African Invertebrates Vol. 45 Pages 287–313 Pietermaritzburg December, 2004 Review of the balantine genus Udeina Michaelsen, 1910 with descriptions of six new species in South Africa (Oligochaeta: Acanthodrilidae, Acanthodrilinae) by Jadwiga Danuta Plisko (Natal Museum, P. Bag 9070, Pietermaritzburg 3200, South Africa and School of Botany & Zoology, University of Natal, P. Bag X01, Scottsville 3209, South Africa; [email protected] or [email protected]) ABSTRACT A brief historical overview of the South African acanthodriline earthworms is given. The taxonomic position of the genus Udeina Michaelsen, 1910 is reviewed. An inclusion of the Udeina species, characterised by balantine arrangement of male reproductive organs in the genus Parachilota Pickford, 1937 as proposed by Ljungström (1969, 1972), is not accepted. Udeina is retained and redefined, and eleven species are now included. The indecisive status in the literature of two species, U. kinbergi (Michaelsen, 1899) and U. reichei (Ude, 1905), is discussed. New material of U. reichei has been studied, and the validity of both species confirmed. The type material of U. montanus Pickford, 1937, U. pliskoae Zicsi, 1998, and U. stuckenbergi Zicsi, 1998 was examined, and new observations, comments, and illustrations are provided. Six new species—U. adriani, U. anneae, U. mapelane, U. nkandla, U. petrosi and U. qudeni—are described and illustrated. Identification keys are provided to the currently recognised genera of South African Acanthodrilinae and to species of Udeina. The species name U. stuckenbergeri erroneously published by Zicsi (1998) is corrected to U. stuckenbergi. INTRODUCTION Although Pickford (1937), Ljungström (1969, 1972), Zicsi & Pajor (1992), and Zicsi (1998) have supplied considerable data on acanthodriline diversity in this country, the indigenous South African species still require substantial study. The known Acanthodrilinae are probably only a minor part of those that may be discovered. The present paper forms a part of a survey of acanthodrilines in South Africa, based on the collection in the Natal Museum. For those Acanthodrilinae monographed by Pickford (1937), features of the male reproductive organs were used to separate genera: 1/ the primitive, acanthodriline arrangement of paired prostatic and male pores; 2/ microscolecine and balantine arrangements manifesting reductions in the number of prostates, and relocation of prostatic and male pores. The morphological nephridial differentiations, described as vesiculate or avesiculate, have been recognised as supportive diagnostic characters separating vesiculate microscolecine species from avesiculate balantine species. For all the acanthodrilines recorded from South Africa, the majority being indigenous species, five genera have been proposed: Eodrilus Michaelsen, 1907, Chilota Michaelsen, 1899b, Microscolex Rosa, 1887, Parachilota Pickford, 1937, and Udeina Michaelsen, 1910. Gates (1959) and Ljungström (1969, 1972) disagreed with the separation of the proandric, avesiculate, balantine species from proandric, avesiculate acanthodriline Parachilota species, suggesting that the balantine reductions might be only variations occurring in those Parachilota species characterised by more primitive 287 288 AFRICAN INVERTEBRATES, VOL 45, 2004 acanthodriline male reproductive organ arrangements. Zicsi (1998) did not follow their suggestion, recognising the genus Udeina for the avesiculate species with the balantine prostatic reduction, known only from South Africa. This supported his interpretation that the genus Yagansia, Michaelsen, 1899 is accepted for those South American vesiculate species with microscolecine prostatic reduction. In the present paper, the validity of Udeina is recognised for South African species, and supported by additional evaluation of its generic features, notably the proandric condition associated with reductions to one pair of spermathecae and one pair of prostates, and constant occurrence of these characters. The taxonomy proposed by Pickford (1937) is accepted here, although following Zicsi’s (1998) recommendation, the South African species assigned by her to Eodrilus are accepted in Eodriloides Zicsi, 1998, supported by similar exclusion of Australian species, including the type-species Eodrilus cornigrave Michaelsen, 1907, from Western Australia, by Jamieson and Dyne (1976). The present paper comprises new taxonomic observations, evaluation of characteristics of the genus Udeina, and descriptions of six new species. MATERIAL AND METHODS During my extended survey of earthworms in South Africa, a collection of acanthodrilines was assembled, and specimens with balantine reductions of the anterior or posterior pair of prostates were noted. An additional substantial material of acanthodrilines donated to the Natal Museum by Dr A. J. Armstrong of the Biodiversity Division, Ezemvelo KZN Wildlife, and by the Mammal Research Institute, University of Pretoria, supplied more individuals with these characters. This stimulated my attention to evaluate South African proandric species, characterised by only one pair of spermathecae, balantine reduction of one pair of prostatic glands, and relocation of male pores. The type material of known species Udeina montanus Pickford, 1937, U. stuckenbergi Zicsi, 1998, and U. pliskoae Zicsi, 1998, was included. No type material of U. kinbergi (Michaelsen, 1899) and U. reichei (Ude, 1905) has been examined. The types of both species were restudied by Michaelsen (1907, 1910, 1912) and Pickford (1937). Both authors confirmed that although the material was in poor condition, balantine reductions were evident. However, the exact locations of prostatic and male pores in the types remain uncertain, as neither author could definitely locate them, and probably further examination could be uninformative due to decomposition of the specimens. A small sample containing two slightly decomposed adults and seven juveniles, collected on a bank of the Vaal River, possibly near the type locality of U. reichei, and identified by Ljungström (1972) as U. kinbergi, was examined by me and found to be U. reichei. The types of new species described below—U. adriani, U. anneae, U. mapelane, U. nkandla, U. petrosi, and U. qudeni—and of already described U. pliskoae and U. stuckenbergi, are preserved in 75% ethanol and housed in the Natal Museum. U. montanus is in the South African Museum, Cape Town, South Africa. The descriptions of all species are based on type specimens, and comparisons with available additional material are made. Description of colour is based on specimens which may have lost intensity during preservation and through storage in alcohol or formalin. The PLISKO: UDEINA IN SOUTH AFRICA 289 number of body segments was counted from the anterior segment in which the prostomium is located, and presented in Arabic numerals. Intersegments and septa location are designated by a slash (e.g. 7/8). Dorsal longitudinal dissections were performed for study of the internal anatomy, and the photographs were taken under a Wild stereo microscope Type 181300. Penial setae were observed in glycerol jelly, permanently mounted in Berlese fluid, and photographed using photo-camera Wild MPS 51 attached to Wild Heerbrugg microscope. Locality data and biological notes were taken from labels; information on the biotopes was often extended by personal comments from Dr A. J. Armstrong, or taken from Pooley and Player (1995). The species U. adriani, U. mapelane, U. nkandla, U. petrosi, U. qudeni and U. stuckenbergi are known from relatively undisturbed locations, suggesting that South African acanthodriline earthworms prefer pristine, natural biotopes, indigenous grasslands and forests. Probably more species will be found in other protected areas of South Africa. Generic characters of South African Acanthodrilinae reported by Pickford (1937) are presented in Table 1. Keys are provided to recognised genera of South African Acanthodrilinae and to the species of Udeina. Glossary, abbreviations and definitions of terms used in the text: 1/n immediately followed by a figure refers to that part of segment covered by clitellar tissue. TABLE 1 Generic characters of South African acanthodriline earthworms presented by Pickford (1937). Genus Microscolex Eodrilus= Chilota Parachilota Udeina Eodriloides Spermiductal/ holandric, or holandric proandric proandric proandric testes funnels proandric with rudimentary funnels in 11 Spermathecae one pair, or two two pairs two pairs two pairs one pair pairs, or absent Spermathecal 7/8 and 8/9; 7/8 and 8/9 7/8 and 8/9 7/8 and 8/9, or 7/8 pores or 7/8 or 8/9; on the anterior or absent part of 8 and 9 Male pores 17 or 18 or 18 or 17/18 18 18 or 19 17 or 18 absent Prostates one pair, or two two pairs two pairs two pairs one pair pairs, or absent Prostatic 17 and 19 17 and 19 17 and 19 17 and 19, 18 or 19 pores or 18 and 20 Male repro microscolecine acanthodriline acanthodriline acanthodriline balantine ductive organs Excretory vesiculate avesiculate vesiculate avesiculate avesiculate system Indigenous to one species 15 species 13 species 52 species two species SA Introduced to two species none none none none SA 290 AFRICAN INVERTEBRATES, VOL 45, 2004 Acanthodriline arrangement of the male reproductive organs, or shortened as one word: acanthodriline – refers to location of two pairs of prostatic pores of the tubular prostates in segments 17 and 19 respectively, and male pores present in segment 18. This condition is usually associated with two pairs of spermathecae, and spermathecal pores occurring in intersegmental furrows 7/8 and 8/9 respectively. This arrangement is considered as primitive in the Acanthodrilidae. Avesiculate – referring to a holoic single nephridium without dilation. The term was used by Pickford (1937) to describe the nephridia observed in acanthodriline species allocated to Eodriloides, Parachilota and Udeina. Balantine – originally the term referred to a reduction of anterior pair of prostates and posterior pair of spermathecae, occurring in species characterised by acanthodriline condition; one pair of prostates occurs in segment 19, and a pair of spermathecae of segment 8 has its pores located in intersegmental furrow 7/8. The present understanding employed in this paper, refers to acanthodriline, avesiculate species, with one pair of prostates, with prostatic pores in 17 or 18 or 19, and with one pair of spermathecae with spermathecal pores in intersegmental furrow 7/8 or 8/9. Male pores may occur in 16 or 17 or 17/18 or 18. Clitellum – the region of the body-wall formed from glandular cells, which secrete material to form a cocoon. It may be saddle-shaped or ring-shaped, although the shape may depend on the phase of sexual maturity. The position of clitellum in most South African Acanthodrilinae species is usually constant, but intraspecific variation occurs in some species. Figures followed by a stroke (for example 7/8) indicate a position between segments or septa. Holandric – refers to number and position of testes and testes funnels; indicates presence of two pairs of testes and two spermiductal funnels occurring in segments 10 and 11 respectively; (compare proandric). Holoic – (old terms: holonephric or meganephric); refers to the paired excretory organs, as simple nephridia in each segment of the body except the pharyngeal segments. Holoic nephridia are observed in all acanthodrilids. Pickford (1937) noted in some species that the single nephridium may possess variable dilation forming terminal bladders, and on this basis, for the species recorded in South Africa, two groups were established: the avesiculate nephridia found in Eodriloides, Parachilota and Udeina, and the vesiculate nephridia in Chilota and Microscolex. JDP – J. D. Plisko. KZN – KwaZulu-Natal province of South Africa. Male pores – ectal ends of vasa deferentia. In majority of the South African acanthodrilines male pores are situated laterally to the b setae; only in two known species (P. vanhoffeni and P. bavenda) approximating towards the mid-ventral line. In some species may be fused with prostatic pores. Microscolecine – originally this term referred to a reduction of posterior pair of prostates and reduction of anterior pair of spermathecae, observed in the species with acanthodriline male reproductive organs. In the vesiculate species recorded in South Africa, the microscolecine characters have been observed only in two introduced, and in one endemic species. Variability in reduction and relocation of male organs were noted. PLISKO: UDEINA IN SOUTH AFRICA 291 NMSA – Natal Museum, Pietermaritzburg, South Africa. Penial seta (pl. penial setae) – specialised seta associated with prostatic pore; development of penial seta depends on maturation of the individual; after the sexual phase, the well developed seta is discharged, being gradually replaced by a new precursory seta, which is progressively modified. During seta formation, precursory setae come to resemble the adult penial setae except in size, and their shapes may be observed. In the majority of known South African acanthodrilines, the penial setae were observed and often described; however, the state of their maturity was not always indicated. The taxonomic importance of penial setae is considerable. Penial setal retractor muscles (pl. retractors) – the muscles originating from the body wall dorso-laterally, usually behind the penial seta and activating them; maturity of retractors depends on maturation of individuals. The site of origin can be of taxonomic value. Proandric – refers to number and position of testes and their funnels; indicates the presence of a single pair of testes and spermiductal funnels in segment 10; (compare holandric). Prostate (pl. prostates) – paired glands associated with male reproductive organs; each consists of a prostatic gland and muscular prostatic duct. In most Acanthodrilidae the prostatic gland is tubular; according to Pickford (1937), it originates as an ectodermal invagination lateral to seta b; the number and position correspond to the number and position of prostatic pores, and for acanthodrilids it is an important taxonomic character. In South African species always tubular. Prostatic duct – extended part of prostatic gland, usually muscular; its ectal part opens into prostatic pore. Prostatic gland – in South African Acanthodrilinae, the glandular, tubular part of prostate; may be confined to one segment or extended through several segments. Prostatic pore (pl. prostatic pores) – ectal part of prostatic duct. In the endemic South African acanthodrilines the pores are situated lateral to the b setae; only in two species (P. vanhoffeni and P. bavenda) the prostatic pores approach the mid-ventral line. South African indigenous mature specimens usually have the pores connected with the male pores by the seminal grooves. Salivary glands – pharyngeal structures; in South African acanthodrilines usually extending backwards to septum 4/5, however, in some species slightly beyond. Segment – the primary unit of segmentation; a portion of the body internally separated by septa, externally by intersegmental furrows. In most South African acanthodriline species segmentation corresponds to external and internal transverse metameric divisions, in clear distinction to indigenous microchaetids, where external annulation is not congruent with internal divisions. Seminal grooves – external elongated depressions connecting male pores with prostatic pores. Seminal vesicle (pl. seminal vesicles) – a part of male reproductive organs; a pocket formed from dilated septum, where the sperm is matured. Septum (pl. septa) – an internal transverse wall between segments, acting as a supporting membrane for internal organs; usually delicate, although in some anterior segments thickened to varying degrees. In the South African species its nature is a supportive taxonomic character. 292 AFRICAN INVERTEBRATES, VOL 45, 2004 Seta (pl. setae) – any chitinous bristle, being a product of a single ectodermal cell at the end of a tubular epidermal ingrowth. Setal formula is expressed in small letters: a = the most ventral seta; b = the seta next lateral to a; c = the seta next lateral to b; d = the seta next lateral to c; aa = median space ventrally between the two setae; ab = space between a and b; bc = space between b and c setae; cd = space between c and d setae; dd = space between the most dorsal setae. In the vast majority of the endemic South African acanthodrilines setae are widely paired; it is not confirmed whether the inter-setal ratio, often being variable, can be applied for taxonomic evaluation. Spermatheca (pl. spermathecae) – an organ in which sperm received from a copulatory partner is stored. Each consists of the ampulla, duct, and diverticulum. The ampulla, usually largest part of the spermatheca, is more or less ovoid, and thin-walled. The duct is variable in shape and size, communicates by its ental part with the ampulla, and through its ectal part with spermiductal pore. The diverticulum can be simple or bi- or multiple-branched, variously shaped; its size and shape are of great taxonomic importance. In South African endemic acanthodrilines, sperm is usually stored in the diverticulum, sometimes also partly in the duct, but has not been observed in the ampulla. The number, position and shape of spermathecae are primary taxonomic characters. Spermathecal pores – external openings at ends of ectal parts of spermathecae. Their number and position are of high taxonomical importance. Spermiductal funnel = testes funnel (pl. spermiductal funnels) – the ental part of sperm duct through which sperm pass from testes into lumen of vas deferens on the way to male pore. U – circumference of the body; in South African Acanthodrilinae the distance between dd is variable. Vas deferens (pl. vasa deferentia) – the sperm duct carrying the sperm from the spermiductal funnel towards the male pore. Vesiculate – a holoic single nephridium, variable in size and shape of dilation forming terminal bladder (a vesicle). A term used by Pickford (1937) for nephridia observed in the species of Chilota and Microscolex. TAXONOMY Key to genera of the subfamily Acanthodrilinae known in South Africa 1 Holandric..............................................................................................................2 – Proandric...............................................................................................................3 2 Excretory system holoic, avesiculate. Male reproductive organs acanthodriline. Two pairs of spermathecae; two pairs of spermathecal pores; each pair in intersegmental furrow 7/8 and 8/9. One pair of male pores in segment 18, or in intersegmental furrow 17/18. Two pairs of prostatic pores; each pair in segment 17 and 19. Gizzard in segment 5; rudimentary, moderately or well developed. Spermathecal, prostatic pores and male pores not approximate towards the mid-ventral line..................... Eodriloides Zicsi, 1998 – Excretory system holoic, vesiculate. Male reproductive organs microscolecine, variable. One or two pairs of spermathecae, or spermathecae absent; one pair of spermathecal pores in intersegmental furrow 7/8 or 8/9, or two pairs, each in segment PLISKO: UDEINA IN SOUTH AFRICA 293 7/8 and 8/9, sometimes absent. One pair of male pores in segment 17 or 18, or absent. One pair of prostatic pores in segment 17, or two pairs, each in 17 and 19. Male pores might be fused with prostatic pores or enter independently. Gizzard in segment 5, rudimentary. In introduced species, a posterior pair of spermiductal funnels might be reduced, or occurs in rudimentary parts. Spermathecal, prostatic pores and male pores not approximate towards the mid-ventral line..................... Microscolex Rosa, 1887 3 Excretory system holoic, vesiculate. Male reproductive organs acanthodriline. Two pairs of spermathecae. Two pairs of spermathecal pores; each pair in intersegmental furrow 7/8 and 8/9. One pair of male pores; male pores in segment 18. Two pairs of prostatic pores; a pair in segment 17 and 19. Gizzard reduced, if present in segment 5, rudimentary or well developed. Spermathecal, prostatic pores and male pores not approximate towards the mid-ventral line.............. Chilota Michaelsen, 1899 – Excretory system holoic, avesiculate....................................................................4 4 Male reproductive organs acanthodriline. Spermathecal, prostatic pores and male pores not approximate towards the mid-ventral line, or situated close to the mid- ventral line. Two pairs of spermathecae. Two pairs of spermathecal pores; each pair in intersegmental furrow 7/8 and 8/9, or on the anterior border of segment 8 and 9. One pair of male pores in segment 18 or 19. Two pairs of prostates; two pairs of prostatic pores, each pair in segment 17 and 19, or in 18 and 20. Gizzard in segment 5 or 6; rudimentary, or moderately or well developed............................. Parachilota Pickford, 1937 – Male reproductive organs balantine. Spermathecal, prostatic pores and male pores not approximating towards mid-ventral line. One pair of spermathecae in 8 or 9. Spermathecal pores in 7/8 or 8/9. One pair of male pores in segment 16 or 17 or 18, or in intersegmental furrow 17/18; in front or behind of prostatic pores, separated, or fused with prostatic pores. One pair of prostatic pores in segment 17 or 18 or 19. One pair of well developed prostates, or one pair of well developed prostates and rudimentary prostatic ducts in segment 17....................Udeina Michaelsen, 1910 Udeina Michaelsen, 1910 emend. Holoscolex Ude, 1905: 421 (preoccupied by Cognetti 1904). Yagansia [partim]: Michaelsen 1899a: 237. Udeina Michaelsen, 1910: 53, 1912: 4, 5; Stephenson 1930: 671, 828; Pickford 1937: 416; Gates 1959: 239; Reynolds & Cook 1976: 62. Udeina [applied for Parachilota]: Ljungström 1969: 370. Udeina [partim]: Ljungström 1972: 100. Udeina Michaelsen, 1890 [sic] Zicsi 1998: 67. Type species: Yagansia kinbergi Michaelsen, 1899a: 443. Revised diagnosis: Setae lumbricine. Prostomium tanylobous. Prostatic pores and male pores not approximating towards the mid-ventral line. Prostates tubular. Proandric. Holoic, avesiculate. One pair of spermathecae in segment 8 or 9. Spermathecal pores in intersegmental furrow 7/8 or 8/9. One pair of male pores in segment 16 or 17 or 18 or in 17/18, in front or behind of prostatic pores, separated or fused with prostatic pores. One pair of prostatic pores in segment 17 or 18 or 19. One pair of well-developed prostates, or with additional vestigial parts of second prostatic pair. Spermathecal diverticulum 294 AFRICAN INVERTEBRATES, VOL 45, 2004 uni- or bilobate. Gizzard variably developed in segment 5. Typhlosole absent. Calciferous glands absent. Distribution: Endemic to South Africa. Known from Free State, North West province, Western Cape, Eastern Cape, KZN. Remarks and historical notes: The first species described with balantine features was Yagansia kinbergi, collected over 150 years ago; the type locality was cited as ‘Caffraria’ or ‘Kaffraria’, possibly a part of western KZN extending into eastern Free State. A single specimen, not clitellate, broken in two pieces, was described by Michaelsen (1899a); subsequently it was restudied by Michaelsen (1907, 1910, 1912), and by Ude (1905) who described the new genus Holoscolex for the new species reichei. Ude (1905) compared these two species, both similarly characterised by balantine male reproductive organs. However, no final clarification of the exact position of the male and prostatic pores was stated. Michaelsen (1910), accepting the generic status suggested by Ude for balantine species known from South Africa, proposed the new generic name Udeina for Holoscolex because it was preoccupied. Subsequently, Michaelsen (1912) synonymised reichei under kinbergi, stating: ‘Gen. Holoscolex Ude: da dieser Name von Cognetti schon eine Glossoscolecinen-Gattung vergeben ist, so bedarf die Udesche Acanthodrilinen-Gattung einer anderen Bezeichnung. Ich nenne sie Udeina, Typus: U. kinbergi (Mich.) = Yagansia Kinbergi Mich. = Holoscolex Reichei Ude.’ To this, Michaelsen also added information on the type locality, suggesting it was in the Free State. In the same paper he wrote: ‘…Südafrika, etwa von der Breite der Kalahari südwärts, charakterisiert durch die Microchätinen-Gattungen Microchaetus und Tritogenia, sowie durch die Acanthodrilinen-Gattungen Chilota und Udeina’. Pickford (1937) also synonymised U. reichei with U. kinbergi. Zicsi (1998), however, considered that U. reichei may be a valid species, based on examination of material collected by himself in the area of Parys on a bank of the Vaal River in the Free State. The third species with balantine features is Udeina montanus Pickford, 1937 described from a small sample from the Langeberg range in the Western Cape, differing from U. kinbergi in many characters (location of prostatic and male pores, fused or separated prostatic and male ducts, number of hearts). This species was accepted as more primitive than U. kinbergi. An independent evolution of both species from different Parachilota species was suggested. The species added by Zicsi (1998), U. pliskoae and U. stuckenbergi, and six new species described in this paper—U. adriani, U. anneae, U. mapelane, U. nkandla, U. petrosi, and U. qudeni—have raised the number of balantine species to eleven. All being proandric, with holoic avesiculate nephridia, having only one pair of spermathecae and one pair of prostatic pores, belong evidently to one species- group, with the balantine condition characterising Udeina. It has been found that the position of the spermathecal pores, male pores, and prostatic pores, are variable within the genus (Table 2). However, the reduction of one prostatic pair, either the anterior or posterior one, is evidently associated with the reduction of one pair of spermathecae and relocation of male pores. The prostatic pores can occur anteriorly or posteriorly to male pores, or be on the same segment as male pores. The ectal parts of the vasa deferentia can enter into the male pores separately, or be fused with ectal parts of prostatic pores. The presence of vestigial parts of the second pair of prostatic ducts observed in U. petrosi may suggest stages in prostate reduction. It is noteworthy that a reduction of PLISKO: UDEINA IN SOUTH AFRICA 295 TABLE 2 Selected characters used for the Udeina species separation. Sperma- Male pores Prostatic Last pair of Species name thecal pores pores pores Intestine hearts Diverticulum adriani 7/8 18 19 16 12 unilobate anneae 7/8 18 19 17 13 unilobate kinbergi 7/8 17 18 16 13 unilobate mapelane 7/8 18 17 16 12 unilobate montanus 7/8 18 19 17 12 unilobate nkandla 7/8 17/18 17 16 12 bilobate petrosi 7/8 18 19 16 12 bilobate pliskoae 7/8 18 17 16 12 unilobate qudeni 7/8 16 17 15 12 unilobate reichei 7/8 17 18 17 13 unilobate stuckenbergi 8/9 17 17 17 12 unilobate one pair of prostates, or relocation of prostatic pores or male pores, are variable in the species currently grouped in Udeina; however, this variability was noted in populations of U. anneae, U. mapelane, U. montanus, U. petrosi, U. pliskoae, and U. stuckenbergi. A similar reduction of the anterior prostatic pair to vestigial prostatic ducts, was observed in three dissected specimens of U. petrosi. Key to species of the genus Udeina Michaelsen, 1910 1 Spermathecal pores in intersegmental furrow 7/8................................................2 – Spermathecal pores in intersegmental furrow 8/9.........stuckenbergi Zicsi, 1998 2 Prostatic pores in segment 17 or 18......................................................................3 – Prostatic pores in segment 19; male pores in 18 ..................................................7 3 Prostatic pores in segment 17...............................................................................4 – Prostatic pores in segment 18; male pores in 17; last pair of hearts in 13..........10 4 Male pores in 16 or in 17/18.................................................................................5 – Male pores in 18 ...................................................................................................6 5 Male pores in segment 16; intestine commences in 15; last pair of hearts in 12 ... qudeni sp. n. – Male pores in intersegmental furrow 17/18; intestine commences in 16; last pair of hearts in 12; spermathecal diverticulum bilobate........................... nkandla sp. n. 6 Spermiductal funnels free; spermathecal duct extended; spermathecal diverticulum large, attached with duct below ampulla; intestine commences in segment 16; last pair of hearts in 12........................................................................mapelane sp. n. – Spermiductal funnels enclosed; spermathecal duct short; spermathecal diverticulum attached to duct at its base; intestine commences in segment 16; last pair of hearts in 12.......................................................................................pliskoae Zicsi, 1998 7 Intestine commences in 16 ...................................................................................8 – Intestine commences in 17 ...................................................................................9 296 AFRICAN INVERTEBRATES, VOL 45, 2004 8 one pair of prostates in segments 17–21; last pair of hearts in 12; spermathecal diverticulum unilobate......................................................................adriani sp. n. – One pair of prostates in segment 19, and additional vestigial parts of prostatic ducts in segment 17; spermathecal diverticulum bilobate ..........................petrosi sp. n. 9 Last pair of hearts in segment 12; spermathecal diverticulum unilobate ............... montanus Pickford, 1937 – Last pair of hearts in 13; spermathecal diverticulum unilobate........ anneae sp. n. 10 Intestine commences in 16 ...................................... kinbergi (Michaelsen, 1899) – Intestine commences in 17; last pair of hearts in segment 13 ................................ reichei (Ude, 1905) Udeina adriani sp. n. Figs 1–3 Etymology: Named in honour of Dr Adrian J. Armstrong of Ezemvelo KZN Wildlife, who supported research on Acanthodrilinae in KZN, and collected this species. Type material: Holotype: clitellate NMSA/Olig.03622 KwaZulu-Natal: Drakensberg, section of Giant’s Castle Game Reserve, Injasuthi [new spelling: Inijasuthi] (29º06'50.671"S:29º26'33.834"E), opposite camp, hillside at approx. 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, A. J. Armstrong. Figs 1–3. Udeina adriani sp. n. 1. Holotype. Right spermatheca, extended, 400 X. A = ampulla; Dv = diverticulum. 2. Left prostate, 180 X. Pg = prostatic gland; Pd = prostatic duct; Ps = penial seta. 3. Ectal parts of penial setae, 500 X.