, 4 ischium with no tubercles on sternal margin 4 4. Male pereopod 7 ischium with concave sternal margin, merus with a protruding lobe at the base bearing short setae (Fig. 7B); male pleopod 1 exopod longer than wide, with a long triangular posterior point (Fig. 7C) A. weberi — Male pereopod 7 ischium with straight sternal margin, merus with basal lobe bearing long setae (Fig. 9B); pleopod 1 exopod wider than long, with a short triangular posterior point (Fig. 9C) A. malaccensis sp. nov. 5. Triangular telson with slightly concave sides (Fig. 29G) A. triangulifera sp. nov. — Semicircular telson 6 6. Pereonite 1 with a lobe at the sides on posterior margin (Fig. 45E); male pereopod 7 ischium distally with a distinct process (Fig. 46A) A. dollfusi — Pereonite 1 with no lobes on posterior margin; male pereopod 7 ischium with no distal process 7 7. Posterior margin of pereonite 1 with a deep incision at the base of epimeron; posterior corner produced backwards in an exceptional manner (Fig. 17B, C) A. travancorensis — Posterior margin of pereonite 1 more or less concave but never incised 8

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8. Dorsum pruinose due to long and triangular scale-spines; male pereopod 7 merus with a distinct conical lobe (Fig. 37C) A. riedeli sp. nov. — Dorsum with inconspicuous scale-spines; male pereopod 7 merus with no conical lobe 9 9. Dorsum more or less distinctly granulated (Fig. 13A) 10 — Smooth dorsum 13 10. One or more pereonites with two paramedian points on male sternites (Fig. 13F) 11 — Male sternites of pereonites with no points A. scabra 11. Male pereopods 2–4 ischium with distinct tubercles on sternal margin (Fig. 15B–D) 12 — Male pereopods 2–4 ischium with no tubercles on sternal margin A. carli sp. nov. 12. Male pereopods 1–4 merus and carpus with distinct setose areas on rostral surface (Fig. 15A–D); pereopod 7 ischium with no lobe at the base (Fig. 16A) A. stebbingi — Male pereopods 1–4 merus and carpus with no setose areas on rostral surface (Fig. 24A–D); pereopod 7 ischium with a protruding lobe at the base (Fig. 25A) A. nilgiriensis sp. nov. 13. Male pereopod 7 merus with a large lamellar semicircular protrusion on sternal margin (Fig. 44B) A. lamellata sp. nov. — Male pereopod 7 merus with no protrusion on sternal margin 14 14. Male pereopods 2–4 ischium with tubercles on sternal margin (Fig. 19B–D); male pereopod 7 ischium with a small lobe at base (Fig. 20A) A. palniensis sp. nov. — Male pereopods 2–4 ischium with no tubercles on sternal margin; male pereopod 7 ischium with no lobe 15 15. Body relatively flattened; antenna reaching the posterior margin of pereonite 5; lateral lobes of cephalon more protruding forwards than carina; male pleopod 1 exopod with long and narrow posterior point bent outwards (Fig. 40D) A. platyperaeon — Body convex; antenna at most surpassing the posterior margin of the pereonite 2; lateral lobes of cephalon less protruding than carina; male pleopod 1 exopod with short posterior point 16 16. Male pereopod 7 ischium distally enlarged to form a rounded lobe (Fig. 27D) A. lobata sp. nov. — Male pereopod 7 ischium distally not particularly enlarged (Fig. 34B) A. pulchra 17. Cephalon with no frontal carina (Fig. 47C, D) Protoradjia 18 — Cephalon with distinct frontal carina (Figs 58C–F, 66B–E) 22 18. Dorsum covered with long piliform scale-spines (Fig. 56A) P. pilosa sp. nov. — Dorsum with inconspicuous scale-spines 19 19. Lateral margin of pereonite 1 anteriorly with a thin but distinct groove (Fig. 47F); male pereopod 7 with no particular modifications (Fig. 48C) P. jacobsoni — Lateral margin of pereonite 1 with no groove; male pereopod 7 with distinct modifications 20

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20. Male pereopod 7 ischium and merus with distinct lobes on rostral surface (Fig. 50F) P. paeninsulae sp. nov. — Male pereopod 7 ischium and merus with no lobes on rostral surface 21 21. Frontal shield bent over vertex in the median part (Fig. 52C); pleopod 1 exopod with triangular posterior point, apically acute (Fig. 53C) P. insularis sp. nov. — Frontal shield not bent over vertex (Fig. 54D); pleopod 1 exopod with posterior point apically rounded (Fig. 55C) P. montana sp. nov. 22. Frontal shield bent over vertex and fused with it in the median part, with two paramedian concavities (Fig. 58D, E) Paratoradjia gen. nov. 23 — Frontal shield interrupted in the middle by deep antennal grooves which continue on the vertex and, more or less developed, on the anterior part of pereonite 1 (Fig. 66A–E) Toradjia 26 23. Smooth dorsum; pereonite 1 with narrow depression parallel to the lateral margin (Fig. 65A) P. sulcata sp. nov. — More or less granulated dorsum; pereonite without lateral depression 24 24. Triangular telson with concave sides (Fig. 58I) P. beroni sp. nov. — Semicircular telson 25 25. Telson with two paramedian longitudinal reliefs (Fig. 62H); male pereopod 6 ischium strongly modified (Fig. 63C); male pereopod 7 ischium with no process on sternal margin P. vietnamensis sp. nov. — Telson with no paramedian reliefs (Fig. 61C); male pereopod 6 ischium without modifications; male pereopod 7 ischium with a distinct longitudinal lamellar process on sternal margin (Fig. 61F) P. indosinensis comb. nov. 6. Dorsum covered with long piliform scale-spines (Fig. 73A) T. hirsuta sp. nov. — Dorsum without piliform scale-spines 27 27. Epimera of pereonite 2 ventrally with tooth-like process (Fig. 69D) T. celebensis — Epimera of pereonite 2 ventrally with transverse lobes (Figs 67A, 71A) 28 28. Anterior part of body dorsally with large granules; male pleopod 1 exopod with distinct triangular posterior point (Fig. 68C) T. gorgona — Dorsum with no granules; male pleopod 1 exopod with short rounded posterior point (Fig. 72A) T. cephalica ACKNOWLEDGEMENTS

The authors wish to thank the following persons and institutions for the loan of material: Ms J. Ellis (BM), Dr L. Forro` (HNHM), Dr D. Defaye and Dr D. Guinot (MNHN), Mrs C. Stocker (NHMB), Dr P. Beron (NNHMS), Prof. L. B. Holthuis and Dr C. H. M. Fransen (NNM), Dr M. Tu¨rkay and Mr A. Allspach (SMF), Dr H. Schmalfuss (SMNS), Dr B. Kensley (USNM), Prof. P.

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Brinck (ZIUL), Mr D. Platvoet (ZMA), Dr J. Just (ZMC), and Prof. G. Hartmann (ZMH). Dr B. Hauser, Geneva (MHNG), is particularly acknowledged also for the facilities provided during two short visits to his institution by one of the authors (F. Ferrara). Many thanks are due to Prof. H. S. Yong (Kuala Lumpur) for his kind help and hospitality during S. Taiti’s visits to Malaysia (1985, 1987). REFERENCES Arcangeli A. 1927. Paraperiscyphis Calegarii Arc. nuova specie di Isopodo terrestre di Sumatra. Bollettino del Laboratorio di Zoologia Generale e Agraria della R. Scuola Superiore di Agricoltura di Portici 20: 79–82. Arcangeli A. 1948. Una specie nuova del genere Toradjia Dollf. (Crostacei Isopodi terrestri). Bollettino del Museo Zoologico dell’Universita` di Torino 1 (1943–48): 29–36, pls I–IV. Arcangeli A. 1952. Le caratteristiche della famiglia Eubelidae. Crostacei Isopodi terrestri. Sue sottofamiglie e suoi generi. Bollettino dell’Istituto e Museo Zoologico dell’Universita` di Torino 3 (1951–52): 61–80. Arcangeli A. 1955. Contributo alle conoscenze sugli Eubelidi di Asia (Crostacei Isopodi terrestri). Bollettino dell’Istituto e Museo Zoologico dell’Universita` di Torino 4 (1953–54): 9–17, pl. I. Budde-Lund G. 1902. A list of Terrestrial Isopods. In: Lanchester WF, ed. On the Crustacea collected during the ‘‘Skeat Expedition’’ to the Malay Peninsula. Proceedings of the Zoological Society of London: 379– 381. Budde-Lund G. 1904. A revision of ‘‘Crustacea Isopoda terrestria’’ with additions and illustrations. 2 Spherilloninae. 3 Armadillo. Kjo benhavn: H. Hagerup, pp. 33–144, pls VI–X. Collinge WE. 1914. On some new terrestrial Isopods from the Andaman Islands and southern India. Records of the Indian Museum 10: 207–210, pls XXIV–XXV. Collinge WE. 1915. Contributions to a knowledge of the Terrestrial Isopoda of India, Pt. I. Records of the Indian Museum 11: 143–151, pls IV–XII. Collinge WE. 1916. Contributions to a knowledge of the Terrestrial Isopoda of India, Pt. II. Records of the Indian Museum 12: 115–128, pls IX–XIX. Dollfus A. 1898. Isopodes terrestres des Indes Ne´erlandaises recueillis par M. le Professeur Max Weber et par M. M. le Docteurs Fritz et Paul Sarasin (de Baˆle). In: Weber W, ed. Zoologische Ergebnisse einer Reise in Niederla¨ndisch Ost-Indien, Leiden 4: 357–382, pls XIII–XV. Ferrara F, Schmalfuss H. 1976. Terrestrial Isopods from West Africa. Part 1: Family ‘‘Eubelidae’’ Budde-Lund, 1899. Monitore Zoologico Italiano (N.S.) Supplemento 7: 1–114. Hoese B. 1981. Morphologie und Funktion des Wasserleitungssystems der terrestrischen Isopoden (Crustacea, Isopoda, Oniscoidea). Zoomorphology 98: 135–167. Hoese B. 1982. Der Ligia-Typ des Wasserleitungssystems bei terrestrischen Isopoden und seine Entwicklung in der Familie Ligiidae (Crustacea, Isopoda, Oniscoidea). Zoologische Jahrbu¨cher. Abteilung fu¨r Anatomie und Ontogenie der Tiere 108: 225–261. International Commission on Zoological Nomenclature. 1985. International Code of Zoological Nomenclature. Third Edition adopted by the XX General Assembly of the International Union of Biological Sciences. (Ed. International Trust for Zoological Nomenclature.) Jackson HG. 1936. Terrestrial Isopods from Malaysia. Bulletin of the Raffles Museum, Singapore, Straits Settlements 12: 77–87. Omer-Cooper J. 1926. A revision of the genus Periscyphis Gerst. (Isopoda Terrestria). Proceedings of the Zoological Society of London 24: 349–400. Richardson Searle H. 1922. Terrestrial Isopoda collected in Java by Dr. Edward Jacobson with descriptions of five new species. Proceedings of the United States National Museum 60: 1–7, pls 1–2. Schmalfuss H. 1980. Die ersten Landasseln aus Dominikanischem Bernstein mit einer systematischphylogenetischen Revision der Familie Sphaeroniscidae (Stuttgarter Bernsteinsammlung: Crustacea, Isopoda, Oniscoidea). Stuttgarter Beitra¨ge zur Naturkunde. Serie B (Geologie und Pala¨ontologie) 61: 1–12. Schmalfuss H. 1986. Die Landisopoden (Oniscidea) Griechenlands. 8. Beitrag: Gattung Kefalloniscus nov. gen. (Scleropactidae). Revue Suisse de Zoologie 93: 279–289. Schmalfuss H. 1989. Phylogenetics in Oniscidea. Monitore Zoologico Italiano (N.S.) Monografia 4: 3–27. Schultz GA. 1982. Terrestrial isopod crustaceans (Oniscoidea) from Mulu Caves, Sarawak, Borneo. Journal of Natural History 16: 101–117. Stebbing TRR. 1911. Indian Isopods. Records of the Indian Museum 6: 179–191. Taiti S, Ferrara F, Schmalfuss H. 1991. Evolution and biogeography of the family Eubelidae (Crustacea, Oniscidea). In: Juchault P, Mocquard JP, eds. The Biology of terrestrial Isopods. III. Proceedings of the Third International Symposium on the Biology of Terrestrial Isopods, Poitiers (France), July 10–12, 1990. Poitiers: Universite´ de Poitiers, 23–30. Vandel A. 1968. I. Isopodes terrestres. In: Mission zoologique belge aux ˆıles Galapagos et en Ecuador (N. et J. 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