Crustacea, Ostracoda, Darwinulidae) from Semiterrestrial Habitats in São Paulo State (Brazil)

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

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 Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020 1Departamento de Zoologia-IBUSP, Rua do Matão, trav. 14, n°101, CEP05508–900 São Paulo–SP, Brazil 2Royal Belgian Institute of Natural Sciences, Freshwater Biology, Vautierstraat 29, 1000 Brussels, Belgium 3University of Ghent, Department of Biology, K. L. Ledeganckstraat 35, B9000 Gent, Belgium

Received March 2002; accepted for publication February 2003

Two new species of semiterrestrial darwinulid ostracods, both belonging to the pagliolii-lineage of the genus Ves- talenula, 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 1997), South Africa (Martens & Rossetti, 1997; Ros- setti & Martens, 1999), New Zealand and Australia Darwinulidae are so-called ancient asexuals (Butlin & (Rossetti, Eagar & Martens, 1998; Rossetti & Mar- Grifﬁths, 1993; Judson & Normark, 1996), organisms tens, 1999; Martens & Rossetti, 2002). This culmi- which have persisted over geologically long periods of nated in a world-wide revision of the group, which time without sexual reproduction. There appear to introduced three new genera, bringing the total num- have been no bisexual populations for at least 150 Myr ber to ﬁve extant genera (Rossetti & Martens, 1998). (Martens, 1998). All living Darwinuloidea belong to one Most work on South American darwinulids dates subfamily, the Darwinulinae Brady & Norman, 1889, from before 1970, with the exception of Rossetti, Mar- although the Palaeozoic–Recent fossil record of this tens & Mourguiart (1996), and only the paper by Pinto superfamily comprises representatives of eight sub- & Kotzian (1961) deals with the detailed taxonomy of families in two families. the Darwinulidae. Martens & Behen (1994) list the For a long time, Recent darwinulids, which are species of darwinulid ostracods at that stage reported exclusively nonmarine, were treated as a marginal from South America (see Martens, Würdig & Behen, group and little attention was devoted to them. How- 1998, for an account of Brazilian darwinulids). How- ever, their recognition as putative ancient asexuals ever, several taxonomic alterations to this list were initiated renewed interest, and provoked a series of suggested by Rossetti & Martens (1998): all species ecological (Van Doninck et al. 2002), molecular (Schön except D. stevensoni were transferred to one of the et al., 1998) and taxonomic studies. Several taxonomic three new genera; Penthesilenula brasiliensis is now papers have presented regional surveys of darwinulid considered a valid species, not a subspecies of P. afri- faunas, e.g. Europe (Martens, Rossetti & Fuhrman, cana and Alicenula serricaudata espinosa was synon- ymized with the nominal subspecies. Taking into

*Corresponding author: E-mail. [email protected] account these alterations, together with the two new

306 R. L. PINTO ET AL. species described here, nine darwinulid species are Muddy patches in a dry streamlet in a rain forest presently known from South America. remnant. The mud was moist at the time of collecting; during wet season standing or running water might be MATERIAL AND METHODS present. Material collected on 15.08.2001 by CEFR and RLP. 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 Type material (CEFR) and J. W. Reid (JWR) in 1999 and by C. E. F. Holotype: an ovigerous, dissected female, with valves Rocha and R. L. Pinto (RLP) in 2001. Ostracods were stored dry in a micropalaeontological slide and dis- collected live from leaf litter, mosses, etc. and killed sected soft parts kept in a sealed slide (MZUSP 15041). and preserved in 70% ethanol. These collections con- Paratypes: an ovigerous female dissected and stored Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020 tained several species from the families Darwinulidae, like the holotype (MZUSP 15043); 2 ovigerous, dis- Cyprididae and Candonidae (Pinto, Rocha & Martens, sected females, with valves used for SEM and stored 2001). The present paper deals with the darwinulid in micropalaeontological slides, and dissected soft genus Vestalenula only, other (new as well as known) parts kept in sealed slides (MZUSP 15042, MZUSP species will be reported on in detail elsewhere. 15044); 3 carapaces used for SEM and stored in micro- Dissections were effected with micro needles under palaeontological slides (MZUSP 15045, MZUSP a stereo-light microscope. Valves were stored dry in 15046, MZUSP 15047); 4 ovigerous females kept in micropalaeontological slides; soft parts were dissected toto in ETOH (MZUSP 15048). in CMC/9AF medium with Rose Bengal and were mounted in permanent slides. Micrographs of valves were made with a Zeiss Scanning Electron Microscope. Derivation of name Deposition of material: all type material is deposited Botocudos is the designation of several South Ameri- in the Museu de Zoologia da Universidade de São can Indian tribes that use a device to enlarge the Paulo (MZUSP). The following abbreviations were lower lip. The present species is characterized by the used in text and figures: fact that the LV forms an enlarged anterior ‘lip’, Valves. Cp, carapace; H, height; L, length; LV, left valve; greatly overlapping the RV. RV, right valve; ms, central muscle scar(s); dv, dorsal view; vv, ventral view; lv, lateral view; iv, internal view. Abbreviated diagnosis Limbs and soft parts. A1, antennula; A2, antenna; Md, Valves elongated (L/H = c. 2.1), with straight parts of mandibula; Mx, maxillula; T1-3, thoracic limbs; CR, dorsal margin slightly sloping towards the front; ante- caudal ramus (previously named furca); exo, exopodite rior overlap LV > RV unusually pronounced and typi- on A2; Ac, ventral aesthetasc clump on A2; s1, s2, t, w, cal; ms relatively large; keel on RV short and rounded x, y, specific setae on limbs; y1–3, aesthetascs. and positioned rather posteriorly; third segment of A1 Nomenclature proposed by Danielopol (1968, 1970), with ventro-apical seta. First endopodal segment of A2 and adapted by Rossetti & Martens (1998) is used in with 2 apical setae; seta z on penultimate segment the description of chaetotaxy of soft parts. For higher claw-like. Md-palp with seta x longer than w. taxonomy, the system proposed by Horne, Cohen & Martens (2002) is followed. Description TAXONOMIC DESCRIPTIONS Cp (Fig. 3I-P) with whitish, smooth surface covered CLASS OSTRACODA LATREILLE, 1806 SUBCLASS with long, widely spaced setae; elongated, in lateral PODOCOPA G.W. MÜLLER, 1894 ORDER PODOCOPIDA view, with straight parts of dorsal (c. 30% of total SARS, 1866 SUBORDER PODOCOPINA SARS, 1866 length) and ventral (c. 55%) margins subparallel, dor- SUPERFAMILY DARWINULOIDEA BRADY & NORMAN, sal margin slightly sloping. LV overlapping RV along anterior, ventral and posterior margins; anterior over- 1889 FAMILY DARWINULIDAE BRADY & NORMAN, 1889 lap very pronounced, LV extending significantly GENUS VESTALENULA ROSSETTI & MARTENS, 1998 beyond RV. Both valves asymmetrical in dorsal and VESTALENULA BOTOCUDA SP. NOV. ventral views, widest in the posterior quarter of the (FIGS 1A–E, 3I–P) carapace, LV bulging over RV on the posterior side, Type locality anteriorly extending significantly beyond RV; central Sítio da Colônia, Mulungu, near Araçoiaba da Serra, part with a weak constriction. Ventral margin in ven- São Paulo State, Brazil. Approximate GPS coordi- tral view strongly sinuous; keel short and rounded, nates: 23∞32¢2≤S, 47∞39¢ 41.1≤W. effectively blocking LV in tightly closed carapaces.

TWO NEW SPECIES OF VESTALENULA (OSTRACODA, DARWINULIDAE) 307 Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020

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.

308 R. L. PINTO ET AL.

LV in lateral (internal) view with caudal margin Mx (not illustrated). Palp two-segmented; proximal nearly evenly rounded, slightly everted towards the segment with an external, subapical seta, 3 terminal dorsal side, anterior margin everted towards the ven- setae (2 straight and 1 curved, the latter shorter and tral side; anterior internal tooth short and small, sit- plumose), and 2 short setae inserted on either side of uated c. 15% of total length from the anterior margin; the terminal segment; terminal segment small, with 1 ms large, positioned nearer the anterior side and con- thin median seta and 2 subequal, stout setae set with sisting of c. 8 scars arranged in a rosette; mandibular a double row of setulae. scar rounded. T1 (not illustrated). Protopodite with branchial RV in lateral (internal) view with caudal margin plate, 2 isolated, unequal setae close to the articula- asymmetrically twisted, ventrally everted towards the tion with the endopodite. Three-jointed endopodite; median rather than dorsal side, anterior margin ﬁrst segment with 2 subapical setae; second seg-

everted towards the ventral side; postero-ventral keel ment with 1 subapical seta; third segment smaller, Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020 short and rounded, situated rather posteriorly, ms with 2 lateral, subequal setae and a strong terminal large, situated nearer the anterior side, consisting of claw. c. 8 scars arranged in a rosette. T2 (not illustrated). Protopodite one-segmented, Both valves with an internal rim along the anterior rather stout, with 3 ventral setae, 1 short proximal margin (= calcified inner margin?); in LV this rim seals and 2 long, unequal distal ones. Endopodite four-seg- the closure of the carapace when overlapping the RV. mented; first segment with 2 unequal ventro-apical A1 (Fig. 1A). First segment stout with dorsal seta, setae and 1 long seta, slightly exceeding the tip of the the second with 2 unequal ventral setae (one c. 2.5 next segment; second and third segment with 1 stout times as long as the other); third segment with 1 ven- ventro-apical seta each; terminal segment short, with tral and 1 dorsal setae (s1); fourth segment with 1 dor- 1 strong apical claw, flanked by 2 subapical claws, the sal seta (s2) and small alpha-seta; fifth podomere ventral longer than the dorsal one. bearing 4 apical setae, 2 subequal, ventral setae (with T3 (not illustrated). Protopodite one-segmented, an alpha-seta near their insertion) and 2 longer ones short, with 2 unequal ventro-apical setae. Endopodite in dorsal position; last podomere with subapical aes- four-jointed; first segment with 1 distal seta approxi- thetasc, 2 long apical setae, the ventral seta about half mately as long as the segment itself; second and third as long as the dorsal, and a dorso-apical ‘alpha’ seta. segments bearing 1 apical seta each, reaching beyond A2 (Fig. 1B). Protopodite two-segmented, first seg- the tip of the next segment; fourth segment with a ment with 2 setae and dorsal hook (h), second seg- long, curved apical claw and 2 subapical setae, the ment with long seta. Exopodite (exo) with long, apical ventral longer than the dorsal one. seta and short, lateral conical spine. Endopodite Caudal rami consisting of a slender base, a shorter three-segmented; first podomere with ventral aes- proximal and a longer apical seta. Postabdomen elon- thetasc clump (Ac) in proximal position and 2 setae gated, lobed and twisted, and bearing a long subapical on the expanded ventro-apical corner; second seta (Fig. 1D, E). 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 Measurements short (x); third endopodite short and slender, with one L = 540–560 mm; H = 250–260 mm (n = 3) 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 pen- Ecology and distribution ultimate segment. Thus far, the species is known from its type locality Md with palp (Fig. 1C) three-segmented, consist- only, where it occurs on patches of moist mud in a rem- ing of basis and two-segmented endopodite; basis nant of the Atlantic Forest in south-eastern Brazil. 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 dis- Differential diagnosis tally widened, medial seta x about 1.5 times as long This species is at once recognizable by its unique com- as the external, subapical seta (w); two internal bination of valve and soft part characters, namely setae present: y very short, z very long, reaching short anterior tooth on the LV and the short keel on halfway the apical claws on the terminal segment; the RV, typical of the pagliolii-lineage, and the post- this podomere narrow, with 5 apically hirsute, abdomen with seta, thus far only found in a represen- unequal claws (3 long, 1 intermediate, 1 short), 1 tative of the danielopoli-lineage. The large anterior short internal seta (c), and a subapical external seta LV > RV overlap is unique amongst extant Darwinul- (b). idae.

VESTALENULA IRAJAI SP. NOV. apical setae and Md-palp with seta x almost twice as (FIGS 2A–D, 3A–H) long as seta w. Type locality Tamoios Road, near Paraibuna, São Paulo State, Description Brazil. Approximate GPS coordinates: 23∞31¢9.6≤S, Carapace (Fig. 3A H) elongated, in lateral view with 45∞32¢21≤W. In small patches of water and amongst - leaves and mosses on a concrete surface; water film straight parts of dorsal (c. 40% of total length) and flowing from rocky wall of this artificial habitat. All ventral (c. 55%) margins parallel. LV overlapping RV material collected on 19.11.1999 by CEFR and JWR. 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, Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020 Type material anterior bluntly beak-like, central part with a weak Holotype: an ovigerous, dissected female, with valves constriction. Ventral margin in ventral view strongly stored dry in a micropalaeontological slide and dis- sinuous; keel short and rounded. sected soft parts kept in a sealed slide (MZUSP LV in lateral (internal) view with caudal margin 15049). evenly rounded, anterior margin asymmetrically pro- Paratypes: 2 ovigerous females dissected and stored duced towards the ventral side; anterior internal tooth like the holotype (MZUSP 15050, MZUSP 15051); 1 short and small, situated c. 1/5 of total length from the ovigerous, dissected female, with valves used for SEM anterior margin; ms large, situated nearer the ante- and stored in a micropalaeontological slide and dis- rior side and consisting of 8 scars arranged in a neat sected soft parts kept in a sealed slide (MZUSP rosette, 3 anterior and 3 posterior, 1 dorsal and 1 ven- 15052); 3 carapaces used for SEM and stored in micro- tral scar; mandibular scar rounded. palaeontological slides (MZUSP 15053, MZUSP RV in lateral (internal) view with caudal margin 15054, MZUSP 15055); 12 females kept in toto in asymmetrically twisted (not visible on illustrations), ETOH (MZUSP 15056). ventrally everted towards the median rather than the dorsal side, anterior margin everted towards the ventral side; postero-ventral keel short and rounded, sit- Other material used for description and illustration uated rather anteriorly, ms large, situated nearer the 1 Sítio da Colonia, Mulungu, near Araçoiaba da anterior side, consisting of 7 scars arranged in a Serra, São Paulo State, Brazil. Approximate GPS rosette (dorsal scar missing). (Garmin) coordinates: 23∞32¢2≤S, 47∞39¢ 41.1≤W (= A1 (Fig. 2A) with 1 dorsal seta on ﬁrst segment and type locality of V. botocuda sp.nov.): One ovigerous 2 ventral setae on second segment, third segment with female collected on 15.08.2001 by CEFR and RLP from a dorsal and a ventral seta. moist mud. A2 (Fig. 2B) exopodite with one long seta and one 2Fazenda Iterei, Miracatu, São Paulo State, Brazil. short spine; ﬁrst endopodal segment of A2 with 1 large Approximate GPS (Garmin) coordinates: 24∞3¢17.8≤S, and one shorter apical seta. 47∞13¢7.6≤W. Two ovigerous females collected on Md-palp (Fig. 2C) with seta z long, reaching beyond 13.9.1999 CEFR and JWR from water and soil of bro- tip of apical segment; seta y short; seta x about twice meliad pouches. 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 api- Derivation of name cal seta, both about equally long. Postabdomen elon- Named after Prof. Dr Irajá Damiani Pinto (Univer- gated, lobed and twisted, without seta (Fig. 2D). sidade Federal do Rio Grande do Sul, Porto Alegre, Other appendages typical of the genus. Brazil), in recognition of his outstanding contribution to the knowledge of South American Ostracoda in general, and Darwinulidae in particular. Measurements L = 450–465 mm; H = 195–200 mm (n = 3)

Abbreviated diagnosis Valves elongated (L/H = 2.3), with straight parts of Ecology and distribution dorsal and ventral margins parallel; ms relatively The species has been found in small patches of water large; keel on RV short and rounded and rather and amongst leaves and mosses from a permanently anteriorly placed; third segment of A1 with ventro- wet grassy exposed area in its type locality. The species apical seta; ﬁrst endopodal segment of A2 with 2 was relatively more abundant (about 40 individuals)

© 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139, 305–313 310 R. L. PINTO ET AL. Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020

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.

© 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139, 305–313 TWO NEW SPECIES OF VESTALENULA (OSTRACODA, DARWINULIDAE) 311 Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020

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.

© 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139, 305–313 312 R. L. PINTO ET AL. in a ﬁlm of water running over a rocky outcrop. Iso- unique and enigmatic. Its carapace morphology is lated specimens were also found from the same habitat unique, as no other extant darwinulid shows a com- as the preceding species (moist mud patches), as well parable anterior LV > RV overlap. In soft part mor- as from mud and water from bromeliad pouches. phology, 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 Differential diagnosis all of the Darwinulidae) with a seta on the postabdo- Vestalenula irajai sp. nov. is most closely related to men. This latter feature is therefore not unique to V. pagliolii, the soft part morphology of both species the danielopoli-lineage within Vestalenula, as was being nearly identical. The valves of V. irajai, however, suggested by Martens & Rossetti (2002). The addi- are signiﬁcantly more elongated with nearly straight tion of two new species to the extant Darwinulidae

dorsal and ventral margins which furthermore run therefore challenges the previous taxonomic classifi- Downloaded from https://academic.oup.com/zoolinnean/article/139/2/305/2629569 by guest on 23 November 2020 parallel to each other. Anterior overlap in V. irajai is cation. The morphology of V. botocuda sp. nov. in clear but less pronounced than in V. botocuda. particular provides useful information on the morphological evolution and phylogeny of this ancient asexual group. First, the presence of a short ventral DISCUSSION keel on the RV together with a seta on the abdomen Terrestrial ostracods have so far been reported from shows that these features have not necessarily East and South Africa and Madagascar (von Daday, coevolved, as was assumed by Martens & Rossetti 1910; Klie, 1939; Harding, 1953; Danielopol & Betsch, (2002). Second, the large frontal overlap of the cara- 1980), Australia (including Tasmania) and New pace is unique in the Darwinulidae and questions Zealand (De Deckker, 1983), the surroundings of remain concerning whether or not this is an adaptive Vladivostok in east Siberia, and the Solomon Islands feature and whether the developmental pathways (Schornikov, 1980) and Europe (Scharf & Keyser, followed to obtain this adult structure are similar to 1991). Terrestricythere ivanovae has (surprisingly) those in, for example, several genera in the Cyprino- been recorded in deep water in a lake. There is as yet tinae (Cyprididae, Cypridoidea). The discovery of no published information on terrestrial ostracods from these morphological features in V. botocuda sp. nov. the Americas. Martens & Behen (1994) postulated suggest exciting possibilities for comparative mor- that this was in all probability due to a lack of phological analysis. research effort rather than actual absence, at least in South America. A preliminary sampling campaign of terrestrial and semiterrestrial habitats in the State of ACKNOWLEDGEMENTS São Paulo in 1999 (see Material and methods) yielded 12 species in eight genera (Pinto et al., 2001), of which Janet W. Reid (Martinsville, VA) is gratefully acknowl- five species in two genera belong to the Darwinulidae. edged for field assistance during the 1999 sampling The diversity of this preliminary survey is surpris- campaign. Dr Dan Danielopol (Mondsee, Austria) and ingly high and holds great promise for further, more an anonymous referee provided useful comments on extensive sampling campaigns elsewhere in Brazil the manuscript. FAPESP allocated a Msc grant to and in the rest of South America. Nevertheless, both RLP and provided financial assistance to KM during species described here are found in semiterrestrial his visits to the University of São Paulo (USP). 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). REFERENCES The present paper describes two new species in the Butlin RK, Griffiths HI. 1993. Ageing without sex? Nature genus Vestalenula, both of which are presently 364: 680. endemic to the eastern part of Brazil. This confirms von Daday E. 1910. Untersuchungen uber die Süsswasser- the relatively high diversity and endemicity of this Mikrofauna Deutsch-Ostafrikas. Zoologica 23 (59): 1–314. genus, at least in the Southern Hemisphere, as postu- Danielopol DL. 1968. Microdarwinula n. g., et quelques lated by Rossetti & Martens (1998). Vestalenula pagli- remarques sur la répartition de la famille Darwinulidae Br. olii, the only other species of Vestalenula known from and Norm (Crustacea, Ostracoda). Annals of Limnology 4 South America, has also been reported from the (2): 153–174. Eemian interglacial of Europe. This species at least Danielopol DL. 1970. Une nouvelle espèce du genre Dar- shows an intercontinental distribution (Martens et al., winula des eaux souterraines de Roumanie et quelques 1997). remarques sur la morphologie des Darwinulidae (Ostracoda- Whereas V. irajai sp. nov. forms part of the Vestal- Podocopida). Travaux de l’Institut Spéléologique ‘Emile Raco- enula pagliolii species cluster, V. botocuda sp. nov. is vitza’ 9: 135–149.

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