BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 66: 51-72, 1996 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGIE, 66: 51-72, 1996 On Korannacythere gen.nov. (Crustacea, Ostracoda), a new genus of temporary poollimnocytherids from southern Africa, with the description of three new species and a generic reassessment of the Limnocytherinae by Koen MART ENS Abstract pour les groupes faisant partie de cette sous-famille (illustre par Ia reduction de Ia structure de Ia charniere et !'augmentation A new genus Korannacythere and three new species, K. devriesi des dimensions de Ia furca dans l'hemipenis). Korannacythere spec. nov., K. ugiensis spec. nov. and K. hamerae spec. nov. are gen.nov. est classe ici parmi les Limnocytherini, mais en raison described from rockpools on the Korannaberg and the de Ia charniere depourvue de dents et de !'absence d'organe Drakensberg (South Africa). The new genus is characterized by prehenseur dans l'hemipenis, il occupe une position isolee au the following main features: absence of pronounced sexual sein de cette tribu. dimorphism in both valve and non-copulatory soft part mor Korannacythere gen.nov. est le second genre de limnocytherides phology, a nearly adont hinge, aberrant chaetotaxy of A2 and apparaissant exclusivement dans des habitats temporaires, Ie simplicity of hemipenis structure (no clasping organ, simple premier etant le genre monospecifique Ovambocythere copulatory process). Korannacythere gen.nov. is here lodged in MARTENS, 1989. Etant donne que les deux genres appartien the Limnocytherini, but, because of the adont hinge and the nent aux tribus differentes et que, entretemps, plusiers autres absence of at least a part of the clasping organ in the hemipenis, limnocytherides ont egalement ete signales dans des habitats non it occupies an isolated position within this tribe. The generic permanents, !'on considere que les Limnocytherinae, en general, taxonomy of the Limnocytherinae is reassessed, and an evolu ont Ia capacite de developper des phases durants lesquelles ils a tionary scenario (exemplified by both a reduction in hinge struc resistent Ia secheresse. L'importance evolutionnaire de cette ture and the increasing size of the furca in the hemipenis) is capacite pour Ia prosperite des lignees d'ostracodes non-marins proposed. est brievement discutee. Un scenario evolutionnaire est elabore a Korannacythere gen.nov. is the second limnocytherid genus partir des informations disponibles concernant Ia morphologie, occurring exclusively in temporary habitats, the other one being Ia distribution et I'ecologie du nouveau genre et des nouvelles the monospecific genus Ovambocythere MARTENS, 1989. As especes. both species belong in different tribes, and as several other lim Mots clefs: taxonomie, morphologie, selection sexuelle, diapause, nocytherids have meanwhile also been reported from non ecologie evolutionnaire. permanent habitats, it is accepted that Limnocytherinae in general have the ability to produce drought-resistant stages. The evolutionary significance of this ability for the success of non marine ostracod lineages is briefly discussed. Key words: taxonomy, morphology, sexual selection, diapause, Introduction evolutionary ecology. Temporary habitats in the semi-arid to arid areas of southern Africa have a remarkably high diversity and endemicity, even at a supra-specific level. With regard to Resume non-marine Ostracoda, the biodiversity of these habitats appears to be comparable only to that of ancient lakes Un nouveau genre Korannacythere et trois nouvelles especes, such as Lake Tanganyika (MARTENS, in press). Some of K. devriesi spec. nov., K. ugiensis spec. nov. et K. hamerae spec. the extensive radiations in these habitats can logically be nov. sont decrits des mares rocheuses du Korannaberg et du expected, such as that of the giant ostracods of the sub Drakensberg (Afrique du Sud). Le nouveau genre se distingue family Megalocypridinae (MARTENS, 1986, 1987), which par Ies principaux caracteres suivants: absence de dimorphisme show special adaptations to life in temporary environ sexuel prononce aussi bien dans Ia valve que dans Ia morphologie ments. Other groups, however, are not even expected to des parties molles non copulatoires, charniere pratiquement occur in such habitats, let alone be speciation prone there. depourvue de dents, chetotaxie aberrante de A2 et structure sim · Ovambocythere milani MARTENS, 1989 (species and genus ple de I'hemipenis (absence d'organe prehenseur, processus copulatoire simple). endemic to northern Namibia) was the first lirnnocytherid Afin de determiner Ia position du nouveau genre, Ia taxonomie ever to be raised from completely desiccated mud, and generique est reevaluee et un scenario evolutionnaire propose the discovery of this taxon actually constituted counter I I 52 K. MARTENS evidence to the long-held belief that species of the super Illustrated (type-)specimens family Cytheroidea were unable to produce drought The following registered specimens have been used for resistant stages. illustrations: The present paper describes a second new genus and three new species of temporary pool limnocytherids (from Korannacythere devriesi gen. et spec. nov. rockpools on both the Korannaberg and the Drakensberg AM. OSTIA (holotype): I(A-K), 2A,B. in South Africa). After a brief generic reassessment of the AM.OSTIB (allotype): 2(C-H) .. post-Cretaceous Limnocytherinae as a whole, it is OC.l866 (3 paratypes) (KM.l889): 7G,H,N,O; established that Korannacythere belongs to a different (KM.l891): 7J; (KM.I892): 71. lineage within the Limnocytherinae than does Ovam OC. 1867 (paratype): 7C,D,M. bocythere: the former belongs in the Limnocytherini, the OC.l868 (para type): 7E,F. latter in the Leucocytherini. This corroborates the opi OC.l869 (paratype): 7A,B. nion that the ability to produce drought- resistent stages Remarks: illustrated specimens of Figs. 7K (KM.l893) in Iimnocytherids is not a freakish event, but is more and 7L (KM. I 894) are lost. widely spread amongst the members of at least this sub family in the Cytheroidea than was previously accepted. Korannacythere ugiensis gen. et spec. nov. The evolutionary implications of this deduction are briefly AM.ECRI60A (holotype): 3C-F, 9C,D. discussed. The new species furthermore shows a number AM.ECRI60B (allotype): 4A-H. of morphological peculiarities, and their potential OC.l871 (paratype): 3B, 9A,B,J. taxonomic significance is also discussed. OC.l872 (paratype): 9G. OC.1873 (2 para types) (KM.2076): 91; (KM.2077): 9H. OC.1874 (2 parat ypes) (KM.2078): 9F; (KM.2079): 9E. OC.1875 (paratype): 3A. Korannacythere hamerae gen. et spec. nov. AM.OST2A (holotype): SA-G. Abbreviations used in text and figures AM.OST2B (allotype): 6A-G, IOG,H. OC.1878 (paratype): !OF. Hemipenis OC.l881 (3 paratypes) (KM.2090): 100; (KM.2091): cp copulatory process lOE; (KM.2092): lOC. cs caudal seta OC.l882 (paratype): lOA,B. dej ductus ejaculatorius OC.I885 (3 paratypes): (KM.2108): ION; (KM.2109) dl distal lobe !OK; (KM.21 10): 10M. f (1-3) furcal setae I - 3 OC.I886 (2 paratypes): (KM.2111): 100; (KM.2112): fu furcal ramus lOL,P. pi proximal lobe OC.l887 (paratype): lOI,J. po entrance pore trab moveable trabecule vesicula ejaculatoria Taxonomic descriptions Other soft parts Subclass Ostracoda LATREILLE, 1806 AI Antennula Order Podocopida G.W. MOLLER, 1894 A2 Antenna Suborder Podocopina G.W. MOLLER, 1894 exo exopodite of A2 Infraorder Cytherocopina GRONDEL, 1967 Md Mandibula Superfamily Cytheroidea BAIRD, 1850 Mx 1 Maxillula Family Limnocytheridae KLIE, 1938 P (1) First walking leg (= first thoracopod Tl) = Subfamily Limnocytherinae KLIE, 1938 P (2) Second walking leg ( T2) Tribe Limnocytherini KLIE, 1938 P (3) Third walking leg (= T3) fu Furca (in female) go Genital operculum (female) Korannacythere gen. nov. Valves H Height of valves Type species: Korannacythere devriesi gen. nov. spec. nov. L Length of valves (here designated). RV Right valve LV Left valve Derivation of name: after the type locality of the type act anterior cardinal tooth species: Korannaberg, north of Bloemfontein, South acs anterior cardinal socket Africa. I' Korannacythere gen. nov. from rockpools 53 Other species: K. hamerae spec. nov.; K. ugiensis spec. HOLOTYPE: a male, with soft parts dissected in glycerine nov. (see below). in a sealed slide and valves stored dry in a cavity-slide (no. AM-OSTIA). Diagnosis: small genus (.:::;: 0.5 mm), with nearly adont hinge (act in RV absent, pet in RV minute or absent, hinge ALLOTYPE: a female, with soft parts dissected in glycerine bar in LV smooth) and almost no sexual dimorphism in in a sealed slide and valves stored dry in a cavity-slide valve shape and structure. Marginal pore canals few, c 10 (no. AM-OSTlB). along anterior margin, short and straight, not branched; fused zone narrow. PARA TYPES: a tube with c 10 specimens (AM-OSTlC); A2 with three curved apical claws, two proximal ones several dissected and dried specimens in slides spinous, in both sexes; exopodite shorter in males than (OC.1866-1870). in females (length of this exopodite in both sexes species specific). Hemipenis simple, with both upper ramus (UR) Deposition of material: the holotype, allotype and ten and lower ramus (LR) of clasping organ (CO) absent; paratypes are deposited in the Albany Museum, furca with a short base and three setae, not hyper Freshwater Biology Section, Grahamstown (South developed, copulatory process (cp) simple, sickle-shaped. Africa). The other paratypes are deposited in the Ostracod Collection of the Royal Belgian Institute of Natural Remarks Sciences, Brussels (Belgium). Of the three endclaws on the A2, the two proximal ones are situated closely together and are sub-equal in shape. Derivation of name: After Mr Chris DE VRIES (Bloem Therefore, even at high magnification with light fontein), who collected the material. microscopy, only two endclaws (one proximal and one distal) are visible in most specimens. Nevertheless, the Abbreviated diagnosis presence of three claws has been confirmed in all three Valves with straight, sloping dorsal margin. Al with apical species with SEM. seta on second segment present. Exopodite on A2 in males not reaching beyond edge of second segment, in females Additional generic features: External valve sculptures well reaching tip of penultimate segment, endclaws short, developed in most populations, with ventral ala and about 1.5 times the length of terminal segment. Glans on dorso-lateral sulci. Width of valves sub-equal in both cp delated, base internally swollen. Apical seta on second ' sexes, females being slightly wider, greatest width in the segment of P(3) reaching tip of penultimate segment. middle or in the anterior half. Three walking limbs (Pl-3) with two dorsal and 1 ventral setae well developed on first Extended diagnosis: small species, with straight dorsal segment. Lateral aesthetasc on penultimate segment of A2 margin sloping in caudal direction; anterior margin widely small, one accompanying seta very large, the other minute. rounded, recurving abruptly towards the ventral margin; external valve sculptures prominent. Al with apical seta Remark: differential diagnoses and ecology of the follow on second segment. A2 with three short and stout end ing species are treated in the general dicussion. claws, c 1.5 times the length of the terminal segment, proximal ones serrate; exopodite twice as long in female as in male: reaching the tip of the penultimate segment in the former, not reaching beyond the edge of the second segment in the latter. Pl the smallest walking limb. Korannacythere devriesi spec. nov. Hemipenis with copulatory process sickle-shaped, with (Figs 1, 2, 7, 8 (A-F)) delated glans and with base additionally widened at inter nal curvature; lobe dl sub-triangular, anterior margin with Type locality: sandstone rockpools at the top of Koran an additional curve in the middle. naberg, Excelcior district, Eastern Freestate. Approx. coordinates: S 28°51'13" E 27°13'51". Altitude: 1710 Description of male mas!. Size of pools ranging from 1-50 m2, depth 5-40 em Valves with dorsal margin straight over 3/4-4/5 of the total and period of inundation from 1 week to over 4 months. length, sloping towards the caudal side, frontal margin The new species was not found in the most short-lived more broadly rounded than caudal margin, frontal margin pools. recurving abruptly towards ventral margin. No calcified Accompanying ostracod fauna: Sarscypridopsis gregaria, inner lamella; posterior fused zone narrower than anterior Jsocypris priomena, Heterocypris incongruens. fused zone, both with few (c 10 each) short, narrow and straight marginal pore canals, line of concrescence not Type material: All material was collected by Mr Ch. DE entirely parallel to valve margin. Four central muscle VRIES, either directly from the inundated rockpools, or scars, two ventral scars oblique. Hinge nearly adont, act raised from dried mud in the lab. Both lots of specimens absent, pet in R V very weakly built; intercardinal bar in were used for descriptions and illustrations. The holotype LV smooth. External valve sculptures prominent, in dor and allotype are both cultured specimens. sal half with two prominent and one smaller bulbus, I I 54 K. MARTENS ~ D H F I - Korannacythere deme. st.g en: et~p eIcd.e mno,v d.,e tmaiall eo. f copulatoryp~ro~e sPs .3 E. Idem, detail of furca. . f . F Mandibular palp. Ig. . A Al. B. A2. C. Helllipems. alp and endites. I. Pl. J. . . . G. M I H Idem chaetotaxy o p '" G Sc.a le X= .33 ,. urn for, A-F, H-K·, 8J .U ffi tOf . I I Korannacythere gen. nov. from rockpools 55 Fig. 2. - Korannacythere devriesi gen. et spec. nov., female. A. LV, internal view. B. RV, internal view. C. AI. D. A2. E. Caudal part of body, showing furcae, genital operculae and caudal seta. F. Pl. G. P2. H. P3. = Scale 33 f.Lill for C-H; 315 f.Lm for A, B. '' 56 K. MAR TENS interspaced with two sulci, in ventral half with an inversed tiona! curvature in the middle of the anterior margin. T-shaped ala. Carapace pointed at both anterior and Medial trabecule nearly straight. posterior side in dorsal view, greatest width situated in Caudal seta in-between both hemipenes (not illustrated) the anterior half; closed valve edges forming a sharp ridge large and stout, consisting of one sub-rectangular base along both frontal and caudal margins, including part of and three apical setulae. dorsal and ventral margins, only absent in medio-dorsal and medio-ventral region. Description of female Eye large and pigmented. Valves (Figs. 2A, B, I A, B, I, J) as in the male, but slightly AI (Fig. lA) with second and third segments each with wider in dorsal view and with anterior margin somewhat one apical seta; penultimate segment with 2 lateral and more broadly rounded (Note: differences in valve shape 4 apical setae, ventra-apical seta being the largest, terminal illustrated here are extremes, most specimens show less and penultimate segments of similar length, terminal seg sexual dimorphism). ment with 2 apical setae and one fused seta/aesthetasc, A 1 (Fig. 2C) as in the male. the seta being the longest, fused part of this structure A2 (Fig. 2D) with exopodite reaching tip of penultimate almost half of the total length of aesthetasc; most setae segment, i.e. about twice as long as in the male; smallest (incl. the one fused with the aesthetasc) claw-like and accompanying seta to lateral aesthetasc on penultimate flagellated. segment as long as the aesthetasc. Other aspects of A2 (Fig. IF) with exopodite only slightly longer than chaetotaxy as in the male. second segment, this segment with one apical seta, almost Mandible and Maxillula as in the male. as long as 3rd (penultimate) segment. This penultimate P(l)-(3) (Figs. 2F-H) with chaetotaxy as in the male, segment with two short dorso-lateral setae, a huge and including relative lengths of setae, P(l) relatively larger serrate ventro-lateral seta, accompanied by a short and than in the male. weak aesthetasc and a degenerate second seta, apically Genital operculae simple. Furcae (Fig. 2E) large, with with one stout seta. Terminal segment small, with three triangular base, one long apical seta and one shorter unusually short and stout claws, proximal ones set with lateral seta, the latter inserted close to the base. Caudal long spines. seta large and stout, consisting of one sub-rectangular Mandible with coxa without special features. Palp (Fig. base and three apical setulae. Abdomen with series of IE) with segments seemingly partly fused. First segment spinuous setae. with one apical seta and respiratory plate. Second seg ment with one huge dorsal seta and two (one long, one Measurements (in ,urn) short) ventral setae. Third segment with a group of five 9 : L = 412-442 (n = 4), H = 273 (n = 2), W = 194-206 = subequal dorsal setae and two longer ventral setae. Fourth (n 2). segment with three apical setae. Fifth segment with three a: L = 406-436 (n = 7), H = 230-261 (n = 5), W = 176-182 = setae. (n 2). Maxillula with respiratory plate (Fig. I G) normal, without hyperdeveloped rays. Three endites and one palp (Fig. I H); first palp segment elongate, with a group of three normal apical setae, and one large seta inserted near the base of the distal palp segment, second palp segment subrectangular, set with three apical setae. Korannacythere ugiensis spec.nov. Three walking limbs with chaetotaxy following the general (Figs 3, 4, 9) limnocytherid pattern and not showing any significant modification. Type locality P(l) (Fig. II) relatively small, with first segment bearing Rockpool on the top of Prentjiesberg, near Ugie two dorsal, one long ventral and two unequal knee-setae, (Southern Drakensberg, Eastern Cape, South Africa) second segment with one short apical seta; terminal claw (RSA/93/033). Collected on 30.3.1993 by K. MARTENS, short and solid. F.C. DE MOOR, H. BARBER-JAMES and F. WEIR. P(2) (Fig. IJ) larger; first segment with 2 dorsal, I long Approx. coordinates: S 31°08'48" E 28°08'18", Altitude ventral and I knee-seta, second segment with one long c. 2000 mas!. pH= 7.0, K = 10 ,uS/em, Temperature= 25 oc. apical seta, as long as the terminal two segments com 25 bined; terminal claw long, solid and curved. Accompanying ostracod fauna: Cyprilla paludum, P(3) (Fig. IK) the largest walking limb, first segment as Plesiocypridopsis spec. in P(2), apical seta on second segment shorter than 3rd segment; apical claw long, weakly sclerified and hirsute. Type material Hemipenis (Fig. IB) of remarkably simple structure. c 50 males and females from the above locality. Furca (Fig. 1D ) with short, subtriangular base and three setae, f2 being the longest. Copulatory process (Fig. I C) HOLOTYPE: a male, with soft parts dissected in glycerine simple, sickle-shaped, with squarish base and dilated in a sealed slide and valves stored dry in a cavity-slide glans. Lobe dl blunt and sub-triangular, with an addi- (no. AM-ECR.l60A). '' Kora!1nacythere gen. nov. from rockpools 57 -- ------ , ..... .... /'"' .......... \ I ', I I \ I I I I I I 1 I I I I a !? Fig. 3. - Korannacythere ugiensis gen. et spec. nov., male. A. R V, internal view. B. A2 with long exopodite (small form). C. A2 with short exopodite (large form). D. Md coxa. E. AI. F. Hemipenis. Scale = 33 .urn for B-F; 315 .urn for A. II 58 K. MARTENS / \ I I I I { I I I I / \ "' / <::::> 0 0 '' Korannacythere gen. nov. from rockpools 59 ALLOTYPE: a female, with soft parts dissected in glycerine species. The holotype belongs to the smaller form with in a sealed slide and valves stored dry in a cavity-slide longer exopodite. (no. AM-ECR.l60B). Measurements (in .urn) PARATYPES: a tube with c 10 specimens (AM 9 : L = 467 (n = 1), H = 273 (n = I) ECR.l60C); several dissected and dried specimens in 0': L = 467-515 (n = 8), H = 273-321 (n = 6), W = 236 = slides (OC.l871-1876). (n 2). Deposition of material: the holotype, allotype and ten paratypes are deposited in the Albany Museum, Korannacytllere llamerae spec.nov. Freshwater Biology Section, Grahamstown (South (Figs. 5, 6, 8 (G-K), 10) Africa). The other paratypes are deposited in the Ostracod Collection of the Royal Belgian Institute of Natural Type locality Sciences, Brussels (Belgium). eChibini minor Tarn, Loteni, KwaZulu Natal, coli. 18.3.1996 {K. MARTENS & M. HAMER) RSN96/030, S = Derivation of name: after the town closest to the type 29°23'01" E 29°32'29", altitude 2400 mas!. K 13.1 25 = = locality, Ugie. .uS/em, temperature 24 °C, pH 7.2. Accompanying ostracod fauna: Isocypris priomena, Abbreviated diagnosis Megalocypris durbani. Remark: the record of the latter Valves with rounded dorsal margin. AI with apical seta taxon is noteworthy, as this is the first time this genus on second segment present. Exopodite on A2 in males and species is found outside of the coastal area of the Cape reaching beyond edge of second segment, in females Province. Although the type of locality is typical of the reaching tip of terminal segment, endclaws about twice species, both the geographical position and the altitude as long as terminal segment. Glans on cp pointed, base are highly remarkable. narrow. Apical seta on second segment of P(3) reaching tip of terminal segment. ·Type material c 50 specimens from the above locality. Extended diagnosis Valves more rectangular, with anterior and posterior HOLOTYPE: a male, with soft parts dissected in glycerine margins in both sexes equally broadly rounded, and with in a sealed slide and valves stored dry in a cavity-slide dorsal margin curved, not straight; external valve (no. AM-OST2A). ornamentation in type population less prominent than in the preceding species. AI with apical seta on second seg ALLOTYPE: a female, with soft parts dissected in glycerine ment. A2 with exopodite in males reaching beyond edge in a sealed slide and valves stored dry in a cavity-slide of second segment, almost halfway the penultimate (no. AM-OST2B). segment in most specimens, exopodite in females reaching tip of terminal segment, lateral seta on penultimate PARATYPES: a tube with c 10 specimens (AM-OST2C); segment slightly less robust, apical claws in both sexes several dissected and dried specimens in slides slightly longer than in the preceding species (c twice as (OC.I878-1887). long as terminal segment). Hemipenis with dl subtriangular, more pointed than in Deposition of material: the holotype, allotype and ten the preceding species, anterior margin almost straight, paratypes are deposited in the Albany Museum, without additional curve, cp sickle-shaped, with pointed Freshwater Biology Section,, Grahamstown (South glans and with base narrow and symmetrically curved Africa). The other paratypes are deposited in the Ostracod (Fig. 3F). P(3) with apical seta on second segment Collection of the Royal Belgian Institute of Natural reaching tip of terminal segment. Sciences, Brussels (Belgium). Derivation ofn ame: the species is named after Dr Michelle Remarks HAMER (Pietermaritzburg), for her continuous help with Two morphological forms of this species were present in field work in KwaZulu Natal and for providing ostracod the type locality: a more abundant and smaller one, where material from various other parts of southern Africa. This males had exopodites on the A2 reaching almost halfway is also in acknowledgement of her significant contribu the penultimate segment (Fig. 3B) and a more rare, larger tion to the knowledge of the biology and taxonomy of form, with slightly shorter male A2 exopodites, which still southern African Anostraca and of temporary pool reach beyond the edge of the second segment (Fig. 3C). ecology in general. There were no differences in other aspects of the mor phology. As this species is at present known from its type Other material investigated locality only, it is difficult to interpret intra-specific RSA/95/012 Crane Tarn 5, Mzimkhulwana Nature variability and both forms are here included in the same Reserve. c I 0 x 15 m, almost entirely on rocks, shallow I' 60 K. MARTENS Fig. 5. - Korannacythere hamerae gen. et spec. nov., holotype male. A. RV, internal view. B. A2. C. AI. D. PI. E. P2. F. P3. G. Hemipenis. = Scale 33 ,urn for B-G; 315 .urn for A.