Contributions to Zoology, 68 (2) 109-141 (1999)

SPB Academic Publishing bv, The Hague

Philosciids with pleopodal lungs from , with description of a new

species (Crustacea, )

Paula+Beatriz Araujo¹ & Andreas Leistikow²

1 de Universidade Federal do Rio Grande do Av. Paulo Gama, Departamento Zoologia, IB, Sul, pr. 12105,

2 CEP 90040-060, Porto Alegre, RS, Brazil, e-mail: [email protected]; Universitdt Bielefeld, Ab-

teilung fur Zoomorphologie und Systematik, Morgenbreede 45, D-33615, Bielefeld, e-mail: leiste@ biologie. uni-bielefeld. de

Keywords : Crustacea, , Isopoda, philosciids, pleopodal lungs, biogeography, Brazil

Abstract Introduction

Several from in species of “philosciid” Oniscidea areknown Brazil, South America is especially rich “philosciid” most of them were found in the southern and eastern parts of Oniscidea, although our knowledge is far from being this The Ferrara & Taiti, 1981, country. genera complete.' Several authors contributed to our knowl- Benthana Budde-Lund, 1908 and Balloniscus Budde-Lund, 1908, in the last but most of the older de- the latter Balloniscidae edge century, considered to represent a separate family

the scriptions do not support phylogenetic evidence. Vandel, 1963, are considered only neotropical philosciids

their bearing respiratory areas on pleopods. Therefore, represen- Budde-Lund (1908) was the first to discriminate

tatives of these are re-examined to shed new light on genera subgenera among the species described as belonging the question whether these species can be considered to be a to Latreille, 1804. His subgenera are now-

monophylum with the autapomorphy “respiratory areas pres- elevated to generic rank. Lemos de Castro ent”, ofthe above-mentioned is adays The phytogeny genera discussed to under contributed our understanding of the morphological and biogeographical aspects. Furthermore, (1958a)

a remarkable and limits of Benthana new species ofAtlantosciaFerrara & Taiti, 1981, specific generic (Budde- which bears pleopodal lungs in all five pairs of pleopods, is Lund, 1908), giving additional information to described. In to this it differs from its sister regard character, of this Gruner’s (1955) revision genus, although species (Van Name, 1940). Additionally, the are rather incomplete. In a later the descriptions hitherto unknown male of Benthana convexa Lemos de work, he clarified the status of Balloniscus sellowii Castro, 1958 will be described; it is evident that this species is different with its from its closest relative B. schubarti Lemos de Castro, (Brandt, 1833) puzzling synonymy (Lemos

1958, which is also found in Brazil, de Castro 1958b, 1976). Balloniscus Budde-Lund,

1885 was separated in its own family by Vandel

that of (1963) mainly on the fact the species this

Contents in all five of genus bear pleopodal lungs pairs

pleopods. Since the occurrence of pleopodal lungs

Abstract 109 in members of the so-called “”, par- Introduction 109 ticularly in the Afro-tropical genus Aphiloscia Systematics 110 is Genus Budde-Lund, 1908, nowadays recognized (Fer- Atlantoscia Ferrara & Taiti, 1981 110

Atlantoscia 110 rara et al. the of one or two genera rubromarginata n. sp. 1994), separation Atlantoscia floridana (Van Name, 1940) 117 in a family is rather unsatisfactory and even use- Genus Balloniscus 1885 123 Budde-Lund, less for reconstruction of phylogeny. Balloniscus sellowii 123 (Brandt, 1833) both Benthana and Balloniscus are typical gen- Genus Benthana Budde-Lund, 1908 131 era for the coastal Atlantic forests of Brazil where Benthana convexa Lemos de Castro, 1958 132

occur with Atlantoscia floridana Phylogenetic relationships 139 they together (Van

Acknowledgements 140 Name, 1940). Four species of these genera are de-

References 140 scribed in detail in the following, and their sys-

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tematic position will be discussed with regards to nus are the shape ofthe dactylar seta, several sen-

and of and with small sub- phylogenetic biogeographical aspects. sory spines carpus merus

The following acronyms are used: MNRJ Museu apical tip (e.g., Fig. 8, Sc4), reduction of the tip

Nacional, Rio de Janeiro, Brazil; SMNS Staatliches on the medial endite of the maxillula, the lateral

Museum fiir Naturkunde, Stuttgart, ; lobe of the maxilla apically truncate.

UFRGS Universidade Federal do Rio Grande do

Sul, Porto Alegre, Brazil; ZMA Zoological Mu-

The Netherlands. Atlantoscia seum Amsterdam, rubromarginata n. sp.

(Figs. 1—5)

Systematics Material, - Holotype O' 10 mm, Brazil, Sergipe State, Barra

dos Coqueiros, 10°54’ S 37°03’ W, Mata Atlantica, on Hoen-

coll. Ermnidada G. bergia ip., Dr. Concenjao Couto, 29.08.1995, Genus Atlantoscia Ferrara & Taiti, 1981 MNRJ 12332; paratypes 2 99 same data as holotype, MNRJ

data 12333; paratypes 1c,399 (partially ovigerous), same

- & 1984 Synonymy. Ocelloscia Schultz Johnson, collection as holotype, A. Leistikow; paratypes 1 o', 19, 19

(ovigerous), Brazil, Sergipe State, Crasto, soil, coll, P.B. Araujo

A and C. SMNS T437. detailed diagnosis of this genus was given when Assis, 02.04.1998,

Additional material: I O', 1 9,1 9 (ovigerous), Brazil, Sergipe the genus was first established (Ferrara & Taiti State, Crasto, soil, coll. P.B. Araujo and C. Assis, 02.04.1998, 1981) to comprise a species from Ascension Is- UFRGS 2319; 3 9 (ovigerous), same data, ZMA Is. 203915. land, which was thought to be identical with Philo- scia floridana Van Name, 1940by Lemos de Castro Color: Dorsally chestnut with lighter median area (1985). Some additions to the diagnosis should be and some non-pigmented spots on tergites, coxal made: Antennula with conical medial set of apex, plates marginally buff, in vivo reddish, cephalotho- aesthetascs paired and graded, antennal flagellum rax chestnut with many non-pigmented muscle with shorter than distal free apical organ article, insertion spots on vertex, pleon as pereion but only

sensilla almost as long as cuticular sheath. Medial two paramedian pale spots on pleonite 1 and 2, endite of maxillula without medial lobe apical tip, ventrally yellowish. of maxilla with three of rows cusps, maxillipedal Cephalothorax: Without linea frontalis, linea su- basis with sulcus setal tuft of lateralis, proximal small lamina fron- pra-antennalis prominent, very of short and seta. Pereio- palp consisting a a long talis, vertex somewhat arched, small lateral lobes, pod I with transverse antenna-grooming of carpus compound eyes composed about 23 ommatidia

brush ornamental seta apically double rostrally, arranged in four rows (Fig. 1, Ctf). fringed with long inner claw, serrate, dactylus Pereion: Tegument rather smooth and shiny with

seta with a small knob, i dactylar terminating only scattered tricorn-like setae, coxal plates I to

V caudally rounded, V to VII caudally pointed, - Atlantoscia alceui Ferrara & Taiti, Type species. bearing sulcus marginalis, few gland pores and 1981 ofPhiloscia Van (junior synonym floridana noduli laterales; nodulus lateralis of coxal plate

Name, 1940), by monotypy IV more dorsally than the others (Fig. 5, Cx4).

Pleon: Set back from pereion, neopleurae of pleon

- Remarks. Atlantoscia Ferrara & Taiti, 1981 be- 3 5 segments to conspicuous, pleotelson with con-

to a line different from of the longs phyletic most cave lateral margins, bearing tricorn-like setae. neotropical philosciid Oniscidea and characterized Antennula: Tri-articulate with distal article coni-

the by shape of the antennula, the structure of the form, almost of half the length of appendage, of the apical organ antenna, and the short noduli medially bearing three sets ofaesthetascs and apical laterales. The of the structure cephalothorax is quite set (Fig. I, Anl). similar to several smaller of the Andes genera (pers. Antenna: Rather slender, flagellum tri-articu-

and all these structures obs.), comprise plesio- late, as long as peduncular article 5 proximal article , morphies. The apomorphic characters of this the distal short ge- longest, joint bearing apical organ

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Fi in 1. An2 Ctf in frontal view; Had habitus dorsal S■ Atlantoscia rubromarginata n. sp. Anl antennula; antenna; cephalothorax

Vl Hal ew; habitus in lateral view; Tel pleotelson. Downloaded from Brill.com10/05/2021 03:16:23PM via free access - Philosciids Brazil 112 P.B. Araujo & A. Leistikow (Isopoda) from

2. Allantoscia 10 Mdl/r left and mandible; with detail of endite Fig. rubromarginata n. sp, Holotype cc mm. right Mxp maxillipede

in rostral Mxl maxillula with detail of of lateral Mx2 maxilla. view; apex endite; Downloaded from Brill.com10/05/2021 03:16:23PM via free access Contributions to Zoology, 68 (2) - 1999 113

Fig 3. Atlantoscia 10 Dac in rostral PEI-4 1-4 caudal rubromarginata n. sp. Holotype cr mm. dactylus view; pereiopods view, etails in rostral Scl ornamental of Sdl seta of Sm4 of view; sensory spine carpus 1; dactylar dactylus 1; sensory spine merus 4; Spl sensory of spine propus 1.

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with long free basal sensilla. Length ratio of pe- “Trachealfeldleiste” of Verhoeff(1920). This term

duncular articles 1 to 5 is 1 : 2 : 2 : 4 : 5, all ar- is more descriptive than “dorsal ridge” as used by

ticles bearing tricorn-like setae (Fig. 2, An2). Unwin (1932), and supports the earlier used term

Mandible:Molar penicil consisting of about four of Verhoeff. Sexual differentiation: Male pleopod

and short 1 than female long two branches, basally fused, pars exopod more pointed exopod 1,

intermedia with left and with two penicils on one on endopod pointed slightly inflated-looking apex,

with of right mandible, as long as additional plumose seta distal third row spines on mediocaudal

(Fig. 2, Mdl/r). surface, minute pectinate scales mediocaudally at

Maxillula: Medial endite of of 2 bearing two long heigth apex genital papilla. Pleopod exopod

penicils and laterally some hair-like setae, lateral with very slender distal part, laterally deeply sinu-

endite with 4+6 five of inner four apically teeth, set ous, bearing sensory spines, endopod surpass-

cleft, laterally with dense fringe of trichiae (Fig. ing exopod, bearing a serrate cuticular plaque

2, Mxl). subapically.

Maxilla: Rather broad, lateral lobe 2.5 times of Uropod: Protopod subquadrangular with lateral

the breadth of medial lobe, densely covered with groove and deep dorsal furrow, endopod of less

hyaline pectinate scales, medial lobe hirsute, thanhalf the length of more distally inserted exopod

10 apically with about cusps (Fig. 2, Mx2). (Fig. 4, UR).

Maxillipede: Basis with sulcus lateralis, proxi- Genital papilla; Ventral shape ovate, genital ori-

mally reduced, bearing tricorn-like setae, palp with fices surpassing slightly ventral shield, apically apical setal tuft, medial tufts composed of about truncate.

four and two setae, respectively, proximal article with two rather long setae, endite lacking setation Remarks. - Superficially, this species resembles a save some setation on rostral ridge and prominent Benthana Budde-Lund, 1908, especially Benthana tooth caudally (Fig. 2, Mxp). albomarginata Lemos de Castro, 1958 due to simi-

Pereiopods: Rather long and slender, especially lar color pattern and overall habitus, but Atlantoscia

of 6 and 4 6 be propus pereiopods 7; pereiopod to rubromarginata n. sp. can distinguished mac- ischium with small pegs rostrodistally, carpus and roscopically by the shape ofmale pleopod 1 exopod,

2 with translucent medial the much merits to 4 scales on shorter uropods and the reddish instead margin, antenna-grooming brush present on propus of white coxal plates. Microscopically it differs in and furrow the cleft carpus 1, forming long transverse on maxillular teeth instead of serrate teeth

therefore medial in Benthana. of this carpus 1, margin concave, orna- An ascription interesting spe- mental sensory spine of carpus 1 with double fringe, cies to Balloniscus has to be rejected, since it dif-

of 3 to 7 with modi- fers in the and some sensory spines merus small neopleurae, the shape setation fied apices, dactylus with long inner claw and of the male pereiopods with three dense rows of slightly shorter inter-ungual seta, dactylar seta with deeply cleft sensory spines in Balloniscus, the shape club-like Dac/PE ofmale apex (Fig. 3, 1-4; Fig. 4, PE 5- pereiopod 7 ischium and the dactylar seta,

7). Sexual differentiation is very faint with male which all agree with the generic diagnosis of pereiopod 7 ischium somewhat stronger. Atlantoscia. Another difference to both Benthana Pleopods: Endopods with straight margins, lack- and Balloniscus is the position of the noduli laterales ing setation, exopods with slight setation margin- with a d/c maximum on coxal plate IV and the

with 5 6 and addi- flat of b/c coordinates. ally, laterally to sensory spines very curve From its only tional in setae pleopod 5 (Fig. 5, PL 1-5), all pleo- congener the new species is at once separated by

with the of podal exopods respiratory area lateroproxi- presence partially covered lungs with ra-

of radial mally: fringe multiple foldings proximally dial foldings instead of an uncovered lung of pleo- bordered by a ridge, exopods appearing laterally pod 1 in Atlantoscia floridana. Another difference broadened. The before-mentioned should is the brush of ridge antenna-grooming carpus 1, which be called “ridge of atrium” the is in A. than lung following more elongate rubromarginata n. sp. terminology used by Hoese (1983) and the term in A. floridana.

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F and tricorn- ‘g- 4. 10 PE5-7 5-7 in caudal Sm6 sensory spine Atlantoscia rubromarginata n. sp. Holotype o' mm. pereiopods view; 'ike seta of Sm7 of UR merus 6; sensory spine merus 7; uropod.

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5. Atlantoscia 10 Cx4 coxal PL 1-5 1-5 with details of 1 Fig. rubromarginata n. sp. Holotype o' mm. plate IV; pleopods endopods

(rostrally) and 2 (caudally); Plx pleopod 1 exopod (paratype, 9 12 mm).

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Atlantoscia floridana (Van Name, 1940) Maxillula: Lateral endite with 4+6 teeth, five

of inner set cleft, one trifid, laterally fringed by

Synonymy. - Philoscia floridana Van Name, 1940. trichiform setae, medial endite with two weak

Chaetophiloscia paulensis Moreira, 1927 (sensuVandel, 1963), penicils, lacking apical point (Fig. 7, Mxl). Ocelloscia floridana comb. n. Schultz & Johnson, 1984, Maxilla: Medial lobe less than half the breadth Atlantoscia alceui Ferrara & Taiti, 1981. of lateral lobe, densely covered with fine trichiform

setae, three rows of lateral lobe - cusps, Material. O’ 8.5 mm, Brazil, Reserva Biologica do Lami, Porto distally

Rio Grande do 1997; several O’O', same data covered with scales 7, Mx2). Alegre, Sul, 99 , sparsely pectinate (Fig.

collection of A. Leistikow and UFRGS as dissected specimen, Maxillipede: Basis with sulcus lateralis, latero- 2429, proximally with cuticular scales, endite lacking

knob-like penicil on rostral surface, some hairs on Color: Rich chestnut with darker median band and a ridge and fine hairs laterally, palp with inner umber band on coxal plates bordered white both and short setal set consisting of long seta, medial medially and laterally, medial umber patches on of tuft with four setae, setae proximal article strong tergites IV to VII. White spots on cephalothorax, (Fig. 7, Mxp). pereion and pleon indicating muscle insertions in Rather Pereiopods: slender, bearing many tri- cuticula. Ventrally pale without distinct markings. corn-like with setae, carpus 1 antennal-grooming Cephalothorax: Vertex arched, linea frontalis device of with brush, antagonistic propus distally lacking, slight lateral lobes, linea supra-antennalis incurved ornamental with setae, sensory spine of 20 present, compound eyes composed up to serrate double fringed apex (Fig. 8, Scl), dactylus ommatidia (Fig. 6, Ctf). with long inner claw and slightly shorter interungual Pereion: Tegument smooth with some small seta, dactylar seta with knob-like apex (Fig. 8, Dac, tubercles, bearing scattered tricorn-like setae, coxal PE 1-4; Fig. 9, PE 5-7). Sexual differentiation: The plates with sulcus marginalis, few (1-3) gland 7 sexes differ in the setation of pereiopod ischium; pores and noduli laterales, inserted most dorsally medial margin with conspicuous sensory spine of on coxal plate IV (Fig. 6, Cxp), caudal margin of half the length of those of lateral edge in the fe- coxal plates bearing a row of distinct quadrangu- in the male. lar male, lacking scales with fringed edge (Fig. 10, Cx3) Pleopods: Exopods obliquely rectangular with Pleon: Narrower than pereion, with small distinct point, lateral margin bearing two to three neopleurae on pleon segments 3 to 5, pleotelson 5 with transverse with lateral tricorn-like sensory spines, pleopod exopod margins concave, bearing caudal row of scales on surface; setae. pectinate endopods

with inconspicuously sinuous lateral margin, small Antennula: Tri-articulate, distal article coniform,

°f respiratory areas lateroproximally on caudal side half the length of antennula, bearing five pairs 10. PL Sexual differentia- °f aesthetascs of exopods (Fig. 1-5). on medial margin to apex (Fig. 6, Anl), tion: Exopod of male pleopod 1 rather triangular,

rounded, lateral margin concave, with- Antenna: Flagellum composed of three articles, medially bent e out spines, endopod laterally with ach almost subequal in length, distal article with sensory

a pical small lateral, subapical protrusion, lateral margin organ, half the length of distal joint, free

sensilla of channel crenulate, medially with row almost as long as cuticular sheath, pedun- spermatic rather cular articles densely covered with tricorn-like of spines. Pleopod 2 exopod elongate, point-

s sinuous with three ctae, length ratio from article 1 to 5 as 1 : 2 : 2 : ed, laterally sensory spines,

3 : 4 (Fig. 6, An2). endopod acute, slightly longer than exopod. of Mandible; Molar penicil consisting of about 5 Uropod: Exopod inserting distally endopod,

with 9, UR). branches, basally fused, pars intermedia protopod laterally grooved (Fig. Ventral shield shorter coniform setae and two penicils on left or one Genital papilla: slightly

Penicil orifices the on right side, additional plumose seta proxi- than genital papilla, on transversely

mity (Fig. 7, Mdl/r). truncate apical margin (Fig. 10, Gen).

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Allantoscia Fig. 6. floridana (Van Name, 1940) cr 8 mm. Anl antennula; An2 antenna; Ctf cephalothorax in frontal view; Cxp position ofnoduli laterales; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson.

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% 7, Atlantoscia left and mandible with detail of intermedia of left floridana (Van Name, 1940) cr 8 mm. Mdl/r right pars one;

iv* x p with detail of lateral Mx2 maxilla. maxillipede with detail of endite in rostral view; Mxl maxillula of apex endite; Downloaded from Brill.com10/05/2021 03:16:23PM via free access 120 P.B. Araujo & A. Leistikow - Philosciids (Isopoda) from Brazil

Fig. 8. Atlantosciafloridana (Van Name, 1940) O' 8 mm. Dac dactylus in rostral view; PEI-4 pereiopods 1-4 caudal view, detail of

1 in rostral Scl ornamental of Sc4 of Sm4 carpus view; sensory spine carpus 1; sensory spine carpus 4; sensory spine of merus 4; Spl

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%. 9. Atlantoscia tricorn-like of basis floridana (Van Name, 1940) a 8 mm. PE5-7 pereiopods 5-7 in caudal view; Sb5 seta 5; Sd5

dactylar seta of Sm6 of UR dactylus 5; sensory spine merus 6; uropod.

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Fig. 10. Atlantoscia floridana (Van Name, 1940) a 8 mm. Cx3 coxal plate III; Gen genital papilla; PL1-5 pleopods 1-5 with details of endopods 1 (rostrally) and 2 (caudally); Plx pleopod 1 exopod.

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Remarks. - Atlantoscia floridana has had a prob- Type species. - Philoscia sellowii Brandt, 1833. lematic nomenclatural history as can be seen by

the It is distributed Remarks. - the described numerous synonyms. widely Among genera herein, in the of semi- coastal forests along the Atlantic shores from Balloniscus is recognized by presence

and Ascension covered in all to Brazil, occurs on Island pleopodal lungs five pairs of pleo-

(Ferrara & Taiti, 1981; Taiti & Ferrara, 1991). In- pods, by the prominent neopleurae, and shape of terestingly, in the northern part of its distribution male pleopod 1 exopod, that are apomorphous char- the maxillular tooth of this is 4+6 pattern species acters of the genus. The pereiopods, being rather teeth with 5 cleft while in the southern ones, part stout with no setal tufts on the laterodistal edge of it sometimes has 4+5 teeth with four cleft ones differ from those carpus and merus, remarkably

(Ferrara & Taiti, 1981; Taiti & Ferrara, 1991; in other but since genera similarly shaped pereio-

et ah, There is a zone of in Araujo 1996). overlap pods are known from several other genera the sys- Santa Catarina State and Rio Grande do Sul States tematic value is still unknown. Another autapo- 6 (Brazil), where specimens with a vestigial tooth morphy is the lack of an ornamental sensory spine

can be found. which in the male pereiopod 1 carpus, is present For characters the moment, the only supposed in female and similar those of the to e.g., Atlanto- to the be autapomorphies are the reduction of lamina scia. A possible synapomorphy with Atlantoscia which is in A. frontalis, present rubromarginata endite of is a straight trifid tooth on the lateral the n. sp., and the color with a dark brown band on maxillula. the coxal plates bordered by light areas.

Balloniscus sellowii (Brandt, 1833) Genus Balloniscus Budde-Lund, 1885 (Figs. 11—16)

Diagnosis. - Cephalothorax without linea fronta- For cf. Lemos de Castro synonymy (1976), lis and lamina frontalis, linea supra-antennalis present. Pleon with well-developed neopleurae, Material. - O' 8.5 mm, Aguas Claras, Viamao, Rio Grande do the less oval giving a more or shape. P.B. 31.V. 11 Buenos Sul, Brazil, leg. Araujo, 1997; 9 mm, prov.

Antcnnula slender, distal article with graded sets Aires, 34°30’44'’S 59°35’05”W, alt. 200 m, “Flachland,

of aesthetascs medially, antennal flagellum tri-arti- ‘Unkrautfeld’ zwischen Rinderweiden und Sojafeldern”, leg.

several M. von Tschirnhaus, 14.2.1996, further specimens, culate, apical organ much shorter than distal arti- collection A. Leistikow. cle, free sensilla as long as cuticular sheath. Man-

dible with dichotomized molar penicil, maxillula Color: Pereion and cephalothorax richly chestnut with the medial endite lacking apical tip, lateral with white patches in median line, tergites cov- endite with 4+6 teeth, inner set cleft, lateral lobe ered with rather coxal of plentiful pale spots, plates maxilla twice as broad as medial lobe, maxilli- bearing two white longitudinal patches enclosing pede basis with sulcus lateralis, endite lacking knob- l'ke an umber band, pleon chestnut with two interrupted, penicil, palp with two setae on proximal tuft. white paramedian lines and some spots. Ventrally Pereiopods slender, densely covered with tricorns,

unpigmented save for the genital papilla bearing antenna-grooming brush transverse, ornamental

an umber ventral shield and brown pereiopods. sensory spine double-fringed serrate, dactylus with Vertex linea Cephalothorax: rather arched, su- long inner claw, dactylar seta simple, coxal plates linea frontalis lacking, with sulcus marginalis, nodulus lateralis, gland pra-antennalis prominent, of the compound with slight Pores along the entire lateral margin. Moduli mcdioventrally eyes lateral lobes and frons from vertex laterales with only slight maximum on coxal plate separated by shallow IV. Pleopods with partially covered lungs in all an angle, frons with two depressions joined

Pairs of on a con- exopods, lateral margin sinuous, apex profrons, forming V, compound eyes

Pointed. Male pleopod 1 exopod circular, with sisting of about 22 ommatidia (Fig. 11, Ctf).

sensory spines along entire distal margin. Uropod Pereion; Tegument rather smooth with some With endopod inserting proximally of exopod. small granula on pereionites, coxal plates smooth

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in frontal view; Had habitus Fig. II. Balloniscus sellowii (Brandt, 1833) a 8.5 mm. Anl antennula; An2 antenna; Ctfcephalothorax in dorsal view; Hal habitus in lateral view; Tel pleotelson.

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with tricorn-like setae, sulcus marginalis border- transverse antenna-grooming brush, deeply fur-

zone of noduli laterales device of ing elongate gland pores, rowed, antagonistic propus consisting present on each coxal plate, d/c-coordinates with of a single row of stout setae, dactylus with long maximum on coxal plate IV. In ovigerous females inner claw and simple dactylar seta, ornamental coxal from of 1 with double plates II to IV separated pereionite by sensory spine carpus serrate fringe

of 4 conspicuous suture (Fig. 16, Cx4). at its apex, rostrodistal margin ischium crenu-

Pleon: Only slightly retracted from pereion, late (Fig. 13, Dac, PE 1-4; Fig. 14, PE 5-7). Sexual neopleurae of pleon segments 3 to 5 prominent, differentiation: Male pereiopods 1 to 4 with me- somewhat with lat- dial of and with three four falciform, pleotelson concave margin carpus merus to eral margins, bordered by tricorn-like setae, and rows of multi-tipped sensory spines (Fig. 13, Sml), some tricorn-like setae on dorsal surface. forming a dense setal comb with loss of charac-

Antennula: teristic ornamental of Articles rather slender, distal one al- sensory spine carpus 1, which

of most as long as proximal two together, bearing is present in pereiopod female (Fig. 16, Scl),

in apical tuft ofaesthetascs and three transverse rows cover of sensory spines carpus and merus 5 to of 6 also in the aesthetascs medially (Fig. 11, Anl). higher male; additionally, merus 6

Antenna: Flagellum tri-articulate, all articles medioproximally swollen. Male ischium 7 proxi- subequal in length, distal joint slightly longer, mally narrowed, looking somewhat funnel-shaped aesthetascs article concave medial on 2 and 3, apical organ one by margin, proximally grooved. fourth of the length of distal article, free sensilla Pleopods: Pleopodal exopods pointed, with lat- as eral sinuous due long as others, peduncle comparatively short, margin very to respiratory field length ratio of article I to 5 is 1 : 2 : 2 : 3 : 4, all of lateroproximal part. Respiratory area laterally densely covered with tricorn-like setae (Fig. II, crenulate with medial area partially covered by

of lateral An2). edge lung atrium, margin bearing many

Mandible: Pars intermedia with coniform setae sensory spines (Fig. 15, PL 1-5). Sexual differ- and entiation: Male 1 two on left or one penicil on right mandible, pleopod exopod rounded with molar whole penicil consisting of 7 to 8 branches, all in- margin bearing sensory spines, endopod long dividualized, additional plumose seta distally of and slender, acute, bearing row of spines medio- molar penicil (Fig. 12, Mdl/r). caudally, some very faint pectinate scales proxi-

Maxillula: Medial endite apically with two slen- mally of interlocking area with apex of genital cone. der penicils, laterodistal margin bearing hair-like Pleopod 2 exopod somewhat more elongate than setae, no lateral point, lateral endite in the exam- exopod of pleopod 3, medial margin with setose ined third almost specimen with 4+6 (1 vestigial) teeth, with area, endopod slender, apical flagelli-

in inner set cleft, one trifid, lateral fringe of com- form, broken off the examined specimens. paratively few, stout trichiform setae (Fig. 12, Mx 1). Uropod: Protopod laterally grooved, subtriangu-

Maxilla: Medial lobe half the width of lateral, lar shaped, endopod slightly weaker than exopod, apically cuspidate, subapically with trichiform setae, inserting more proximally (Fig. 14, UR). lateral lobe with pectinate scales caudally (Fig. Genital appendage: Ventral shield coniform, 12, Mx2). mouths of ductus ejaculatorii somewhat surpassing

basis with sulcus its setation discernible in Maxillipede: lateralis, medi- apex, no light microscope ally of it cuticula with at 400 x scaly, palp very inconspicu- magnification (Fig. 15, Gen). ous articulation between distal and medial article, setal tuft of medial article reduced, bearing four Remarks. - Vandel (1963) established a new fam- setae, proximally of it a pair of one long and one ily Balloniscidae for the hitherto known species short seta; setae of proximal article long, almost of South American philosciid species with pleopodal

He the subequally shaped, epipod medially with some lungs. mentioned that lungs are not closed scales and pectinate scales (Fig. 12, Mxp). in several species, which furthermore lack a linea

Fereiopods: Long but fairly stout appendages frontalis - the Balloniscus Budde-Lund, 1885 s.

- the bearing many tricorn-like setae, carpus 1 with long str. and separated species with linea fronta-

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12. sellowii and with detail intermedia ofleft Fig. Balloniscus (Brandt, 1833) cf 8.5 mm. Mdl/rleft right mandible of pars one; Mxp

with detail of endite in maxillula with detail Mx2 maxilla. maxillipede rostral view; Mxl of apex of lateral endite;

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p ig- 13. caudal Balloniscus sellowii (Brandt, 1833) o’ 8.5 mm. Dac dactylus in rostral view; PEI-4 pereiopods 1-4, view, with detail °f 1 and distal of ischium in rostral of 1; Sml propus to carpus margin 4 view, respectively; Scl sensory spines carpus sensory spine °f meral of 1. brush 1; Spl sensory spine propus Downloaded from Brill.com10/05/2021 03:16:23PM via free access 128 P.B. Araujo & A. Leistikow - Philosciids (Isopoda) from Brazil

Fig. 14. Balloniscus sellowii 8.5 PE5-7 Sc7 of Sm7 (Brandt, 1833) cr mm. pereiopods 5-7; sensory spine carpus 7; sensory spine of merus 7; UR uropod.

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Fig. 15. Balloniscus sellowii (Brandt, 1833) O' 8.5 mm. Gen genital papilla; PL 1-5 pleopod 1-5

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16. Balloniscus note Fig. sellowii (Brandt, 1833) 9 11 mm (ovigerous). Cx4 coxal plate 4, longitudinal sulcus; PEI, PE7 pereiopods

1 and 7 PL1-2 Scl of 7 of and (rostrally) (caudally); pleopods 1-2; sensory spine carpus 1; Sml, sensory spines merus 1 7.

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in the Plataoniscus 1 with transverse antenna- lis and covered lungs new genus pods slender, carpus inner Vandel, 1963. Reca (1970) reconsidered the sta- grooming brush, dactylus with claw long, coxal bear- tus of several South American species of Balloni- simple dactylar seta rather short, plates

the nodulus and scidae; he came to conclusion that the generic ing lateralis sulcus marginalis, gland

in limits fluent and a re-examination of the whole with area are pores lacking. Pleopods respiratory family has to be considered. Lemos de Castro (1976) exopod 1, male exopod triangular with laterally

the conclusion that several are in of the came to species directed subapical protrusion most spe- sellowii merely junior synonyms of Balloniscus cies. Uropod with endopod inserting proximally exhaustive list. and gave an synonymy Although of exopod.

Balloniscus sellowii is rather well studied, com-

with other South American representatives - pared Type species. Philoscia picta (Brandt, 1833) of Oniscidea, there are no considerations about

its status. Morphologically it is quite teeth is phylogenetic Remarks. - The shape of the maxillular

both and Atlantoscia. Some similar to Benthana of this the other known rather typical genus; only the of evidence for a common ancestry are shape philosciids with serrate teeth are Ctenoscia the antennula with stepped of aesthetascs groups Verhoeff, 1928 from southern , Benthanops

on the distal article, the shape of the medially Barnard, 1932 from South , Benthanoides antenna-grooming brush of pereiopod 1 carpus, Lemos de Castro, 1958, Benthanoscia Lemos de the which is rather narrow and elongate and shape Castro, 1958, both from eastern South America, of the of the setal fields in the males sensory spines and Alboscia Schultz, 1995 from and of Balloniscus andBenthana. Actually Balloniscus South of Brazil (Araujo, in press). Pectinate teeth shows several more derived characters, i.e., the known from members of are equally the Ligiidae long short antenna, stout pereiopods neopleurae, this be the Brandt & Ratzeburg, 1831. Thus, may and the morphology of the pleopodal lungs, but at plesiomorphic character state for the philosciid least the first characters can be evidenced as not Crinochaeta, as is the additional stalk, which is evolving independently, as they are in functional also found in Ligiidae and Scleropactidae Verhoeff, connection with more clinging habits as opposed 1938. In Benthana the maxillular teeth are arranged , to the runner-strategies of long-limbed species (cf. different from that in the in a manner quite pre- Schmalfuss, 1979 and 1998). of the lateral endite is ceding genera. The apex

in broader and the teeth are standing a semicircle

laterally, similar to Ligia Fabricius, 1798. From Genus Benthana Budde-Lund, 1908 this point of view, the structural features found on

the maxilla are useless for phylogenetic reconstruc- Diagnosis. - Cephalothorax with linea supra- tion. The generic independence of Benthana and antennalis; lineafrontalis and lamina frontalis lack- Benthanoides has to be doubted; it seems as if the ing, slight lateral lobes, ommatidia of compound of Benthanoides are the more basal coni- species simply eyes arranged in four rows. Antennula with characteristic dentiform species which lack the pro- form distal article, bearing graded sets ofaesthetascs trusion of the pleopod 1 exopod. However, prior medially, antenna with tri-articulate flagellum,

to this a detailed than distal free establishing synonymy, phylo- apical organ much shorter article, of all the hitherto known has cuticular genetic study species sensilla as long as sheath. Molar penicil to be undertaken. An apomorphy of Benthana s. dichotomized, maxillula with medial endite lack-

str. is the shape of pleopod 1 exopod with a lateral ing apical tip, lateral endite with 4+6 teeth and

dentiform The of the ornamen- additional stalk, five teeth of inner set pectinate, protrusion. shape

tal of 1 is similar to that of maxilla with lateral lobe two times broader than sensory spine carpus

1928 in medial endite, maxillipede basis with sulcus Ischioscia Verhoeff, although differing several details. have been evolved inde- lateralis, endite without knob-like penicil, proxi- It may

mal setal tuft of palp comprising two setae. Pereio- pendently.

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Benthana convexa Lemos de Castro, 1958 knob-like penicil, longitudinal ridge bearing some

(Figs. 17—22) setae, laterally very fine setation, palp with proxi-

mal setal tuft composed of long and short seta,

Material: cf 9 Alto de Mosela,Petropolis, Rio de Janeiro, mm, medial tuft bearing four setae (Fig. 18, Mxp).

Brazil, leg. Daley, 1.36, MNRJ 8194; several 99, same data. Pereiopods: All articles bearing several tricorns,

carpus 1 with transverse antenna grooming brush, Color; All specimens had faded out to a yellowish which almost reaches lateral covering a groove buff; muscle insertions on cephalothorax slightly of of it some cuticular margin carpus, proximally darker. scales and crest of trichiform setae; ornamental Cephalothorax: Like the preceding species with sensory spine with hand-like apex. Ischium 7 with small lateral lobes and linea supra-antennalis, linea 5 prominent sensory spines grouped laterally, dac- frontalis lacking, vertex arched, compound eyes tylus with simple dactylar seta, inner claw promi- consisting of about 20 ommatidia (Fig. 17, Ctf). nent, longer than inter-ungual seta (Fig. 19, Dac, Pereion: Slightly convex in cross section, tegu- PE 1-4; Fig. 20, PE 5-7). Sexual differentiation: ment rather smooth, coxal plates with sulcus Male 1 4 with and pereiopods to carpus menus marginalis and noduli laterales, gland pores lack- 3 4 medially bearing to rows of deeply cleft sen- ing, slight maximum of d/c-coordinates of noduli sory spines, ischium of pereiopod 7 with a more IV laterales on coxal plate (Fig. 22, Cx4; Fig. 17, concave medial margin in the male, male merus 7 Cxp). strongly concave medially, giving it an arcuate Pleon: Conspicuously retracted from pereion with with cuticular scales. appearance, mediodistally small 3 ncopleurae on plconsegments to 5, pleo- Pleopods: Male pleopodal exopods laterally with telson with only slightly concave lateral margins, 5 in 5 8 in sensory spines pleopod to spines pleo- caudally not surpassed by protopods of uropods. pod 3, endopods laterally sinuous, convex in pleo- Antennula: Tri-articulate with coniform distal pod 5, small respiratory area at least in pleopod 1 article bearing two pairs of aesthetascs medially (Fig. 21, PL 1-5). Sexual differentiation: Pleopod and one apically (Fig. 17, Anl). 1 exopod inverted heart-shaped, laterally with Antenna: Fairly slender with tri-articulate fla- 3 subapical protrusion, proximally of it about sen- gellum, flagellar articles subequal in length, proxi- sory spines, endopod acute, laterally incurved, mal one the longest, apical organ short, one third caudomedial row of spines only in medial part of of the length of distal article, with free sensilla as normal extension present. Pleopod 2 exopod slightly long as cuticular sheath (Fig. 17, An2). more pointed than exopod 3, with same number Mandible: Pars intermedia with dense cushion

of sensory spines laterally, endopod pointed, dis- of coniform setae, two penicils and basally some tal part flagelliform, apex unfortunately broken in longer setae on left mandible, right side with fewer dissected specimen. setae and single penicil, additional plumose seta Uropod and genital appendage: As in generic proximally, molar penicil consisting of about 9 diagnosis (Fig. 20, UR; Fig. 21, Gen). individualized branches (Fig. 18, Mdl/r).

Maxillula: Medial endite pointed, with two slen-

Remarks. - The based der penicils apically, lateral endite apically bear- original description was on

ing 4+6 teeth, five of inner set pectinate, lateral only three females from Cantareira, Sao Paulo

fringe composed of rather stout setae compared (Lemos de Castro, 1958a). The above given

the characterization with the with other genera (Fig. 18, Mxl). description completes male traits. The differs from its closest Maxilla: Medial lobe half as broad as lateral lobe, species

B. schubarti Lemos de 1958 covered with trichiform setae, apically cuspidate, relative, Castro, by

the much and the of the lateral endite almost lacking setation, some very longer antennae shape

fine setae laterally (Fig. 18, Mx2). sublateral tooth of the maxillula, which is of half

B. Lemos Maxillipcdc: Basis with sulcus marginalis, proxi- the length of the lateral one in convexa

mally cuticula slightly scaled, endite with strong de Castro, 1958 and much smaller in B. schubarti

tooth and two laterodistal hooks, rostrally lacking (Lemos de Castro, 1958a). These two species seem

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Fig. 17. Benthana convexa Lemos de Castro, 1958 o■ 9 mm. Anl antennula; An2 antenna; Ctf cephalothorax in frontal view; Cxp position of noduli laterales; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson.

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18. Benthana left and with of ofleft Fig. convexa Lemos de Castro, 1958 cr 9 mm. Mdl/r right mandible detail pars intermedia one;

with detail of endite rostral Mxl with of of lateral Mx2 maxilla. Mxp maxillipede in view; maxillula detail apex endite;

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Fig. 19. Benthana convexa Lemos de Castro, 1958 o' 9 mm. Dac dactylus in rostral view; PEI-4 pereiopods 1-4 caudal view, detail

of 1 in rostral Scl ornamental ofbrush of Sdl seta of carpus view; sensory spine and sensory spine carpus 1; dactylar dactylus 1; Sm2 sensory spine of meral brush 2.

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Sc5 Fig. 20. Benthana convexa Lemos de Castro, 1958 a 9 mm. PE5-7 pereiopods 5-7; Sb5 tricom-like seta of basis 5; sensory spine of carpus 5; Sm6 sensory spine of merus 6; UR uropod.

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Fig. 21. Benthana convexa Lemos de Castro, 1958 O• 9 mm. Gen genitalpapilla; PL1-5 pleopod 1-5.

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Fig. 22. Benthana convexaLemos de Castro, 1958 9 11 mm. Cx4 coxal plate 4; PEI, PE7 pereiopods 1 and 7 (caudally), with detail

of 1 PL 1-2 1-2 Scl of 1. pereiopod rostrally; pleopod exopods; sensory spines carpus

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to be close to each other, while their position within of the shape of the antennal organ these species

Benthana the of the depends on interpretation are excluded from the monophyletic group to which character of the rather short Ischioscia and other state uropods: uro- several genera belong (cf. pods with the endopod inserting proximally of the Leistikow, in press); most probably this character exopod, while in all the other species, the protopod is also plesiomorphic. Nonetheless, some charac-

and in all but B. inserts more caudally werneri ters indicate a close relationship of Benthana to

Lemos de Castro, 1958 the insertion of the rami is Atlantoscia, i.e., the shape of the maxilla with the

the level. Verhoeff at same (1941, 1951) stressed lateral lobe apically concave, the position of the that the uropods with the protopod reaching dis- noduli laterales being gradually more dorsal from tally of the pleotelson and the endopod inserting coxal plates 1 to IV, and the lateral border steeply at the same level are similar to the condition found narrows on coxal plate V. in Ligiidae. This type of uropod is a symplesio- Both genera seem to have some relationship to

and while morphy of Ligiidae Benthana, B. con- Balloniscus, although they are more distant from

B. schubarti and B. werneri an that than from each other. vexa, acquired genus they are In par- uropodal shape similar to other philosciid species ticular Benthana has similar carpal brushes in the independently. This might be reasonable, for the males on pereiopods 1 to 3. Since similar carpal

of Crinochaeta for brushes in endopods higher act removing occur other genera, it is important to

in the water from the pleoventral area and the water stress the similarity even shape of the sen-

for these conducting system (Hoese, 1981), although not sory spines forming brushes, e.g., totally water intake as stated by Verhoeff (1920). Thus, different to those seen on Ischioscia that bears car- there be selective brushes formed cuticular scales instead may a pressure favoring proxi- pal by of

On the other since For the evidence of mally inserting endopods. hand, sensory spines. a postulated the fine structure of the uropods of Benthanoides synapomorphy of Benthana and the Balloniscidae, is of the evolution of this common philosciid type, a re-examination of the second genus of Ballonis- uropodal type is explained most parsimoniously cidae, Plataoniscus Vandel, 1963, is desirable. by identical genetic information; they seem to be However, the treatment of Plataoniscus and Bal-

the more basal than Benthana species because of loniscus as a family on its own is unsatisfactory, the male covered in plesiomorphic pleopod 1 exopod. Thus, since pleopodal lungs occur the Afro- an with the rami the level uropod inserting at same tropical genus Aphiloscia, and even in Atlantoscia

be an of a within the their might autapomorphy group rubromarginata n. sp.; respective closest rela- genus Benthana and the three above-mentioned tives lack such respiratory organs. Until the phy- the basal within its If this species are most genus. logenetic relationships can be established, the

is a cleft maxillular tooth Balloniscidae should be within the assumption right, single lumped para-

in Benthanoides instead of all pectinate might be phyletic “Philosciidae”. In this regard the follow- an of this and the should be mentioned. The dentation autapomorphy group, generic ing of the lateral

identity can be supported. endite of the maxillula is quite similar in Balloniscus

and the southern populations ofA. floridana. This

feature could represent a synapomorphy of these

Phylogenetic relationships genera.

In the genera dealt with, the pereiopods struc-

The three genera discussed above share several turally resemble those of Aphiloscia. However, at characters. Due to the lack of an elaborate phylo- the present state of knowledge it is not clear if genetic of the Criochaeta the these characters analysis and, thus, could support any close relation- of it lack a is difficult inter- of all these In there is ground plan quite to ship genera. particular, no

several the status pret characters, e.g., of the pec- biogeographic evidence for a close relationship; tinate teeth of Benthana. All species bear rather, Aphiloscia is mainly East African in distribution.

which is it is remarkable big compound eyes, a plesiomorphic However, that all genera discussed character East compared to other Oniscidea. By virtue above are mainly of Brazilian distribution and

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23. Known distribution of dealt with in this work. Fig. philosciid genera

found in the Atlantic region (Lemos de Castro, 1976; References

Ferrara & Taiti, 1981; Taiti & Ferrara, 1991) (Fig.

23). The fauna of this region is related to the Afro- Araujo PB. (in press). Two new species ofAlboscia Schultz, 1995 from Rio Grande do Sul, Brazil (Isopoda, Oniscidea, tropical region although both tectonic plates have Philosciidae). Crustaceana. been separated since the Late Cretaceous (Emiliani, Araujo PB, Buckup L, Bond-Buckup G. 1996. Isopodos 1995). So it can not be excluded that there exists terrestres (Crustacea, Oniscidea) de Santa Catarina e Rio the Brazilian an ancient connection between and Grande do Sul. Iheringia, Ser. Zool. 81: 111-138.

African the of Budde-Lund G. 1908. von und Ostafrika genera. Although philosciid genera Isopoda Madagaskar

rail Diagnosen verwandter Arten. IViss. Ergebn. Reise the African tropics are rather well studied Ostafrika 2: 263-308, pis. XII-XVIII. (Schmalfuss & Ferrara, 1978; Taiti & Ferrara, 1980),

Emiliani C. 1995. Planet Earth - Cosmology, Geology and it is too early to establish phylogenetic relation- Evolution ofLife and Environment. New York: Cambridge between those ships genera. Univ. Press.

Ferrara F, Taiti S. 1981. Terrestrial isopods from Ascen-

sion Island. Monilore Zool. hal. (N.S.) 14: 189-198. Acknowledgements Ferrara F, Paoli P, Taiti S. 1994. Philosciids with pleopodal

The of the lungs? case genus Aphiloscia Budde-Lund,

The authors wish their thanks Erminda da 1908 with to express to Dr, (Crustacea, Isopoda, Oniscidea) a description

Conceigao G. Couto, Universidade Federal de Sergipe, Brazil, of six new species. J. Nat. Hist. 28: 1231-1264, who donated the material ofAtlantoscia Gruner HE. 1955. Die Benthana 1908. rubromarginata n. sp., Gattung Budde-Lund,

Dr. P. Young (MNRJ), Brazil, for the loan ofmaterial, and Dr. Zool. Jb„ Syst. 83: 441-452.

S. Taiti, CNR, Florence, , for providing material of Hoese B. 1981. Morphologie und Funktion des Wasser-

Aphiloscia vilis Budde-Lund, 1908. leitungssystems der terrestrischen Isopoden (Crustacea,

Downloaded from Brill.com10/05/2021 03:16:23PM via free access Contributions to Zoology, 68 (2) - 1999 141

Isopoda, ). Zoomorphology 98: 135-167. nae in terrestrial isopods. J. Crust. Biol. 18: 10-24.

Hoese B. 1983. Morphologic und Evolution der Lungen bei Schmalfuss H, Ferrara F. 1978. Terrestrial isopods from

den terrestrischen Isopoden (Crustacea, Isopoda, Oni- west Africa. Part 2: Families Tylidae, Ligiidae, Trichoni- Philo- scoidea). Zool. Jb., Anal. 109: 487-501. scidae, Styloniscidae, Rhyscotidae, Halophilosciidae, Porcellionidae, Leistikow A. (in press). Three new species of terrestrial sciidae, Platyarthridae, Trachelipidae,

isopods from (Crustacea; Isopoda: Oniscidea). Armadillidiidae. Monitore Zool. Ital. 11: 15-97.

crusta- Stud. Neotrop. Fauna Environm. 33. Schultz GA, Johnson C. 1984. Terrestrial isopod

Castro A. 1958a. do Benthana from Florida Lemos de Revisao genero ceans (Tylidae, Ligiidae, Halophilosciidae,

Budde-Lund, 1908 (Isopoda, Oniscidae). Arq. Mus. Nac. Philosciidae, Rhyscotidae). J. Crust. Biol. 4: 154-171.

46: 85-118. Taiti S, Ferrara F. 1980. The family Philosciidae (Crusta-

in Africa south of the Sahara. Monitore Lemos de Castro A. 1958b, On the systematic position of cea, Oniscoidea)

some American species of Philoscia Latreille, 1804. Amer. Zool. Ital. (N.S.)(Suppl.), 12: 53-98.

Mus. Nov. 1908: 1-10. Taiti S, Ferrara F. 1991. Two new species of terrestrial

sinonimia . 25: 901- Lemos de Castro A. 1976. Consideraqoes sobre a isopods from J. Nat. Hist.

e a distribuigao geografica de Balloniscus sellowii (Brandt, 916.

Bras. Biol. 36: Unwin EE. 1932. On the of the 1833) (Isopoda, Balloniscidae). Revta. structure respiratory organs

392-391. of the terrestrial Isopoda. Pap. Proc. Roy. Soc.

Lemos de Castro A. 1985. Consideraqoes sobre Atlantoscia 1931: 37-104.

alceui Ferrara & Taiti, 1981 (Isopoda, Oniscoidea, Philo- Vandel A. 1963. Isopodes terrestres receuillis en Amerique

Biol. 45: 417-422. du Sud C.D. Debouteville. In: Debouteville sciidae). Revta. Bras. par CD, Rapo-

de Australe. Paris: Edi- Morcira C. 1927. Duas especies novas de crustaceos isopodes port EH eds. Biologie VAmerique

terrestres do Brasil. Bolm. Biol. Lab. Parasit. Fac. Med. tions du Centre Nationale de la Recherche Scientifique,

S. Paulo 10: 194-200. 2; 63-100.

Name GW. van 1940. A supplement to the American land Vcrhocff KW. 1920. Uber die Atmung der Landasseln,

Kenntnis der der and freshwater isopod Crustacea. Bull. Am. Mus. Nat. Hist. zugleich ein Beitrag zur Entstehung

77: 109-142. Landtiere. Z. \Viss. Zool. 118: 365-447.

Reca AR 1970. Oniscoideos argentinos. 1. Sobre la posicion Verhoeff KW. 1941. Zur Kenntnis siidamerikanischer Oni-

sistematica de Philoscia Giambiagi, 1939. Physis scoideen. Zool. Anz. 133: 114-126.

29: 423-429. Verhoeff KW. 1951. Landisopoden aus Siidamerika. Fur-

Schmalfuss H. 1979. Ligia simoni. A model for the evolu- ther Zool. Res. Swed. Antarkt. Exp. 4: 1-19.

tion of terrestrial isopods. Stuttg. Beitr. Naturk. Ser. A

317: 1-5.

Schmalfuss II. 1998. Evolutionary strategies of the anten- Received: 28 September 1998

Downloaded from Brill.com10/05/2021 03:16:23PM via free access