Zootaxa, On Amphibolocypris arida sp.nov. (Crustacea, Ostracoda ...

Zootaxa 2408: 47–58 (2010) www.mapress.com / zootaxa/

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On Amphibolocypris arida sp.nov. (Crustacea, Ostracoda), from rock pools in Botswana (southern Africa) MERLIJN JOCQUE1,4, LUC BRENDONCK1, BRUCE J RIDDOCH2 & KOEN MARTENS3 1 K.U.Leuven, Laboratory of Aquatic Ecology and Evolutionary Biology, De Bériotstraat 32, 3000 Leuven, Belgium and Central Laboratory of General Ecology, Bulgarian Academy of Sciences, 2 Yurii Gagarin Street, 1113 Sofia, Bulgaria. 2 School of Life Sciences, Oxford Brookes University, Headington Campus, Gipsy Lane, Oxford, OX3 0BP, U.K. 3 Koninklijk Belgisch Instituut voot Natuurwetenschappen, Afdeling Zoetwaterbiologie (Royal Belgian Institute of Natural Sciences, Freshwater Biology), Vautierstraat 29, B-1000 Brussels, Belgium and University of Ghent, Biology, K.L. Ledeganckstraat 35, B9000 Gent, Belgium 4 Corresponding author. Laboratory of Aquatic Ecology and Evolutionary Biology, Katholieke Universiteit Leuven, Ch. De Bériotstraat 32, B-3000 Leuven, Belgium. phone: ++ 32-16-323966, fax: ++ 32-16-32.45.75, E-mail: [email protected]

Abstract The ostracod fauna of southern Africa remains ill-known, in spite of the fact that the temporary pools of this zoogeographical region hold degrees of endemicity comparable only to those of the ancient lakes of East Africa. The present contribution describes a new species of the cypridid genus Amphibolocypris, A. arida sp.nov., and announces the existence of at least two further new species from the same area. The genus, up to now presumed monospecific, might constitute an extensive radiation across southern Africa. Unusually large species-specific differences in hemipenis outline morphology appear to indicate that speciation occurred through sexual, rather than through natural selection. The occurrence of the claw-like subapical seta on the walking limb in at least four genera could be a case of convergent evolution, at least in one, maybe even in two cases between species of different genera and even subfamilies. Key words: phylogeny, taxonomy, temporary pools, sexual selection, convergent evolution, western Cape

Introduction Martens et al. (2008) estimated that there are about 2000 described species in c 200 genera of extant nonmarine ostracods world-wide. However, the ostracod fauna of southern Africa remains relatively ill-known, this in spite of more than a century of taxonomic research (for overviews, see Sars 1924, Martens 1984, 2001). Martens (2001) retained about 150 described species in southern Africa (region as defined by Day et al. 2001), but the current limited coverage of this region implies that many more as yet undescribed species await description or discovery. Since the 2001 review, several other new species and genera have been described (e.g. Martens 2007, Savatenalinton & Martens 2009) and some presumed extinct taxa were found again, notably Liocypris grandis (by Martens 2003, Matzke-Karasz & Martens 2005) and Afrocypris barnardi (by Matzke-Karasz & Martens 2007). These giant ostracods are confined to temporary (rock-) pools, habitats already noted by Martens (1998) to harbour a rich and largely endemic ostracod fauna. With regard to levels of generic endemicity, these pools in the (mainly western) part of southern Africa are comparable only to those of the ancient lakes in East Africa (Tanganyika, Malawi) (Martens 1998). Here, we describe a new species of the subfamily Isocypridinae. Amphibolocypris arida sp.nov. is only the second representative described from the genus. The present description is based on material from a wider survey, comparing the biodiversity in rock pools in three separate geographical areas: southern Africa (Jocque et al. 2006), Western Australia (Jocque et al. 2007a) and Utah (USA) (Jocque et al. 2007b).

Accepted by R. Matzke-Karasz: 23 Jan. 2010; published: 24 Mar. 2010

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Material and methods Material: The material described here was collected during a fieldtrip between 15 January and 2 March 2003 to the Hardveld zone of south-eastern Botswana, near the village of Thamaga (see coordinates below and Figure 1). Rainfall in the area is mostly between October and April, which includes the sampling period, and mean annual rainfall is 475 mm (Jocque et al. 2006), but varies both within and between wet seasons. The sampling protocol was described by Jocque et al. (2006). Some additional material, used for comparative purposes, was collected by various explorers in Namibia in the 1980s.

FIGURE 1. Map of the type locality of Amphibolocypris arida sp.nov. (after Jocque et al. 2006).

Samples were preserved in 70% ethanol and sorted with a Leica Wild M-10 binocular microscope. Soft parts were dissected in glycerine and sealed on a glass slide. Valves were stored dry in micropalaeontological slides. Drawings of soft parts were made using camera lucida. Valves were observed and illustrated using Scanning Electron Microscopy (Philips XL30 SEM). All type specimens are stored in The Ostracod Collection of the Royal Belgian Institute of Natural Sciences (Brussels, Belgium). Abbreviations used in text and figures A1 = Antennula. A2 = Antenna. Cp = carapace. CR = Caudal Ramus. H = height of valves. L = length of valves. ls= lateral shield of hemipenis. LV = left valve. Md = Mandibula. ms= medial shield of hemipenis. Mx1 = Maxillula. RV = right valve. T1 = first thoracopod. T2 = second thoracopod. T3 = third thoracopod. Chaetotaxy of the limbs follows the model proposed by Broodbakker & Danielopol (1982), revised for the A2 by Martens (1987) and for the T3 by Meisch (2000). Higher taxonomy of the Ostracoda follows the synopsis by Horne et al. (2002).

Results Taxonomic descriptions Class Ostracoda Latreille, 1806

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Subclass Podocopa G.W. Müller, 1894 Order Podocopida Sars, 1866 Suborder Cypridocopina Baird, 1845 Superfamily Cypridoidea Baird, 1845 Family Cyprididae Baird, 1845 Subfamily Isocypridinae Rome, 1965 Synonymy: Amphibolocypridinae Hartmann & Puri, 1974; Platycypridinae Hartmann & Puri, 1974

Diagnosis: Medium-sized ostracods, with laterally flattened and elongated valves, posterior calcified inner lamella either narrow or completely absent. Walking limbs (T2) large, with long setae and claws, and elongated segments 3, 4a and 4b; these limbs mostly sticking out of the carapace. Remarks: Rome (1965) defined the subfamily Isocypridinae (with genera Isocypris G.W. Müller, 1908, Platycypris Herbst, 1957 and Amphibolocypris Rome, 1965) based on a single character, namely the fact that the posterior inner margin coincides with the posterior valve margin (“... le bord interne coincide avec le bord postérieur.”). However, in Isocypris nivea Sars, 1924, as well as in the present new species of Amphibolocypris, the posterior calcified inner lamella is narrow, but present. The diagnosis of the subfamily is therefore here amended. More defining characters for the present subfamily might be found in the distal chaetotaxy of the coxal plate of the Mx1, but this would require a revision of the variability of this character across several subfamilies, and this is beyond the scope of the present paper.

Key to the genera of Isocypridinae (after Rome 1965) 1. 2. -

5th (terminal) segment of T2 with a short, subapical seta ...................................................... Isocypris (cosmopolitan) 5th (terminal) segment of T2 with claw-like subapical seta, reaching beyond the middle of the end claw .................. 2 CR with a proximal seta............................................................................................................. Platycypris (Australia) CR without proximal seta .............................................................................................. Amphibolocypris (SW Africa)

Amphibolocypris Rome, 1965 Type species: Amphibolocypris exigua Rome, 1965 (by original designation)

Origin of name: ‘Amphibolos’, from the Greek 'Amphibolein' = ambiguous, as this genus shares characters of both Isocypris and Platycypris, together with 'Cypris'. Diagnosis (amended from Rome 1965): Carapace elongated, narrow in dorsal view. Third endite of Mx1 with 2 large, smooth Zahnborsten. T2 (walking limb) with dorsal seta on 4th segment long and claw-like. CR without proximal seta.

Amphibolocypris arida sp.nov. Jocque & Martens (Figs 2–5) Type locality: Rock pools near Thamaga, south-eastern Botswana (24˚ 40’ 30” S, 25˚ 31’00” E). The pools are situated on granite rocky outcrops, surfacing in a mainly savannah matrix of grasslands and shrubs. The pools are generally shallow (max 20 cm deep) and small (max. 2 m radius), with water temperature reaching up to ON AMPHIBOLOCYPRIS ARIDA SP.NOV.

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37.1 °C at mid day. Based on measurements in 8 pools, habitat waters were characterized by a basic pH varying between 7.3 and 8.8 and were always low in conductivity with values ranging from 27.6 to 69.1 µSm/cm (Jocque, unpubl.). Accompanying ostracod fauna: Heterocypris sp.nov., Heterocypris incongruens, Sarscypridopsis cf gregaria, Potamocypris spec.

FIGURE 2. Amphibolocypris arida sp.nov., male and female. A. Male, LV, exterior view (OC.3167). B. Idem, RV, interior view. C. Female, LV, interior view (OC.3166). D. Idem, RV, exterior view. Specimen slightly damaged. E. Female, RV, exterior view, detail (OC.3168). F. Idem, higher magnification. Scale: A-D,G = 500 μm, E = 20 μm, F = 5 μm.

Type material: Holotype: a male, with soft parts dissected in glycerine in a sealed slide and with valves stored dry (after use for SEM) in a micropalaeontological slide (OC.3167) Allotype: a female, with soft parts dissected in glycerine in a sealed slide and with valves stored dry (after use for SEM) in a micropalaeontological slide (OC.3168) Paratypes: 2 males and 2 females, with soft parts dissected in glycerine in a sealed slide and with valves lost (OC.3166, OC.3169-OC.3171) Derivation of name: as the species is thus far only known from rock pools in a (semi-) arid area, the specific name refers to this characteristic of its habitat. Diagnosis: Valves elongated, with anterior margin more broadly rounded, posterior margin bluntly pointed, posterior calcified inner margin narrow, but present. External surface set with elongated microridges. Valves sexually dimorphic, with female valves being more elongated. Natatory setae on A2 of medium length. T2 with subapical seta claw-like, reaching middle of elongated end claw. CR without proximal seta. Hemipenis with one shield with one club-like expansion, the other shield with two elongated lobes, one pointed, basally inserted, and one distally rounded, more apically inserted. Prehensile palps almost symmetrical, relatively short, with broad base and sickle-shaped.

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FIGURE 3. Amphibolocypris arida sp.nov., male and female. A. Male, A1 (OC.3167), B. Male, A2 (OC.3167). C. Female, A2 (OC.3166). D. Female, Md, detail of coxal plate (OC.3166). E. Female, Md-palp, chaetotaxy of terminal segment not shown (OC.3166). E’. Female, Md-palp, terminal segment (OC.3166). Scale: A = 100 μm, B-E’ = 78 μm. ON AMPHIBOLOCYPRIS ARIDA SP.NOV.


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FIGURE 4. Amphibolocypris arida sp.nov., male and female. A. Female, Mx1: palp and 3 endites, respiratory plates not shown (OC.3166). B. Female, T2 (OC.3166). C. Female, T3 (OC.3166). C’. female, T3, detail of distal pincer (OC.3171). D. Female, attachment of CR (OC.3166). E. Male, CR (OC.3167). F. Female, T1 (OC.3166). Scale : A-C,D = 78 μm, C ’= 29 μm, E,F = 146 μm.

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FIGURE 5. Amphibolocypris arida sp.nov., all male. A. Hemipenis (OC.3167). B. Left, prehensile palp (OC.3167). C. Right T1, with prehensile palp (OC.3167). Scale: A-C = 78 μm.

Additional description of male: Valves in lateral view (Figs 2A,B) elongated, weakly calcified, externally set with longitudinal micro-ridges and sparse rimmed pores (Figs 2E,F = from female valves). with rounded dorsal margin, anterior margin more broadly rounded than posterior margin, ventral margin slightly sinuous. Valve outline antero-dorsally with a dent. Anterior calcified inner lamella broad, dorsally ending at dent; posterior calcified inner lamella narrow, but present. Muscle scars situated in front of the middle. Carapace in dorsal view very narrow, lancet shape. No overlap between LV and RV. A1 (Fig. 3A) with 7 segments. First segment with 2 ventral setae, no dorsal seta. Second segment with one medio-dorsal seta, Rome and Wouters organs not visible. Third segment more than twice as long and wide, with one shorter ventral and one longer dorsal seta. Fourth segment almost as long as wide, with 2 shorter ventral setae and 2 long dorsal natatory setae. Fifth segment slightly longer than wide, also with two shorter ventral and 2 longer dorsal natatory setae. Sixth segment almost twice as long as wide, with 1 short ventral setae and 4 long natatory setae. Terminal (seventh) segment more than twice as long as wide, with 1 shorter seta, one long aesthetasc Ya (not drawn) and 2 long natatory setae. A2 (Fig. 3B) with exopodite reduced to a small plate, bearing 1 long and 2 short setae. Endopodite 3segmented. First segment elongated and stout, aesthetasc Y short and club-like, distal part swollen. Natatory setae of unequal length, but all relatively short, mostly not reaching tip of second segment; ventral-most seta the longest. Second segment with 2 dorso-lateral and 3 ventro-lateral setae (2 long, 1 short), distal chaetotaxy typical of male Cyprididae, with setae z2 and z3 changed into claws, z1 a seta and claw G1 turned into a long seta, extending beyond tips of all claws, these claws stout, relatively short and set with strong spines. Terminal segment with claws GM and Gm, seta g and aesthetasc y3, basally fused with an accompanying seta. Md (compare to Figs 3D,E,E’ = female) with coxal plate distally set with rows of spines and small setae. Palp with alpha-seta short, narrow and smooth, beta-seta short, stout and hirsute, gamma-seta short, broad and

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hirsute in distal fourth of its length. Second segment dorsally with a group of 3 smooth setae, ventrally with 3 long and smooth and 1 shorter and hirsute setae. Third segment dorsally with 4 subapical setae, ventrally with 1 subapical seta, distally with 3 normal setae and a gamma-seta. Terminal segment with 3 stout claws and 3 small setae. Mx1 (compare to Fig. 4A = female) with second palp-segment slightly spatulate, Zahnborsten on third endite smooth. T1 (Figs 5A, B) with 1 a-seta, 1 b- and 1 d-setae. Distal chaetotaxy of coxal plate consisting of 16 setae of sometimes very different shape and length. Prehensile palps almost symmetrical; first segments with large ventral protuberance, set with 1 large sensory organ; second segments sickle-shaped, tapering towards the end, distally with one long and stout sensory organ. T2 (compare to Fig. 4B= female) with elongated segments and unusually long setae. First segment with seta d1 reaching into second segment, this (knee-) segment with seta d2 missing. Third segment with 1 long ventro-apical seta, reaching almost to fifth segment. Fourth segment divided into two elongated subsegments: segment 4a with a relatively short ventro-apical seta, not reaching tip of segment 4b, this latter segment with 2 subequal, ventro-apical setae. Fifth (terminal) segment with 1 subapical seta and 1 subapical claw (generic character) and 1 long and thin apical claw. T3 (compare to Figs 4C,C’= female) a cleaning limb. First segment with 3 setae. Second segment with 1 long apical seta. Third segment with 1 short lateral seta. Distal part of third and fourth segment fused to a pincer shaped organ, bearing 1 long seta, 1 seta of medium length, set with two rows of setulae and 1 very short seta. CR (Fig. 4E) distally with 2 claws and 1 apical seta, proximal seta missing (generic character). Attachment (Fig. 4D = female) slender, with simple distal bifurcation. Hemipenis (Fig. 5C) with a complex of ventral lobes of medial and lateral shields: one shield with one club-like expansion, the other shield with two elongated lobes, one pointed, basally inserted, one distally rounded, more apically inserted. Additional description of female: Female valves more elongated, dorsal margin bluntly pointed, with greatest height less than ½ the length and situated almost in the middle. Anterior margin more rounded than posterior margin, but both less so than in the male. Calcified inner lamellae and position of muscle scars as in the male. A1, Md (Figs 3D,E,E’), Mx1 (Fig. 4A) and T3 (Fig. 4C,C’) as in the male. A2 (Fig. 3C) with natatory setae slightly longer than in the male, z1-3 all setae, reaching up to or beyond tips of end claws, the latter more slender and longer than in the male and set with more delicate spines. T1 (Fig. 4F) with palp short and wide, distally with 3 relatively short setae. T2 (Fig. 4B) with segments and setae slightly longer and more slender than in the male. CR slightly more slender than in the male. Remarks: Most species in the subfamily Isocypridinae are known to have weakly calcified valves. In the present study, most specimens of A. arida sp.nov. have either weakly calcified, or completely decalcified valves. This means that only few specimens had valves reflecting the true shape, hence the limited number of illustrations of these valves. The longitudinal ridges on the valve surfaces (Figs 2E,F) could function as reinforcement of weakly calcified valves. Measurements (in mm): Male, holotype: RV, L = 1.69, H= 0.65; LV, L= 1.67, H=0.65 Female, allotype: RV, L= 1.58, H= 0.63; LV, L = 1.58, H= 0.64 Differential diagnosis: Amphibolocypris arida sp.nov. differs from the other described species in the genus, the type species A. exigua, by the more elongated valves, with more pointed posterior margin, especially in the female; by the shape of the hemipenis (one large rounded lobe and one narrower lobe in A. exigua, one subquadrate lobe and three more elongated lobes in A. arida) and of the prehensile palps (proximal segments with more elongated ventral protuberances and more elongated and narrower distal segments with shorter distal sensory organs in A. exigua). Two further, as yet undescribed, species from Namibia (Martens, unpubl.) differ from both A. exigua and A. arida sp.nov. in the shape of the hemipenis (see below).

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FIGURE 6. Hemipenis outlines of four species of Amphibolocypris. Not to scale. A. A. arida sp.nov. (OC.3167). B. A. exigua Rome, 1965. C. A. sp. A. sp.nov. (KM.630). D. A. sp. B. sp.nov. (KM.657). B redrawn after Rome (1965).

Ecology and distribution: The species is presently known from its type locality only, namely rock pools on a granite outcrop in south-eastern Botswana. From other, as yet unpublished records of other species of ON AMPHIBOLOCYPRIS ARIDA SP.NOV.


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Amphibolocypris from Namibia (see above), it would seem that this once monospecific genus might be more speciose than was previously assumed, in which case the individual species could have rather limited distributions. Other species investigated: As a comparison, two further new species of Amphibolocypris from the western part of southern Africa are mentioned here, and the outlines of the hemipenis given. Due to limitation of material (single specimen, completely decalcified valves), these species are left in open nomenclature.

Amphibolocypris sp. A. sp.nov. (Fig. 6C) Locality: Depression in granite in northern Namib desert, South of Hoanib River, Namibia. Coordinates: 19 ˚31’S 13 ˚ 05’E. Collected by J.A. Day on 24.12.1979 and 11.1.1980. Accompanying ostracod fauna: Plesiocypridopsis spec., Sarscypridopsis cf pygmaea. Material: 1 male, with valves lost and soft parts dissected in glycerine in a sealed slide. Fig. 6C shows the outline of the hemipenis of this specimen.

Amphibolocypris sp. B. sp.nov. (Fig. 6D) Locality: Shallow granite rainpool, near Etjokule, Waterberg plateau, Namibia. Coordinate: 20˚ 22’S, 17˚ 22’E (SE2017Ad). Collected by B. Curtis on 19.02.1983. Material: 1 male, with valves lost and soft parts dissected in glycerine in a sealed slide. Fig. 6D shows the outline of the hemipenis of this specimen.

Discussion Morphology and sexual selection The present new species is only the second one to be described in the genus Amphibolocypris Rome, 1965. The discovery of A. arida sp.nov. confirms the validity of the genus, as it shows that A. exigua, the type species, is not just a freak species of Isocypridinae. Unpublished records from Namibia (Martens unpubl.) comprise at least a further two species belonging to this genus (as identified by the hemipenis morphology), so that a substantial radiation of the genus in southern Africa appears to exist. Unfortunately, the almost invariably decalcified material makes it difficult to formally describe the species in this genus. The hemipenis morphology appears to differ considerably within the genus (Figure 6A-D), suggesting a high level of sexual selection. The situation is comparable to that in the giant ostracod genus Sclerocypris, where males also have different hemipenis morphologies, and where, similarly, several endemic species occur in the temporary habitats of the western part of southern Africa (Martens 1986, 1988, 2001). In other South African ostracod groups, such as the phylogenetically more distant Limnocytheridae, sexual selection has also been invoked as a driving force in speciation of endemic species flocks (Martens 2000). Convergent evolution One of the morphological characters defining the genus Amphibolocypris is the claw-like subapical seta on the distal segment of the walking leg (T2). Because of this transformation from seta to claw, the species in this genus have two distal claws on the T2. The exclusively southern African Amphibolocypris shares this character with the genus Platycypris, which is endemic to Australia and mostly occurs in saline to hyper saline lakes (Herbst 1957, DeDeckker 1983). If this shared character is a result of common descent, than the origin of both genera must date back to the time that Africa and Australia were still connected in Gondwanaland, i.e.

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most likely more than 100-150 Myr ago (Hay et al. 1999). This is not impossible, and would show that genera in Cyprididae can indeed be of Mesozoic age, and may experience significant morphological stasis in somatic (as opposed to reproductive) characters over long periods of time (Martens et al. 1998). However, it is also possible that Amphibolocypris and Platycypris are not sister taxa and are less closely related than it would seem, because a double end claw on T2 also occurs in more distantly related genera, namely Limanocypris Schornikov, 1961 (in subfamily Limanocypridinae Hartmann & Puri, 1974) from Siberia, the Spanish endemic Candelacypris Baltanás, 2001 (in subfamily Eucypridinae Bronshtein, 1947) (both monospecific) and in Scottia (in S. pseudobrowniana Kempf, 1971 - in subfamily Scottiinae Bronshtein, 1947). In any case, the appearance of the claw-like seta in two different subfamilies within the Cyprididae indicates an instance of clear convergent evolution, possibly suggesting that developmental switching of this character may represent some recurrent mutations across taxa. Such a mechanism might render the character unreliable as a taxonomic tool. It is at present impossible to deduce in which way the additional claw on the walking limb could constitute an adaptive character, as the taxa in which this character occurs live in different habitats. Platycypris, Limanocypris and Candelacypris have all been found in saline to hypersaline lakes, mostly with soft bottoms (clay or sand). Amphibolocypris exigua is known only from its type locality (Leeuw pan in Kruger park, South Africa, also a soft bottomed pan, but temporary and mostly with freshwater), while A. arida sp.nov. is only known from temporary, freshwater rock pools. One aspect that all of these habitats might have in common is the presence of filamentous algae, in which case the double claw could be used to glide over this substratum. However, at this stage, this is entirely speculative. Some species of Scottia, moreover, are (semi-) terrestrial (Martens et al. 2004) and thus have a completely different ecology.

Acknowledgements We thank Arne Van Den Bogaerde for the help with the sampling and Sukonthip Savatenalinton for offering valuable comments on an earlier version of the manuscript. Two anonymous referees provided useful feedback. MJ was supported by an IWT (Institute for the Promotion of Innovation by Science and Technology in Flanders) PhD scholarship. This research was financially supported by project G.0118.03 of the FWO (Fund for Scientific Research, Flanders). Julien Cillis and Claudine Behen (R.B.I.N.Sc, Brussels) offered technical assistance with the SEM illustrations and with the line drawings, respectively. Ricardo Pinto (Brasilia) also kindly assisted with the production of the SEM illustrations.

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