The tardigrada of Southern Africa, with the description of minibiotus harrylewisi, a new species from KwaZulu-Natal, South Africa (Eutardigrada: Macrobiotidae).
Abstract: Nine species of tardigrades (Phylum Tardigrada) were found in moss and lichen samples in KwaZulu-Natal, South Africa, with two species being also present in Lesotho. Macrobiotus richtersi, Minibiotus intermedius, Hypsibius convergens, Ramazzottius theroni, Milnesium tardigradum, and Echiniscus duboisi were previously known to be part of the southern African fauna. Macrobiotus iharosi has not been reported before from this region. One species, Macrobiotus cf. echinogenitus, could be identified only to species complex because eggs were not found. One South African species, Minibiotus harrylewisi sp. n., is new to science and is described and illustrated. It differs from other Minibiotus species in its adult cuticle and its egg ornamentation. It most closely resembles M. furcatus, from which it differs in having smooth lunules on leg IV, smaller eyes, smaller and more rounded posterior cuticular pores, and eggs with a smooth shell and much longer, non-bifurcate processes. These new records bring to 61 the number of tardigrade species reported from southern African mosses, lichens and soil.

KEY WORDS: Tardigrada, Lesotho, South Africa, KwaZulu-Natal, new species, new records, check list.
Article Type: Report
Subject: Zoology (Identification and classification)
Zoology (Research)
Authors: Meyer, Harry A.
Hinton, Juliana G.
Pub Date: 12/01/2009
Publication: Name: African Invertebrates Publisher: The Council of Natal Museum Audience: Academic Format: Magazine/Journal Subject: Zoology and wildlife conservation Copyright: COPYRIGHT 2009 The Council of Natal Museum ISSN: 1681-5556
Issue: Date: Dec, 2009 Source Volume: 50 Source Issue: 2
Topic: Event Code: 310 Science & research
Geographic: Geographic Scope: South Africa Geographic Code: 6SOUT South Africa
Accession Number: 219376288
Full Text: INTRODUCTION

The first investigations of tardigrades (Phylum Tardigrada) in southern Africa were made in the early decades of the twentieth century, when Murray (1907, 1913) found 16 species in South Africa. Since then, studies of tardigrades in southern Africa (herein defined as encompassing modern Angola, Zambia, Malawi, Namibia, Zimbabwe, Botswana, Swaziland, Mozambique, Lesotho, and South Africa) have been sporadic. Currently, 14 papers report the presence in southern Africa of 16 genera and 60 species, including those identified only to genus or species complex (Murray 1907, 1913; Marcus 1936; da Cunha & do Nascimento Ribeiro 1964; Dastych 1980, 1992, 1993; Binda 1984; Pilato et al. 1991; Van Rompu et al. 1995; J0rgensen 2001; Middleton 2003; Kaczmarek & Michalczyk 2004; Kaczmarek et al. 2006).

Tardigrades have not been reported from Zambia and Swaziland. Most southern African tardigrades have been collected from cryptogams (Table 1). Soil has only rarely been sampled, and freshwater only once.

In this paper we report tardigrades collected from cryptogams in Lesotho and KwaZulu-Natal, South Africa.

MATERIAL AND METHODS

All samples were collected in midwinter on 9-11 July, 2008. Foliose lichens were collected from a stone fence in Sani Pass (SP), Lesotho and from Acacia trees at Hluhluwe-Imfolozi Game Park (HGP) and Tala Private Game Reserve (TPGR) in KwaZulu-Natal Province, South Africa. In KwaZulu-Natal ground moss was collected in Underberg. Eight samples were stored in paper envelopes until laboratory processing. Moss and lichen samples were soaked overnight in tap water and examined using a stereoscopic microscope. Specimens were mounted on slides in polyvinyl lactophenol and examined with phase contrast microscopy.

Tardigrades were identified using keys and descriptions in Nelson & McInnes (2002) and Ramazzotti & Maucci (1983), and by reference to the current literature. Anatomical terminology, global species lists, and current taxonomy and nomenclature are given according to Guidetti & Bertolani (2005) and Degma & Guidetti (2007). Comments on tardigrade zoogeography are based primarily on McInnes (1994).

All morphological measurements are given in micrometres. The pt index is the ratio of the length of a given structure to that of the buccal tube expressed as a percentage (Pilato 1981). Buccal tube length and the point of stylet support insertion were measured from the anterior margin of the stylet sheaths to the base of the tube. Buccal tube widths were measured as the external diameter of the buccal tube at the level of the stylet support insertion. The lengths of the primary and secondary branches of the claws were measured from the base to the apex, including accessory points.

The material has been deposited in the following collections: SMLA - W.A.K. Seale Museum, Department of Biology and Health Sciences, McNeese State University, Lake Charles, Louisiana, USA; NMSA - Natal Museum, Pietermaritzburg, South Africa.

TAXONOMY

Class Heterotardigrada Marcus, 1927

Order Echiniscoidea Richters, 1926

Family Echiniscidae Thulin, 1928

Genus Echiniscus Schulze, 1840

Twelve of the 158 species of Echiniscus have been reported in southern Africa (Table 1).

Echiniscus duboisi Richters, 1902

Figs 1-7, Table 2

Description: Descriptions of this species indicate considerable variation in the shape, length, and number of lateral and dorsal spines (Ramazzotti & Maucci 1983). In Richter's (1902) description and illustration, lateral spines B through E are serrated, and have the same length as the smooth dorsal spines [C.sup.d] and [D.sup.d]. Specimens from Cape Colony (Murray 1913), Brazil (de Barros 1942), and New Guinea (Iharos 1967) lack dorsal and lateral spines at B. Illustrations of E. duboisi vary considerably in their depiction of its habitus, cuticle, and spines, and only de Barros (1942) and Iharos (1967) provide limited morphometric data.

Table 2 presents morphometric data for twelve specimens of Echiniscus duboisi. Cuticular ornamentation consists of widely-spaced depressions ranging in diameter from 1.0-3.1 (Figs 1, 2). Cirri and spines are shorter, and the number of teeth on the dentate collars fewer, in South African than in Brazilian specimens. Three specimens lack lateral spine C, seven lateral spine D, and one dorsal spine Dd. Dorsal spines, especially Dd, are robust and serrated, with wide variation in number and size of teeth (Figs 3, 4). Lateral spines C and D are either serrated or smooth; E is always serrated (Figs 5-7).

[FIGURES 1-2 OMITTED]

Material examined: SOUTH AFRICA: KwaZulu-Natal: 24 specimens, TPGR, from foliose lichens on Acacia trees, 17 deposited at SMLA (accession numbers 9340-9346, 9349) and 7 at NMSA (accession number 9347).

[FIGURES 3-7 OMITTED]

Distribution: E. duboisi has not been found in the Nearctic or Palearctic realms, but has been widely reported elsewhere, including South Africa (Table 1).

Class Eutardigrada Richters, 1926

Order Parachela Schuster, Nelson, Grigarick & Christenberry, 1980

Family Hypsibiidae Pilato, 1969

Genus Hypsibius Thulin, 1928

Four of the 41 species of Hypsibius are known from southern Africa (Table 1). Hypsibius convergens (Urbanowicz, 1925)

Material examined: SOUTH AFRICA: KwaZulu-Natal: 36 specimens, HGP, in foliose lichens from Acacia trees (SMLA accession numbers 9361, 9362, 9364). Milnesium tardigradum, Minibiotus intermedius, and Macrobiotus richtersi were present in the same samples.

Distribution: H. convergens is a cosmopolitan species, previously found in Gauteng Province, South Africa (Table 1).

Genus Ramazzottius Binda & Pilato, 1986

Three of the 24 species of Ramazzottius are known from southern Africa (Table 1).

Ramazzottius theroni Dastych, 1993

Material examined: SOUTH AFRICA: KwaZulu-Natal: 31 specimens and 1 egg, HGP, in foliose lichens on Acacia trees (SMLA accession numbers 9340, 9341, 9353, 9355, 9357; NMSA accession number 9347). Echiniscus duboisi, Macrobiotus iharosi, Minibiotus harrylewisi sp. n., Min. intermedius, and Milnesium tardigradum were present in the same samples.

Distribution: Hitherto this species was known only from its type locality in Western Cape Province, South Africa.

Family Macrobiotidae Thulin, 1928

Genus Macrobiotus Schulze, 1834

Species of Macrobiotus cannot be identified without their eggs. Older literature often did not employ the rigorous standards of identification used today (Pilato & Binda 2001; Guidetti & Bertolani 2005). Up to now, 10 of the 161 species in this genus have been reported in southern Africa (Table 1).

Macrobiotus cf. echinogenitus Richters, 1904

Material examined: SOUTH AFRICA: KwaZulu-Natal: 1 specimen, Underberg, in moss (SMLA accession number 9359).

Distribution: Murray (1907) reported the presence of M. echinogenitus, a cosmopolitan species, in Cape Colony, South Africa. He did not record where in Cape Colony the specimens were collected but presumably they were from the modern provinces of Eastern Cape or Western Cape.

Remarks: The specimen keys to Macrobiotus echinogenitus in Ramazzotti & Maucci (1983). Its buccal and claw structure are consistent with M. echinogenitus, but in the absence of eggs, further identification is not possible.

Macrobiotus iharosi Pilato, Binda & Catanzaro, 1991

Material examined: SOUTH AFRICA: KwaZulu-Natal: 36 specimens and 8 eggs, TPGR, in foliose lichens on Acacia trees (SMLA accession numbers 9340, 9346, 9356, 9357; NMSA accession number 9347). Ramazzottius theroni, Milnesium tardigradum, Minibiotus harrylewisi sp. n., Min. intermedius, and Echiniscus duboisi were found in the same samples.

Distribution: The known distribution of M. iharosi has hitherto been restricted to Tanzania and the Democratic Republic of Congo (Pilato et al. 1991, 2003).

Macrobiotus richtersi Murray, 1911

Material examined: SOUTH AFRICA: KwaZulu-Natal: 5 specimens and 1 egg, TPGR, in foliose lichen on an Acacia tree (SMLA accession numbers 9363, 9364). Hypsibius convergens and Minibiotus intermedius were present in the same sample.

Distribution: This cosmopolitan species has previously been found in Angola, South Africa, and Zimbabwe (Table 1).

Genus Minibiotus Schuster, Nelson, Grigarick & Christenberry, 1980

Minibiotus currently consists of 42 species, and four of them are known to occur in southern Africa (Table 1). Minibiotus africanus occurs elsewhere in sub-Saharan Africa (Binda et al. 2001).

Guidetti et al. (2007) listed a suite of 10 characters shared by most, though not all, species of Minibiotus. These characters are: (1) anteroventral mouth; (2) oral cavity teeth absent or strongly reduced; (3) rigid, narrow buccal tube (width pt=12); (4) buccal tube thickened below the point of insertion of stylet supports; (5) stylet supports inserted at a considerable distance from the posterior end of the buccal tube (pt=73); (6) short ventral lamina (pt=73); (7) short macroplacoid row length (pt=42); (8) double curvature of the buccal tube; (9) first macroplacoid located very close to the pharyngeal apophyses; and (10) three almost rounded macroplacoids.

Minibiotus harrylewisi sp. n.

Figs 8-17; Tables 3, 4

Etymology: The specific epithet honors the first author's father, Harry Lewis Meyer, in recognition of his unstinting support of his son's academic career. Description (adult measurements are given from holotype unless otherwise indicated; pt index in brackets and italics):

Body length 336.0. Body white or transparent. Eyespots present in posterior position. Cuticle without sculpture or granulation (including legs), with 10 bands of circular, oval, trefoil, or quadrifoil pores (no stellate pores); bands 2-6 pores wide (Figs 8, 11-13). Circular and oval pores more common on anterior region of body; trefoil and quadrifoil more common on posterior body and also on legs. Pores significantly larger on posterior half of body (Fig. 11) than on anterior half (Fig. 12) (1.3, n=32 on anterior end; 1.8, n=27 on posterior end; p<0.0001; combined data from 10 specimens), and on legs (Fig. 13) than on rest of body (1.9, n=37 on legs; 1.5, n=60 on body; p<0.001; combined data from 10 specimens).

Peribuccal papillae absent. Oral cavity armature absent or not visible with light microscopy. Mouth anteroventral. Buccal tube rigid, 27.9 long and 2.6 [9.3] wide (Fig. 14), with double curvature (Fig. 14). Stylet support inserted on buccal tube at 17.7 [63.4]. Short ventral lamina, 17.9 [63.4] long. Pharyngeal apophyses triangular and close to first macroplacoid. First macroplacoid round and 2.7 [9.7] long; second pearshaped, 2.0 [7.2] long; third also pear-shaped, 2.5 [9.0] long. Macroplacoid row 9.8 [35.7] long. Indistinct, rod-shaped microplacoid 1.0 [3.6] long. Placoid row 11.7 [39.5] long. Pharyngeal bulb round, 35.5x36.0.

Claws short and slender. Primary branches of all claws with distinct accessory points. All claws with smooth-edged (not dentate or crenate) lunules but well developed only on leg IV. Leg I: primary branch with basal claw 7.5 [26.9] long, secondary branch 5.6 [20.7] long. Leg II: pb with bc 7.8 [28.0] long, sb 4.7 [76.8] long. Leg III: pb with bc 8.6 [28.7] long, sb 7.3 [26.2] long (Fig. 15). Leg IV pb with bc 9.3 [33.0] in length, sb 6.6 [23.7] (Fig. 16).

[FIGURES 8-10 OMITTED]

[FIGURES 11-13 OMITTED]

Eggs white/colourless, laid freely, with smooth surface (Fig. 17), with 32-41 long, thin, non-membranous processes around circumference. Processes densely packed, in the form of elongated, tapering cones with bulbous base. Size variable (Table 4).

Comparison: The presence of three macroplacoids in M. harrylewisi sp. n. differentiates it from species of Minibiotus with two. About half of the species in the genus have sculptured, granulated, or smooth cuticles; M. harrylewisi sp. n. differs from these in having a cuticle without sculpture or granulation, but having numerous pores. Among species with pores, the eggs of several have short processes covered with membranes, unlike the long, thin, non-membranous processes of M. harrylewisi sp. n. Among species with pores and conical egg processes, the new species lacks the large leg pore or large stellate pores characteristic of M. constellatus Michalczyk & Kaczmarek, 2003, M. siderius Pilato, Binda & Lisi, 2003, M. gumersindoi Guil & Guidetti, 2005, or M. eichhorni Michalczyk & Kaczmarek, 2004. Unlike M. vinciguerrae Binda & Pilato, 1992, the pores of M. harrylewisi sp. n. are arranged in distinct bands. The new species differs from M. pustulatus Ramazzotti, 1959 in having much smaller caudal pores (mean of caudal pores 1.8 in the new species and 6.7 in M. pustulatus) and longer egg processes, and from M. bisoctus Horning, Schuster & Grigarick, 1978 in possessing eyes, smaller pores (1.3-1.9 in the new species and 3 in M. bisoctus), and lack of leg granulation. Unlike M. ethelae, the caudal cuticle of M. harrylewisi sp. n. is not thickened. The new species most closely resembles M. furcatus. The lunules of Leg IV in M. harrylewisi sp. n. are not dentate. The pores of the new species, especially those on the anterior body, are smaller and more rounded. The egg processes in M. harrylewisi sp. n. are twice as long as in M. furcatus, are never bifurcated, and the egg shell is never punctate. Variation: Microplacoid often absent on smallest specimens. Division between the two most anterior and between the two most posterior bands of pores not distinct in some specimens.

[FIGURES 14-16 OMITTED]

[FIGURE 17 OMITTED]

Holotype: SOUTH AFRICA: KwaZulu-Natal: TPGR, 11.vii.1988, foliose lichen on Acacia trees, H.A. Meyer, deposited in W.A.K. Seale Museum, Department of Biology and Health Sciences, McNeese State University, Lake Charles, Louisiana, USA (accession number 9349).

Paratypes and other material examined: 6 paratypes and 7 eggs, same data as holotype (accession numbers 9340, 9343, 9346, 9349); 2 additional eggs, 109 additional specimens, same collection data as holotype; 1 paratype and 1 egg, same collection data as holotype, deposited in the Natal Museum, Pietermaritzburg, South Africa (accession numbers 9347, 9348); 1 cluster of three connected eggs found; 1 embryonated egg found with buccal apparatus and placoids.

Distribution: The new species is know from the type locality only. Echiniscus duboisi, Ramazzottius theroni, Macrobiotus iharosi, Minibiotus intermedius, and Milnesium tardigradum were present in the same lichens.

Minibiotus intermedius (Plate, 1888)

Material examined: SOUTH AFRICA & LESOTHO: 19 specimens, SP, HGP and TPGR, foliose lichens (SMLA accession numbers 9352, 9358, 9362, 9365). Also present in the same samples were Hypsibius convergens, Ramazzottius theroni, Macrobiotus iharosi, M. richtersi, Minibiotus harrylewisi sp. n., Milnesium tardigradum, and Echiniscus duboisi.

Distribution: A cosmopolitan species, Minibiotus intermedius has previously been found in Angola, Botswana, Namibia, and South Africa (Table 1).

Order Apochela Schuster, Nelson, Grigarick & Christenberry, 1980

Family Milnesiidae Ramazzotti, 1962

Genus Milnesium Doyere, 1840

Thirteen recent species of Milnesium are known to science. Of these, only one has been found in southern Africa (Table 1).

Milnesium tardigradum Doyere, 1840

Material examined: SOUTH AFRICA & LESOTHO: 15 specimens, SP, HGP and TPGR, in foliose lichens (SMLA accession numbers 9340, 9342, 9343, 9349, 9352, 9356-9358, 9361, 9362). Present in the same samples were Hypsibius convergens, Ramazzottius theroni, Macrobiotus iharosi, Minibiotus harrylewisi sp. n., Minibiotus intermedius, and Echiniscus duboisi.

Distribution: Milnesium tardigradum is a cosmopolitan species and has previously been found in Angola, Botswana, Lesotho, Namibia, South Africa, and Botswana (Table 1).

DISCUSSION AND CONCLUSIONS

In samples from KwaZulu-Natal and Lesotho nine tardigrade species were found. Two species were present in Lesotho and all nine were present in KwaZulu-Natal. Six species (M. richtersi, M. intermedius, H. convergens, R. theroni, M.tardigradum, and E. duboisi) were previously known from southern Africa, but M. iharosi represents a new record for the region. One species, Macrobiotus cf. echinogenitus, could not be identified beyond species complex, because no eggs were found. One South African species, M. harrylewisi sp. n., is new to science.

Pilato and Binda (2001) categorized 6.8% of all non-marine tardigrade species as cosmopolitan in distribution, i.e., reported in five or more biogeographical regions. In southern Africa 21 (40.4 %) of the fully identified species are cosmopolitan (Table 1). In comparison, only 17.6% of Nearctic tardigrade species are cosmopolitan (Meyer & Hinton 2007). While the proportion of southern African tardigrade fauna that is cosmopolitan is relatively high, the endemic proportion is correspondingly low. The known distribution of 11 species (21.2 %) is limited to southern Africa (Table 1), compared to 30.0 % endemic to North America (Meyer & Hinton 2007). On a global scale, 68.4 % of all terrestrial tardigrade species are found in only one biogeographic realm (Pilato & Binda 2001).

This study should be considered a preliminary survey of tardigrades in KwaZulu-Natal, South Africa. No doubt additional species remain to be found. Tardigrade distributions are extremely patchy at fine spatial scales (Meyer 2006), and only the rigorous sampling programs employed by All Taxa Biological Inventories (e.g. Bartels & Nelson 2006) can estimate their true diversity.

Guil and Cabrero-Sanudo (2007) noted that almost none of the authors of papers on tardigrades from the Afrotropical and Indomalayan realms are from those areas themselves. Among the authors of papers about southern African tardigrades, all except Middleton (2003) have been from Europe or North America. Our understanding of the distribution and abundance of tardigrades in southern Africa will benefit when researchers from the region play a greater role in their study.

ACKNOWLEDGEMENTS

We wish to thank Lindsay Fox for helping with sample processing, Stephen and Kathleen Jones for providing logistical support, and Harry L. Meyer, Lukasz Kaczmarek and an anonymous referee for valuable comments on earlier versions of the manuscript.

REFERENCES

Bartels, P.J. & Nelson, D.R. 2006. A large-scale, multihabitat inventory of the Phylum Tardigrada in the Great Smoky Mountains National Park, USA: a preliminary report. Hydrobiologia 558: 111-118.

Binda, M.G. 1984. Notizie sui tardigrade dell'Africa meridionale con descrizione di una nuova specie di Apodibius (Eutardigrada). Animalia 11: 5-15.

Binda, M.G., Pilato, G., Moncada, E. & Napolitano, A. 2001. Some tardigrades from Central Africa with the description of two new species: Macrobiotus ragnonesei and M. priviterae (Eutardigrada Macrobiotidae). Tropical Zoology 14: 233-242.

Da Cunha, A.X. & do Nascimento Ribeiro, F. 1964. Tardigrados de Angola: Contribuicao a seu etudo. Garcia de Orta (Lisbon) 12: 397-406.

Dastych, H. 1980. Hypsibius szeptycki sp. n., a new species of Tardigrada from South Africa. Bulletin de l'Academie Polonaise des Sciences, Serie des sciences biologiques 27: 505-508.

--1992. Paradiphascon manningi gen. n. sp. n., a new water bear from South Africa, with the erecting of a new subfamily Diphasconinae (Tardigrada). Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut 89: 125-139.

--1993. A new genus and four new species of semiterrestrial water-bears from South Africa (Tardigrada). Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut 90: 175-186.

Degma, P. & Guidetti, R. 2007. Notes to the current checklist of Tardigrada. Zootaxa 1579: 41-53.

de Barros, R.1942. Tardigrados do Estado do Sao Paulo, Brasil. I. Introducao. Generos "Echiniscus" e "Pseudechinscus." Revista Brasileira de Biologia 2: 257-269.

Guidetti, R. & Bertolani, R. 2005. Tardigrade taxonomy: an updated check list of the taxa and a list of characters for their identification. Zootaxa 845: 1-46.

Guidetti, R., Bertolani, R. & Degma, P. 2007. New taxonomic position of several Macrobiotus species (Eutardigrada: Macrobiotidae). Zootaxa 1471: 61-68.

Guil, N. & Cabrero-Sanudo, F.J. 2007. Analysis of the species description process for a little known invertebrate group: the limnoterrestrial tardigrades (Bilateria, Tardigrada). Biodiversity and Conservation 16: 1063-1086.

Iharos, G. 1967. Beitrage zur Kenntnis der Tardigradefauna von Neuguinea. Opuscula Zoologica Budapest 7: 111-116.

Jorgensen, A. 2001. Graphical presentation of the African tardigrade fauna using GIS with description of Isohypsibius malawiensis sp. n. (Eutardigrada: Hypsibiidae) from Lake Malawi. Zoologischer Anzeiger 240: 441-449.

Kaczmarek, L. & Michalczyk, L. 2004. Notes on some tardigrades from South Africa, with the description of Diphascon (Diphascon) zaniewi sp. n. Zootaxa 576: 1-6.

Kaczmarek, L., Beasley, C. & Michalczyk, L. 2006. The first record of the genus Haplohexapodibius Pilato & Beasley, 1987 in Africa with notes on synonymy of Hexapodibius beasleyi Maucci, 1988 with Haplohexapodibius seductor Pilato & Beasley, 1987. African Zoology 41: 290-293.

MARCUS, E. 1936. Tardigrada. Das Tierreich 66: 1-340.

McInnes, S.J. 1994. Zoogeographic distribution of terrestrial/freshwater tardigrades from current literature. Journal of Natural History 28: 257-352.

Meyer, H.A. 2006. Small-scale spatial distribution variability in terrestrial tardigrade populations. Hydrobiologia 558: 133-139.

Meyer, H.A. & Hinton, J.G. 2007. Limno-terrestrial Tardigrada of the Nearctic realm. Journal of Limnology 66 (Suppl. 1): 97-103.

Middleton, R.C. 2003. Tardigrades in southern Africa. African Journal of Ecology 41: 280-282.

Murray, J. 1907. Some South African Tardigrada. Journal of the Royal Microscopical Society 5: 515-524.

--1913. African Tardigrada. Journal of the Royal Microscopical Society 2: 138-144.

NELSON, D. R. & MCINNES, S. J. 2002. Tardigrada. In: Rundle, S.D., Robertson, A.L. & Schmid-Araya, J.M., eds, Freshwater meiofauna: Biology and ecology. Leiden: Backhuys, pp. 177-215.

Pilato, G. 1981. Analisi di nuovi caratteri nello studio degli Eutardigradi. Animalia 8: 51-57.

Pilato, G. & Binda, M.G. 2001. Biogeography and limno-terrestrial tardigrades: Are they truly incompatible binomials? Zoologischer Anzeiger 240: 511-516.

Pilato, G., Binda, M.G. & Catanzaro, R. 1991. Remarks on some tardigrades of the African fauna with the description of three new species of Macrobiotus Schultze 1834. Tropical Zoology 4: 167-178.

Pilato, G., Binda, M.G. & Lisi, O. 2003. Notes on some tardigrades from Central Africa, with the description of a new species of Hypsibiidae. Zootaxa 241: 1-7.

Ramazzotti, G. & Maucci, W. 1983. Il Philum Tardigrada. Memorie dell'Istituto Italiano di Idrobiologia 41: 1-1011.

Richters, F. 1902. Neue Moosbewohner. Bericht der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main 1902: 23-26.

Van Rompu, E.A., De Smet, W.H. & Coomans, A. 1995. Some terrestrial tardigrades from Zimbabwe. Bio logisch Jaarboek Dodonaea 62: 48-55.

Harry A. Meyer and Juliana G. Hinton Department of Biology and Health Sciences, McNeese State University, Lake Charles, Louisiana, 70609 USA;hmeyer@mcneese.edu
TABLE 1

Distribution of tardigrade species collected in southern Africa.
Abbreviations. Nations: AO-Angola, BW-Botswana, LS-Lesotho,
MW-Malawi, MZ-Mozambique, NA-Namibia, SA-South Africa, ZW-Zimbabwe.
South African provinces: CC-Cape Colony (where the
original reference does not permit assigning to modern provinces),
GT-Gauteng, KN-KwaZulu-Natal, LP-Limpopo, MP-Mpumalanga,
NW-North West, WC-Western Cape. Biogeographical regions:
Af-Afrtotropical, An-Antarctic and subantarctic, Au-Australian, I
-Indomalayan, Na-Nearctic, Nt-Neotropical, NZ-New Zealand, O
-Oceania, P-Palearctic. Substrates: M-moss, L-lichen,
C-unspecified cryptogam, S-soil, A-aquatic sediment. References:
B-Binda (1984), [D.sup.1]-Dastych (1980), [D.sup.2]-Dastych (1992),
D3-Dastych (1993), DN-da Cunha & do Nascimento Ribeiro (1964),
J-Jorgensen (2001), KB-Kaczmarek et al. (2006), KM-Kaczmarek &
Michalczyk (2004), Ma-Marcus (1936), Mi-Middleton (2003),
[Mu.sup.1]-Murray (1907), [Mu.sup.2]-Murray (1913), P-Pilato et
al. (1991), V-Van Rompu et al. (1995).

Species                          Substrate   Location and Reference

Bryodelphax parvulus Thulin,         L       AO(DN)
  1928
Echiniscus sp.                       C       LP(Mi), NW(Mi), ZW(Mi)
Echiniscus africanus Murray,         M       AO(DN), CC([Mu.sup.1]),
  1907                                       LP([Mu.sup.2])
Echiniscus cf. africanus             C       LS(Mi)
Echiniscus angolensis da Cunha       L       AO(DN), BW(Mi)
  & do Nascimiento Ribeiro,
  1964
Echiniscus arctomys Ehrenberg,       M       CC([Mu.sup.1])
  1853
Echiniscus bigranulatus              C       BW(Mi)
  Richters, 1907
Echiniscus crassispinosus            M       CC([Mu.sup.1])
  Murray, 1907
Echiniscus duboisi Richters,         M       CC([Mu.sup.2])
  1902
Echiniscus kerguelensis                      NA(Ma)
  Richters, 1904
Echiniscus limae da Cunha &          L       AO(DN)
  do Nascimento Ribeiro, 1964
Echiniscus longispinosus             M       CC([Mu.sup.1]), WC(B,P)
  Murray, 1907
Echiniscus merokensis                M       AO(DN)
  Richters, 1904
Echiniscus perarmatus Murray,        M       CC([Mu.sup.1])
  1907
Echiniscus pusae Marcus, 1928                CC(Ma)
Pseudechiniscus bispinosus           M       CC([Mu.sup.1])
  (Murray, 1907)
Pseudechiniscus jiroveci             M       WC(B,P)
  Bartos, 1963
Pseudechiniscus suillus              M       AO(DN), LP([Mu.sup.2])
  (Ehrenberg, 1853)
Doryphoribius citrinus               M       WC(B)
  (Maucci, 1972)
Hypsibius sp.                        C       LS(Mi)
Hypsibius convergens                 M       GT(KM)
  (Urbanowicz, 1925)
Hypsibius dujardini                  M       WC(B)
  (Doyere, 1840)
Hypsibius maculatus                  M       GT/MP(KM)
  (Iharos, 1969)
Hypsibius scabropygus                        NA(Ma)
  Cuenot, 1929
Isohypsibius deconincki              S       KN(B)
  Pilato, 1971
Isohypsibius kristenseni             S       MZ(P)
  Pilato, Catanzaro & Binda,
  1989
Isohypsibius malawensis              A       MW(J)
  Jorgensen, 2001
Isohypsibius nodosus                 M       CC([Mu.sup.1])
  (Murray, 1907)
Isohypsibius pseudoundulatus         M       AO(DN)
  (da Cunha & do Nascimento
  Ribeiro, 1964)
Isohypsibius sattleri                M       AO(DN), WC(B)
  (Richters, 1902)
Isohypsibius schaudinni              S       ZW(V)
  (Richters, 1909)
Ramazzottius sp.                     C       LS(Mi)
Ramazzottius oberhaeuseri            L       AO(DN)
  (Doyere, 1840)
Ramazzottius szeptycki               M       LP([D.sup.1])
  (Dastych, 1980)
Ramazzottius theroni Dastych,        L       WC([D.sup.3])
  1983
Diphascon (Adropion) scoticum        M       CC([Mu.sup.2])
  Murray, 1905
Diphascon (Diphascon) zaniewi        M       NW(KM)
  Kaczmarek & Michalczyk, 2004
Paradiphascon manningi             M, S      WC([D.sup.2])
  Dastych, 1992
Astatumen trinacriae               M, S      WC(B), ZW(V)
  (Arcidiacono, 1962)
Haplomacrobiotus seductor            M       GT/MP(KB)
  Pilato & Beasley, 1987
Calcarobiotus filmeri                S       MP([D.sup.3])
  Dastych, 1993
Calcarobiotus occulti              M, S      KN([D.sup.3])
  Dastych, 1993
Macrobiotus areolatus                M       MZ(B)
  Murray, 1907
Macrobiotus drakensbergi             M       LP([D.sup.3])
  Dastych, 1993
Macrobiotus echinogenitus            M       CC([Mu.sup.1])
  Richters, 1904
Macrobiotus furciger                 M       WC(B)
  Murray, 1906
Macrobiotus hufelandi              M, L      AO(DN), CC([Mu.sup.1]),
  C.A.S. Schultze, 1834                      LP([Mu.sup.2])
Macrobiotus cf. hufelandi            C       BW(Mi), LS(Mi)
Macrobiotus nuragicus Pilato         S       KN(B)
  & Sperlinga, 1975
Macrobiotus occidentalis             M       AO(DN)
  Murray, 1910
Macrobiotus richtersi              M, S      AO(DN), GT([Mu.sup.2]),
  Murray, 1911                               WC(B), ZW(V)
Macrobiotus cf. richtersi            M       LS(Mi), GT/MP(KM),
                                             WC(B)
Macrobiotus sapiens Binda &          L       NA(P)
  Pilato, 1984
Minibiotus sp.                       C       LS(Mi)
Minibiotus crassidens              M, L      CC([Mu.sup.1])
  (Murray, 1907)
Minibiotus furcatus                  L       AO(DN)
  (Ehrenberg, 1859)
Minibiotus harrylewisi sp. n.        L       KN(this paper)
Minibiotus hufelandioides            M       LP([Mu.sup.2])
  (Murray, 1910)
Minibiotus intermedius             M, L      AO(DN), BW(Mi), NA(Ma),
  (Plate, 1888)                              GT/MP(KM), WC(B,P)
Milnesium tardigradum              M, L      AO(DN), BW(Mi), LS(Mi),
  Doyere, 1840                               NA(Ma), CC([Mu.sup.1]),
                                             LP(Mi), NW(Mi), ZW(Mi)
Apodibius nuntius Binda, 1984      M, S      MZ(B), ZW(V)

Species                          Biogeography

Bryodelphax parvulus Thulin,     Na, Nt, P, Af, I
  1928
Echiniscus sp.
Echiniscus africanus Murray,     Af, I
  1907
Echiniscus cf. africanus
Echiniscus angolensis da Cunha   Nt, Af
  & do Nascimiento Ribeiro,
  1964
Echiniscus arctomys Ehrenberg,   Na, Nt, P, Af, I, O,
  1853                           Au, NZ, An
Echiniscus bigranulatus          Nt, P, Af, I, NZ
  Richters, 1907
Echiniscus crassispinosus        Nt, P, Af
  Murray, 1907
Echiniscus duboisi Richters,     Nt, Af, I, Au
  1902
Echiniscus kerguelensis          Na, Nt, P, Af, Au, NZ, An
  Richters, 1904
Echiniscus limae da Cunha &      Af
  do Nascimento Ribeiro, 1964
Echiniscus longispinosus         Af
  Murray, 1907
Echiniscus merokensis            Na, Nt, P, Af
  Richters, 1904
Echiniscus perarmatus Murray,    Na, Nt, Af, O
  1907
Echiniscus pusae Marcus, 1928    Af, I
Pseudechiniscus bispinosus       Nt, Af
  (Murray, 1907)
Pseudechiniscus jiroveci         P, Af
  Bartos, 1963
Pseudechiniscus suillus          Na, Nt, P, Af, I, O,
  (Ehrenberg, 1853)              Au, NZ, An
Doryphoribius citrinus           P, Af, I
  (Maucci, 1972)
Hypsibius sp.
Hypsibius convergens             Na, Nt, P, Af, I, O,
  (Urbanowicz, 1925)             Au, NZ, An
Hypsibius dujardini              Na, Nt, P, Af, I, O,
  (Doyere, 1840)                 NZ, An
Hypsibius maculatus              P, Af
  (Iharos, 1969)
Hypsibius scabropygus            Na, P, Af
  Cuenot, 1929
Isohypsibius deconincki          P, Af
  Pilato, 1971
Isohypsibius kristenseni         P, Af
  Pilato, Catanzaro & Binda,
  1989
Isohypsibius malawensis          Af
  Jorgensen, 2001
Isohypsibius nodosus             Na, Nt, P, Af, I, O, NZ
  (Murray, 1907)
Isohypsibius pseudoundulatus     Af
  (da Cunha & do Nascimento
  Ribeiro, 1964)
Isohypsibius sattleri            Na, Nt, P, Af, I, Au,
  (Richters, 1902)               NZ, An
Isohypsibius schaudinni          Na, Nt, P, Af, I
  (Richters, 1909)
Ramazzottius sp.
Ramazzottius oberhaeuseri        Nt, Nt, P, Af, I, O,
  (Doyere, 1840)                 NZ, An
Ramazzottius szeptycki           Af
  (Dastych, 1980)
Ramazzottius theroni Dastych,    Af
  1983
Diphascon (Adropion) scoticum    Na, Nt, P, Af, I, O,
  Murray, 1905                   Au, NZ, An
Diphascon (Diphascon) zaniewi    Af
  Kaczmarek & Michalczyk, 2004
Paradiphascon manningi           Af
  Dastych, 1992
Astatumen trinacriae             Na, P, Af, I
  (Arcidiacono, 1962)
Haplomacrobiotus seductor        Na, Af
  Pilato & Beasley, 1987
Calcarobiotus filmeri            Af
  Dastych, 1993
Calcarobiotus occulti            Af
  Dastych, 1993
Macrobiotus areolatus            Na, Nt, P, Af, I, O,
  Murray, 1907                   Au, NZ
Macrobiotus drakensbergi         Af
  Dastych, 1993
Macrobiotus echinogenitus        Na, Nt, P, Af, I, Au,
  Richters, 1904                 NZ, An
Macrobiotus furciger             Na, Nt, P, Af, I, NZ, An
  Murray, 1906
Macrobiotus hufelandi            Na, Nt, P, Af, I, O,
  C.A.S. Schultze, 1834          Au, NZ, An
Macrobiotus cf. hufelandi
Macrobiotus nuragicus Pilato     P, Af, I
  & Sperlinga, 1975
Macrobiotus occidentalis         Na, Nt, P, Af, O, Au, NZ
  Murray, 1910
Macrobiotus richtersi            Na, Nt, P, Af, I, O,
  Murray, 1911                   Au, NZ
Macrobiotus cf. richtersi
Macrobiotus sapiens Binda &      P, Af, I
  Pilato, 1984
Minibiotus sp.
Minibiotus crassidens            P, Af, O, Au
  (Murray, 1907)
Minibiotus furcatus              Na, Nt, P, Af, I
  (Ehrenberg, 1859)
Minibiotus harrylewisi sp. n.    Af
Minibiotus hufelandioides        P, Af, O, Au
  (Murray, 1910)
Minibiotus intermedius           Na, Nt, P, Af, I, O,
  (Plate, 1888)                  Au, NZ, An
Milnesium tardigradum            Na, Nt, P, Af, I, O,
  Doyere, 1840                   Au, NZ, An
Apodibius nuntius Binda, 1984    Af

TABLE 2

Morphometric data of selected structures on 12 specimens of
Echiniscus duboisi from KwaZulu-Natal, South Africa. Ranges
and means are lengths in [micro]m. Range refers to the smallest
and largest structure found among all measured specimens
(a zero indicates the structure was not present on some
specimens). Abbreviations: N--number of specimens measured,
SD--standard deviation.

Character                    N       Range        Mean     SD

Body                        12    176.6-292.9    219.4    40.8
Internal cirrus              8      6.5-15.5      11.2     2.7
Cephalic papilla             9      3.9-8.4        6.0     9.5
External cirrus              9      9.1-23.7      16.7     4.4
Cirrus A                    12     19.6-33.5      28.8     4.0
Clava                       11      3.3-7.2        5.2     1.3
Lateral spine C              9        0-16.4      10.9     3.0
Lateral spine D              5        0-16.7      13.7     2.8
Lateral spine E             12      4.7-17.6      12.8     3.2
Dorsal spine C (d)          11        0-15.9      11.7     4.2
Dorsal spine D (d)          12     12.2-25.1      20.3     4.1
Leg I spine                  8      1.3-2.5        2.0     0.4
Leg IV papilla              11      2.2-4.1        3.2     0.6
No. of teeth on dentate     12        6-11         8.6     1.3
  collar, Leg IV
Dentate teeth, Leg IV        4      1.7-4.6        3.4     1.0

TABLE 3

Morphometric data and pt values of selected structures of 10
specimens of Minibiotus harrylewisi sp. n. All lengths in
[micro]m. Range refers to the smallest and largest structure
found among all measured specimens (a zero indicates the
structure was not present on some specimens). Abbreviations:
N--number of specimens measured, SD--standard deviation,
n.a.--not applicable, pt--pt index.

                                               Range

Character                        N     Length         pt

Body                             10    200.3-386.6    n.a.
Buccal tube                      10    19.7-32.9      n.a.
Stylet support insertion         10    12.1-21.8      61.4-67.6
Buccal tube width                10    1.9-3.4        6.7-11.2
First macroplacoid               10    1.3-3.7        5.2-12.4
Second macroplacoid              10    1.2-2.6        6.0-8.6
Third macroplacoid               10    1.2-2.9        4.7-10.0
Macroplacoid row                 10    5.6-10.9       26.3-36.6
Microplacoid                     7     0-2.6          0-7.9
Placoid row                      7     10.4-34.7      34.6-46.0
Primary claw branch, Leg I       10    5.5-9.7        23.1-32.6
Secondary claw branch, Leg I     10    4.0-7.0        16.4-23.1
Primary claw branch, Leg II      10    6.3-10.5       27.4-35.2
Secondary claw branch, Leg II    10    3.6-7.9        16.8-27.1
Primary claw branch, Leg III     10    6.5-9.8        27.9-34.3
Secondary claw branch, Leg III   10    3.4-8.0        16.0-26.8
Primary claw branch, Leg IV      10    6.7-13.6       26.9-45.6
Secondary claw branch, Leg IV    10    4.5-9.9        19.3-33.2

                                                 Mean

Character                              Length         pt

Body                                   304.2          n.a.
Buccal tube                            28.1           n.a.
Stylet support insertion               18.1           64.1
Buccal tube width                      2.6            9.3
First macroplacoid                     2.2            8.0
Second macroplacoid                    2.0            7.2
Third macroplacoid                     2.2            7.7
Macroplacoid row                       9.1            32.3
Microplacoid                           1.4            4.5
Placoid row                            11.8           39.4
Primary claw branch, Leg I             7.7            27.6
Secondary claw branch, Leg I           5.6            19.9
Primary claw branch, Leg II            8.3            29.7
Secondary claw branch, Leg II          5.8            20.5
Primary claw branch, Leg III           8.8            31.1
Secondary claw branch, Leg III         6.4            22.4
Primary claw branch, Leg IV            9.9            35.2
Secondary claw branch, Leg IV          6.9            2.0

                                                 SD

Character                              Length         pt

Body                                   65.7           n.a.
Buccal tube                            4.3            n.a.
Stylet support insertion               3.1            3.8
Buccal tube width                      0.5            1.3
First macroplacoid                     0.7            2.1
Second macroplacoid                    0.4            0.9
Third macroplacoid                     0.6            1.8
Macroplacoid row                       1.8            3.7
Microplacoid                           0.7            2.2
Placoid row                            1.2            1.0
Primary claw branch, Leg I             2.9            6.4
Secondary claw branch, Leg I           1.0            1.9
Primary claw branch, Leg II            2.6            8.5
Secondary claw branch, Leg II          1.5            3.6
Primary claw branch, Leg III           1.2            2.4
Secondary claw branch, Leg III         2.4            1.6
Primary claw branch, Leg IV            2.1            4.7
Secondary claw branch, Leg IV          24.4           4.5

TABLE 4

Morphometric data of selected structures on seven eggs of
Minibiotus harrylewisi sp. n. Ranges and means are in [micro]m.
Range refers to the smallest and largest structure found among
all measured specimens. Abbreviations: N--number of specimens
measured, SD--standard deviation.

Character                     N    Range         Mean     SD

Diameter without processes    7    66.1-80.0     70.7     5.4
Diameter with processes       7    78.2-101.9    88.5     7.2
Height of process             9    7.6-12.8      10.2     1.4
Width of process base         9    3.2-5.6       4.3      0.7
Distance between processes    9    1.3-4.1       2.8      4.0
Number of processes           7    32.0-41.0     36.6     3.2
  around circumference
Number of processes in        5    94.0-166.0    118.8    32.6
  optical section
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