ISSN 0067-9208

NATURAL SCIENCES

VOLUME 29, PART 2 NOVEMBER 2013

First records of the -eating Dasypeltis confusa Trape & Mané, 2006 in Nigeria and , with range extensions for Ghana

by

Michael F. Bates

Department of Herpetology, National Museum, P.O. Box 266, Bloemfontein 9300, South E-mail: [email protected]

ABSTRACT

Bates, M.F. 2013. First records of the egg-eating snake Dasypeltis confusa Trape & Mané, 2006 in Nigeria and Chad, with range extensions for Ghana. Navors. nas. Mus., Bloemfontein 29(2): 17-27. New records are provided for the egg-eating snake Dasypeltis confusa from three West African countries. These include the first records of this for Nigeria, based on specimens from four localities (Jos and Zonkwa areas) in the centre of the country; and the first record for Chad, based on a specimen from Mayo-Kebbi in the south-west of the country. In addition, the known range in Ghana is extended to Yabraso in the central part of the country and the town of Wa in the north-west. The absence of records in several areas of apparently suitable savanna habitat in west and suggest that this species may have a much more extensive range than currently known. (Dasypeltis confusa, distribution, , Nigeria, Chad, Ghana)

ISBN 978-1-86847-154-6 18 Navors. nas. Mus., Bloemfontein, Volume 29, Part 2

CONTENTS

INTRODUCTION ...... 18 MATERIALS AND METHODS ...... 20 RESULTS AND DISCUSSION ...... 20 Scalation ...... 20 Colour pattern ...... 21 Geographical distribution ...... 23 ACKNOWLEDGEMENTS ...... 25 REFERENCES ...... 25

INTRODUCTION

Trape & Mané (2006a) described three new species of Dasypeltis Wagler, 1830 from West Africa, including Dasypeltis confusa. Trape et al. (2012) subsequently confirmed the status of these species based on a molecular analysis of West African Dasypeltis. These authors also described Dasypeltis parascabra, raised Dasypeltis gansi latericia Trape & Mané, 2006 to full species status, and confirmed that (Linnaeus, 1758) does not occur in West Africa. There are now 12 recognized species of egg-eating ( Dasypeltis) in Africa and the (Gans 1959, 1964; Trape et al. 2012).

Dasypeltis confusa occurs mainly in savanna areas of sub-Saharan Africa south of 14oN latitude. It appears to be largely restricted to the Savanna, from southern Senegal to Guinea, southern Mali, southern Ivory Coast, Ghana, Togo and Benin, southwards to , Gabon, Republic of the Congo, Angola, western Zambia and parts of Democratic Republic of the Congo, then eastwards through to Southern and (Gans 1959, 1964―‘D. scabra’ with ‘5L’ dorsal colour pattern; Chirio & LeBreton 2007; Pauwels & Vande werghe 2008; Bates & Broadley 2008a,b; Trape & Mané 2006a,b; Segniagbeto et al. 2011; Trape et al. 2012; Bates et al. 2012; Broadley & Bates in prep.). In Cameroon it has also been recorded from savanna- forest mosaic (Chirio & LeBreton 2007), and it occurs in both savanna and coastal in Gabon (Pauwels & Vande werghe 2008). However, many areas within its general range are poorly surveyed and lack any records (see Trape et al. 2012).

Dasypeltis confusa is characterised by its ‘5L’ dorsal colour pattern (see Gans 1959) which consists of large dark saddles, oval to more-or-less diamond-shaped, joined for the most part to the dark lateral bars on either side of the body, on a grey to pale brown back (Fig. 1; Trape & Mané 2006a,b). The dark markings often have a distinct white margin that tends to emphasise the linked pattern (e.g. pl. 8, fig 6 in Gans 1959; p. 403 in Chirio & LeBreton 2007; fig. 205 in Pauwels & Vande weghe 2008). Also helpful for distinguishing D. confusa from other species of Dasypeltis―at least in West Africa―is the combination of 23-26 midbody scale rows; ventrals 213-227 (males), 224-242 (females); and subcaudals 65-73 (males), 53-67 (females) (Trape & Mané 2006a; repeated by Trape et al. 2012).

2013 M.F. Bates – New records of Dasypeltis confusa in West Africa 19

Figure 1: Dorsal view of the holotype (MNHN 2006.0303, adult female; Ibel, south-eastern Senegal) of Dasypeltis confusa at the Muséum national d’Histoire naturelle, Paris. The specimen was photographed while submerged in ethanol. (Photo: M.F. Bates)

Most of Gans’ (1959) Nigerian Dasypeltis are referable to the forest species Dasypeltis fasciata A. Smith, 1849 (but see discussion below). Although he did not record any Dasypeltis with a ‘5L’ dorsal colour pattern (= D. confusa) from this country, Gans did report a ‘5N’ D. scabra from Lagos which Trape et al. (2012) later referred to their new species, Dasypeltis parascabra. However, Gans (1959) did record ‘D. scabra’ with ‘5L’ dorsal pattern (= D. confusa) from the Batouri area (East Province) in adjacent Cameroon. In a table (appendix B) showing the occurrence of snake species in savanna of Africa, Hughes (1983) recorded D. scabra from Nigeria (no specific localities given), while Chirio & LeBreton (2007) list Nigeria in the range of D. confusa without providing detailed localities. This species has not previously been recorded from Chad (see Gans 1959, 1964; Trape & Mane 2006a,b; Trape et al. 2012). Although D. confusa has been recorded from south-eastern Ghana (as ‘5L D. scabra’, Gans 1959; see also Trape et al. 2012), there are no records of this species in the central and north-western parts of the country.

During September 2011 the author visited the Natural History Museum in London and the Muséum national d’Histoire naturelle in Paris to examine specimens of Dasypeltis as part of a revision of the and distribution of the genus. The opportunity was taken to examine in detail the holotype of D. confusa in Paris (MNHN 2006.0303) and several specimens with the ‘5L’ dorsal colour pattern (see Gans 1959), including a few specimens from West Africa (Nigeria, Chad and Ghana). Specimens examined included the first records of D. confusa from Nigeria and Chad, and the first records of this species from north-western Ghana.

20 Navors. nas. Mus., Bloemfontein, Volume 29, Part 2

MATERIALS AND METHODS

Characters important for identification purposes were examined, for the most part with the aid of a stereo-microscope, according to the methods of Gans (1959). Ventrals were counted according to Dowling (1951) with additional typical ventrals (often called pre- ventrals) indicated after a plus sign (see below). The total count was used by Gans (1959; i.e. all distinctly widened ventral plates behind the second pair of chin shields, excluding the anal plate). Both sides of the head were examined and scale counts are presented for one side only unless there was variation between left and right sides. Dorsal saddles (equivalent to Gans’ 1959 ‘pattern cycles’) were counted from the nape to above the vent. Sex was determined by presence of testes or hemipenes (males) or (female).

An additional specimen (USNM 223931) of Dasypeltis confusa from Ghana was located at the National Museum of Natural History, Smithsonian Institution (Washington, D.C.). For this specimen, measurements as well as ventral, subcaudal and midbody scale row counts, were provided by J. Jacobs. Detailed photographic images of this specimen (male, right hemipenis everted) were provided by J. Poindexter II, and these allowed accurate examination of head shields.

RESULTS AND DISCUSSION

Scalation For all specimens of Dasypeltis confusa (locality details follow below): nasal divided below the nostril (divided above and below nostril on both sides of head in BM 1966.265, and on right side only in BM 1977.755); temporals on either side of head 2+3 or 2+4 (3+3 on right side in MHNH 1965.37); preoculars 1; postoculars 2 (but one on left side of BM 1973.2250, and one on right side of USNM 223931); supralabials usually 7 (3rd and 4th enter orbit), but 6 (3+4) in BM 1973.2250; frontal shield smooth except for a single row (two rows in BM 1982.433) of shallow pits on the outer margins; inter-prefrontal sulcus usually weakly developed with prefrontals merely in contact, but moderately developed in BM 1966.265 and BM 1973.2251; 2-4 rows of lateral dorsal scales serrated (usually 3rd to 5th or 6th row, but 4th and 5th rows only in BM 1966.265, and 4th to 6th rows in BM 1977.755).

Nigeria BM 1977.755 – female (gravid with nine eggs, six evident in body plus three found in jar) – Vom, SSW of Jos, Plateau State (09°44’N, 08°47’E, c. 1250-1320 m a.s.l.), collected 25 November 1976 by D.P. Britt. Ventrals 227+1, paired subcaudals 57, midbody scale rows 27, dorsal saddles 54.

BM 1966.265 – male (swollen testes) – Zonkwa, Kaduna State (09°47’N, 08°17’E, c. 856 m a.s.l.), collected by G.T. Dunger. Ventrals 217+1, paired subcaudals 75, midbody scale rows 25, dorsal saddles 59.

BM 1973.2250 – male (inverted left hemipenis) – Naraguta village, NNE of Jos, Plateau State (09°59’N, 08°54’E, c. 1100-1120 m a.s.l.), collected by G.T. Dunger. Ventrals 219+1, paired subcaudals 71, midbody scale rows 24, dorsal saddles 56.

2013 M.F. Bates – New records of Dasypeltis confusa in West Africa 21

BM 1973.2251 (Fig. 2) – male (inverted right hemipenis) – Jos, Plateau State (09°56’N, 08°53’E, c. 1170-1220 m a.s.l.), collected in 1966 by G.T. Dunger. Ventrals 220+1, paired subcaudals 67, midbody scale rows 25, dorsal saddles 54.

Chad MNHN 1965.37 – male (inverted right hemipenis) – Mayo-Kebbi (as discussed below), collected in 1964 by Sister Marie Reginald. Ventrals 223+1, paired subcaudals 69, midbody scale rows 25, dorsal saddles 59.

Ghana BM 1982.433 – male (everted hemipenes) – Wa, Upper West (10°04’N, 02°30’W, 300-340 m a.s.l.), collected 9 March 1981 by S. Spawls. Ventrals 212+1, paired subcaudals 46+ (tip broken), midbody scale rows 25, dorsal saddles 52.

USNM 223931 (Fig. 3) – male (right side hemipenis everted) – Yabraso, Brong-Ahafo Region (08º04’N, 01º48’W, c. 195 m a.s.l.), collected 9 April 1968 by J.C. Geest. Snout- vent length 470 mm, tail length 106 mm. Ventrals 225+1 (most anterior plate paired), paired subcaudals 75, midbody scale rows 25; dorsal saddles 50.

The midbody scale row count of 27 for the Vom female (BM 1977.755) is higher than the range (23-26) for the species in West Africa as recorded by Trape & Mané (2006a), but for ‘5L’ specimens (= D. confusa), Gans (1959) gives a range of 22-27 for Ghana and 23-26 for Cameroon (these are the only ‘West African’ countries where he records ‘5L D. scabra’). The subcaudal count of 75 for the Zonkwa (BM 1966.265) and Yabraso (USNM 223931) males slightly exceeds the highest recorded for West Africa (range given as 65-73 by Trape & Mané 2006; and 64-74 by Gans 1959 for Ghana, with no data for males from Cameroon).

Colour pattern Despite variation, all specimens can be classified as having a ‘5L’ dorsal pattern (Gans 1959; Trape & Mané 2006a). However, in some specimens the saddles and lateral bars are often, or even usually, not actually linked. For the Zonkwa specimen (BM 1966.265) the dorsal saddles were in the shape of narrow, elongated diamonds, mostly linked to the lateral bars, with distinct pale inter-saddle spaces. BM 1982.433 and BM 1977.55 were similar to BM 1966.265, but BM 1977.55 had larger diamond-shaped saddles. However, in four snakes (BM 1973.2250 & .2251, USNM 223931, MNHN 1965.37) the lateral bars were for the most part not in contact with the (mostly oval to rectangular) dorsal saddles, although they were usually positioned alongside them (Figs 2 & 3) rather than between them as in typical D. scabra (e.g. Gans 1959). This is similar to the pattern in Gans’ (1959) image (pl. 8, fig. 5) of a specimen from Brazzaville, and also similar to the D. confusa from Bourrofaye in Senegal illustrated in Trape & Mané (2006a). In contrast, Gans’ (1959) image (pl. 8, fig. 6) of a specimen from Semliki in Uganda shows “the characteristic linked scabra pattern” with large, elongated, diamond-shaped dark saddles joined at the sides to dark lateral bars, with distinct pale borders and pale inter-saddle spaces. These pale borders and spaces were usually not distinct (e.g. Fig. 2) in the specimens examined, probably due to long periods of preservation. In BM 1973.2250 and USNM 223931 (anteriorly, see Fig. 3) some of the saddles and lateral bars were arranged in the ‘5N’ unlinked pattern, i.e. lateral bars distributed between rather than linked to (or lying alongside) the saddles. Such variation in the linked pattern as described above may cause confusion and result in 22 Navors. nas. Mus., Bloemfontein, Volume 29, Part 2

Figure 2: Dorsal view of male Dasypeltis confusa (BM 1973.2251) from Jos, northern Nigeria. (Photo: P. Campbell, Natural History Museum, London)

Figure 3: Dorsal view of male Dasypeltis confusa (USNM 223931) from Yabraso, central Ghana. (Photo: J. Poindexter II, National Museum of Natural History, Smithsonian Institution, Washington, D.C.)

2013 M.F. Bates – New records of Dasypeltis confusa in West Africa 23 specimens of D. confusa being identified as D. scabra. This may have been the case with some of the Benin material examined by Hughes (2013). The venter of most specimens was cream (pale green in BM 1973.2250 & .2251 from the Jos area) with small dark specks, and often also occasional larger spots, at the edges, but in BM 1966.265 the anterior part of each ventral plate was dark.

Geographical distribution The new Nigerian localities (Fig. 4) are the first detailed and confirmed records of D. confusa for this country. These records are situated about 800 km east of the nearest other D. confusa records in Benin and over 400 km west of the nearest records in northern Cameroon (Chirio & LeBreton 2007; Trape et al. 2012). There are also two specimens (CMNH 92743 & 92744) referable to D. confusa (photographic images examined, dorsal pattern similar to Fig. 1, venter cream with some brown stippling at edges) in the Carnegie Museum of Natural History (Pittsburgh) labeled ‘Nigeria ?’, collected in either Nigeria or Cameroon in the period 1960 to early 1980s (S. Rogers in litt. 16 July 2013). As most of Nigeria is covered by savanna, it is perhaps surprising that there appear to be so few specimens of D. confusa from there in museum collections, and that this species was not previously confirmed as occurring in the country. However, there are also several records of this species in Cameroon on the border with Nigeria (Chirio & LeBreton 2007), so it is likely to be much more widespread in the latter country. The author has examined a female D. confusa (Zoologische Museum, Universität Hamburg: ZMH R08929) from Kuti (06º28’N, 11º26’E, c. 766 m a.s.l.) in Cameroon, about 13 km south of the border with Nigeria (SVL 467 mm, tail length 80 mm, ventrals 216+1, paired subcaudals 55, midbody scale rows 23, 56 dark dorsal saddles linked for the most part to dark lateral bars).

The Mayo-Kebbi record in Chad (Fig. 4) represents a range extension of at least 250 km north-east of the nearest other D. confusa record in northern Cameroon (Chirio & LeBreton 2007). The latter authors also mapped a locality between Bélel and Bocaranga on the eastern border of Cameroon—slightly further away and south of Mayo-Kebbi—which probably corresponds with a record plotted at the border between Cameroon, Chad and Central African Republic by Trape et al. (2012). From 1960 to 2002, Chad was divided into 14 prefectures, one of which was called ‘Mayo-Kebbi’, an area in the south-west of the country bordering northern Cameroon. The country was later divided into ‘regions’, with Mayo-Kebbi being subdivided into Mayo-Kebbi Est and Mayo-Kebbi Ouest. While it is not possible to determine exactly where the ‘Mayo-Kebbi’ specimen was collected, the area lies largely within the western block of the East , habitat compatible for this species.

Trape et al. (2012) plotted records of D. confusa in south-eastern Ghana, apparently based on specimens of ‘D. scabra’ with a ‘linked’ (‘5L’) dorsal pattern (i.e. D. confusa) as listed by Gans (1959). The latter author recorded definite ‘5L’ specimens from three localities in south-eastern Ghana: Accra (5o33’N, 0o12’W), ‘Kete’ (= Kete Krachi) (7o48’N, 0o1’W) and Somanya (6o6’14”N, 0o0’54”W), and also possible ‘5L’ specimens (listed as ‘5L?’) from Achimota (5o37’N, 0o14’W) and Legon Hill, Achimota (5o38’N, 0o13’W), both near the coast in the south-east of the country. The Wa record represents a range extension of 370 km north-west of Kete Krachi. The nearest record to Wa outside Ghana is at Mamarabougou in southern Mali (Trape et al. 2012), 348 km to the north-west. The Yabraso record extends the known range to the central parts of the country, and it now seems likely that D. confusa is widespread in savanna in most parts of Ghana outside the 24 Navors. nas. Mus., Bloemfontein, Volume 29, Part 2

forested region of the south-west. In Ghana, D. gansi has also been recorded at Wa (see Bates et al. 2013) and at Achimota School (see Bates & Ineich 2012).

Figure 4: Distribution of Dasypeltis confusa in Nigeria and Chad. For the latter country the only known record is at ‘Mayo-Kebbi’, the name of a former prefecture or region (shaded black). The recorded range of this species in the parts of Cameroon shown in the figure is indicated by dark grey shading (according to Chirio & Le Breton 2007). The approximate extent of forest is indicated in medium grey, Sudan-Guinea Savanna in light grey, and the in white.

Regarding the northern limits of D. confusa, it can be noted that Gans (1964) reported on a

‘5L D. scabra’ from Diafabare in Mali which he plotted (his fig. 3) south-east of Bandiagara near the border with Burkina Faso. Trape et al. (2012) listed the same specimen as D. confusa, but the locality name was preceded by a question mark and it was not plotted on their map, perhaps because the given locality appears to be situated in the arid Sahel, habitat more typical of D. sahelensis Trape & Mané, 2006, and because of the discrepancy between where Gans (1964) plotted the locality (near the border with Burkina Faso) and its actual position on a map.

Two of Gans’ (1959) Nigerian localities for D. fasciata were not plotted on the map for this species in Trape et al. (2012). The first is Wushishi (near Zungeru) in the central-western part of the country. This is a savanna area where D. fasciata is not expected to occur (Trape & Mane 2006a), although Gans’ (1959) data (male, 244 ventrals, 75 subcaudals, typical fasciata-like dorsal pattern) fits the latter species. The other locality is Gadana (apparently an old place name in northern Cameroon), south of Lake Chad, also in savanna habitat. For this male specimen with 227 ventrals (no subcaudal count given), Gans (1959) was somewhat uncertain about the dorsal pattern, but he referred it (questionably) to D. fasciata.

2013 M.F. Bates – New records of Dasypeltis confusa in West Africa 25

For males from West Africa, this ventral count is at the high end of the range for D. confusa (213-227) and at the low end of the range for D. fasciata (227-245), but well within the range of Dasypeltis gansi Trape & Mané, 2006 (221-240), a savanna species recently recorded from Lake Chad (Trape & Mane 2006a; Trape et al. 2012; Bates & Ineich 2012). Dorsally patterned D. gansi have in the past been confused with D. fasciata (see Trape & Mane 2006a), so the Gadana specimen may indeed represent the former species. The maps of both Trape & Mané (2006a) and Trape et al. (2012) plot a record for D. gansi in this region (south of Lake Chad), but as they do not make specific mention of the record, it is not possible to say whether it represents the same specimen examined by Gans (1959).

Although D. confusa appears to have a vast range, its distribution has not been investigated in detail, especially in areas outside West Africa (Trape & Mané 2006a,b; Trape et al. 2012) and adjacent Cameroon (Chirio & LeBreton 2007). Future investigations may show that D. confusa is even more widespread in the savannas of western and central Africa.

ACKNOWLEDGEMENTS

I thank the director and council of the National Museum (Bloemfontein) for the opportunity to conduct and publish this work; Patrick Campbell at the Natural History Museum (London) and Ivan Ineich at the Muséum national d’Histoire naturelle (Paris) for facilitating my visits in September 2011; Jeremy Jacobs and James Poindexter II for respectively examining, and supplying photographs of, specimens at the National Museum of Natural History, Smithsonian Institution (Washington, D.C.); Steve Rogers for photographs and information pertaining to specimens in the Carnegie Museum of Natural History (Pittsburgh); Alexander Haas for the loan of ZMH R08929 (Kuti, Cameroon); Steve Spawls for providing photographs of Dasypeltis confusa from Wa, Ghana; and Aaron Bauer for copies of literature. Patrick Campbell also supplied photographs of BM 1973.2250 & .2251. Craig Barlow (National Museum) is thanked for preparing the map (Fig. 4).The author’s trip to London and Paris was funded by the Speciation Project (a South African National Research Foundation [NRF] grant to Krystal A. Tolley, South African National Biodiversity Institute, Cape Town). Dr Donald G. Broadley (Natural History Museum of Zimbabwe, Bulawayo) and an anonymous reviewer are thanked for their comments on the manuscript.

REFERENCES

BATES, M.F. & BROADLEY, D.G. 2008a. Geographical Variation in Egg-eating Snakes of the Genus Dasypeltis in Africa south of Latitude 12o S. Sixth World Congress of Herpetology, Manaus, Brazil (17-22 August 2008) (Abstracts).

BATES, M.F. & BROADLEY, D.G. 2008b. Geographical variation in scalation and colour pattern in the egg- eating snakes of the genus Dasypeltis in Africa south of Latitute 12o S. Ninth Conference of the Herpetological Association of Africa, Sterkfontein Dam, Harrismith, South Africa (26-30 November 2008) (Abstracts).

BATES, M.[F.], BROADLEY, D.[G.], BARLOW, A., WÜSTER, W. & TOLLEY, K.[A.] 2012. Taxonomy and distribution of the African egg-eating snakes of the genus Dasypeltis. Seventh World Congress of Herpetology, Vancouver, Canada (8-14 August 2012) (Abstracts). African Herp News 58: 27-28.

BATES, M.F. & INEICH, I. 2012. A new size record for the West African egg-eating snake Dasypeltis gansi Trape & Mane, 2006, with new distribution records. Navors. nas. Mus., Bloemfontein 28(3): 41-48.

BATES, M.F., TRAPE J.-F. & SPAWLS, S. 2013. Geographical distribution: Dasypeltis gansi. African Herp News 60: 30-32.

CHIRIO, L. & LeBRETON, M. 2007. Atlas des du Cameroun. Paris: Muséum national d’Histoire naturelle, IRD.

26 Navors. nas. Mus., Bloemfontein, Volume 29, Part 2

DOWLING, H.G. 1951. A proposed standard system of counting ventral scales in snakes. Brit. J. Herpetol. 1(5): 97-99.

GANS, C. 1959. A taxonomic Revision of the African Snake Genus Dasypeltis (Reptila : Serpentes). Ann. Mus. Roy. Congo Belge, Tervuren, Ser. 8, 74: 1-237.

GANS, C. 1964. Further comments on the forms of the African snake genus Dasypeltis (Reptilia : Serpentes). Rev. Zool. Bot. Afr. 69(3,4): 279-295.

HUGHES, B. 1983. African snake faunas. Bonn. Zool. Beitr. 34(1-3): 311-356.

HUGHES, B. 2013. Snakes of Bénin, West Africa. Bull. Soc. Herp. Fr. 2012 [2013] 144: 101-159.

PAUWELS, O.S.G. & VANDE WEGHE, J.P. 2008. Les reptiles du Gabon. Washington: Smithsonian Institution.

SEGNIAGBETO, G.H., TRAPE, J.-F., DAVID, P., OHLER, A., DUBOIS, A. & GLITHO, I.A. 2011. The snake fauna of Togo: systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema 33(3): 325-360.

TRAPE, J.-F. & MANÉ, Y. 2006a. Le genre Dasypeltis Wagler (Serpentes : ) en Afrique de l’Ouest : description de trois espèces et d’une sous-espèce nouvelles. Bull. Soc. Herp. Fr. 119: 27-56.

TRAPE, J.-F. & MANÉ, Y. 2006b. Guide des serpents d’Afrique occidentale: Savane et désert. Paris: IRD Editions: Institut de Recherche pour le Développement.

TRAPE, S., MEDIANNIKOV, O. & TRAPE, J.-F. 2012. When colour patterns reflect phylogeography: New species of Dasypeltis (Serpentes: Colubridae: Boigini) from West Africa. C. R. Biologies 335: 488-501.

2013 M.F. Bates – New records of Dasypeltis confusa in West Africa 27

EDITORIAL COMMITTEE

Editor: M.F. Bates, Ph.D. (Stellenbosch); Co-editor (Natural Sciences): R.J. Nuttall, M.Sc. (Natal). Co-editors (Human Sciences): J. Haasbroek, D.Phil. (UOFS); S. Moodley, M.A. (Wits).

Consulting Editors: Prof. C. Chimimba (Department of Zoology and Entomology, University of Pretoria, South Africa); Dr J. Deacon (South African Heritage Resources Agency, Cape Town, South Africa – retired); Dr A. Dippenaar-Schoeman (ARC – Plant Protection Research Institute, Pretoria, South Africa); Dr A. Kemp (Northern Flagship Institution, National Museum of Natural History, Pretoria, South Africa – retired); Dr D.T. Rowe-Rowe (Ezemvelo KZN Wildlife, Pietermaritzburg, South Africa – retired); Prof. B.S. Rubidge (Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Johannesburg, South Africa); Prof. A.E. van Wyk (Department of Botany, University of Pretoria, South Africa); Prof. A. Wessels (Department of History, University of the Free State, Bloemfontein, South Africa).

Orders to: National Museum, P.O. Box 266, Bloemfontein 9300, South Africa E-mail: [email protected]