On two new species of the genus Vestalenula Rossetti & Martens ...

Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The nean Society of London, 2003? 2003 1392 305313 Original Article

Lin-

TWO NEW SPECIES OF VESTALENULA (OSTRACODA, DARWINULIDAE)R. L. PINTO ET AL.

Zoological Journal of the Linnean Society, 2003, 139, 305–313. With 3 figures

On two new species of the genus Vestalenula Rossetti & Martens, 1998 (Crustacea, Ostracoda, Darwinulidae) from semiterrestrial habitats in São Paulo State (Brazil) R. L. PINTO1,*, C. E. F. ROCHA1 and K. MARTENS2,3 1

Departamento de Zoologia-IBUSP, Rua do Matão, trav. 14, n°101, CEP05508–900 São Paulo–SP, Brazil Royal Belgian Institute of Natural Sciences, Freshwater Biology, Vautierstraat 29, 1000 Brussels, Belgium 3 University of Ghent, Department of Biology, K. L. Ledeganckstraat 35, B9000 Gent, Belgium 2

Received March 2002; accepted for publication February 2003

Two new species of semiterrestrial darwinulid ostracods, both belonging to the pagliolii-lineage of the genus Vestalenula, are described here. Vestalenula botocuda sp. nov., collected from moist mud in a rain forest remnant, is an enigmatic species, as it combines valve characters of the boteai-lineage with soft part features of the danielopoli-lineage within the genus. Vestalenula irajai sp. nov., found in several types of semiterrestrial habitats, is closely related to V. pagliolii in its soft part morphology, but has more elongated valves, with straight dorsal and ventral margins running parallel to each other. The description of these new species confirms the relatively high levels of endemicity and diversity in Southern Hemisphere Vestalenula and challenges earlier classifications of this group. © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139, 305-313.

ADDITIONAL KEYWORDS: ancient asexuals – morphology – South America – Southern Hemisphere – taxonomy.

INTRODUCTION Darwinulidae are so-called ancient asexuals (Butlin & Griffiths, 1993; Judson & Normark, 1996), organisms which have persisted over geologically long periods of time without sexual reproduction. There appear to have been no bisexual populations for at least 150 Myr (Martens, 1998). All living Darwinuloidea belong to one subfamily, the Darwinulinae Brady & Norman, 1889, although the Palaeozoic–Recent fossil record of this superfamily comprises representatives of eight subfamilies in two families. For a long time, Recent darwinulids, which are exclusively nonmarine, were treated as a marginal group and little attention was devoted to them. However, their recognition as putative ancient asexuals initiated renewed interest, and provoked a series of ecological (Van Doninck et al. 2002), molecular (Schön et al., 1998) and taxonomic studies. Several taxonomic papers have presented regional surveys of darwinulid faunas, e.g. Europe (Martens, Rossetti & Fuhrman, *Corresponding author: E-mail. [email protected]

1997), South Africa (Martens & Rossetti, 1997; Rossetti & Martens, 1999), New Zealand and Australia (Rossetti, Eagar & Martens, 1998; Rossetti & Martens, 1999; Martens & Rossetti, 2002). This culminated in a world-wide revision of the group, which introduced three new genera, bringing the total number to five extant genera (Rossetti & Martens, 1998). Most work on South American darwinulids dates from before 1970, with the exception of Rossetti, Martens & Mourguiart (1996), and only the paper by Pinto & Kotzian (1961) deals with the detailed taxonomy of the Darwinulidae. Martens & Behen (1994) list the species of darwinulid ostracods at that stage reported from South America (see Martens, Würdig & Behen, 1998, for an account of Brazilian darwinulids). However, several taxonomic alterations to this list were suggested by Rossetti & Martens (1998): all species except D. stevensoni were transferred to one of the three new genera; Penthesilenula brasiliensis is now considered a valid species, not a subspecies of P. africana and Alicenula serricaudata espinosa was synonymized with the nominal subspecies. Taking into account these alterations, together with the two new

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species described here, nine darwinulid species are presently known from South America.

MATERIAL AND METHODS Material was collected during a preliminary sampling campaign of terrestrial and semiterrestrial habitats in the State of São Paulo, conducted by C. E. F. Rocha (CEFR) and J. W. Reid (JWR) in 1999 and by C. E. F. Rocha and R. L. Pinto (RLP) in 2001. Ostracods were collected live from leaf litter, mosses, etc. and killed and preserved in 70% ethanol. These collections contained several species from the families Darwinulidae, Cyprididae and Candonidae (Pinto, Rocha & Martens, 2001). The present paper deals with the darwinulid genus Vestalenula only, other (new as well as known) species will be reported on in detail elsewhere. Dissections were effected with micro needles under a stereo-light microscope. Valves were stored dry in micropalaeontological slides; soft parts were dissected in CMC/9AF medium with Rose Bengal and were mounted in permanent slides. Micrographs of valves were made with a Zeiss Scanning Electron Microscope. Deposition of material: all type material is deposited in the Museu de Zoologia da Universidade de São Paulo (MZUSP). The following abbreviations were used in text and figures: Valves. Cp, carapace; H, height; L, length; LV, left valve; RV, right valve; ms, central muscle scar(s); dv, dorsal view; vv, ventral view; lv, lateral view; iv, internal view. Limbs and soft parts. A1, antennula; A2, antenna; Md, mandibula; Mx, maxillula; T1-3, thoracic limbs; CR, caudal ramus (previously named furca); exo, exopodite on A2; Ac, ventral aesthetasc clump on A2; s1, s2, t, w, x, y, specific setae on limbs; y1–3, aesthetascs. Nomenclature proposed by Danielopol (1968, 1970), and adapted by Rossetti & Martens (1998) is used in the description of chaetotaxy of soft parts. For higher taxonomy, the system proposed by Horne, Cohen & Martens (2002) is followed.

TAXONOMIC DESCRIPTIONS CLASS OSTRACODA LATREILLE, 1806 SUBCLASS PODOCOPA G.W. MÜLLER, 1894 ORDER PODOCOPIDA SARS, 1866 SUBORDER PODOCOPINA SARS, 1866 SUPERFAMILY DARWINULOIDEA BRADY & NORMAN, 1889 FAMILY DARWINULIDAE BRADY & NORMAN, 1889 GENUS VESTALENULA ROSSETTI & MARTENS, 1998 VESTALENULA BOTOCUDA SP. NOV. (FIGS 1A–E, 3I–P) Type locality Sítio da Colônia, Mulungu, near Araçoiaba da Serra, São Paulo State, Brazil. Approximate GPS coordinates: 23∞32¢2≤S, 47∞39¢ 41.1≤W.

Muddy patches in a dry streamlet in a rain forest remnant. The mud was moist at the time of collecting; during wet season standing or running water might be present. Material collected on 15.08.2001 by CEFR and RLP.

Type material Holotype: an ovigerous, dissected female, with valves stored dry in a micropalaeontological slide and dissected soft parts kept in a sealed slide (MZUSP 15041). Paratypes: an ovigerous female dissected and stored like the holotype (MZUSP 15043); 2 ovigerous, dissected females, with valves used for SEM and stored in micropalaeontological slides, and dissected soft parts kept in sealed slides (MZUSP 15042, MZUSP 15044); 3 carapaces used for SEM and stored in micropalaeontological slides (MZUSP 15045, MZUSP 15046, MZUSP 15047); 4 ovigerous females kept in toto in ETOH (MZUSP 15048).

Derivation of name Botocudos is the designation of several South American Indian tribes that use a device to enlarge the lower lip. The present species is characterized by the fact that the LV forms an enlarged anterior ‘lip’, greatly overlapping the RV.

Abbreviated diagnosis Valves elongated (L/H = c. 2.1), with straight parts of dorsal margin slightly sloping towards the front; anterior overlap LV > RV unusually pronounced and typical; ms relatively large; keel on RV short and rounded and positioned rather posteriorly; third segment of A1 with ventro-apical seta. First endopodal segment of A2 with 2 apical setae; seta z on penultimate segment claw-like. Md-palp with seta x longer than w.

Description Cp (Fig. 3I-P) with whitish, smooth surface covered with long, widely spaced setae; elongated, in lateral view, with straight parts of dorsal ( c. 30% of total length) and ventral (c. 55%) margins subparallel, dorsal margin slightly sloping. LV overlapping RV along anterior, ventral and posterior margins; anterior overlap very pronounced, LV extending significantly beyond RV. Both valves asymmetrical in dorsal and ventral views, widest in the posterior quarter of the carapace, LV bulging over RV on the posterior side, anteriorly extending significantly beyond RV; central part with a weak constriction. Ventral margin in ventral view strongly sinuous; keel short and rounded, effectively blocking LV in tightly closed carapaces.


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Figure 1. Vestalenula botocuda sp. nov. A. A1, lv (MZUSP 15044). B. A2, lv (MZUSP 15041). C. Md-palp, lv (idem). D. CR and postabdomen, vv (MZUSP 15042). E. CR and postabdomen, lv (MZUSP 15041). Scale bars = 20 mm.


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LV in lateral (internal) view with caudal margin nearly evenly rounded, slightly everted towards the dorsal side, anterior margin everted towards the ventral side; anterior internal tooth short and small, situated c. 15% of total length from the anterior margin; ms large, positioned nearer the anterior side and consisting of c. 8 scars arranged in a rosette; mandibular scar rounded. RV in lateral (internal) view with caudal margin asymmetrically twisted, ventrally everted towards the median rather than dorsal side, anterior margin everted towards the ventral side; postero-ventral keel short and rounded, situated rather posteriorly, ms large, situated nearer the anterior side, consisting of c. 8 scars arranged in a rosette. Both valves with an internal rim along the anterior margin (= calcified inner margin?); in LV this rim seals the closure of the carapace when overlapping the RV. A1 (Fig. 1A). First segment stout with dorsal seta, the second with 2 unequal ventral setae (one c. 2.5 times as long as the other); third segment with 1 ventral and 1 dorsal setae ( s1); fourth segment with 1 dorsal seta (s2) and small alpha-seta; fifth podomere bearing 4 apical setae, 2 subequal, ventral setae (with an alpha-seta near their insertion) and 2 longer ones in dorsal position; last podomere with subapical aesthetasc, 2 long apical setae, the ventral seta about half as long as the dorsal, and a dorso-apical ‘alpha’ seta. A2 (Fig. 1B). Protopodite two-segmented, first segment with 2 setae and dorsal hook ( h), second segment with long seta. Exopodite (exo) with long, apical seta and short, lateral conical spine. Endopodite three-segmented; first podomere with ventral aesthetasc clump (Ac) in proximal position and 2 setae on the expanded ventro-apical corner; second podomere with 2 ventral aesthetascs (y1 and y2) and short, claw-like ventro-apical seta (t), subapical seta (z) and 4 distal claws: 2 long, 1 intermediate and 1 short (x); third endopodite short and slender, with one ventral aesthetasc (y3) and 2 apical claws, one short and about half the length of the second one, the latter reaching beyond the tips of the long claws of the penultimate segment. Md with palp (Fig. 1C) three-segmented, consisting of basis and two-segmented endopodite; basis typical of Darwinulidae, i.e. set with branchial plate and fan of large setae (possibly used in filter-feeding – not illustrated). First podomere of endopodite distally widened, medial seta x about 1.5 times as long as the external, subapical seta (w); two internal setae present: y very short, z very long, reaching halfway the apical claws on the terminal segment; this podomere narrow, with 5 apically hirsute, unequal claws (3 long, 1 intermediate, 1 short), 1 short internal seta (c), and a subapical external seta (b).

Mx (not illustrated). Palp two-segmented; proximal segment with an external, subapical seta, 3 terminal setae (2 straight and 1 curved, the latter shorter and plumose), and 2 short setae inserted on either side of the terminal segment; terminal segment small, with 1 thin median seta and 2 subequal, stout setae set with a double row of setulae. T1 (not illustrated). Protopodite with branchial plate, 2 isolated, unequal setae close to the articulation with the endopodite. Three-jointed endopodite; first segment with 2 subapical setae; second segment with 1 subapical seta; third segment smaller, with 2 lateral, subequal setae and a strong terminal claw. T2 (not illustrated). Protopodite one-segmented, rather stout, with 3 ventral setae, 1 short proximal and 2 long, unequal distal ones. Endopodite four-segmented; first segment with 2 unequal ventro-apical setae and 1 long seta, slightly exceeding the tip of the next segment; second and third segment with 1 stout ventro-apical seta each; terminal segment short, with 1 strong apical claw, flanked by 2 subapical claws, the ventral longer than the dorsal one. T3 (not illustrated). Protopodite one-segmented, short, with 2 unequal ventro-apical setae. Endopodite four-jointed; first segment with 1 distal seta approximately as long as the segment itself; second and third segments bearing 1 apical seta each, reaching beyond the tip of the next segment; fourth segment with a long, curved apical claw and 2 subapical setae, the ventral longer than the dorsal one. Caudal rami consisting of a slender base, a shorter proximal and a longer apical seta. Postabdomen elongated, lobed and twisted, and bearing a long subapical seta (Fig. 1D, E).

Measurements L = 540–560 mm; H = 250–260 mm (n = 3)

Ecology and distribution Thus far, the species is known from its type locality only, where it occurs on patches of moist mud in a remnant of the Atlantic Forest in south-eastern Brazil.

Differential diagnosis This species is at once recognizable by its unique combination of valve and soft part characters, namely short anterior tooth on the LV and the short keel on the RV, typical of the pagliolii-lineage, and the postabdomen with seta, thus far only found in a representative of the danielopoli-lineage. The large anterior LV > RV overlap is unique amongst extant Darwinulidae.



VESTALENULA IRAJAI SP. (FIGS 2A–D, 3A–H)

NOV.

Type locality Tamoios Road, near Paraibuna, São Paulo State, Brazil. Approximate GPS coordinates: 23∞31¢9.6≤S, 45∞32¢21≤W. In small patches of water and amongst leaves and mosses on a concrete surface; water film flowing from rocky wall of this artificial habitat. All material collected on 19.11.1999 by CEFR and JWR.

Type material Holotype: an ovigerous, dissected female, with valves stored dry in a micropalaeontological slide and dissected soft parts kept in a sealed slide (MZUSP 15049). Paratypes: 2 ovigerous females dissected and stored like the holotype (MZUSP 15050, MZUSP 15051); 1 ovigerous, dissected female, with valves used for SEM and stored in a micropalaeontological slide and dissected soft parts kept in a sealed slide (MZUSP 15052); 3 carapaces used for SEM and stored in micropalaeontological slides (MZUSP 15053, MZUSP 15054, MZUSP 15055); 12 females kept in toto in ETOH (MZUSP 15056).

Other material used for description and illustration 1 Sítio da Colonia, Mulungu, near Araçoiaba da Serra, São Paulo State, Brazil. Approximate GPS (Garmin) coordinates: 23∞32¢2≤S, 47∞39¢ 41.1≤W (= type locality of V. botocuda sp.nov.): One ovigerous female collected on 15.08.2001 by CEFR and RLP from moist mud. 2 Fazenda Iterei, Miracatu, São Paulo State, Brazil. Approximate GPS (Garmin) coordinates: 24∞3¢17.8≤S, 47∞13¢7.6≤W. Two ovigerous females collected on 13.9.1999 CEFR and JWR from water and soil of bromeliad pouches.

Derivation of name Named after Prof. Dr Irajá Damiani Pinto (Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil), in recognition of his outstanding contribution to the knowledge of South American Ostracoda in general, and Darwinulidae in particular.

Abbreviated diagnosis Valves elongated (L/H = 2.3), with straight parts of dorsal and ventral margins parallel; ms relatively large; keel on RV short and rounded and rather anteriorly placed; third segment of A1 with ventroapical seta; first endopodal segment of A2 with 2

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apical setae and Md-palp with seta x almost twice as long as seta w.

Description Carapace (Fig. 3A-H) elongated, in lateral view with straight parts of dorsal (c. 40% of total length) and ventral (c. 55%) margins parallel. LV overlapping RV along anterior, ventral and posterior margins; anterior overlap clear. Both valves asymmetrical in dorsal and ventral views, widest in the posterior quarter of the carapace, posterior side with LV bulging over RV, anterior bluntly beak-like, central part with a weak constriction. Ventral margin in ventral view strongly sinuous; keel short and rounded. LV in lateral (internal) view with caudal margin evenly rounded, anterior margin asymmetrically produced towards the ventral side; anterior internal tooth short and small, situated c. 1/5 of total length from the anterior margin; ms large, situated nearer the anterior side and consisting of 8 scars arranged in a neat rosette, 3 anterior and 3 posterior, 1 dorsal and 1 ventral scar; mandibular scar rounded. RV in lateral (internal) view with caudal margin asymmetrically twisted (not visible on illustrations), ventrally everted towards the median rather than the dorsal side, anterior margin everted towards the ventral side; postero-ventral keel short and rounded, situated rather anteriorly, ms large, situated nearer the anterior side, consisting of 7 scars arranged in a rosette (dorsal scar missing). A1 (Fig. 2A) with 1 dorsal seta on first segment and 2 ventral setae on second segment, third segment with a dorsal and a ventral seta. A2 (Fig. 2B) exopodite with one long seta and one short spine; first endopodal segment of A2 with 1 large and one shorter apical seta. Md-palp (Fig. 2C) with seta z long, reaching beyond tip of apical segment; seta y short; seta x about twice as long as seta w; apical segment with 3 long claws, one shorter and one very short near seta c. Caudal rami consisting of a slender base and an apical seta, both about equally long. Postabdomen elongated, lobed and twisted, without seta (Fig. 2D). Other appendages typical of the genus.

Measurements L = 450–465 mm; H = 195–200 mm (n = 3)

Ecology and distribution The species has been found in small patches of water and amongst leaves and mosses from a permanently wet grassy exposed area in its type locality. The species was relatively more abundant (about 40 individuals)


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Figure 2. Vestalenula irajai sp. nov. A. A1, lv (animal lost). B. A2, lv (MZUSP 15052). C. Md-palp, lv (animal lost). D. CR and postabdomen, lv (idem). Scale bars = 20 mm.



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Figure 3. A-H: Vestalenula irajai sp. nov. I-P: Vestalenula botocuda, sp. nov. A. LV, iv (MZUSP 15052). B. RV, iv (idem). C. LV, iv, detail of ms (idem). D. LV, iv, detail of anterior tooth (arrow) (idem). E. RV, iv, detail of ms (idem). F. RV, iv, detail of posterior keel (arrow) (idem). G. Cp, dv (MZUSP 15054). H. Cp, right lv (MZUSP 15053). I. Cp, right lv (MZUSP 15045). J. LV, iv (MZUSP 15042). K. RV, iv (idem). L. Cp, dv, detail of anterior overlap (MZUSP 15046). M. Cp, right lv, detail of anterior overlap (MZUSP 15045). N. Cp, vv, detail of posterior tooth (arrow) blocking LV (MZUSP 15047). O. RV, iv, detail of posterior keel (arrow) (MZUSP 15042). P. Cp, dv (MZUSP 15046). Scale bars: A, B, G-K, P = 100 mm; F, L-O = 20 mm; C-E = 10 mm.


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in a film of water running over a rocky outcrop. Isolated specimens were also found from the same habitat as the preceding species (moist mud patches), as well as from mud and water from bromeliad pouches.

Differential diagnosis Vestalenula irajai sp. nov. is most closely related to V. pagliolii, the soft part morphology of both species being nearly identical. The valves of V. irajai, however, are significantly more elongated with nearly straight dorsal and ventral margins which furthermore run parallel to each other. Anterior overlap in V. irajai is clear but less pronounced than in V. botocuda.

DISCUSSION Terrestrial ostracods have so far been reported from East and South Africa and Madagascar (von Daday, 1910; Klie, 1939; Harding, 1953; Danielopol & Betsch, 1980), Australia (including Tasmania) and New Zealand (De Deckker, 1983), the surroundings of Vladivostok in east Siberia, and the Solomon Islands (Schornikov, 1980) and Europe (Scharf & Keyser, 1991). Terrestricythere ivanovae has (surprisingly) been recorded in deep water in a lake. There is as yet no published information on terrestrial ostracods from the Americas. Martens & Behen (1994) postulated that this was in all probability due to a lack of research effort rather than actual absence, at least in South America. A preliminary sampling campaign of terrestrial and semiterrestrial habitats in the State of São Paulo in 1999 (see Material and methods) yielded 12 species in eight genera (Pinto et al., 2001), of which five species in two genera belong to the Darwinulidae. The diversity of this preliminary survey is surprisingly high and holds great promise for further, more extensive sampling campaigns elsewhere in Brazil and in the rest of South America. Nevertheless, both species described here are found in semiterrestrial rather than fully terrestrial habitats, such as leaf litter, etc. The first fully terrestrial darwinulid remains to be described (R. L. Pinto et al. unpubl. data). The present paper describes two new species in the genus Vestalenula, both of which are presently endemic to the eastern part of Brazil. This confirms the relatively high diversity and endemicity of this genus, at least in the Southern Hemisphere, as postulated by Rossetti & Martens (1998). Vestalenula pagliolii, the only other species of Vestalenula known from South America, has also been reported from the Eemian interglacial of Europe. This species at least shows an intercontinental distribution (Martens et al., 1997). Whereas V. irajai sp. nov. forms part of the Vestalenula pagliolii species cluster, V. botocuda sp. nov. is

unique and enigmatic. Its carapace morphology is unique, as no other extant darwinulid shows a comparable anterior LV > RV overlap. In soft part morphology, its closest relative is V. matildae Martens & Rossetti, 2002 from Australia, as this is the only other species in Vestalenula (and for that matter in all of the Darwinulidae) with a seta on the postabdomen. This latter feature is therefore not unique to the danielopoli-lineage within Vestalenula, as was suggested by Martens & Rossetti (2002). The addition of two new species to the extant Darwinulidae therefore challenges the previous taxonomic classification. The morphology of V. botocuda sp. nov. in particular provides useful information on the morphological evolution and phylogeny of this ancient asexual group. First, the presence of a short ventral keel on the RV together with a seta on the abdomen shows that these features have not necessarily coevolved, as was assumed by Martens & Rossetti (2002). Second, the large frontal overlap of the carapace is unique in the Darwinulidae and questions remain concerning whether or not this is an adaptive feature and whether the developmental pathways followed to obtain this adult structure are similar to those in, for example, several genera in the Cyprinotinae (Cyprididae, Cypridoidea). The discovery of these morphological features in V. botocuda sp. nov. suggest exciting possibilities for comparative morphological analysis.

ACKNOWLEDGEMENTS Janet W. Reid (Martinsville, VA) is gratefully acknowledged for field assistance during the 1999 sampling campaign. Dr Dan Danielopol (Mondsee, Austria) and an anonymous referee provided useful comments on the manuscript. FAPESP allocated a Msc grant to RLP and provided financial assistance to KM during his visits to the University of São Paulo (USP).

REFERENCES Butlin RK, Griffiths HI. 1993. Ageing without sex? Nature 364: 680. von Daday E. 1910. Untersuchungen uber die SüsswasserMikrofauna Deutsch-Ostafrikas. Zoologica 23 (59): 1–314. Danielopol DL. 1968. Microdarwinula n. g., et quelques remarques sur la répartition de la famille Darwinulidae Br. and Norm (Crustacea, Ostracoda). Annals of Limnology 4 (2): 153–174. Danielopol DL. 1970. Une nouvelle espèce du genre Darwinula des eaux souterraines de Roumanie et quelques remarques sur la morphologie des Darwinulidae (OstracodaPodocopida). Travaux de l’Institut Spéléologique ‘Emile Racovitza’ 9: 135–149.


TWO NEW SPECIES OF VESTALENULA (OSTRACODA, DARWINULIDAE) Danielopol DL, Betsch JM. 1980. Ostracodes terrestres de Madagascar: Systématique, origine, adaptations. Revue de l’Écologie et de Biologie du Sol 17: 87–123. De Deckker P. 1983. Terrestrial ostracods in Australia. Australian Museum Memoir 18: 87–100. Harding JP. 1953. The first known example of a terrestrial ostracod, Mesocypris terrestris sp. nov. Annals of the Natal Museum 12: 359–365. Horne DJ, Cohen A, Martens K. 2002. Taxonomy, morphology and biology of Quaternary and living Ostracoda. In: Chivas A, Holmes J, eds. The Ostracoda: applications in Quaternary research. AGU Geophysical Monograph Series, No. 131: 5–36. Judson OP, Normark BB. 1996. Ancient asexual scandals. Trends in Ecology and Evolution 11: 41–52. Klie W. 1939. Ostracoden aus dem Kenia-Gebiet, vornehmlich von dessen Hochgebirgen. Internationale Revue der Gesamte Hydrobiologie und Hydrogeographie 39: 99–161. Martens K. 1998. Sex and ostracods: a new synthesis. In: Martens K, ed. Sex and parthenogenesis – evolutionary ecology of reproductive modes in non-marine ostracods. Leiden: Backhuys, 295–321. Martens K, Behen F. 1994. A checklist of the non-marine ostracods (Crustacea, Ostracoda) from South-American inland waters and adjacent islands. Travaux scientifiques du Musée d’Histoire naturelle de Luxembourg No. 22. Martens K, Rossetti G. 1997. On two new species of Darwinula Brady & Robertson, 1885 (Crustacea, Ostracoda) from South African dolomitic springs. Bulletin van het Koninklijk Belgisch Instituut voor Natuurwetenschappen, Biologie 67: 57–66. Martens K, Rossetti G. 2002. On the Darwinulidae (Crustacea, Ostracoda) from Oceania, with the description of Vestalenula matildae n. sp. Invertebrate Systematics 16: 195–208. Martens K, Rossetti G, Fuhrmann F. 1997. Pleistocene and Recent species of the family Darwinulidae Brady & Norman, 1889 (Crustacea, Ostracoda) in Europe. Hydrobiologia 357: 99–116. Martens K, Würdig NL, Behen F. 1998. Maxillopoda. nonmarine Ostracoda. In: Young PS, ed. Catalogue of Crustacea

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of Brazil. Rio de Janeiro: Museu Nacional, Série Livros 6: 45–65. Pinto ID, Kotzian CB. 1961. Novos Ostrácodes da Família Darwinulidae e a variação das impressões musculares. Boletim do Instituto de Ciências Naturais, Porto Alegre 11: 5–64. Pinto R, Rocha CEF, Martens K. 2001. Terrestrial and semi-terrestrial ostracods (Crustacea, Ostracoda) from Brazil (South America). 14th International Ostracod Symposium, Shizuoka, Japan. Abstracts, 78. Rossetti G, Eagar S, Martens K. 1998. On two new species of the genus Darwinula (Crustacea, Ostracoda) from New Zealand. Italian Journal of Zoology 65: 325–332. Rossetti G, Martens K. 1998. Taxonomic Revision of the Recent and Holocene representatives of the Family Darwinulidae (Crustacea, Ostracoda), with a description of three new genera. Bulletin van het Koninklijk Belgisch Instituut voor Natuurwetenschappen, Biologie 68: 55–110. Rossetti G, Martens K. 1999. Contribution to taxonomy and distribution of the genus Vestalenula Rossetti & Martens, 1998 (Crustacea, Ostracoda, Darwinulidae), with the description of two new species. Bulletin van het Koninklijk Belgisch Instituut voor Natuurwetenschappen, Biologie 69: 57–66. Rossetti G, Martens K, Mourguiart P. 1996. On Darwinula incae Delachaux, 1928. A Stereo-Atlas of Ostracod Shells 23: 35–40. Scharf B, Keyser D. 1991. Living and subfossil Ostracoda (Crustacea) from lac de Bouchet (France, Auvergne). Documents de CERLLAT, Mémoirs 2: 387–391. Schön I, Butlin RK, Griffiths HI, Martens K. 1998. Slow evolution in an ancient asexual ostracod. Proceedings of the Royal Society, London B 265: 235–242. Schornikov EI. 1980. Ostracodes in terrestrial biotopes. Zoologicheskii Zhurnal 59: 1306–1319. (In Russian with English abstract). Van Doninck K, Schön I, De Bruyn L, Martens K. 2002. General purpose genotype in an ancient asexual. Oecologia 132: 205–212.
