Eleven new South African earthworms (Oligochaeta: Microchaetidae) with new information on some known species, and an inventory of the microchaetids of KwaZulu-Natal
Abstract
Plisko, Jadwiga Danuta (2003): Eleven new South African earthworms (Oligochaeta: Microchaetidae) with new information on some known species, and an inventory of the microchaetids of KwaZulu-Natal. African Invertebrates 44 (2): 279-325, DOI: 10.5281/zenodo.7666260
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African Invertebrates Vol. 44 (2) Pages 279–325 Pietermaritzburg December, 2003 279 Eleven new South African earthworms (Oligochaeta: Microchaetidae) with new information on some known species, and an inventory of the microchaetids of KwaZulu-Natal by Jadwiga Danuta Plisko (Natal Museum, P. Bag 9070, Pietermaritzburg 3200, South Africa; School of Botany and Zoology, University of Natal, P.O. Box X01, Scottsville, 3209 South Africa; [email protected]g.za) ABSTRACT Eleven new South African indigenous earthworm species in three genera of Microchaetidae are described and illustrated: Microchaetus distasmosus, idechoritus, quaerus; Proandricus adami, amphius, oresbiosus (assigned to the lesothoensis species-group), lovuus (assigned to the belli species-group); Tritogenia debbieae, hiltonia, qudeni (assigned to the sulcata species-group); for the fourth new species, tetrata, a new speciesgroup is proposed. Of 137 currently known microchaetid species 77 are known from KwaZulu-Natal (KZN): 15 Microchaetus, 30 Proandricus, 31 Tritogenia, and one Michalakus. The exceptional diversity and endemicity of microchaetid species in this province is noted. Attention is drawn to the threat of extinction of several indigenous species. Species distribution in KZN is mapped. New localities are recorded for Microchaetus: natalensis, parvus, pondoanus, tuberosus; Proandricus: bourquini, beddardi, setosus, adriani, babanango, bergvillensis, entumeni, gracilis, jasoni, ortyi, thornvillensis; Tritogenia: grisea, kruegeri, ngomensis, douglasi, herbana, howickiana, karkloofia, lunata, shawi, sulcata, alveata; and Michalakus initus. New taxonomic or distributional information on certain species is added. INTRODUCTION The objectives of this paper are: first, to describe eleven new species in three genera of Microchaetidae: three species in Microchaetus (distasmosus, idechoritus, quaerus); four in Proandricus (adami, amphius, oresbiosus and lovuus); four in Tritogenia (debbieae, hiltonia, qudeni and tetrata); secondly, to present an annotated inventory of the microchaetid taxa known to occur in KZN, with indication on their indigenous status; and thirdly, to add new taxonomical or distributional information for selected species. As a result of my intensive collecting and study of earthworms in KZN during the past 15 years, and examining new material recently collected in protected areas of this province by Dr A. J. Armstrong from the Biodiversity Division, KZN Wildlife, a large and diverse fauna has been revealed. It is now known than at least 77 species in four genera— Microchaetus (15), Proandricus (30), Tritogenia (31) and Michalakus (1)—occur in this province (Kinberg 1867, Rosa 1891, Benham 1886a b 1892, Beddard 1907, Michaelsen 1907 1913a b c 1928, Plisko & Zicsi 1991, Zicsi & Pajor 1992, Plisko 1991 1992 1993a b 1995a b 1996a b 1997 1998 2000a b 2002a b). This is nearly 57% of the total number of South African microchaetid species described to date (Tables 1–3). Of these 77 species, 69 appear to be endemic to KZN. In addition, the province has a significant, though poorly documented fauna of indigenous Acanthodrilidae, and a considerable number of allochthonous, introduced earthworm species which sometimes occur in dense populations. The high level of such provincial endemism of megadrile earthworms, something that is
280 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 usually associated with the limited distribution of individual species, raises concern about their conservation status. To facilitate an understanding of the conservation needs of these characteristic and important members of the soil fauna, an annotated inventory of all the microchaetid species recorded in KZN is given. MATERIAL AND METHODS Type material and other studied specimens are housed in the Natal Museum. Descriptions are based on specimens preserved in alcohol or formalin. Internal anatomy was observed after conducting a dorsal incision longitudinally. Morphological and anatomical terminology referring to species assigned to Microchaetus and Michalakus follows previous publications (Plisko 1992 1993b 1996a 1998 2002b). Proandric species and their assignment to speciesgroups in Proandricus follow earlier publications (Plisko 1993a 1994 1996b 2000b 2002a). Three new species of Tritogenia—debbieae, hiltonia and qudeni—are assigned to the earlier proposed sulcata species-group (Plisko 1997). For the fourth new species, T. tetrata, a new tetrata species-group is proposed. The keys for earlier known microchaetid species can be consulted in my earlier publications (Plisko 1997 1998 2000b). Species previously recorded from KZN, not examined in the present study, are also listed and various remarks are added. Each map showing the collection sites in KZN has in its corner a map of South Africa, with the present names of the provinces. Collection sites of species Microchaetus and Michalakus are combined in one map (Fig. 1). Separate maps are presented for species of Proandricus and Tritogenia (Figs 7, 14 respectively). Co-ordinates provided in brackets refer to place names, or to the localities recorded on the original labels. Photographs of all new species of Microchaetus (Figs 2–6), Proandricus (Figs 8–13), Tritogenia debbieae (Fig. 15) and Tritogenia qudeni (Fig. 17) were taken using a Wild photomicroscope. Tritogenia hiltonia and Tritogenia tetrata were scanned using a high-resolution flatbed scanner (Figs 16, 18). Abbreviations: AJA A. J. Armstrong BK B. Kasseepursad BRS B. R. Stuckenberg cl clitellate DG D. Gwala DMc D. McCulley HM H. Murray JDP J. D. Plisko juv juvenile KZN KwaZulu-Natal NMSA Natal Museum, Pietermaritzburg, South Africa NMSA/Olig. Oligochaeta Collection in Natal Museum OB O. Bourquin PN P. Ngwenya SA Republic of South Africa TCD T. C. Dlamini TL T. Liversage TN T. Ndlovu ZMH Zoologisches Museum und Institut in Hamburg, Germany
PLISKO: NEW SPECIES OF MICROCHAETIDAE 281 Microchaetus Rapp, 1849 Microchaetus Rapp, 1849: 142. Type species Lumbricus microchaetus Rapp, 1849 by monotypy. Type depository: Natural History Museum in Vienna. Forty nine species recorded from SA and Swaziland (Table 1); 15 known from KZN, 12 being endemic. Species distribution in KZN is shown in Fig. 1. Forty eight holandric and one metandric, with one oesophageal gizzard in segment 7, and one pair of nephridia per segment. Three new species, distasmosus, idechoritus and quaerus, are described below. For the species earlier recorded from KZN, natalensis, parvus, pondoanus and tuberosus, new collection data are provided. For benhami, caementerii, papillatus and vernoni, notes are given. For species: ambitus Plisko, 1998, griphus Plisko, 2002, herberti Plisko, 2002, mkuzi Plisko, 1992 and zaloumisi Plisko, 1992, not examined in this paper, earlier known collection sites are mapped (Fig. 1). alipentus = Western Cape ambitus* benhami = Eastern Cape braunsi = Eastern Cape caementerii* circulatus = Northern Cape crousi = Western Cape davidi = Northern Cape decipiens = Eastern Cape distasmosus* franciscus = Western Cape griphus* guntheri =Western Cape hamerae = Western Cape herberti* idechoritus* imitatus = Western Cape klopperi = Eastern Cape ljungströmi = Western Cape madidus = Eastern Cape metandrus = Western Cape microchaetus = Eastern Cape mkuzi* montanus = Eastern Cape namaensis = Western Cape natalensis** = SA: Mpumalanga, Limpopo; SWAZILAND obscurus = Western Cape occidualis = Western Cape occiduus = Western Cape papillatus* parvus** = Mpumalanga pauli = Western Cape pearsonianus = Western Cape pentheri = Eastern Cape pentus = Western Cape peringueyi = Western Cape pondoanus** = Eastern Cape quaerus* rappi = Eastern Cape ritae = Eastern Cape rivus = Eastern Cape rosai = Western Cape sandersi = Eastern Cape senarius = Western Cape sophiae = Western Cape stuckenbergi = Eastern Cape tuberosus* vernoni* zaloumisi* TABLE 1 The distribution of Microchaetus species. Species marked with one asterisk are endemic to KwaZulu-Natal; those marked with two asterisks occur in KZN but have broader distributions as indicated. Those with no asterisk have not been recorded from KZN but occur in the provinces indicated. TAXONOMY
282 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 Microchaetus distasmosus sp.n. (Figs 2–3) Etymology: G. distasmos = uncertainty. Refers to the unusual combination of internal characters observed in this species. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.02532 Mapelane Nature Reserve (28º54'S:31º55'E) at the edge of forest and grassland, from moist, sandy soil, 26 November 1996, JDP, TL. Paratypes: NMSA/Olig.02377, 6 not fully mature, with Fig. 1. The distribution of species of Microchaetus and Michalakus in KZN. Microchaetus: 1 = ambitus; 2 = caementerii; 3 = distasmosus; 4 = griphus; 5 = herberti; 6 = idechoritus; 7 = mkuzi; 8 = natalensis; 9 = papillatus; 10 = parvus; 11 = pondoanus; 12 = quaerus; 13 = tuberosus; 14 = vernoni; 15 = zaloumisi; 16 = Michalakus initus. Water bodies Protected areas
PLISKO: NEW SPECIES OF MICROCHAETIDAE 283 tubercula pubertatis, collected together with holotype; NMSA/Olig.00626a Umfolozi Game Reserve (28º22'S: 31º52'E) on high bank of White Mfolozi River, grassland, sandy moist soil, 24 February 1990, 20 cl + 10 juv, JDP [associated with natalensis]. Description based on holotype and paratypes. External characters: General: body cylindrical, extended during preservation. Colour: holotype and paratypes preserved in alcohol for over six years do not reveal pigmentation; grey. Dimensions: holotype preserved, much extended, 75 mm long, 2 mm wide at 10, 2.5 mm wide at tubercula pubertatis; paratypes 55–80 mm long, 2 mm wide at 10, 3–4 mm at tubercula pubertatis. Segment number: holotype 197+ (complete number is uncertain, due to poor preservation); paratype 163. Prostomium: prolobous, small. Segmentation: secondary annulation present on preclitellar segments; 1 and 2 short, simple, with obvious intersegmental furrow 1/2, both segments with irregular longitudinal grooves; 3 simple; 4–8 with two ringlets, first longer than second; 9 with two ringlets, second short; 10 and the following simple, postclitellarly randomly demarcated. Setae: minute, closely paired. Nephridial pores: not observed. Female pores: obvious on paratypes, paired in 14 below c setae. Male pores: externally not observed; possibly in intersegmental furrow 18/19, as vasa deferentia enter body wall in 18. Spermathecal pores: externally not observed. Clitellar region (Figs 2–3): Clitellum: saddle-shaped, white, elongated, clearly segmented, on 1/n11,12–28,29; anterior and posterior borders not clearly marked; anteriorly ventral borders extend to line of papillae located on ab setae; on 18–27 bordering tubercula pubertatis. Tubercula pubertatis: on holotype and paratypes well developed glandular tubercles, segmented, on 18,19–27,28; ventrally extending to ab setae. Papillae: different sizes and shapes, glandular, encircling ab setae on first ringlets of 5–7 and also on 11–18 (Fig. 3). Internal characters: Septa: 4/5 thin, 5/6 and 6/7 most thickened; 7/8 and 8/9 little thickened. Gizzard: cylindrical, large, muscular, in 7. Calciferous glands: in 10 large, separated dorsally and ventrally. Intestine: commences in 13. Typhlosole: in holotype commences in 20 as thin tube, slightly enlarging backwards, terminating in segment 75. Dorsal blood vessel: 4–8 double, separated; 9 double, close, cordiform. Paired dorsoventral vessels: 4–8 enlarged vessels; 9–11 moniliform. Nephridia: meganephridia; little coiled loops with Figs 2–3. Microchaetus distasmosus sp. n. 2. Holotype, anterior part of body, ventral view with clitellum and tubercula pubertatis. 3. Paratype, clitellar region, enlarged. [C = clitellum, T = tuberculum pubertatis].
284 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 thin, extended caeca. Spermiductal funnels: holandric arrangement; two pairs, each closely connected posteriorly with anterior parts of seminal vesicles, in 10 and 11 respectively. Vasa deferentia: difficult to observe due to softened body wall and decomposed tissues; probably enter body wall at posterior part of segment 18. Seminal vesicles: two pairs of large sacs commencing at septa 10/11 and 11/12 respectively. Spermathecae: small, elongated thecae in 8 and 9; 1–2 at each side, close to intersegmental furrow 8/9 and 9/10 respectively; ectal parts probably in 8/9 and 9/10, although anterior not confirmed. Ovaries: not observed. Genital glands: glandular tissues in 15, and 18–27 associated with internal glands of tubercula pubertatis. Biological notes: Mapelane Nature Reserve comprises coastal dune forests extending southwards from the Mfolozi riverbank with a variety of woodland patches and grasslands. Holotype and some paratypes were found at the edge of large patches of grassland, about 10 km south from the river mouth. The other paratypes were collected north-west of the holotype site on the bank of the White Mfolozi River, covered by high indigenous grasses, and were associated with natalensis. Both localities are similar in vegetation and soil. Transportation of cocoons or individuals by water or in mud can be expected. Distribution: Known only from the Mfolozi River banks (Fig. 1). Discussion: A distinctive species although similar and probably related to quaerus described below. Both species have spermathecae in segments 8 and 9. The location of spermiductal pores in intersegmental furrow 8/9, however, was not confirmed in distasmosus. M. distasmosus also differs from quaerus in the length and position of the clitellum, shape of tubercula pubertatis, and thickness of preclitellar septa. In the other species of Microchaetus the most anterior position of spermathecal pores was observed in namaensis (known only from the Northern Cape) which has the anterior pores in 9/10 and the posterior in intersegmental furrows 10/11 11/12 respectively. In the other two holandric genera Tritogenia and Michalakus, the most anterior positions of spermathecal pores are in intersegmental furrows 10/11 11/12 in Tritogenia shawi and T. soleata, and in 10/11 11/12 12/13 in Michalakus initus. It is noteworthy that in some members of the genus Proandricus assigned to the lesothoensis-species group, the utmost anterior spermathecal pores occur in 9/10, or between ringlets in segment 9 (Proandricus amphius sp. n. and P. oresbiosus sp. n.). Microchaetus idechoritus sp. n. (Figs 4–5) Etymology: G. ide = forest, G. chorites = indigenous. Refers to the species habitat. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.02188 Ngele Forest (30º35'S:29º41'E) indigenous forest, from first 20 cm of black, moist soil, 24 November 1995, JDP, BRS. Paratype NMSA/Olig.03716 one not fully mature, abscised postclitellarly, collected with holotype. Description based on holotype and paratype. External characters: General: body cylindrical, contracted during preservation. Colour: holotype and paratype preserved in alcohol dorsally violet, ventrally yellowish-grey. Dimensions: holotype
PLISKO: NEW SPECIES OF MICROCHAETIDAE 285 preserved, slightly contracted 115 mm long, 5 mm wide at 10, 7 mm wide at tubercula pubertatis; paratype abscised 94+ mm long, 5 mm wide at 10, 7 mm at tubercula pubertatis. Segment number: holotype 198. Prostomium: prolobous, moderate. Segmentation: secondary annulation present on preclitellar segments; 1 and 2 short, with clear intersegmental furrow 1/2, both segments with irregular longitudinal grooves; 3 simple, long as 1+2; 4–8 with two ringlets similar in size and appearance; 9 with two ringlets, second shorter than first; 10 and following simple. Setae: closely paired; ab = cd, aa > bc. Nephridial pores: obvious, in c or little below c setae. Female pores: paired in 14, above b setae. Male pores: obvious, in 18 above b setae, with elongated small swellings. Spermathecal pores: externally not observed; locations of spermathecae indicate intersegmental furrows 13/14 and 14/15 respectively. Clitellar region (Figs 4–5): Clitellum: saddle-shaped, white, elongated, clearly segmented, on 12–21 with anterior and posterior borders clearly marked; on 12–15 ventral borders extend to ab setae; on 16–18 bordering tubercula pubertatis; on 19–21 covering only dorsal parts of segments. Tubercula pubertatis: on holotype elongated glandular, segmented, yellowish tissues on 16–18; ventrally extending to b setae, dorsally below nephridial pores. Papillae: paired, small, glandular swellings on 10 and 12, encircling ab setae. Internal characters: Septa: 4/5 5/6 and 6/7 thin; 7/8 and 8/9 thickened moderately, muscular, both similar in size and appearance; 9/10 and other septa thin. Gizzard: cylindrical, large, muscular, in 7. Calciferous glands: in 9, dorso-ventral, separated dorsally and ventrally. Intestine: commences in 13. Typhlosole: in holotype commences in 13 as large U-shaped tube, enlarging backwards, and terminates in 136. Dorsal blood vessel: 6–7 partly double, partly separated; 8 double, separated, 9 close, cordiform. Paired dorsoventral vessels: 5–8 thin; 9–11 moniliform. Nephridia: meganephridia; coiled loops with extended Vshaped caeca. Spermiductal funnels: holandric arrangement; two pairs of large funnels in 10 and 11 respectively, both closely connected at their posterior parts with seminal vesicles; anterior pair iridescent, posterior not iridescent. Vasa deferentia: two obvious ducts closely connected, commencing at lateral side of posterior pair of spermiductal funnels, run backwards to anterior part of segment 18. Seminal vesicles: one large pair of sacs commencing at septum 10/11, closely attached to spermiductal funnels, extends into segment 11 and terminates at 11/12. Spermathecae: in holotype, elongated clubFigs 4–5. Microchaetus idechoritus sp. n., holotype. 4. Dorsal view of clitellar region. 5. Lateral view, with enlarged area of tuberculum pubertatis. [C = clitellum, T = tuberculum pubertatis].
286 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 shaped thecae in 14 and 15; anteriorly paired; posteriorly much smaller, 3 at left side, 2 at right side. In aclitellate paratype developing tubercula pubertatis, spermathecae were not observed. Ovaries: not observed. Genital glands: flat, small glands in 9–12; in 14– 17 finger-shaped composite of 2–3 parts; in 18 composite of 4 long finger-shaped glands. Biological notes: Ngele Forest, part of the much larger forest complex of Weza State Forest, incorporates natural vegetation of Podocarpus trees, thick bush and shrubs characteristic of afromontane forests. Although this complex was surrounded by plantations of pine, cypress and poplar trees, these were absent at the collecting site. Thick layers of decomposing litter and black moist soil were abundantly inhabited by microchaetid Tritogenia ngelensis Plisko, 1997, indigenous acanthodrilid of the genus Parachilota, and by introduced Dendrodrilus rubidus (Savigny, 1826) of Lumbricidae. However, only two idechoritus specimens were collected. Distribution: Known only from the indigenous part of the Weza State Forest (Fig. 1 Ngele Forest), in southern KZN. Discussion: A distinct species with spermathecal pores in intersegmental furrows 13/14 and 14/15, and male pores in segment 18. Only three other species of this genus, herberti, natalensis and parvus, have their spermathecal pores in these furrows. However, all four species differ in general appearance, size, position of clitellum, and shape of tubercula pubertatis. Microchaetus quaerus sp.n. (Fig. 6) Etymology: L quaero = seek to learn. Refers to the unusual conjunction of internal characters noted in this species. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.03659 Hluhluwe Game Reserve, Mbombe Forest (28º03'15"S:32º03'12"E) at 484 m, forest valley, slightly damp sandy-clay next to water-seep, 30 September 2002, AJA, DG. Paratypes: NMSA/ Olig.03664, 2 cl + 1 juv collected with holotype. Description based on holotype and paratypes. External characters: General: body cylindrical. Colour: holotype, clitellate and juvenile paratypes preserved in alcohol for only a few days do not reveal pigmentation; grey-pinkish dorsally and ventrally. Dimensions: holotype preserved, slightly contracted, 104 mm long, 6 mm wide at 10, 8 mm wide at tubercula pubertatis; clitellate paratype 94 mm long, 6 mm wide at 10, 7 mm at clitellum; juvenile paratype 42 mm long, 3 mm wide. Segment number: holotype 180, clitellate paratype 170. Prostomium: prolobous, small, retracted into buccal cavity. Segmentation: secondary annulation present on preclitellar segments; 1 and 2 short, simple, with clear intersegmental furrow 1/2, both with irregular longitudinal grooves; 3 simple; 4–5 with two ringlets similar in size and appearance; 6 with two ringlets, first longer than second; 7–8 subdivided incompletely, 9 only partly demarcated; 10 and following simple, postclitellarly randomly demarcated. Setae: minute; in front of clitellum aa < bc, ab = cd. Nephridial pores: not observed. Female pores: obvious, paired, in 14 above b setae. Male pores: externally not observed; possibly in intersegmental
PLISKO: NEW SPECIES OF MICROCHAETIDAE 287 furrow 18/19, as indicated by vasa deferentia entering body wall in segment 18. Spermathecal pores: externally difficult to trace, observed only at one side of one paratype, in intersegmental furrow 8/9; ectal parts of spermathecae were observed during dissection in intersegmental furrows 8/9 and 9/10 respectively. Clitellar region (Fig. 6): Clitellum: whitish-grey; elongated, clearly segmented; on 11–27, anterior and posterior borders are not clearly marked; anteriorly ventral borders extend to line of papillae located on ab setae; on 19–24 bordering tubercula pubertatis; thin clitellar tissues also cover ventral parts of clitellar segments. Tubercula pubertatis: on holotype and paratypes glandular, segmented, elongated strips, on 19–24; ventrally extending to ab setae. Papillae: prominent, glandular, encircling ab setae on 7–10 and 25–27; on 11–18 small globular blisters on ab setae. Internal characters: Septa: 4/5 thin, 5/6–6/7 slightly thicker than anterior; 7/8 and 8/9 thin. Gizzard: cylindrical, large, muscular, in 7. Calciferous glands: in 10, large, with only slightly marked separation dorsally; ventral separation little wider, although not apparent. Intestine: commences in 13; 10 pairs of yellowish, prominent, twice-folded dorso-lateral intestinal folds in segments 23–32. Typhlosole: commences in 17 as thin, flat tube; enlarging backwards changes into large U-shaped. In paratype terminates in segment 75. Dorsal blood vessel: 4–8 double, separated; 9 double, close, cordiform. Paired dorsoventral vessels: 4–8 enlarged vessels; 9–11 much enlarged, moniliform. Nephridia: meganephridia; little coiled loops with extended, ribbon-like caeca. Spermiductal funnels: holandric arrangement; two pairs, free, in 10 and 11 respectively. Vasa deferentia: commencing on lateral sides of spermiductal funnels in 10 and 11 respectively, extend and run to segment 18; penetrations into body wall occur probably at intersegmental furrow 18/19, but it was not confirmed. Seminal vesicles: two pairs of large sacs commencing at septa 10/11 and 11/12 respectively. Spermathecae: small, elongated thecae in 8 and 9; 2–4 at each side, with ectal parts in intersegmental furrows 8/9 and 9/10 respectively. Ovaries: not observed. Genital glands: glandular tissues in 19–24 associated with tubercula pubertatis. Fig. 6. Microchaetus quaerus sp. n., holotype, ventral view with tubercula pubertatis and papillae. [P = papilla, T = tuberculum pubertatis]. PP 5 mm T
294 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 of the area earlier known as Transkei. P. londti described from garden soil in the vicinity of Pretoria (Gauteng), then discovered in older material from Masite mountains. P. zicsi is known only from type locality in the Port Elizabeth district in the Eastern Cape. P. crookesi is known only from the Vernon Crookes Nature Reserve, and no new material is available. Fig. 7. The distribution of Proandricus species in KZN. 1 = adami; 2 = adriani; 3 = alatus; 4 = amphius; 5 = armstrongi; 6 = babanango; 7 = beddardi; 8 = bergvillensis; 9 = bourquini; 10 = bulwerensis; 11 = colletti; 12 = crookesi; 13 = entumeni; 14 = gracilis; 15 = hedothecus; 16 = hlatikulu; 17 = injasuti; 18 = ivari; 19 = jasoni; 20 = lovuus; 21 = marleyi; 22 = martensi; 23 = michelleae; 24 = modestus; 25 = oresbiosus; 26 = ortyi; 27 = pajori; 28 = richerti; 29 = setosus; 30 = thornvillensis. Water bodies Protected areas
PLISKO: NEW SPECIES OF MICROCHAETIDAE 295 The lesothoensis species-group To this group, characterised by spermathecal pores occurring in one intersegmental furrow 10/11, or in two intersegmental furrows 9/10 and 10/11, or between ringlets of segment 9 and in intersegmental furrow 10/11, are assigned: lesothoensis (Reinecke & Ryke, 1969), adami* sp. n., amphius* sp. n., oresbiosus* sp. n., bourquini*, pajori*, and sani Plisko, 2002. Species marked with an asterisk are known from the Drakensberg escarpment in KZN. P. sani and lesothoensis were collected at high altitudes in the Drakensberg, and in adjacent Lesotho: sani near South African border, lesothoensis from the Maluti mountains. P. pajori is known only from the type material (Plisko 1993a). Collection sites in KZN are shown in Fig. 7. Proandricus adami sp. n. (Fig. 8) Etymology: Named for Adam Shuttleworth in recognition of his interest in earthworms. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.02530 Drakensberg, Royal Natal National Park, on the bank of Devil’s Hoek River (28º42'S:28º54'E), grassland, among roots, 4 December 1996, JDP, TL. Paratypes: NMSA/Olig.02531, 6 juv, collected with holotype; NMSA/Olig.00918, 1 cl, and NMSA/Olig.00919, 2 abscised cl + 8 juv, collected at Drakensberg foothills, Little Berg [south of the type locality] 2 March 1989, JDP, BRS. Description based on holotype and paratypes. External characters: General: body cylindrical. Colour: in life and alcohol-preserved not pigmented, whitishgrey. Dimensions: preserved and slightly contracted holotype 60 mm long, 5 mm wide at 10, 6 mm at tubercula pubertatis. Segment number: holotype 92; paratype 103. Prostomium: prolobous, small, separated from first segment. Segmentation: secondary Fig. 8. Proandricus adami sp. n., holotype, ventral view with clitellar tissues and tubercula pubertatis. [C = clitellum, T = tuberculum pubertatis]. T 5 mm C
296 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 annulation present on preclitellar segments: 1 and 2 simple, with irregular longitudinal grooves; 3 simple; 4–8 with two simple ringlets similar in size and appearance; 9 with two ringlets, second shorter than first; 10, clitellar and postclitellar simple. Setae: moderate, closely paired; first pairs on 3; preclitellarly in regular rows, posteriorly selected pairs of ab sporadically shifted into inter-setal space between bc; on 28 ab = cd, aa > bc. Nephridial pores: obvious, in ab setal lines; first pair in 2/3. Female pores: paired, in 14, slightly below cd setae, deeply in clitellar tissues. Male pores: in 17, in tubercula pubertatis area, small, simple openings. Spermathecal pores: externally not observed; however, during dissection it was noticed that ectal parts of first spermathecal pair enter intersegmental furrow 9/10; ectal parts of second pair located in segment 10 enter into intersegmental furrow 10/11. Clitellar region (Fig. 8): Clitellum: segmented, white, on 11,12–20,21; anteriorly clitellar tissues extending dorsally on part of 11; posterior border at intersegmental furrow 20/21 or 21/22; on some paratypes extending fully on 21; ventrally partially covering segments 12–15. Tubercula pubertatis: elongated oval strips, not covered by clitellar tissues, on 16–20, between cd–ab setae, separated from clitellar tissues. Papillae: little swellings, paired or single, on 10–13, 22, 23, around ab setae. Internal characters: Septa: 4/5 thickened, muscular; 5/6 and 6/7 not thickened, thin; 7/8 and 8/9 thickened, muscular; other septa thin, firm. Gizzard: globular, muscular, in 7. Calciferous glands: in 9, laterally; moderate, globular, distinctly separated dorsally and ventrally. Intestine: commencing in 13, developing into folded tube in following segments; in 21–27 dorsally covered by irregularly twisted paired folds. Typhlosole: commences in 21 or 22, terminates in holotype in 53. Dorsal blood vessel: double, close to each other in 5 and 7; slightly separated in 6, widely in 8; 9 enlarged, double, cordiform; 10 and following segments enlarged, simple; crossing septa simple. Paired dorsoventral vessels: 5–8 thin vessels, 9– 11 enlarged, moniliform. Nephridia: Meganephridia; thin, elongated coiled loops with extended elongated caeca. Spermiductal funnels: proandric arrangement; one pair of large, iridescent funnels in 10, closely connected posteriorly with seminal sacs. Vasa deferentia: single, anteriorly thick ducts, commencing laterally in 10, extend through 11 and run over flat genital glands of segments 12–14, where become embedded in body wall. Ectal parts were not observed. Seminal vesicles: one pair of very large, composite sacs, irregular in shapes, commencing at septum 10/11 extending into 11 and pushing neighboring septum 11/12 expand into 12. Both sacs composed of variable in shape and size granulated lobes, communicating with base at 10/11, closely connected with spermiductal funnels. Spermathecae: in 9 and 10, one at each side; both pairs iridescent, with sperm. Anterior spermathecae narrow, club-shaped, much smaller than posteriors, with ectal parts entering body wall at intersegmental furrow 9/10. Posterior spermathecae very large, oval, with long curved ectal ducts entering intersegmental furrow 10/11. Ovaries: not observed. Genital glands: small, flat, twiceor three-folded glands, in some of segments 9–13. Biological notes: The species was collected in the central Drakensberg mountains, in an area protected by KZN Wildlife, in indigenous grassland slope, and in river-bank beyond the boundaries of the park. Mature and juvenile specimens were collected in both sites during summer. Iridescence observed in spermathecae confirm species sexual reproduction.
PLISKO: NEW SPECIES OF MICROCHAETIDAE 297 Discussion: The location of spermathecae in segments 9 and 10 is diagnostic for the lesothoensis species-group. The species is similar to amphius described below. The shape and position of clitellum and tubercula pubertatis resemble those of amphius. The typhlosole and intestine folds are also similarly located. The species differ in the postclitellar arrangement of setae and location of anterior pair of spermathecal pores. In adami pores are in the intersegmental furrow 9/10, and their activity is confirmed by the presence of sperm in the anterior spermathecae. In amphius the spermathecae are empty with their ectal parts entering the intra-segmental body wall in segment 9, a condition presently known only in amphius and sani. Distribution: Known from the Drakensberg mountain range in western KZN (Fig. 7). Proandricus amphius sp. n. (Fig. 9) Etymology: Gr. amphi = around, on both sides. Refers to the clitellum encircling the segments. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.03620 Drakensberg, section of Giant’s Castle Game Reserve, Injasuthi (29º06'51"S:29º26'34"E) opposite camp, hillside at ca. 1540 m, at south-facing slope, wooded grassland at edge of dry drainage in Protea savannah, 2–10 cm below surface among roots of medium-tall Themeda grass, 14 June 2002, AJA. Paratypes: NMSA/Olig.03621, 9 cl; NMSA/Olig.03627, 10 juv. All collected with holotype. Description based on holotype and paratypes. External characters: General: body cylindrical, compact. Colour: in life, and preserved in alcohol, whitishgrey, not pigmented. Dimensions: preserved and slightly contracted holotype 66 mm long, 4 mm wide at 10, 5 mm at tubercula pubertatis; clitellate paratypes 55–67 mm long. Segment number: holotype 120, paratypes 126–131. Prostomium: prolobous, small. Segmentation: secondary annulation present on preclitellar segments; 1 and 2 simple, with irregular longitudinal grooves; 3 simple; 4–9 with two simple ringlets similar in size and appearance; 10 and 11 simple, longer than anterior ringlets; clitellar simple, similar in size, postclitellar simple, shorter than clitellar. Setae: minute, closely paired Fig. 9. Proandricus amphius sp. n., holotype, ventral view with clitellum, tubercula pubertatis and papillae. [C = clitellum, T = tuberculum pubertatis, P = papilla of segment 16]. CT 5 mm P
298 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 in regular rows; preclitellarly ab = cd, aa > bc, at posterior end aa = bc. Nephridial pores: conspicuous, below cd setal lines. Female pores: paired, in anterior part of 14, laterally, slightly below cd setae. Male pores: in 17, in tubercula pubertatis area between ab and cd setae. Spermathecal pores: externally not observed. However, during dissection it was observed that ectal parts of first spermathecal pair were entering into thick body wall near furrow located between ringlets of segment 9. Pores were not noted. Ectal parts of second pair located in 10, although also deeply embedded in wall tissues, were extended into intersegmental furrow 10/11. Clitellar region (Fig. 9): Clitellum: on holotype and mature paratypes segmented, white, encircling segments 1/n11,12–20,1/n21, leaving uncovered by clitellar tissues only area of tubercula pubertatis and some small ventral parts of segment 12 in area of ab setae; anterior border not well marked, with clitellar tissues extending only partly dorsally on 11; posterior border at intersegmental furrow 20/21, on some paratypes extending partially on 21. On less developed paratypes clitellar tissues ventrally were much thinner than those of mature individuals. Tubercula pubertatis: well developed elongated strips, not covered by clitellar tissues, on 1/n16–1/n20, between cd and ab setae, with shallow grooves separating each strip from clitellar tissues. Papillae: prominent, sucker-like swellings, paired, on 16 around ab setae (Fig. 9); both papillae associated with genital glands. On one paratype papillae are displaced: right one on segment 16, left one on 17. Internal characters: Septa: 4/5 5/6 thickened moderately, elastic, strong; 6/7 not thickened, thin; 7/8 most thickened, 8/9 thickened less than 7/8; other septa thin, firm. Gizzard: globular, muscular, in 7. Calciferous glands: in 9, small, globular, distinctly separated dorsally and ventrally. Intestine: commences in 13, in following segments developing into folded tube; in 21– 27 dorsally covered by irregularly twisted folds. Typhlosole: commences in 21 or 22, terminates in holotype in 53. Dorsal blood vessel: 5–7 thin, double, close; 8 double, separated; 9 enlarged, double, cordiform; in 10 and following segments enlarged and simple; simple when crossing septa. Paired dorsoventral vessels: 5–8 thin vessels, 9– 11 enlarged, moniliform. Nephridia: meganephridia; coiled loops with extended Vshaped caeca. Spermiductal funnels: proandric arrangement; one pair of large, iridescent funnels, enclosed in sacs closely connected posteriorly at septum of 10 with seminal sacs. Vasa deferentia: distinct single ducts commencing at lateral parts of spermiductal funnels, run backwards at each side of the body to posterior parts of 16, entering body wall in anterior part of 17. Connections with male pores are covered by body wall, however, externally marked male pores indicate on their termination in 17. Seminal vesicles: one pair of composite sacs, commencing at septum 10/11 extending into 11, where pushing, although not interrupting septum 11/12, occupies space of 12. Sacs, variable in shape and size, consist of granulated lobes connected with spermiductal funnels at their base. Spermathecae: in 9 and 10; anterior pair differ in size and shape from posterior pair, being elongated duct, with no sperm, and entering body wall in 9; it is however not clear if ducts penetrate whole thickness of the body tissues, or ducts are ending blindly. Large, elongated-oval, iridescent ampullae with long, curved ducts of posterior pair enter intersegmental furrow 10/11. In all dissected mature specimens posterior pair was full of sperm, although anterior pair was always empty. Ovaries: not
PLISKO: NEW SPECIES OF MICROCHAETIDAE 299 observed. Genital glands: one pair in 16 associated with papillae; both glands flat, twiceor three-folded, moderate in size. Biological notes: The species was collected at Injasuthi, an area of the Giant’s Castle Game Reserve, in the Drakensberg foothills, protected by KZN Wildlife, where there are large open areas of indigenous grassland and patches of indigenous bushes. Mature and juvenile specimens were found in winter, at an altitude over 1500 m. Iridescent posterior spermathecae and spermiductal funnels observed in mature individuals indicate maturity and sexual activity in late summer, extended into late autumn and possibly also in early winter. The species was found in association with an indigenous acanthodrilid of the genus Udeina. Discussion: The location of spermathecae in segments 9 and 10 suggest assignation to the lesothoensis species-group. The species is similar to adami sp. n. having clitellum encircling body, although in the last species encircling is not complete. However the species differ in the location of the pores of the anterior spermathecae. Similar to sani, having anterior pair of spermathecae in segment 9 with ectal parts entering intrasegmental body wall, a condition rarely observed in microchaetids. In the other species of the lesothoensis species-group, the spermathecal pores are in intersegmental furrows 9/10 and 10/11 (adami sp. n., lesothoensis and bourquini) or only in 10/11 (pajori). P. amphius differs from sani by having the clitellum encircling the clitellar segments dorsally and ventrally, with the exception of tubercula pubertatis and small anterior portion of segment 12 in the area of ab setae and papillae. The lack of sperm in anterior spermathecae and entirely filled posterior pair observed in amphius suggests that during copulation the sperm may not enter anterior spermathecae. It is possible that ectal parts of the first spermathecal pair are not completely developed. Distribution: The species is known only from the type locality (Fig. 7), in an area where some other species of the lesothoensis species-group are known to occur. Proandricus oresbiosus sp. n. (Figs 10–11) Etymology: G. oresbios = living in or on mountains. The species name refers to the type locality situated in the Drakensberg mountain range. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.02440, Drakensberg, Royal Natal National Park, north side of Devil’s Hoek River (28º41'S:28º53'E) grassland, under rock, 4 December 1996. Paratypes: NMSA/Olig.02434 ca. 28 km north-west of Bergville (28º42'S:29º22'E) primary grassland, between roots of various plants, from sandy moist soil, 5 December 1996, 3 cl +1 juv. All collected by JDP and TL. Description based on holotype and paratypes. External characters: General: body cylindrical. Colour: in life and alcohol-preserved, grey, not pigmented. Dimensions: preserved and slightly contracted holotype 90 mm long, 4,5 mm wide at 10, 6 mm at tubercula pubertatis; clitellate paratypes 75–102 mm long. Segment number: holotype 106, paratypes 117–138; juvenile 120. Prostomium: prolobous. Segmentation: secondary annulation present on preclitellar segments; 1 and 2 simple, with irregular
300 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 longitudinal grooves; 3 simple; 4–6 with two simple ringlets similar in size and appearance; 7–9 with two ringlets, second shorter than first; 10, 11, clitellar and postclitellar simple. Setae: minute, closely paired in regular rows, although ab setae on some segments are slightly shifted medially; first pairs on 3. Nephridial pores: obvious in cd setal lines; first pair in intersegmental furrow 2/3. Female pores: not observed. Male pores: probably in intersegmental furrow 16/17. Spermathecal pores: externally not observed. Clitellar region (Figs.10–11): Clitellum: saddle-shaped, segmented, white; on 1/n11,12– 20,21; ventral edges extend slightly below cd setae. Tubercula pubertatis: glandular oval pads; segmented, rimmed dorsally and ventrally; on 1/n16,17–20, dorsally bordering clitellum at cd setae. Papillae: glandular, moderate in size, paired, encircling ab setae on segments 10–12; on holotype additionally on 16 small blisters (Fig. 10); on two paratypes a pair of prominent, yellowish papillae on 18 (Fig. 11). Internal characters: Septa: 4/5 7/8 8/9 thickened moderately, slightly muscular, firm; 5/6 6/7 not thickened, although strong. Gizzard: globular, muscular, in 7. Calciferous glands: in 9, large, distinctly separated dorsally and ventrally. Intestine: commences in 12. Typhlosole: commences in 22, terminates in paratype in 56. Dorsal blood vessel: 5–8 partly separated, partly double; 9 enlarged, double, cordiform; in 10 and following segments enlarged, simple; crossing septa simple. Paired dorsoventral vessels: 5–8 thin vessels, 9–11 enlarged, moniliform. Nephridia: meganephridia; coiled loops with elongated caeca. Spermiductal funnels: proandric arrangement; one pair, free in 10. Vasa deferentia: single ducts commence at lateral parts of spermiductal funnels and run backwards at each side of body to posterior part of 16, where enter body wall, and probably into male pores. Seminal vesicles: one pair of large sacs in 11, commencing at septum 10/11. Spermathecae: in 9 and 10; different in size and shape; small ampullae with elongated necks on glandular basis, or simple, elongated ampullae. In 9 two pairs, located in unusual arrangement, i.e. in anterior and posterior parts of segment; anterior pair small, with its ectal ends entering body wall between ringlets of segment 9, with no confirmed connection with spermathecal pores; ectal parts of posterior pair located behind anterior pair, enter into intersegmental furrow 9/10. In segment 10 one pair with ectal parts entering into intersegmental furrow 10/11. No iridescence in spermathecae of both segments was observed; thick, white, mucus was found in three dissected thecae. Figs 10–11. Proandricus oresbiosus sp. n. 10. Holotype, ventral view with clitellum, tubercula pubertatis and papilla of segment 16. 11. Paratype, ventral view with clitellum and papillae of segment 18. [C= clitellum, T = tuberculum pubertatis, P = papilla].
PLISKO: NEW SPECIES OF MICROCHAETIDAE 301 Ovaries: not observed. Genital glands: in holotype large, flat glands in 10, 12, 16; in paratype one pair of large, composite, round, flat glands in segment 18, associated with papillae of same segment. Biological notes: The Royal Natal National Park, situated in the central Drakensberg mountain range, between altitude ranges from 1340–3048 m, has different geological formations and several habitat types (Pooley and Player 1995). The vegetation comprises grassland with short grasses and herbs, ferns, lichens and mosses, stream and riverbank woodland, sage-wood scrub, Protea savannah, bushes, and Podocarpus-dominated forests. A large part of the park has been under protection for nearly 50 years. The holotype was collected at the eastern side of the park, in a broad valley, covered with rocks, riverine bush and tall grasses. The paratypes were found outside the park nearly 28 km from the holotype locality, in primary grassland at much lower altitude. Discussion: The locations of spermathecae in segments 9 and 10 suggest assignation to the lesothoensis species-group. Similar to sani and to amphius sp. n., having spermathecal ectal parts entering intra-segmental body wall. The presence of two distinct pairs of spermathecae in segment 9 distinguishes oresbiosus from other species of this group. Distribution: Known from Drakensberg escarpment of western KZN (Fig. 7). Proandricus bourquini Plisko, 1996 Proandricus bourquini Plisko, 1996b: 298. Material examined: KwaZulu-Natal: Ncandu Forest Reserve (27º45'30"S:29º42'30"E): at ca. 1830 m, grassland plateau near Ulumbi River, from moist soil, NMSA/Olig.02357 29 January 1996, 1 juv, JDP; NMSA/Olig.02363 31 January 1996, 1 juv, JDP; NMSA/ Olig.02343 on bank of Ulumbi River, from wet soil, 29 January 1996, 1 juv, JDP; NMSA/Olig.03489 Ncandu Forest (27º54'03"S:29º41'36"E) valley, grassland, 23 October 2001, 1 cl, AJA , PN. Remarks: This species is known from the type locality and its close vicinity in foothills of Drakensberg mountain range in KZN (Fig. 7). The belli species-group To this group, characterized by spermathecal pores occurring only in two intersegmental furrows 11/12 and 12/13, and three variably thickened septa 4/5 7/8 8/9, are assigned: belli (Benham, 1892), lovuus* sp. n., beddardi*, hlatikulu* Plisko, 2002, modestus**, setosus**, magdae Plisko, 1993, perolofi Plisko, 1994, and skeadi Plisko, 1993. Although the anatomical characters found in the species assigned to this group show similarity, their geographical distribution does not confirm a unity of the group. The species marked with one asterisk are known only from KZN: lovuus is known from the area of Richmond and Pietermaritzburg, hlatikulu and beddardi from limited number of sites in the north-eastern KZN. The species marked with two asterisks occur in KZN and also in the other provinces: setosus in the central and northern KZN and southern Mpumalanga; modestus is known from numerous sites of South Africa (see remarks under P. modestus). It occurs in natural biotopes, and in agricultural fields into which it was probably introduced by humans.
302 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 No new material for hlatikulu has been collected since species description. The occurrence of species known from KZN is shown in Fig. 7. P. belli, magdae, perolofi and skeadi are known only from the Eastern Cape province. Proandricus lovuus sp. n. (Figs 12–13) Etymology: Named after the Lovu River on the bank of which the holotype was found. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.00482 on the bank of Lovu River in Richmond (29º52'S:30º16'E) area, in moist soil, 7 December 1989, JDP. Paratypes: NMSA/Olig.03769, 6 cl and NMSA/Olig.03690, 1 cl, Pietermaritzburg, Scottsville (29º35'S:30º25'E) on bank of Umsindusi River near tennis court, in 1–15 cm of sandy soil, 8 January 1989, JDP. Other material: NMSA/Olig.00802, 25 cl + 34 juv collected in close neighbourhood of paratype locality, 8 January 1989, JDP. Description based on holotype and paratypes. External characters General: cylindrical along the whole body length. Colour: alcohol-preserved yellowishwhite. Dimensions: preserved holotype 74 mm long, 3 mm wide at 10, 4 mm in region of tubercula pubertatis; paratypes 55–79 mm long, 3–4 mm wide. Segment number: holotype 144, paratype 165–176. Prostomium: prolobous. Segmentation: secondary annulation present on preclitellar segments; 1 and 2 short, with no clear segmental separation, both with irregular longitudinal grooves; 3 simple, as long as 1 and 2; 4–8 with 2 ringlets, similar in size and appearance; 9 with 2 ringlets, first longer than second; 10–11 and clitellar simple; postclitellar short, simple. Setae: moderate in size, paired; first pairs on 3, subsequent pairs of 4–9 on first ringlet. Preclitellarly aa > bc, postclitellarly ab = cd. Nephridial pores: obvious, in c setal lines; first pair in intersegmental furrow 3/4. Female pores: not observed. Male pores: externally not observed; probably in 14/15 intersegmental furrow. Spermathecal pores: two in intersegmental furrows 11/12 and 12/13. Clitellar region (Figs 12–13): Clitellum: on 11,12–21,22; saddle-shaped, segmented, whitish-grey; on 12–13 ventral edges extend to d setal lines, on 14–1/n19 to dorsal border of tubercula pubertatis; on 1/n19–21 extend below cd setae; on 11 only dorsally, thin layer of clitellar tissues. Tubercula pubertatis: on 1/n14,15–1/n19 below ventral Figs 12–13. Proandricus lovuus sp. n. 12. Holotype, dorso-lateral view with row of small papillae. 13. Paratype, enlarged lateral side with tuberculum pubertatis. [C = clitellum, T = tuberculum pubertatis, P = papilla of segment 18].
PLISKO: NEW SPECIES OF MICROCHAETIDAE 303 clitellar edges; oval swellings with thin grooves at their ventral borders (Fig.13). Papillae: small swellings, paired, in ab setal lines on 10–22 (Fig. 12). Internal characters Septa: 4/5 most thickened; 7/8 and 8/9 thickened moderately; other septa thin. Gizzard: large, muscular. Calciferous glands: in 9; laterally, widely separated dorsally and ventrally. Intestine: commences in 12. Typhlosole: in holotype commences abruptly as thick U-shaped in 19; terminates in 126. Dorsal blood vessel: double in 5–7, close; 8 double, separated; 9 cordiform; 10 and following simple; simple when crossing septa. Paired dorso-ventral vessels: thin tubes in 4– 8; 9–11 moniliform, not much enlarged. Nephridia: Meganephridia, paired small tufts of coiled tubules, with thin, elongated V-shaped caeca. Spermiductal funnels: proandric arrangement; one pair in 10, closely connected at septum 10/11 with vesiculae seminales. Seminal vesicles: one large pair commencing at septum 10/ 11, extending equally into 10 and 11. Spermathecae: one pair per segment; elongated ampullae, with short, thin necks; ectal parts enter body wall close to intersegmental furrows 11/12 and 12/13 respectively; posterior pair slightly larger than anterior, iridescent. Vasa deferentia: not observed, being not fully developed or deeply embedded in body wall. Ovaries: not observed. Genital glands: flat glands, in 14– 18; partially associated with internal glands of tubercula pubertatis. Biological notes: The holotype was collected in pasture on the bank of small stream. The paratypes were found at considerable distance from the type locality, on bank of the Umsindusi River. Degradation of both areas, and water pollution were in evidence. Discussion: Three thickened septa and spermathecal pores in 11/12 and 12/13 suggest species assignation to the belli species-group, although septum 4/5 is not thickened as much as is observed in the other species of this group. P. lovuus resembles the similarly sized setosus. Both species have spermathecae matching in shape. They differ notably in the shape and location of the seminal vesicles, commencement and extension of typhlosole. The moderate size of setae in lovuus also distinguishes it from setosus. Distribution: The species is known only from two sites, both in Pietermaritzburg area (Fig. 7). Proandricus beddardi (Benham, 1886) Microchaetus beddardi Benham, 1886b: 78; Benham 1892: 142; Beddard, 1895: 672; Michaelsen 1900: 449, 1918: 327; Reynolds & Cook, 1976: 7. Proandricus beddardi; Plisko, 1992: 357. Material examined: KwaZulu-Natal: NMSA/Olig.03658 Hluhluwe Game Reserve, Mbombe Forest (28º03'15"S:32º03'12"E) at 484 m, in forest valley, slightly damp sandyclay, next to water-seep, 30 September 2002, 4 cl + 4 juv, AJA, DG [associated with Microchaetus quaerus sp.n.] Remarks: The type locality was broadly indicated as ‘Natal’. Collected by Plisko (1992) from a few sites in north-eastern KZN. Present records confirm species occurrence in close neighbourhood of the earlier recorded sites (Fig. 7).
310 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 dry soil, 14 November 1974, 3 cl + 15 juv, JDP. Gauteng: NMSA/Olig.02500 Brits (25º38'S:27º47'E) cultivated field, 18 February 1997, 2 cl, W. J., Meyer. Pretoria North (24º32'S:30º24'E) on the bank of Bon Accord Dam, in soaked soil, 20 November 1974: NMSA/Olig.02168, 11 cl + 28 juv; NMSA/Olig.02170, 4 cl + 25 juv; NMSA/Olig.02171, 7 cl + 23 juv; all collected by JDP. NMSA/Olig.02176 Groenpont Gevangenis near Vereeniging (26º40'S:27º56'E) cultivated field, 2 cl, W. J. Meyer. Mpumalanga: NMSA/ Olig.02663 Witbank (25º52'S:29º15'E) on bank of Klipspruit, in moist soil, 23 November 1997, 2 cl + 1 juv, I. Meyer. NMSA/Olig.02401 26 km north of Lydenburg (25º06'S:30º28'E) in savannah dry soil, 29 November 1996, 7 cl + 2 juv, JDP, TL. Remarks: This species has been known from Free State and North West provinces (Michaelsen 1902 1912 1918, Plisko & Zicsi 1991). Recent data confirm species presence in North West and extends the known range into Gauteng and Mpumalanga provinces. In KZN this species was not found. Tritogenia kruegeri Plisko & Zicsi, 1991 Tritogenia kruegeri Plisko & Zicsi, 1991: 121. Reynolds & Cook 1993: 17. Material examined: Mpumalanga: Kruger National Park, Crocodile Bridge (25º21'S:31º54'E) under Wild Fig tree: NMSA/Olig.01708, 1 cl, NMSA/Olig.01709, 1 cl, NMSA/Olig.01722, 1 cl, NMSA/Olig.01723, 1 cl, all collected 3–13 February 1976 by A. J. Reinecke. NMSA/Olig.02207, 7 km north of Amersfoort (27º00'S:29º51'E) on bank of swamp, in moist soil, 5 December 1995, 4 cl + 10 juv, JDP, TL. NMSA/ Olig.02211 Genezareth Camp near Lake Chrissie (26º18'S:30º13'E) in moist soil, fallow ground near lake, 5 December 1995, 2 cl + 2 juv, JDP, TL. NMSA/Olig.02288 Sabie (25º02'S:30º58'E) area, near Thalalanati hotel, grassland, moist soil, 5 cl + 2 juv, JDP, TL. Limpopo: NMSA/Olig.02273 Louis Trichardt (23º01'S:29º56'E) area, near Clouds End, grassland, dry soil, 3 cl + 5 juv, 11 December 1995, JDP, TL. Remarks: This species is known from grassland habitats in Limpopo province, forested areas, riverbanks and swamps in Mpumalanga province. Not collected in KZN. Tritogenia ngomensis Plisko, 1992 Tritogenia ngomensis Plisko, 1992: 370; Reynolds & Cook 1993: 18. Material examined: KwaZulu-Natal: NMSA/Olig.03419 Ophathe Game Reserve (28º26'28"S:31º24'09"E) hillside, grassland, between roots of grasses and various plants, 9 April 2001, 3 cl + 7 juv, AJA, PN. Remarks: The species has been described from Ngome mistbelt forest. Current material was found southwards of the type locality. In recent individuals the clitellum was longer than observed in the type material on segments 13–22, but was extended over segments 13–1/n23, 24. The sulcata species-group To this group, characterised by spermathecal pores occurring in two intersegmental furrows 10/11 and 11/12, or 11/12 and 12/13, or 12/13 and 13/14, are assigned: sulcata* Kinberg, 1867, debbieae* sp. n., hiltonia* sp. n., qudeni* sp. n., annettae* Plisko, 1997, curiosa* Plisko, 1997, curta* Plisko & Zicsi, 1991, diversa* Plisko, 1997,
PLISKO: NEW SPECIES OF MICROCHAETIDAE 311 douglasi*, howickiana*, karkloofia*, lunata*, miniseta* Plisko, 1997, ngelensis* Plisko, 1997, shawi*, soleata* Plisko, 1997, ataxia** Plisko, 1997, herbana** Plisko, 1997, and palusicola** Plisko, 1997. Most of the species, all marked with one asterisk, occur in KZN (Fig. 14), although always in small areas. T. ataxia, herbana and palusicola, marked with two asterisks, occur in KZN and in neighbouring province Mpumalanga. For douglasi, herbana, howickiana, karkloofia, lunata, shawi, and sulcata new data are provided. No new material available for annetteae, curiosa, curta, diversa, miniseta, ngelensis, soleata. Tritogenia debbieae sp. n. (Fig. 15) Etymology: This species is named for Mrs Debbie Jennings, the Collection Manager at the Natal Museum, in appreciation for her extended friendly help. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.02448 Mooi River (29º12'S:30º01'E) primary grassland near Road M3, from moist soil between grassroots, 5 December 1996, JDP, TL. Paratypes: NMSA/Olig.02449, 3 cl + 6 juv, collected with holotype. Description based on holotype and paratypes. External characters: General: body cylindrical, compact after preservation. Colour: grey in life, preserved in formalin whitish-grey. Dimensions: preserved and contracted holotype 48 mm long, 4 mm wide at 10, 5 mm at tubercula pubertatis; paratypes 58–60 mm long, 4–6 mm wide. Segment number: holotype 73; paratype 94. Prostomium: prolobous, clearly separated from first segment. Segmentation: secondary annulation present on preclitellar segments in holotype and paratypes; 1–3 simple, with irregular longitudinal grooves; 4–9 with 2 simple ringlets, similar in size and appearance; 10, clitellar and postclitellar segments simple. Setae: minuscule, closely paired, preclitellarly only ab noted, postclitellarly not observed. Nephridial pores: not observed. Female pores: paired, minute openings in anterior part of 14 in front of ab setae. Male pores: in intersegmental furrow 18/19. Spermathecal pores: not observed externally; during dissection noted close to intersegmental furrows 11/12 and 12/13. Fig. 15. Tritogenia debbieae sp. n., holotype, ventral view with tubercula pubertatis and row of papillae. [P = papilla of segment 14, T = tuberculum pubertatis]. PT 5 mm
312 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 Clitellar region (Fig. 15): Clitellum: not well developed, with thin layer of clitellar tissues; saddle-shaped, segmented, whitish-grey; holotype 13–23; on 13–17 clitellar tissues cover segments dorsally extending only to dorsal edges of tubercula pubertatis. Tubercula pubertatis: on 18,1/n18–1/n22, below ventral edges of clitellum; oval glandular pads, randomly grooved by irregular shallow furrows; ventrally widely separated. Papillae: on 10–18 paired or single, round swellings encircling ab setae, different sizes. Internal characters: Septa: 4/5–7/8 thin; other septa in preclitellar segments also thin. Gizzard: large, muscular, in 6–7; commencing abruptly in part of segment 6, extends and occupies whole of 7, terminating at septum 7/8. Calciferous glands: in 9–10, large, dorsolateraly; ventrally separated widely, dorsally by narrow groove. Intestine: commences in 13. Typhlosole: commences in 17 (or 18?), slim, V-shaped, gradually enlarging into U-shaped; terminates in 50. Dorsal blood vessel: double in 4–11; 10–11 enlarged; double when crossing septa 4/5–10/11; in following segments simple. Paired dorsoventral commissural vessels: 4–8 thin, gradually enlarging; 9–11 thick, moniliform. Nephridia: two pairs per segment; small serpentine loops, one pair ventrally, second pair dorsally. Spermiductal funnels: holandric arrangement; two pairs of large, iridescent funnels in 10 and 11 respectively; in holotype right sided sacs slightly smaller than left sided; each pair enclosed in separated sac, connected with vesiculae seminales at septa 10/11 and 11/12 respectively. Seminal vesicles: two pairs of small sacs, linked with testis sacs; in 10 and 11 respectively; posterior pair slightly extending into 12. Spermathecae: in clitellate holotype paired; large, spherical with short neck; ectal parts entering intersegmental furrows 11/12 and 12/13 respectively. In aclitellate paratype with well-developed tubercula pubertatis three spermathecae per segment: 1 at left side, 2 at right side; all globular, different size, iridescent. Ovaries: not observed. Genital glands: in holotype, in some of segments associated with papillae, only at left side of the body, flat, glandular, round structures; in paratype glands not observed. Biological notes: The species was collected in grassland between the roots of tall grasses. Clitellate, aclitellate and juvenile individuals in different states of development were found in moist soil in the middle of summer. Iridescent sperm in spermathecae was observed in the clitellate individuals and in those with no clitellum. The species was associated with abundant Microchaetus parvus and Microchaetus natalensis. Distribution: Known only from the type locality, located north-west of Pietermaritzburg (Fig. 14). Discussion: Having spermathecal pores in two intersegmental furrows 11/12 and 12/13, debbieae is assigned to the sulcata species-group. Lacking thickened septa in preclitellar segments, this species evidently differs from the other congeners. Tritogenia hiltonia sp.n. (Fig. 16) Etymology: Named after the name of the type locality, Hilton. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.03534 Hilton College (29º30'48"S: 30º18'03"E) at 1119 m, Mistbelt grassland, with many flowering herbs, collected 2–30 cm between roots of various plants, 13 December 2001, AJA, HM. Paratypes: NMSA/Olig.03644, 7 cl, collected together with holotype. Other material:
PLISKO: NEW SPECIES OF MICROCHAETIDAE 313 NMSA/Olig.03645, 12 juv and NMSA/Olig.03646, 2 cocoons collected with holotype. Hilton College (29º30'47"S:30º18'02"E) at 1109 m, medium-tall grassland with many herbs, dry soil, at ca. 30 cm, 22 January 2002: NMSA/Olig.03546, 1 cl and NMSA/ Olig.03547, 4 juv. All material collected by AJA, HM. Cedara (29º32'S:30º17'E) experimental agricultural field, sample 1 and 2: NMSA/Olig.02856 and NMSA/ Olig.02857, 1 juv and 2 semi-mature, 25 February 2000, R. J. Haynes. Description based on holotype and paratypes. External characters: General: body cylindrical, compact after preservation. Colour: grey in life, preserved in formalin whitish-grey. Dimensions: preserved and contracted holotype 56 mm long, 4 mm wide at 10, 5 mm at tubercula pubertatis; clitellate paratypes 38–56 mm long, 4– 6 mm wide; juvenile 28–40 mm long. Segment number: holotype 90; paratypes 70–97. Prostomium: prolobous, not clearly separated from first segment. Segmentation: secondary annulation present on preclitellar segments; 1–3 simple, with irregular longitudinal grooves; 4–8 with 2 simple ringlets, similar in size and appearance; 9 with 2 ringlets, second shorter than first; segment 10, clitellar and postclitellar simple. Setae: minuscule, closely paired, in irregular rows; in preclitellar segments difficult to trace; postclitellarly ab closely paired in slightly irregular, longitudinal rows, cd sporadically observed only in some segments. Nephridial pores: not observed. Female pores: paired, minute openings in anterior part of segment 14. Male pores: externally not observed; probably in intersegmental furrow 19/20, where are small swellings. Spermathecal pores: conspicuous externally in intersegmental furrows 11/12 and 12/13; one or two small pores with round swellings. Clitellar region (Fig. 16): Clitellum: saddle-shaped, segmented, whitish-grey; on holotype on 13–23; on 13–17 clitellar tissues developed only dorsally; on 18–23 ventral edges extend to dorsal borders of tubercula pubertatis. Tubercula pubertatis: on 18–22 between ventral edges of clitellum and median line; nearly square glandular swellings, randomly grooved by irregular shallow furrows; ventrally on 21–22 separated medially by narrow triangle invagination; on 17 rounded single swelling above groove separating tubercula. Papillae: on 10–14, paired or single, round swellings around ab or cd setae. Fig. 16. Tritogenia hiltonia sp. n., holotype, anterior part of the body ventrally, showing secondary annulation of segments 4–9, and tubercula pubertatis.
314 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 Internal characters: Septa: 4/5 little thickened, 5/6 6/7 thickened very much, muscular, similar in thickness and appearance; 7/8 8/9 thickened less than anterior; 9/10 aborted; other septa in preclitellar segments thin; in posterior segments slightly thicker. Gizzard: large, muscular, in 6; commencing abruptly at septum 5/6 extends and occupies whole of 6, terminating abruptly at septum 6/7. Calciferous glands: in 9–10, stalked; separated widely ventrally, dorsally by short distance. Stalks extend along the whole length of calciferous glands. Intestine: commences in 13. Typhlosole: commences in 16, slim, V-shaped; in following segments gradually enlarging, becoming square, medially furrowed; terminates in area of 54. Dorsal blood vessel: double in 4–11; double when crossing septa 4/5–10/11; in 12 slightly enlarged, simple; in following segments simple. Paired dorsoventral commissural vessels: 4–8 thin, gradually enlarging; 9–11 thick, moniliform. Nephridia: in posterior segments two pairs; ventral pairs close to median body line, second pairs dorsally. Spermiductal funnels: holandric arrangement; two pairs similar in size and appearance, in 10 and 11 respectively; each pair enclosed in separate sac connected with vesiculae seminales; in dissected holotype sperm was observed in all funnels. Seminal vesicles: two pairs of small sacs in 10 and 11 respectively; one pair per segment, each linked with testis sacs at septa 10/11 and 11/12 respectively. Spermathecae: in 12 and 13, iridescent; 1–2 at each side; irregular in shape, thin, elongated with slightly enlarged ampullae; close to genital glands 12 and 13. Ovaries: paired, in 13, large, funnel-like. Genital glands: paired; in 12–15 small, flat, round; in 21 finger-shaped, elongated, 3–4 branched. Biological notes: Found between roots of various plants at depth 2–30 cm. Several individuals at different stages of development and two cocoons were present in the same sample. Sperm observed in the spermiductal funnels and spermathecae confirm breeding activity during summer. Distribution: Known only from the type locality in the area of Pietermaritzburg (Fig. 14). Discussion: The species belongs to the sulcata species-group having spermathecal pores in two intersegmental furrows 11/12 and 12/13. Similar to sulcata but distinguished by the position and shape of tubercula pubertatis, the number and position of genital papillae, and the arrangement of setae. Tritogenia qudeni sp. n. (Fig. 17) Etymology: Named after the type locality, Qudeni Forest Reserve. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.03441 Qudeni Forest Reserve (28º39'40"S:30º47'53"E), valley, short grassland, from top soil, 11 April 2001, AJA, PN. Paratypes: NMSA/Olig.03647, 6 cl + 2 juv, collected with holotype; NMSA/ Olig.03436 Qudeni Forest Reserve (28º39'42"S:30º47'53"E) hillside, short grassland in top soil, 11 April 2001, 2 with tubercula pubertatis, AJA, PN; NMSA/Olig.03434 Qudeni Forest Reserve (28º39'43"S:30º47'54"E) hillside, grassland, between roots, 11 April 2001, 2 with tubercula pubertatis, AJA, PN. Other material: NMSA/Olig.02093, 6 km south-east of Qudeni (28º38'S:30º52'E) near road, grassland, moist soil, 23 January
PLISKO: NEW SPECIES OF MICROCHAETIDAE 315 1995, 3 semi-mature with tubercula pubertatis + 8 juv, JDP, BRS. Description based on holotype and paratypes. External characters: General: body cylindrical, compact after preservation. Colour: grey in life, preserved in formalin whitish-grey. Dimensions: preserved and contracted holotype 36 mm long, 3 mm wide at 10, 4 mm at tubercula pubertatis; clitellate paratypes 24–40 mm long, 3– 4 mm wide; juvenile 14–18 mm long. Segment number: holotype 79, paratypes 80–84. Prostomium: prolobous, not clearly separated from first segment. Segmentation: secondary annulation present on preclitellar segments; 1–3 simple with irregular longitudinal grooves; 4–9 with 2 simple ringlets, second ringlet shorter than first; 10, clitellar and postclitellar simple. Setae: minuscule, closely paired; in preclitellar segments noted only ab on first ringlets of 4–9, and on 10–17; postclitellarly ab and cd observed sporadically. Nephridial pores: not observed. Female pores: externally not observed. Male pores: probably in intersegmental furrow 18/19 or in anterior part of segment 19, where are paired, small invaginations. Spermathecal pores: conspicuous, in intersegmental furrows 11/12 and 12/13, paired small invaginations with round swellings. Clitellar region (Fig. 17): Clitellum: saddle-shaped, segmented, whitish-grey; on holotype on 13–23; 13–17 clitellar tissues extending to ab seta; on 18–23 ventral edges run parallel to borders of rims separating clitellum and tubercula pubertatis. On paratypes clitellar borders not clearly marked. Tubercula pubertatis: on 18–22, below clitellar rims, ventrally distantly separated (Fig. 17); oblong glandular swellings randomly grooved by irregular shallow furrows running parallel with intersegmental furrows; longitudinally with shallow furrows separating clitellar rims from glandular tubercula. Papillae: on 10–17, 23, paired, round swellings around ab setae. Internal characters: Septa: 4/5 5/6 6/7 thickened very much, muscular, similar in thickness and appearance; 7/8 8/9 thickened less than anterior; 9/10 aborted; other septa in preclitellar segments thin; in posterior segments slightly thicker. Gizzard: large, muscular, in 6–7; commencing abruptly at septum 5/6, extends and occupies whole of 6 and partly 7; septum 6/7 attached Fig. 17. Tritogenia qudeni sp. n., holotype, ventral side of clitellar region [C = clitellum, T = tuberculum pubertatis]. CT 3 mm
316 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 to gizzard in its middle portion, extending posteriorly, and ending with termination of gizzard. Calciferous glands: stalked, dorsal; in 9–10; separated widely ventrally, by very short distance dorsally; stalks extend through 9–10. Intestine: commences in 13. Typhlosole: commences gradually as thick, V-shaped in area of 20, enlarging increasingly into large tube, terminates in area of 48. Dorsal blood vessel: 4–11 double; double when crossing septa 4/5–10/11; 12 slightly enlarged, simple; in following segments simple. Paired dorsoventral commissural vessels: 4–8 thin, gradually enlarging; 9–11 thick, moniliform. Nephridia: two pairs: ventral pairs close to median body line, second pairs dorsally. Spermiductal funnels: holandric arrangement; iridescent, similar size and appearance in 10 and 11 respectively; each pair enclosed in mutual sac connected with seminal vesicles; in dissected holotype sperm was observed in all four funnels. Seminal vesicles: two pairs of small sacs, linked with testis sacs; in 10 and 11 respectively. Spermathecae: very small, thin, elongated; close to genital glands of 12 and 13. Ovaries: paired, in 13; large, funnel-like. Genital glands: paired, small, flat, round, in 12–15, associated with papillae. Biological notes: Qudeni Forest Reserve, located almost at the northernmost limit of the midland mistbelt area, contains semi-coastal forest mixed with Podocarpus and other species of trees characteristic of afromontane forests, and also with grassland patches. The species was found between roots of various plants in topsoil, in the forest patches of grassland, and also in open grassland outside of the forest. Mature and juvenile specimens were collected together in early autumn. In some sites the species was associated with alveata and juvenile Proandricus sp. Distribution: Known from the type locality and few sites in its close neighbourhood, all situated in central KZN (Fig. 14). Discussion: The species belongs to the sulcata species-group having spermathecal pores in two intersegmental furrows 11/12 and 12/13. Similar to ataxia, but distinguished by the shape of tubercula pubertatis, the spermathecae, and the arrangement of setae. Tritogenia douglasi Plisko, 1997 Tritogenia douglasi Plisko, 1997: 255; Plisko 2000b: 76. Material examined: KwaZulu-Natal: Eshowe (28º50'S:31º28'E) area: NMSA/ Olig.03161 Rockyridge Farm, under wattle trees, 15 March 2000, 1 juv; NMSA/ Olig.03027 Rockyridge Farm, under pine tree, 19 April 2000, 1 juv; NMSA/Olig.03184 Mistyridge Farm, veld, 19 June 2000, 30 cl + 50 juv; all collected by TCD. Remarks: Known from the Dlinza Forest Reserve and from its neighbouring fields recently incorporated into farming fields (Fig. 14). Tritogenia herbana Plisko, 1997 Tritogenia herbana Plisko, 1997: 257. Material examined: KwaZulu-Natal: NMSA/Olig.01251 Platberg (28º20'S:29º59'E) Elandslaagte, Mawelawela Game Farm (Mr G. Mitchell Innes Farm) grassland, 7 December 1992,1 cl + 4 semi-mature, JDP, BRS. Remarks: The external and internal characters are in accord with the original description (Plisko, 1997), although the genital glands were observed only in segment 12, a variation
PLISKO: NEW SPECIES OF MICROCHAETIDAE 317 possibly related to the degree of maturity. Known from KZN, and from the north-eastern Drakensberg escarpment in Mpumalanga province. Present record extends the known range of species into central KZN, and suggests much broader occurrence than is usually observed in the other species of this genus. The collection sites in KZN are shown in Fig. 14. Tritogenia howickiana (Michaelsen, 1913) Microchaetus sulcatus var. howickianus Michaelsen, 1913c: 432; Reynolds & Cook 1976: 115. Tritogenia howickiana; Michaelsen 1918: 333; Plisko & Zicsi 1991: 112; Plisko 1992: 368, 1997: 280. Material examined: KwaZulu-Natal: NMSA/Olig.02749 Howick (29º27'S:30º14'E) Amberfield, from garden soil, 15 May 1997, 4 cl, J. A. Pringle. NMSA/Olig.02330 Otto’s Bluff, The Craig’s Farm, on the bank of local stream, 6 December 1989, 1 cl + 5 juv, JDP. Remarks: Known only from the type locality and its close vicinity (Fig. 14). Tritogenia karkloofia Plisko & Zicsi, 1991 Tritogenia karkloofia Plisko & Zicsi, 1991: 115; Reynolds & Cook 1993: 16. Material examined: KwaZulu-Natal: Nkandla State Forest, hillside, grassland, between roots of grasses and various plants: NMSA/Olig.03424 (28º44'35"S:31º08'57"E) 11 with tubercula pubertatis + 11 juv; NMSA/Olig.03423 (28º44'35"S:31º08'57"E) 2 juv; NMSA/Olig.03422 (28º44'35"S:31º08'57"E) 5 with tubercula pubertatis; all collected 10 April 2001 by AJA, PN. NMSA/Olig.02172 Entumeni Nature Reserve (28º53'S:31º19'E) indigenous forest, from the litter, 1 February 1989, 1 cl, JDP, BRS. NMSA/Olig.03355 Wagendrift Nature Reserve (29º02'31"S:29º50'13"E) from top layer of soil and among grass-roots in summit grassland, 27 February 2001, 6 cl, AJA, PN; NMSA/Olig.03454 Wagendrift Nature Reserve (29º02'32"S: 29º50'15"E) medium-high grass and small shrubs, in top layer of soil above rocks, 4 May 2001, 11 cl, AJA, BK; NMSA/Olig.03455 Wagendrift Nature Reserve (29º02'32"S:29º50'15"E) in top layer of soil above rocks, medium-high grass and small shrubs, 4 May 2001, 2 cl + 2 juv, AJA, BK; NMSA/Olig.00932 Karkloof Nature Reserve, Safari World (29º25'S:30º18'E) on bank of Umgeni River, at ca. 740 m, from sandy soil, 1 December 1988, 4 semimature + 4 juv, JDP; NMSA/Olig.00489 Karkloof Nature Reserve, Safari World (29º25'S:30º18'E) at ca. 800 m, on bank of Umgeni River, from moist, sandy soil, 4 January 1989, 10 cl + 14 juv, JDP; NMSA/Olig.00715 Karkloof Falls Nature Reserve, Safari World (29º25'S:30º18'E) primary grassland, 28 January 1991, 25 cl + 2 juv, JDP. NMSA/Olig.02334 Otto’s Bluff (29º30'S:30º23'E) the Craig’s Farm, on bank of local stream, 6 December 1989, 1 cl, JDP. Remarks: Known from central KZN. The species was associated with mucosa in some collection sites(Fig. 14). Tritogenia lunata Plisko, 1997 Tritogenia lunata Plisko, 1997: 259. Material examined: KwaZulu-Natal: Karkloof Nature Reserve, Melmoth section: NMSA/Olig.03473 (29º16'51"S:30º16'52"E) hillside, rocky grassland, among molerat mounds, 9 October 2001, 2 cl + 7 juv; NMSA/Olig.03476 (29º16'57"S:30º16'33"E)
318 AFRICAN INVERTEBRATES, VOL 44 (2), 2003 hillside south-west, valley, grassland, collected on surface, 9 October 2001, 1 cl [larger than type material]; NMSA/Olig.03475 (29º17'00"S:30º16'31"E) valley above stream, grassland, 9 October 2001, 2 cl + 2 juv; NMSA/Olig.03477 (29º17'09"S:30º16'24"E) valley, edge of vlei, grassland, from peat soil, 10 October 2001, 1 cl; all collected by AJA, PN. Karkloof (29º18'S:30º13'E) Mr Geekie’s Farm Benvie: NMSA/Olig.00235 at ca. 1260 m, forest edge of lower part of afromontane forest, collected under moss and between roots of various plants, on rocks and stones along stream, 22 February 1989, 1 aclitellate, JDP, BRS; NMSA/Olig.01004 at ca. 1260 m, mixed Podocarpus forest edge, lower part, from first 1–20 cm of moist soil, 20 December 1988, 5 cl + 21 juv, JDP, BRS; NMSA/Olig.00240 at ca.1250 m, on the side of road, on bank of muddy stream between roots in muddy black soil, 22 February 1989, 1 aclitellate, JDP, BRS; NMSA/OLIG.00247 at ca. 1260 m, higher part of afromontane forest edge, from litter and top soil, 22 February 1989, 2 with tubercula pubertatis + 7 juv, JDP, BRS. Doreen Clark Nature Reserve: NMSA/Olig.03322 (29º34'41"S:30º17'21"E) grassland, mediumtall grass, 15 January 2001, 1 not fully developed, AJA; from grassland with tall Digiteria eviantha (29º34'42"S:30º17'20"E) NMSA/Olig.03445 2 cl [much smaller than type material], and NMSA/Olig.03446 2 cl abscised, collected 25 April 2001 by AJA, BK. NMSA/Olig.03481 Marwaqa Nature Reserve (29º48'43"S:29º44'55"E) plateau, grassland, short thick grass, 15 October 2001, 2 cl, AJA, PN. Remarks: Known from numerous sites in the Midlands of KZN (Fig. 14). Occurs in afromontane forests, patches of grassland, indigenous bush and forest, and in open primary grasslands. Variation in size was observed: there were mature individuals much larger or smaller than those known from the type locality. The species collected in the Doreen Clark Nature Reserve was associated with introduced megascolecid Amynthas aeruginosus (Kinberg, 1867). Tritogenia shawi Plisko & Zicsi, 1991 Tritogenia shawi Plisko & Zicsi, 1991: 117; Plisko 1992: 371; Reynolds & Cook 1993: 20. Material examined: KwaZulu-Natal: NMSA/Olig.02107 Pietermaritzburg (29º35'S:30º25'E) Bisley Park, under log in the picnic site bordered with Acacia bush, 22 December 1993, 1 cl, K. R. Cradock. Remarks: Known only from the region of Pietermaritzburg (Fig. 14). Tritogenia sulcata Kinberg, 1867 Tritogenia sulcata Kinberg, 1867: 97; Perrier 1886: 876; Michaelsen 1899a: 415, 1900: 453, 1918: 338; Reynolds & Cook 1976: 176; Plisko 1992: 373. Megachaeta (Tritogenia) sulcata; Michaelsen 1891: 50. Megachaeta? sulcata; Michaelsen 1891: 50. Tritogenia sulcata [part.]; Michaelsen 1908: 31. Microchaetus sulcatus f. typicus Michaelsen, 1913c: 431. Tritogenia morosa Cognetti, 1906: 13; Michaelsen 1913c: 431, 1918: 338. Microchaetus sulcatus; Reynolds & Cook 1976: 176. Megachaeta sulcata; Reynolds & Cook 1976: 176. Material examined: KwaZulu-Natal: NMSA/Olig.00467 on bank of Umgeni River (29º28'S:30º29'E) at crossroad to Safari World, after rain, from top layer of moist soil, 6 December 1989, 1 semi-mature + 5 juv, JDP. Pietermaritzburg (29º35'S:30º25'E) Town
PLISKO: NEW SPECIES OF MICROCHAETIDAE 319 Bush: NMSA/Olig.00688 from litter, 22 March 1988, 1 cl + 1 juv; NMSA/Olig.00968 Town Bush, at left side of the road leading to Government Nursery, from dry black soil, 12 October 1988 3 juv. NMSA/Olig.00908 Orient Park (30º31'S:30º13'E) near Midmar Dam, grassland, 31 January 1991, 22 cl + 34 aclitellate. All collected by JDP. Remarks: The type locality was cited as Port Natal [now Durban]. The recent collection sites confirm species distribution in eastern and central KZN (Fig. 14). The mucosa species-group To this group, characterised by spermathecal pores occurring in three intersegmental furrows 10/11 11/12 12/13, or 11/12 12/13 13/14, or 13/14 14/15 15/16, are assigned: mucosa* Plisko & Zicsi, 1991, alveata*, insolita* Plisko, 1997, koilia* Plisko, 1997, liversagei** Plisko, 1997, silvicola Plisko, 1997, turneri Plisko, 1997. Species marked with one asterisk are known only from KZN indigenous forests, grasslands and undisturbed areas. T. liversagei is known from northern KZN, and the Drakensberg mountain range in Mpumalanga, occuring in primary grassland, grassland utilised as pastures, and also in the soil under endemic Acacia trees. T. silvicola and turneri are known only from forested areas in northern Limpopo province. New records are obtainable for alveata; no new material avaiable for insolita, koilia, mucosa, and liversagei. Tritogenia alveata Plisko, 1997 Tritogenia alveata Plisko, 1997: 268. Material examined: KwaZulu-Natal: Qudeni Forest Reserve: NMSA/Olig.03440 valley, short grassland from top soil (28º39'40"S:30º47'53"E) 2 cl; NMSA/Olig.03435 (28º39'42"S:30º47'53"E) hillside, short grassland from top soil, 1 cl; NMSA/Olig.03433 (28º39'42"S: 30º47'54"E) hillside, grassland between roots,1 cl; all collected 11 April 2001 by AJA, PN. Remarks: Described and known from the Qudeni Forest Reserve and its close vicinity (Fig. 14). The tetrata species-group This newly proposed species-group is established for the species characterised by four rows of multiple spermathecae in segments 12–15, and spermathecal pores in intersegmental furrows 12/13 13/14 14/15 15/16. To date no other known Tritogenia species is characterised in this way. Tritogenia tetrata sp. n. (Fig. 18) Etymology: G. tetra = four. Refers to the arrangement of spermathecal pores in four intersegmental furrows. Material examined: KwaZulu-Natal: Holotype NMSA/Olig.03528 Nkandla Forest Reserve (28º42'59"S:31º08'05"E) at 1178 m of forest edge, under recently burnt short grass, between roots of various plants, 21 November 2001, AJA, HM. Paratypes: NMSA/ Olig.03529, 2 cl + 3 with developing tubercula pubertatis + 3 juv, collected together