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A revision of the egg-eating snakes of the genus Dasypeltis Wagler (Squamata: Colubridae: Colubrinae) in north-eastern Africa and south-western Arabia, with descriptions of three new species 

Michael F. Bates1 & Donald G. Broadley2† 

1Department of Herpetology, National Museum, P.O. Box 266, Bloemfontein 9300, South Africa

2 Department of Herpetology, Natural History Museum of Zimbabwe, P.O. Box 240, Bulawayo, Zimbabwe

1 Corresponding author: E-mail: herp@nasmus.co.za

Bates & Broadley – Revision of Dasypeltis in NE Africa and SW Arabia

CONTENTS

Abstract
Introduction
Materials and Methods
Character analysis
Systematics
Dasypeltis medici (Bianconi)
Dasypeltis confusa Trape & Mané
Dasypeltis scabra (Linnaeus)
Dasypeltis bazi Saleh & Sarhan
Dasypeltis taylori Bates & Broadley sp. nov.
Dasypeltis atra Sternfeld
Dasypeltis fasciata A. Smith
Dasypeltis abyssina (Duméril, Bibron & Duméril)
Dasypeltis crucifera Bates sp. nov.
Dasypeltis arabica Broadley & Bates sp. nov.
Multivariate analyses
Summary of species
Discussion
Key to the genus Dasypeltis in north-eastern Africa and south-western Arabia
Acknowledgements
References
Appendix

Abstract

The genus Dasypeltis Wagler, 1830 in north-eastern Africa (from latitude 8ºS to the Mediterranean Sea, east of 29°E) and south-western Arabia (west of longitude 45°E) is reviewed. Ten species are recognised in this region on the basis of morphological characters, including colour pattern. Dasypeltis fasciata A. Smith, 1849 inhabits lowland forest in western and southern Uganda, while Dasypeltis medici (Bianconi, 1859) occurs in coastal forest/savannah mosaic in southern Somalia, Kenya and northern Tanzania. We show that Dasypeltis medici lamuensis Gans, 1957 is a junior synonym of D. medici. Dasypeltis atra Sternfeld, 1912—in a variety of colour phases/patterns—is widespread in the region, occurring in montane forest, moorland, forest/savannah mosaic, and savannah along the border between Tanzania and Kenya. Dasypeltis scabra (Linnaeus, 1758) is widely distributed in East African savannahs, except in parts of South Sudan, Uganda, western Kenya and Rwanda where Dasypeltis confusa Trape & Mané, 2006 infiltrates from the west, and in northern Somalia and Djibouti between 600 and 1370 m where it is replaced by Dasypeltis taylori Bates & Broadley sp. nov. Dasypeltis bazi Saleh & Sarhan, 2016 occurs as an isolated population in the Fayoum Depression west of Cairo, Egypt, with a possible isolate at Erkowit in the Red Sea Hills of north-eastern Sudan. We designate a lectotype for Dasypeltis abyssina (Duméril, Bibron & Duméril, 1854)—previously known only from the type description—and demonstrate that it is a distinct and valid species restricted to open savannah in the highlands (1800–2450 m) of north-western Ethiopia and central Eritrea; the paralectotype from Sudan is referable to D. scabra. Dasypeltis crucifera Bates sp. nov. is known only from two localities at lower elevations (600–1417 m) in northern Eritrea. Dasypeltis arabica Broadley & Bates sp. nov. inhabits the highlands of south-western Arabia (Saudi Arabia and Yemen) at elevations of 1600–2200 m. Multivariate analyses of scale characters and numbers of pattern cycles provides support for the above taxonomy. We provide a map detailing the ranges of the various species. The Horn of Africa, with at least six species (and another in Arabia), appears to be a centre of diversity for the genus Dasypeltis.

Key words: Dasypeltis, taxonomy, morphology, zoogeography, north-eastern Africa, south-western Arabia

† Donald G. Broadley passed away on 10 March 2016.

INTRODUCTION

Carl Gans (1959) noted that the egg-eating snakes of the genus Dasypeltis Wagler 1830 had attracted considerable attention because of their exclusive diet of bird eggs and the related adaptations of their skull and anterior vertebrae, but that identification of specimens was often difficult. This, he suggested, was because previous analyses had been limited to relatively small areas—and therefore geographical variation in colour pattern and scalation features was poorly understood. With this in mind he conducted a detailed taxonomic analysis on a large sample (over 1000 specimens) representing the entire range of the genus, i.e. sub-Saharan Africa, Egypt and south-western Arabia. His aim was firstly to distinguish valid species, and secondly to describe geographical and individual variation within each of these. Because of a scarcity of material from several large areas, his emphasis was on describing variation rather than attempting to validate existing subspecies or describe new taxa, and his taxonomic conclusions were therefore deliberately conservative. Gans (1959) found that colour pattern was a very useful character and used this, in combination with various other morphological features, to separate species and distinguish populations.

Gans (1959) identified nine basic dorsal colour patterns, some with sub-types. Most of these patterns, including a few of the sub-types, are now considered diagnostic of various species. For the widespread Dasypeltis scabra (Linnaeus, 1758) he documented a few distict variations in dorsal colour pattern. Some of these patterns were largely or wholly restricted to particular regions. For example, Gans’ (1959) ‘brown’ grouping included, for central and eastern Africa, three types: brown (‘2B’), black (‘2M’) and marked (‘2Bx’; typically with dark bands—often incomplete—across the back), these being referable to Dasypeltis atra Sternfeld, 1912 (see also Gans 1964; Broadley & Bates 2009). Specimens with the linked ‘5L’ colour pattern, consisting of dark, more-or-less diamond-shaped saddles confluent with dark lateral bars—such that they formed bands over the body—occur in western and central Africa, and these populations are now referable to Dasypeltis confusa Trapé & Mané, 2006 (see also Bates 2013). Another variation was ‘5I’ where black rectangular saddles and lateral bars form complete closure around the pale inter-saddle areas. However, most other populations of D. scabra throughout its extensive range had the variable non-linked ‘5N’ dorsal pattern consisting of dark markings in the form of large ‘saddles’ (varying in size and shape) along the middle of the back, alternating with lateral bars.

In his taxonomic revision of Dasypeltis, Gans (1959) recorded only three species from north-eastern Africa: the widespread D. scabra, two subspecies of Dasypeltis medici (Bianconi, 1859)—Dasypeltis medici medici (pattern ‘8’: usually dark rectangular saddles separated by narrow pale interspaces, with adjacent narrow dark lateral bars) in southern Kenya and Tanzania, and Dasypeltis medici lamuensis Gans, 1957 (pattern ‘3’: usually unmarked or faintly marked) along the coast from Somalia to the Kenya-Tanzania border, and Dasypeltis fasciata A. Smith, 1849 (pattern ‘4’: checker board pattern on back) which occurred marginally in western Uganda. However, in 1964 Gans recognised Dasypeltis atra Sternfeld, 1912 as a full species in the montane forests of the region. Broadley & Howell (1991), in a key to the Tanzanian species of Dasypeltis, separated ‘2Bx’ patterned specimens of D. atra (see Gans 1959, 1964) from D. scabra on the basis of their higher numbers of pattern cycles (85–107 versus 42–79). Largen & Rasmussen (1993) recognised only a single species of egg-eater, namely D. scabra, in Ethiopia and Eritrea. Hughes (1997) mapped the distribution of Dasypeltis in Uganda, based on material in the Natural History Museum (London) and the World Museum (Liverpool), but doubted whether the ‘2Bx’ colour phase of Gans (1959) should be assigned to D. atra.

In their revision of West African Dasypeltis, Trape & Mané (2006), in an aside, revived Dasypeltis abyssina (Duméril, Bibron & Duméril, 1854) of Ethiopia from the synonymy of D. scabra, after examining the syntypes in Paris. However, they provided only scant details as to why they considered this taxon valid. Broadley & Bates (2009) documented range extensions of the patterned (‘2Bx’) phase of D. atra into savannah areas of northern Tanzania, and the dorsally uniform black and brown phases into south-western Tanzania. Largen & Spawls (2010) recorded D. scabra from Ethiopia and Eritrea, and D. atra from Ethiopia, but also illustrated a D. abyssina (labelled as D. scabra, see below) from Mulu (actually at Keriyo hamlet, S. Spawls in litt. 5 April 2011) where it is sympatric with D. atra. An isolated population in the Fayoum Depression in Egypt was recently described as Dasypeltis bazi Saleh & Sarhan, 2016.

There are currently 13 recognised species of Dasypeltis, namely D. scabra, D. palmarum (Leach), D. inornata Smith, D. fasciata, D. abyssina, D. medici (D. m. medici and D. m. lamuensis), D. atra, D. confusa, D. gansi Trape & Mané, 2006, D. latericia Trape & Mané, 2006, D. sahelensis Trape & Mané, 2006, D. parascabra Trape, Mediannikov & Trape, 2012, and D. bazi (see Gans 1959, 1964; Trape & Mané 2006; Trape et al. 2012; Saleh & Sarhan 2016).

Since Gans’s (1959) revision, significant collections of specimens have become available for most parts of the study area, and this allowed for a more detailed evaluation of variation and patterns of geographical distribution. Several taxonomic problems for populations in north-eastern Africa and south-western Arabia require resolution. For example, there has been much uncertainty about the status and distribution of D. atra due to the wide variety of colour morphs and confusion with patterned D. scabra (see Hughes 1997; Broadley & Bates 2009). Examination of most of the available Dasypeltis material from this region also yielded new records of D. abyssina—a species which until now has not been properly diagnosed from its congeners—and recognition of easily identifiable and apparently closely related populations from the Keren-Agordat area, and across the Red Sea in the highlands of the south-western Arabian Peninsula, both described here as new species. In addition, an isolated population in Somaliland and Djibouti is described as a new species.

MATERIALS AND METHODS

Study area.—The study area is defined as Egypt, other parts of Africa north of latitude 8° S and east of longitude 29° E, and the Arabian Peninsula west of longitude 46° E. It therefore includes all eastern populations of D. atra, and the eastern-most populations of D. confusa and D. fasciata. To the north the area extends to the Mediterranean Sea and encompasses the isolated Egyptian population of D. bazi (see Gans 1959; Saleh & Sarhan 2016). The countries and regions covered by this revision are therefore: south-western Saudi Arabia, western Yemen, Egypt, the eastern parts of Sudan and South Sudan, Eritrea, Ethiopia, Djibouti, Somalia, Uganda, Kenya, Rwanda (which includes a small area west of 29° E), Burundi and northern Tanzania.

Material examined and characters used.—The starting point for this study was the relevant scalation, size and colour pattern data published in Appendix 1 of Gans (1959), together with the appropriate plates. Data in Gans (1964: 293) was also used. Most of the relevant material in the Natural History Museum in London was examined by us, together with several additional important specimens in other European, North American and African museums. The institutional abbreviations used below are from Sabaj (2016), except we use ‘BM’ for Natural History Museum in London, ‘MUZM’ for Makerere University Zoology Museum in Kampala, ‘AJL’ for the private collection of Angelo Lambiris in Durban (to be incorporated into the Durban Natural Science Museum), and ‘CG’ refers to the Carl Gans collection (see Gans 1964).

AJL                 Angelo Lambiris private collection, Durban

AMNH            American Museum of Natural History, New York

AAU               Addis Ababa University, Department of Biology

BM                  Natural History Museum, London

CAS                California Academy of Sciences, San Francisco

CG                  Carl Gans collection (Gans 1964)

CM                  Carnegie Museum of Natural History, Pittsburgh

FMNH            Field Museum of Natural History, Zoology Department, Chicago

IRSNB                        Intitut Royal des Sciences Naturelles de Belgique, Brussels

LACM            Natural History Museum of Los Angeles County, Los Angeles

LIV                 World Museum, Liverpool

MCZ               Museum of Comparative Zoology, Harvard University, Cambridge

MHNG            Muséum d’Histoire naturelle, Genève [Geneva]

MNHN            Muséum national d’Histoire naturelle, Paris

MRAC                        Musée Royal de l’Afrique Centrale, Tervuren

MSNG            Museo Civico di Storia Naturale “Giacomo Doria”, Genova [Genoa]

MSNM            Museo Civico di Storia Naturale di Milano [Milan]

MSNPV          Museo di Storia Naturale dell’Università di Pavia [Pavia]

MUZM            Makerere University Zoology Museum, Kampala

MZUF             Museo di Storia Naturale di Firenze, Sezione di Zoologia “La Specola”, Firenzi [Florence]

MZUT             Museo di Zoologia, Università di Torino [Turin]

NMBO            National Museum, Bloemfontein

NMK                           National Museums of Kenya, Nairobi

NMW              Naturhistorisches Museum, Wien [Vienna]

NMZB            Natural History Museum of Zimbabwe, Bulawayo

PEM                Port Elizabeth Museum, Bayworld

RMNH            Naturalis Biodiversity Centre, Leiden

TMP                Ditsong National Museum of Natural History, Pretoria

SMNS             Staatliches Museum für Naturkunde, Stuttgart

TCWC             Biodiversity Research and Teaching Collections, Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station

UMMZ            University of Michigan Museum of Zoology, Ann Arbor

USNM            National Museum of Natural History, Smithsonian Institution, Department of Vertebrate Zoology, Washington D.C.

ZFMK             Zoologisches Forschungsmuseum Alexander Koenig, Bonn

ZMB               Museum für Naturkunde, Berlin

ZMUC            Kobenhavns Universitet, Zoologisk Museum [Zoological Museum, University of Copenhagen], Copenhagen

Scalation and size data was also obtained from the curators of various museums (see Acknowledgements) and B. Hughes (for Ugandan material in the World Museum, Liverpool [LIV]). In several cases we recorded or confirmed the dorsal, and in some cases also ventral, colour patterns of specimens by examining colour images of specimens from various museums (e.g. World Museum, Liverpool; Museo Civico di Storia Naturale “Giacomo Doria”, Genova [Genoa]; Museo Regionale di Scienzi Naturali, Turin; National Museums of Kenya, Nairobi). Data is available for a total of 640 specimens (Appendix 1).

Material listed under ‘Localities’ comprises specimens examined by the authors, or for which data and/or photographs were provided and allowed positive determinations. In this section countries are listed from north to south, while specific localities are listed alphabetically. The mapping notation (e.g. SE 0135B3) refers to quarter-degree units (e.g. Minter et al. 2004: 48; but for the last part of the code we use numbers 1 to 4 instead of letters A to D), although on the map (Fig. 3) we plot these as half-degree units.

Most of the characters used by Gans (1959) were employed and evaluated in the same way in the present study, with modifications (or explanations) as indicated below. Specimens were examined under a binocular dissecting microscope or with the aid of magnifying lenses for some of the larger specimens. Ventrals were counted from the first scale following the posterior pair of chin shields, up to but excluding the anal plate (first ventral is always wider than long and usually similar to the next ventral; small extranumerary scales sandwiched between posterior chin shields were not included); subcaudals were counted from the first adjoining pair behind the vent up to the pair in contact with the terminal spine (spine excluded); longitudinal dorsal scale rows were counted at or near midbody (halfway between tip of snout and vent), usually by starting the count caudad and reversing in the same way at the dorsal ridge (vertebral area), but taking special care to follow the inclination of the lateral rows (usually 3–4) of serrated and reduced scales; pattern cycles refers to the number of blotches or bands (‘saddles’) on the back, starting after the first (often chevron) marking on the nape (which often extends onto the back of the head) up to the marking above the vent (in cases of bilateral asymmetry, fusion of dorsal markings, or when these markings were unclear, lateral bars were used as guiding criteria); texture of the frontal scale was scored as indicated under ‘Character analysis’; depth of sulcus formed by the pair of prefrontal scales was scored as deep, moderate or weak (scales merely in contact) as illustrated by Gans (1959, Fig. 6). Anterior temporals are the (usually) elongated scales posterior to, and in contact with, the postoculars; the uppermost one is in contact with a parietal scale, while the lowermost one is in contact with a supralabial. Posterior temporals are in contact behind the anterior temporals; the uppermost in contact with a parietal and lowermost in contact with a supralabial. Occasional supranumerary scales between anterior and posterior temporals were excluded from counts. The temporal formula refers to the number of anterior temporals plus the number of posterior temporals (e.g. 2+3). Snout-vent length (SVL) (tip of snout to vent) and tail length (vent to tip of tail) were measured with the snake aligned against a millimetre rule, but SVL of curled specimens was measured using a length of string. Measurements of truncated tails were not included in calculations.

In some cases material reported on by Gans (1959) was also examined by the authors and our scale and size data were used (Appendix 1). Because of known differences in numbers of ventral and subcaudal scales between males and females (e.g. Gans 1959), specimens were sexed. Sex was determined by presence/absence of (one or both) inverted hemipenes (slit open to reveal spines if not obvious) in the tail base, presence of everted hemipenes, and/or presence of internal reproductive structures (eggs, ovaries, oviducts, testes, seminiferous tubules). Only sexed specimens were used for the analysis of numbers of ventral and subcaudal scales (except MSNG 6738 from Keren in Eritrea, one of only four specimens representing a new species, taken as female based on high ventral and low subcaudal counts; see below).

For the descriptive account of D. bazi five specimens from Egypt were examined, comprising three of the four specimens from this region reported on by Gans (1959, but excluding ZMB 5865), as well as a specimen (ZMB 51552) discovered by Barry Hughes among undetermined material at the museum in Berlin (Frank Tillack in litt. 9 April 2014), and a specimen (NMW 26879/1) in the Naturhistorisches Museum in Vienna. A set of colour photographs of the Berlin specimens, showing dorsal, ventral and lateral aspects of the body and tail, as well as close-up views of the head (including left and right sides), of these specimens, was also later consulted. For ZMB 5865, SVL and tail measurements, and number of pattern cycles on the back, follow Gans (1959), while numbers of midbody scale rows and pairs of subcaudals were determined by F. Tillack (these corresponded with Gans’ 1959 counts); other morphological data for this specimen was determined after examining the colour images. Data for specimen ZFMK 38415 (Erkowit, Sudan), which we refer to as Dasypeltis cf. bazi, was provided by W. Böhme, but colour photographs of this specimen were also examined. The ratio greatest frontal length/greatest width was determined after performing measurements on enlarged photographic images.

According to Nicola Maio (in litt. 8 September 2014) the two Egyptian specimens of Rachiodon scaber recorded by Panceri (1878), in the Comparative Anatomy section of Naples Museum, were transferred to the Museo Zoologico at the same university after the Second World War, but now appear to be lost. There is also no record of these specimens having been transferred to Museo Civico di Storia Naturale di Milano (S. Scali in litt. 10 September 2014). However, the old catalogue of the museum in Milan does record a few specimens from Egypt identified by Prof. Gasco in 1876, including one ‘Rachiodon scaber’ (catalogue number 1622 – photographic image of catalogue examined), but unfortunately none of these specimens still exist at the museum, possibly having been destroyed by fire in 1943 during the Second World War (S. Scali in litt. 26 September 2014). The Cairo specimen reported on by Jan (1863) is also no longer available in the Milan collection (S. Scali in litt. 26 September 2014).

The description of the new species from Somaliland and Djibouti (see below) was based mainly on data in Gans (1959) and details available from a set of high-resolution colour photographs of all proposed type specimens showing dorsal, ventral and lateral aspects of the body and tail, as well as close-up views of the head (including left and right sides), provided by the Natural History Museum (London) and Museum National d’Histoire Naturelle (Paris), although two proposed paratypes (BM 1992.501 and MNHN 84–707) were also examined by the first author while visiting these museums in September 2011 and October 2015, respectively. While the proposed holotype of the new species from Eritrea (see below) was examined by the first author at the natural history museum in Berlin, a set of close-up photographs was also supplied subsequently; a similar set of photographs of the proposed paratypes in Italian museums—which could not be examined personally—were also supplied, together with sex, SVL and tail length measurements, and ventral, subcaudal and midbody scale counts. For the new ‘5I’-patterned species from Arabia (see below), similar photos were provided of the proposed holotype (and one proposed paratype, BM 1987.1018) to allow for a detailed description, although these and other proposed types were also examined by both authors at the museum in London. Greatest frontal length/greatest width for most of the above specimens was determined by performing measurements on enlarged photographic images.

In addition to the data presented in this paper, comparative data consulted for the diagnoses of species (see below) were obtained from Duméril et al. (1854), Mertens (1954, 1955), Gans (1959, 1964), FitzSimons (1962), Broadley (1990), Trape & Mané (2006), Trape et al. (2012), Bates & Ineich (2012), Bates (2013) and Saleh & Sarhan (2016). In the case of D. atra, to avoid inadvertently including data for mainly brown or uniform phase specimens of other species of Dasypeltis, the only extralimital data used was for specimens from eastern Democratic Republic of the Congo (Mount Kabobo and the ‘districts’ of Oriental and Kivu) with ‘2M’, ‘2B’ and ‘2Bx’ dorsal colour patterns as identified by Gans (1959) in his appendix. An exceptionally low female subcaudal count of 50 for D. gansi (see Bates & Ineich 2012) was excluded from the ranges listed under ‘Diagnosis’ below, as were ventral and subcaudal counts (250 and 55 respectively) for a specimen of D. bazi identified as a male by Saleh & Sarhan (2016) but which is probably a female (see discussion below; these data were also excluded from the diagnostic key). Morphological data for D. parascabra were derived from the small type series together with a few additional specimens—for which data is available in Gans (1959)—referred to this species by Trape et al. (2012); ranges are as follows—males: ventrals 210–223, subcaudals 67–74; females: ventrals 224–237, subcaudals 57–64; both sexes: midbody scale rows 22–26, pattern cycles 56–70. Photographs of the holotype (SMF 46642) of D. s. loveridgei were available at: http://www.biologie.uni-ulm.de/cgi-bin/query_all/details.pl?id=107169&stufe=7&typ=ZOO&sid=T&lang=e (accessed 26 September 2014).

The following type specimens were examined for comparative purposes by the first author in September/October 2011: holotype (ZMB 1856) and two paratypes (ZMB 1857–8) of Dasypeltis scaber var. mossambicus Peters, 1864 (= D. scabra); holotypes of Coluber Palmarum (= D. palmarum, BM 1946.1.2.35), Dasypeltis inornatus (= D. inornata, BM 1946.1.2.47), Dasypeltis scaber var. breviceps Peters, 1864 (= D. scabra, ZMB 3886), Dasypeltis scaber var. fasciolata Peters, 1864 (= D. medici, ZMB 5737), D. lineolata Peters, 1878 (= D. atra, ZMB 9224, see below), D. confusa (MNHN 2006.0315) and D. sahelensis (MNHN 2006.0315); paratypes of D. medici lamuensis (= D. medici, see below, BM 1951.1.3.72–3, BM 98.1.8.15, BM 87.11.3.31), D. gansi (MNHN 2006.0308) and D. latericia (MNHN 2006.0314); and syntypes of Rachiodon abyssinus (= D. abyssina, see below, MNHN 6567, 7227) and Dasypeltis macrops Boulenger (= D. fasciata, BM 1946.1.5.17–8). Photographs of the syntypes (ZMB 22516A, ZMB 22516B , ZMB 27761) of D. scabra var. atra (= D. atra), provided by F. Tillack, were also examined.

Statistical and multivariate analyses.—Univariate analysis was conducted using the computer software package Statistica v. 6. Two kinds of ordination comparisons were conducted in order to determine whether samples were distinguishable using a multi-dimensional approach. For Principal Components Analysis (PCA), IBM SPSS Statistics v. 20 was used, while Statistica Six Sigma was used for Discriminant Function Analysis; PAleontological STatistics v. 3.08 was used for PERMANOVA and to produce graphs. As all population groups (= species) discussed below are considered real geographical entities, it was considered appropriate to conduct discriminant function analysis to determine if the groups are distinguishable. Because of known sexual dimorphism in numbers of ventral plates and pairs of subcaudal scales (Gans 1959; see below), males and females were analysed separately. Eleven meristic variables (characters as defined above and by Gans 1959, and discussed below) were used in the analyses: numbers of ventrals, subcaudals, midbody scale rows, supralabials (counts for left and right sides of the head combined), preoculars (combined count), postoculars (combined count), anterior temporals (combined count), posterior temporals (combined count) and pattern cycles; and the interval variables: frontal pitting (coded as 0 if smooth, 1 if present only as one or two rows at the edges, and 2 if extensive), and depth of the inter-prefrontal sulcus (0 if weakly marked, 1 if fair/medium, and 2 if deep). Only specimens with full sets of data were included (marked by an asterisk in Appendix 1). Analyses were conducted for the entire study area, but also separately for the Horn of Africa (Ethiopia, Eritrea, Djibouti, Somalia) + Egypt (excluding D. medici, represented by only a single Somalian specimen with a full set of data) to determine whether the new species were distinguishable from all other congeners in that region.

Species concept and species delimitation.—We apply a lineage-based species concept whereby a species is represented by an independently evolving metapopulation lineage (see Frost & Hillis 1990; De Queiroz 1998, 2007). Morphological characters were the operational criteria for species delimitation.

Broadley (1983, also 1990) relegated Dasypeltis scabra loveridgei Mertens, 1954 from Namibia to the synonymy of D. scabra, but the former has a distinctive dorsal colour pattern (hourglass-shaped dorsal saddles alternating with more-or-less square to rectangular lateral bars; ‘6N’ of Gans 1959) and almost always six supralabials (2nd and 3rd in contact with the orbit) on either side of the head (Mertens 1954; Bates et al. 2009; Bates et al. unpubl. data). Apart from Mertens’ (1954) types, Gans (1959) provided data for additional Namibian ‘6N’ specimens, all but a few having the 6[2,3] supralabial formula. This combination of characters is unique to D. s. loveridgei which we therefore consider a good evolutionary species and formerly elevate to species status as Dasypeltis loveridgei Mertens, 1954 comb. nov. It should be noted that one of Mertens’ (1954) paratypes has the ‘5N’ dorsal pattern and 7[3,4] supralabial formula and is referable to D. scabra (Gans 1959: 95, 219; Bates et al. in prep.).

Character analysis

  1. Colour phases.—Gans (1959, see his Figs 9 & 10) recognised 13 colour phases (dorsal colour patterns) for the genus Dasypeltis, with only ‘inornata’ (‘1’), ‘loveridgei’ (‘6’) and ‘palmarum’ (‘7’) not found in north-eastern Africa. The ‘montane’ form [= atra] included melanistic (‘2M’), uniform brown (‘2B’), and patterned (‘2Bx’, dorsal saddles often confluent, or in partial contact, with lateral markings) phases (Fig. 17A–M). Gans (1959) recorded the ‘lineolata’ (‘9’, dorsum dark with narrow, interrupted, pale transverse bands) phase [= D. atra] only from central Kenya (Fig. 17D), but some specimens from other areas have similar markings (see Fig. 17E, Ethiopia). ‘Dasypeltis scabra’ included true D. scabra (‘5N’, variable, but dorsal saddles not linked to lateral bars, see Gans 1959) (Figs 5A–H & 6); D. bazi (5N-like but with large, cranio-caudally elongate, brown saddles, usually bifurcated at each end with a fairly shallow but distinct notch, and spot or blotch-like lateral markings; Gans 1959, Pl. 6, Fig. 1) (Figs 7A, 9A, 11 & 12); D. taylori sp. nov. (‘5N’-like but with large, closely-spaced, square to slightly rectangular saddles often separated by narrow, sharply-defined white crossbars; Gans 1959, Pl. 6, Fig. 6) (Figs 13 & 14); D. confusa (‘5L’, saddles and lateral bars linked; Gans 1959, Pl. 8, Fig. 6) (Fig. 4A–D); and D. arabica sp. nov. (‘5I’, rectangular saddles and lateral bars enclose pale inter-saddle areas; Gans 1959, Pl. 6, Fig. 4) (Figs 27A & 29A). Dasypeltis fasciata (‘4’) was distinguished by its ‘chequerboard’ pattern being expressed only on the interstitial skin and lower edges of the scales (Gans 1959, Pl. 3, Figs 1–4) (Figs 19A & C). Dasypeltis medici, distinguished by the darkly pigmented edges of the paired apical pits on its dorsal scales, occurred as both patterned (‘8’, rectangular saddles with distinct white interspaces usually in contact with elongate lateral bars) and uniform or largely uniform but feebly marked (‘3’ – subspecies lamuensis) phases (Gans 1959, Pl. 13) (Fig. 2A–E). Dasypeltis crucifera sp. nov. has a unique dorsal pattern consisting of dark brown oval saddles, and elongated lateral bars, which together almost completely enclose pale cross-markings that form the inter-saddle areas (Figs 23A & 25A). Gans (1959) did not recognise D. abyssina, which is now known to occur in both (dorsally) patterned (‘5N’-like but with mostly oblique dark saddles, usually wider than long, and often confluent or forming zigzags on the back and tail) and uniform grey-brown phases (see below and Fig. 20A, C–G).
  2. Number of pattern cycles (Table 1).This is a very variable character, but patterned atra, D. abyssina, D. arabica sp. nov., D. medici and D. fasciata usually have counts of 70 or above, and are usually distinguishable from D. bazi, D. taylori sp. nov., D. crucifera sp. nov., D. confusa and often also D. scabra in this regard. In the study area, high numbers of cycles (108–141) in D. fasciata distinguish it from all other species except D. atra and a couple of specimens of D. medici; while the lowest counts are for D. bazi (39-49, 38 in a specimen reported by Saleh & Sarhan 2016) and D. taylori sp. nov. (44-59).

INSERT TABLE 1

  1. Number of ventrals (Table 2). arabica sp. nov. and D. abyssina have the highest counts, followed by D. fasciata and D. medici. Due to wide variation within species, and extensive overlap of ranges between species, this character on its own does not help to distinguish D. atra, D. bazi, D. taylori sp. nov., D. crucifera sp. nov., D. confusa and D. scabra. In D. medici, males and females often have similar ventral counts.
  2. Number of subcaudals (Table 2).The two most arboreal species, fasciata and D. medici, have the highest subcaudal counts, averaging over 68 in both sexes. There is considerable geographical variation in D. scabra, with the highest counts in the south (Tanzania). Dasypeltis confusa, D. scabra, D. bazi (based only on specimens examined, i.e. excluding data from Saleh & Sarhan [2016] as some of their specimens may have been incorrectly sexed), D. taylori sp. nov., D. atra, D. abyssina, D. crucifera sp. nov. and D. arabica sp. nov. have similar mean counts (59–63 in males, 48–56 in females).

INSERT TABLE 2 (FULL PAGE)

  1. Type of head pitting with reference to the frontal shield.—Gans (1959) recognised three states: N = none, M = marginal pitting (his Fig. 2B), and A = entire surface pitted (his Fig. 2A). Pits may take the form of swollen pimple-like protrusions on the frontal, especially centrally. In the present study this scheme has been refined to show five states: 0 = smooth, no pitting; 1 = a single row of pits on the posterior and lateral margins of the frontal; 2 = more extensive marginal pitting (Fig. 2B of Gans 1959); 3 = extensive pitting, but with centre of frontal still smooth (Fig. 2A of Gans 1959); 4 = entire frontal pitted and centre of shield often rugose or corrugated. In the study area Dasypeltis medici and taylori sp. nov. almost always have extensively pitted frontals (States 3 or 4), while most other species usually have marginal pitting (States 1–2), but in D. scabra and D. bazi the pitting varies from absent (State 0) to extensive (State 4). The frontal shields of five specimens of D. bazi illustrated in Saleh & Sarhan (2016, photographs in Fig. 5) appear to be extensively pitted and sometimes also corrugated (States 3 or 4).
  2. Depth of inter-prefrontal sulcus.—This character was classified in the same way as by Gans (1959). The sulcus was moderately to deeply sunken in taylori sp. nov., D. arabica sp. nov. and some populations of D. scabra (but usually moderately sunken to absent elsewhere), whereas it was usually weakly marked (scales merely in contact) or moderately developed in all other species.
  3. Number of supralabial shields.—In north-eastern Africa and Arabia there are normally seven supralabials, the third and fourth entering the orbit. The 6[2,3] formula (meaning six supralabials on either side of the head, with the 2nd and 3rd in contact with the orbital) is very rare except in loveridgei comb. nov. and a few other populations of the D. scabra species complex in southern Africa (Mertens 1954; Bates & Broadley in prep.). Occasional variations (on one or both sides of the head) in the study area were: 5[2,3], 5[3], 6[2,3], 6[3], 6[3,4], 7[2,3,4], 7[3,4,5], 8[3,4], 8[3,4,5] and 8[4,5].
  4. Number of postoculars.—In the study area most specimens have a single preocular and two postoculars on either side of the head, but a single postocular is commonly found in western populations of atra (e.g. 93.0% of Rwanda/Burundi snakes [N = 43] had a single postocular on either side; by contrast 75.0% of Ethiopian snakes [N = 28] had two postoculars on either side). The occurrence of a single postocular is also high in extralimital populations―in a sample of 126 specimens of D. atra from eastern Democratic Republic of the Congo, examined by the first author at the Royal Museum for Central Africa (Tervuren, Belgium), as many as 73.8% had one postocular on either side of the head, 14.3% had one postocular on one side and two on the other, while only 11.9% had two postoculars on both sides of the head.
  5. Number of anterior and posterior temporals.—A temporal formula of 2+3 was common for all species, with 2+2 and 2+4 occuring frequently. Two anterior temporals on either side of the head was by far the most common condition for most species in the study area; as was three posterior temporals on either side, although counts of two or four were common. Although Saleh & Sarhan (2016) also gave this formula as 2+3 for bazi, in the single Fayoum specimen examined for this study, one side of the head had three anterior temporals, while another Egyptian specimen had a formula of 3+3 on both sides of the head.
  6. Number of dorsal scale rows at midbody.—There was considerable overlap in numbers of midbody scale rows between the different species. However, northern species ( arabica sp. nov., D. abyssina, D. crucifera sp. nov., D. bazi and D. taylori sp. nov.) never had more than 25 rows (21–25 versus 22–28 rows in other species). The highest means (≥25) were for D. scabra and D. confusa, and the lowest (≤23.5) were for D. abyssina, D. bazi (based only on specimens examined, i.e. excluding data from Saleh & Sarhan [2016], see below), D. crucifera sp. nov. and D. fasciata.
  7. Relative tail length (Fig. 1).—Although there was much overlap in this character for most species (see species accounts), including those for which reasonable samples were available as shown in the figure below, Dasypeltis medici males and females usually had distinctly longer tails relative to SVL compared to males and females, respectively, of other species, while fasciata tended to have a slightly longer tail than the other species. Figure 1 also illustrates sexual dimorphism in tail length, with males usually having longer tails relative to SVL; as well as sexual dimorphism in body size, with the largest females exceeding the size of the largest males.

INSERT FIG. 1

Systematics

Apart from the species discussed and compared below, two available names currently in the synonomy of D. scabra (see Gans 1959; Broadley 1990) deserve mention in light of the fact that material from three populations currently identifiable as, or referable to, D. scabra are here to be described as new species. Dasypeltis scabra mossambica (Tete and Boror in Mozambique) and Dasypeltis scabra breviceps (‘Cafferland’, i.e. Eastern Cape Province, South Africa) were described by Peters (1864). The three syntypes (ZMB 1856, juvenile male; paratypes 1857 and 1858, both adult females) of the former and holotype (ZMB 3886, adult female) of the latter were examined for this study. The types of D. s. mossambica have ‘5N’-type dorsal patterns with moderate-sized saddles generally round to oval in shape, nasal divided below the nostril, supralabial formula 7[3,4] on either side of head, postoculars two on either side, frontal smooth in juvenile but fully pitted (State 4) in adults. For other characters: ZMB 1856, inter-prefrontal sulcus weakly marked, ventrals 207, subcaudals 67, midbody scale rows 25, pattern cycles 57; ZMB 1857: sulcus deeply marked, 236, 60, 25, 50; ZMB 1858: sulcus moderately marked, 237, 59, 27, 61. The holotype of D. scabra breviceps has a ‘5N’-type dorsal pattern but the saddles and lateral bars are somewhat narrow and irregular, nasal divided below the nostril, supralabial formula 7[3,4] on either side of head, postoculars 2 on either side, frontal fully pitted (State 4), inter-prefrontal sulcus deeply marked, ventrals 203, subcaudals 61, midbody scale rows 27, pattern cycles 63. Peters (1864) noted that the ground colour of D. scabra mossambica was yellowish. Both of the above names are clearly synonyms of D. scabra.

Squamata: Colubridae: Boigini: Dasypeltis Wagler, 1830

Dasypeltis medici (Bianconi)

Eastern Forest Egg-eater

Fig. 2A–F

Dipsas medici Bianconi, 1859: 277. Type locality: ‘Mozambique’ [= Inhambane].

Dasypeltis abyssina ? (not Duméril, Bibron & Duméril) Stejneger, 1893: 733 (Mt Kilimanjaro, Tanzania).

Dasypeltis scabra var. F Boulenger, 1894: 356 (part, Mt Kilimanjaro).

Dasypeltis scaber (not Linnaeus) Loveridge, 1929: 28 (part) & 1936: 256 (Mt Mbololo, Lamu Island); Uthmöller, 1937: 356 (part, Gomberi, Mt Kilimanjaro); Scortecci, 1939: 276 (part, Belet Amin, Somalia).

Dasypeltis medici medici Gans, 1957: 3 & 1959: 157; Malonza et al. 2011: 160 (Kitobo Forest, Kenya); Göthel, 2015a: 19 & 2015h: 89.

Dasypeltis medici lamuensis Gans, 1957: 5. Type locality: Lamu Island, Kenya, holotype MCZ 40581; Loveridge, 1957: 289 (check-list); Gans, 1959: 159 & 1961: 74; Spawls, 1978: 10; Lanza, 1983: 223 (Alessandra Island near Gelib, zone of Mareri SW of Gelib, Belet Amin); Broadley & Howell, 1991: 32; Spawls et al., 2002: 416; Spawls et al., 2006: 129; Largen & Spawls, 2010: 490; Göthel, 2015a: 19, 2015c: 30, 2015d: 50, 2015f: 65 & 2015h: 89.

Dasypeltis medici Gans, 1971: 461; Largen & Parker, 2004: 26; Branch, 2005: 59 & 2014: 68; Vets & Safari, 2014: 43 (University of Dodoma).

Description. Dorsal colour pattern varies considerably. Patterned specimens may be similar to D. scabra: grey to tan with narrow, darker, rectangular saddles (longer than wide) separated by narrow cream to white interspaces, with irregular narrow dark lateral bars situated for the most part adjacent to the pale interspaces. The saddles may be much elongated and closely set and give the appearance of a broad dark vertebral stripe broken by more-or-less equally spaced pale flecks. Pattern cycles 57–108 (mean 74.0, N = 24); 1–5 chevrons (pointing forward) on nape and anterior part of body (N = 13). Some specimens are uniform pink, orange, red, grey, fawn or brown above (Spawls et al. 2002), but specimens with intermediate patterns also occur, e.g. reddish (Fig. 2C–D) or grey (Fig. 2E–F) with fairly vague dark dorsal markings and mere indications of whitish interspaces (e.g. CAS 135751, FMNH 251325, ZMUC 6111); NML 1699 is reddish-brown with 3–4 vague chevrons only (anterior part of body), and a cream venter. Edges of apical pits on dorsal scales darkly pigmented. Venter usually cream, often with fine grey stippling (may be extensive as in NMW 26926); but occasionally yellowish-cream (e.g. NMK S 3956, Mount Warges; ZFMK 63582, near Mombasa [with extensive grey speckling]); and salmon-coloured with central part of each plate dark brown anteriorly in FMNH 142635 (vicinity of Nairobi).

Preoculars usually one on either side of head, but one on one side and two on the other in NMW 9964/18 and ZMB 8597, and two on either side in ZMUC 6143 (N = 61); postoculars usually two on either side of head, but one in 10.3% of specimens, and one on one side and two on the other in 4.6% of specimens (N = 87); frontal shield usually extensively pitted (States 3–4), but marginally pitted (States 1–2) in 10.5% of specimens (including three snakes ≥422 mm SVL) and smooth (State 0) in one adult (MSNG 42710b) (N = 57); inter-prefrontal suture usually weakly marked, but moderately marked in 18.9% of specimens, and deeply marked in AMNH 143717 (N = 53); supralabials 12–15, usually 14 (both sides of head), usually 7 [3rd and 4th entering orbit] on either side of head, but variations are: 6[2,3] on both sides in NMK 1699, 6 [2,3] on right side in MSNG 427210b and USNM 16755, 6[3,4] on both sides in NMK 2535 & 3249, 7 [2,3,4] on right side in ZMUC 6124, 8 [3,4] on left side in SMNS 4216A, and 8[3,4,5] on left and 7 [3,4,5] on right side in SMNS 4216F (N = 85); anterior temporals usually two on either side of head (70.5%), occasionally three (14.8%), 3 left:2 right or 2:3 (13.1%) and 4:3 in USNM 16755 (N = 61); posterior temporals usually three on either side of head (60.7%), occasionally 3:4 or 4:3 (14.8%), four (14.8%), 3:2 or 2:3 (4.9%), 5:4 in MCZ 40580, five in ZMB 17453 (N = 61), and two (left side, right damaged) in ZMB 48150; temporal formula on either side of head usually 2+3, occasionally 2+4 or 3+3; midbody scale rows 22–27 (mean 24.8 ±0.97, N = 57); ventrals 220–252 in males (N = 29), 218–252 in females (N = 49); subcaudals 75–109 in males (N = 27), 61–90 in females (N = 47). Ratio SVL: tail length 3.5–4.7 in males (N = 27), 4.0–6.1 in females (N = 44); total length: tail length 4.5–5.7 in males (N = 27), 5.0–7.1 in females (N = 44).

INSERT FIG. 2

Size. Largest male (NMK 2631 – Gede, Kenya) 679 mm snout-vent length [SVL] + 146 mm tail length = 825 mm total length; largest female (FMNH 142635 – ‘vicinity of Nairobi’, Kenya) 912 + 149 = 1061 mm.

Habitat. Coastal and montane forest and savannah at elevations from sea level to about 2500 m on Mount Warges (Kenya). In southern Africa it occurs in lowland evergreen forest (Broadley 1990).

Distribution. Widely distributed in the eastern parts of sub-Saharan Africa, from Mareri in southern Somalia southwards through eastern and central Kenya, eastern Tanzania, Malawi, extreme eastern Zimbabwe, Mozambique, and KwaZulu-Natal in South Africa as far south as St Lucia (see Broadley 1990; Spawls et al. 2002; Bates et al. 2014; Fig. 3). There is also an isolated Zambian record (AJL 3027) from riparian forest on the Mululangana River in the Zambezi Valley, 26 km SW of Chirundu [SE 1628B1] (Broadley et al. 2003).

Sympatry. Sympatric with D. scabra at Belet Amin (Somalia), Kilifi (Kenya), and Amani, Dodoma, Morogoro and Zanzibar (Tanzania); sympatric with D. atra at Nyambene Hills (Kenya) and Moshi (Tanzania); sympatric with both of these species at Mt Kilimanjaro in Tanzania (Appendix 1).

Localities. SOMALIA. Belet Amin [NE 0042B2] MSNG 42710b; near Mareri (Labadaah to Bula Yaag) [NE 0042D1] CAS 148077, MZUF 27155. KENYA. Chyulu Hills [SE 0037D2] NMK 402; Diani Beach, Ukanda [SE 0439B3] Fig. 2E–F; Gede [SE 0340A3] AMNH 88631, NMK 2597, 2619, 2631, 3200; Giriama [SE 0339D3] NMK 480; Kikambala [SE 0339D4] NMK 1050; Kilifi [SE 0339D2] BM 51.1.3.72–3; Lamu Island [SE 0240B4] MCZ 40581–2 [holotype and allotype of D. medici lamuensis]; Malindi [SE 0340A1] NMK 1561, 2522, 2526–7, 2529–33, 2535, 2538, 2587, 3290, 3292, ZMB 50191; Malindi, Sabaki River [SE 0340A1] photographic record (dorsum and top of head plain pale orange, edges of apical pits on dorsal scales darkly pigmented) and bottom photograph in Spawls et al. 2002: 417; near Mombasa [SE 0439B1] ZFMK 63582; Mombasa, 30 km south of [SE 0439B3] NMW 26926/1; Mount Mbololo [SE 0338A4] MCZ 40580; Maungu, Teita [SE 0338D2] BM 98.1.8.15; Mount Warges [NE 0037C4] NMK 3956; ‘vicinity of Nairobi’ [SE 0136B4] FMNH 142635; Nguni, Mwingi [SE 0137D2] NMK 2377; Nyambene Hills, Meru [NE 0037B2] NMK 1699; Taita [SE 0338A4] ZMB 9244; Takaungu [SE 0339D2] ZMB 17496; Tana River, Mchelelo Forest [SE 0140A3] NMK 4131; Taveta, Kitobo Forest [SE 0337B3] NMK 4121; Tiwi, Kwale [SE 0439B1] CAS 135751; Voi to Ndi [SE 0338D3] BM 98.1.8.14; Watamu [SE 0340A3] (Fig. 2C). TANZANIA. Amani [SE 0538B1] MCZ 23274, ZMB 37598, ZMUC 6124; Bombani, East Usambara Mtns [SE 0538B1] ZMUC 6123, 6133; Dar-es-Salaam [SE 0639C4] NMW 9964/18; Dar-es-Salaam, vicinity of [SE 0639C4] (Fig. 2A); Dodoma, University of Dodoma campus [SE 0635B1] photographic record in Vats & Safari (2014); East Usambara Mtns (Fig. 2B); FMNH 251325, ZMUC 6143; Gomberi, Mount Kilimanjaro [SE 0337A2] (Uthmöller 1934); Kibognoto [SE0337 A1] ZMB 37600; Kimboza Forest [SE 0737B2] ZMB 48150; Longido [SE 0236D1] ZMUC 6144; Mafia Island [SE 0739D4] MCZ 54827, NMZB 11265, SMF 19460; Makindu River [SE 0738C3] MCZ 18206; Mlinga Forest Reserve [SE 0538B1] ZMB 17318; Morogoro [SE 0637D3] AMNH 143717, USNM 62901, 62904, 62906, ZMB 27616; Moshi [SE 0337A4] SMNS 4216A–F; Mount Kilimanjaro [SE 0337A2] BM 87.11.3.31, USNM 16755; Mount Kwaraha [SE 0435B2] ZFMK 82342; Msolwa, Rubeho Mtns [SE 0736B4] ZMUC 6121; Nyange, Uluguru Mtns [SE 0637D4] MCZ 23273; Tanga [SE 0539A1] Fig. 2D and middle photograph of reddish-brown, weakly marked specimen in Spawls et al. 2002: 417; Usa River near Arusha [SE 0336B3] CM 36586; Usambara Mountains, ZMB 17453, and Spawls et al. 2002: 417; Zanzibar Island [SE 0639A2] BM 50.1.5.40, NMK 3249, ZMB 5737, 17219, ZMUC 6111; Zanzibar Coast, ZMB 8597.

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it appears to be widespread and abundant and should be regarded as Least Concern.

Remarks. The northern-most populations of D. medici were separated as a subspecies, D. medici lamuensis, by Gans (1957). This taxon was diagnosed by the lack of the typical medici colour pattern (saddles and lateral bars) on the body (although this may be present but poorly defined, and Gans [1964] later noted that a faint typical medici pattern may be present in juveniles) and lower numbers of ventrals in both sexes. However, medici (marked), uniform, and ‘intermediate’ patterns are found in many specimens from south-eastern Kenya, especially at Malindi and Gede (AMNH, NMK and ZMB material not examined by Gans; Appendix 1: Malindi: in alcohol usually uniform and mostly pale brown [e.g. NMK 1561, 2619], occasionally grey [e.g. NMK 2532], but two specimens [NMK 2526, grey; NMK 2530, pale brown] have very weakly marked darker saddles, while two young snakes [NMK 2540, 422 mm SVL; ZMB 50191, 461 mm SVL] have the typically marked [but not bold] medici pattern; Gede: uniform pale brown, but NMK 2631 is grey with very vague indications of darker vertebral blotches and whitish interspaces). Reddish, largely uniform specimens, with faint vertebral blotches and whitish interspaces—from Watamu (SE Kenya) and Tanga (N Tanzania)—and a greyish, largely uniform specimen, with similar markings—from Diani Beach (SE Kenya)—are illustrated in Fig. 2. Weakly marked specimens with indications of dorsal and lateral markings are also occasionally found further south, e.g. Mlola Forest on Mafia Island in Tanzania (NMZB 11265) and Bazaruto Island off the southern coast of Mozambique (NMZB 12037). The average ventral counts are indeed lower in northern populations (Somalia + Kenya), but there is considerable overlap with typical southern populations (Appendix 1), and ventral counts are actually lowest in the southern-most population in KwaZulu-Natal, South Africa: 206–215 in three males (TMP 28779, 46041, 56547), 214–223 in three females (TMP 44652, 47840, 78994) (D.G. Broadley unpubl. data). In light of this variation in colour pattern and ventral counts, we here relegate Dasypeltis medici lamuensis Gans, 1957 to the synonomy of Dasypeltis medici (Bianconi, 1859).

INSERT FIG. 3 (FULL PAGE)

Dasypeltis confusa Trape & Mané

Diamond-back Egg-eater

Fig. 4A–D

Dasypeltis scabra scabra (not Linnaeus) Loveridge, 1955: 46 (South Sudan).

Dasypeltis scabra (not Linnaeus) Gans, 1959: 141 (part, 5L specimens from South Sudan and Uganda); Pitman 1974: 116 (part, Uganda); Hughes, 1997: 68 (part, Uganda); Behangana & Goodman, 2002: 66 (part, Uganda).

Dasypeltis confusa Trape & Mané, 2006: 28. Type locality: Ibel, Senegal, holotype MNHN 2006.0303; Göthel 2015h: 79.

Description. Dorsum grey with darker markings, usually in the form of rhombic saddles (often diamond-shaped with sharpest points aligned antero-posteriorly) which are confluent with the lateral bars (i.e. the ‘5L’ linked pattern of Gans 1959, Pl. VIII, Fig. 6); dark markings usually have a cream to white margin around them, emphasising the pattern; pattern cycles 50–79 (mean 61.7, N = 30); dorsal saddles are not always joined to the lateral bars, so that the dorsal pattern may be similar to that of D. scabra (‘5N’) over small portions of the back in some specimens. A few specimens from South Sudan (six of the 14 from Torit – see Gans 1959: 203) and Uganda (MCZ 47801, 47830, UMMZ 88530) are plain brown dorsally. Venter cream-white, often with grey stippling at the edges.

Preoculars one on either side of head (N = 40); postoculars usually two on either side of head, but only one on left side in FMNH 62216 (N = 45); frontal shield for the most part smooth, usually with 1–2 rows of shallow pits at the edges (States 1–2), but occasionally completely smooth and without pits (State 0) (ZMB 56436, UMMZ 88529) (N = 38); inter-prefrontal suture usually weakly marked, but moderately marked in 21.7% of specimens, and well marked (deep) in BM 1954.1.12.9 (N = 23); supralabials 11–14, usually 14 (both sides of head), usually 7 [3rd and 4th entering orbit] on either side of head, but variations are: 6 [2,3] (BM 1954.1.12.8), 6 [3,4] on left side of head (FMNH 58336), and 6 [3,4] on left and 5 [3] on right (ZMB 77441) (N = 42); anterior temporals usually two on either side of head (95.3%), but 1 left: 2 right in BM 1954.1.12.8 and 2: 1 in BM 1954.1.12.9 (N = 43); posterior temporals usually three on either side of head (62.8%), two in NMW 9964/7, occasionally 3:4 or 4:3 (11.6%), four (11.6%), 2:3 or 3:2 (9.3%), and five in MCZ 47815 (N = 43); temporal formula on either side of head usually 2+3, occasionally 2+4 or 2+2; midbody scale rows 23–27 (mean 25.0 ±1.03, N = 40); ventrals 199–224 in males (N = 14), 212–239 in females (N = 30); subcaudals 51–67 in males (N = 12), 48–61 in females (N = 24). Ratio SVL: tail length 4.7–5.8 in males (N = 12), 4.9–7.5 in females (N = 20); total length: tail length 5.7–6.8 in males (N = 12), 5.9–8.5 in females (N = 20).

INSERT FIG. 4

Size. Largest male (NMK 489 – Kacheliba, Kenya) 424 + 74 = 498 mm; largest female (FMNH 58496 – Juba, South Sudan) 575 + 92 = 667 mm.

Habitat. Found in savannah at elevations of 500 m to 1200 m.

Distribution. From Senegal to Nigeria in West Africa, and from Cameroon south to Angola and north-western Zambia, then eastwards through the Central African Republic, northern Democratic Republic of the Congo (‘5L’ patterned specimens in Gans 1959; Trape & Mané 2006; Trape et al. 2012; Bates 2013; Broadley & Bates unpubl. data), southern South Sudan, western and northern Uganda, to the Rift Valley at Kacheliba and Mumias in western Kenya, with an isolated record at Lake Kivu in western Rwanda (Fig. 3). It may also extend into south-western Ethiopia, based on two specimens with imprecise/questionable locality data (see below), but this requires confirmation.

Sympatry. Sympatric with D. atra 10 km S of Juba (South Sudan), Busingiro and Gulu (Uganda) and Kacheliba (Kenya); sympatric with D. scabra at Lake Kivu (Rwanda). (Appendix 1).

Mimicry. The dorsal pattern of D. confusa closely resembles that of the night adders Causus rhombeatus (Lichtenstein), C. bilineatus Boulenger, C. defilippii (Jan) and C. maculatus (Hallowell) which all occur in north-eastern Africa (Spawls et al. 2002; Rasmussen 2005; Dobiey & Vogel 2007).

Localities. SOUTH SUDAN. Juba [NE 0431D3] FMNH 58496, ZFMK 29694; Logutoic, MCZ 53401; Lokwi [NE 0432A2] FMNH 62216; Mongalla [NE 0531D1] MCZ 53402; Nimule [NE 0332C1] MCZ 53403; ‘Sudan’ (probably East Equatoria state, or vicinity of Wadelai, Uganda; see Remarks under D. scabra) NMW 9964/7; Torit [NE 0432B1] FMNH 58361–3, 62207–14, 62217, MCZ 53404–5; Yei [NE 0430B1] FMNH 58336. ETHIOPIA. No precise locality, NMW 9964.2; ‘Addis Ababa’, ZMB 56436. UGANDA. Ajai Game Reserve [NE 0231C] MUZM —; Bisu, Kigezi [NE 0131C] BM 51.1.3.69; Bugoma Forest [NE 0130B4] LACM 39043; Busingiro, Budongo Forest [NE 0131D3] MCZ 47815; Gulu, Acholi [NE 0232C4] MCZ 47830; Kyambura Game Reserve [SE 0030A1] NMZB 14333, 15411; Lira, Lango [NE 0232B4] BM 51.1.5.39, MCZ 47801, UMMZ 88529–30; Moyo, West Madi [NE 0331D1] BM 51.1.3.84; Nakosa [SE 0030A3] LIV 1960.51.2; Nyanshimbo [SE 0030A1] LIV 1962.106.19; Sana [SE 0030B4] LIV 1960.67.2; Semliki [NE 0130A2] BM 1954.1.12.8 (‘120’) & 1954.1.12.9 (‘121’); Serere, Teso [NE 0133C2] BM 1976.848. KENYA. Kacheliba [NE 0135A3] NMK 485, 489; Mumias [NE 0034A4] CAS 141784. RWANDA. Lake Kivu [SE 0229A1] ZMB 77441.

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it appears to be widespread and should be regarded as Least Concern.

Remarks. The plain phase specimens from South Sudan and Uganda reported on here (see above) are the first of their kind to be documented. Dorsally uniform, or mostly unpatterned, specimens (brown or grey) are also known for D. scabra (southern Africa: e.g. De Waal 1978; Broadley 1990), D. medici (Gans 1959; Spawls et al. 2002; this study), D. gansi (Trape & Mané 2006; Bates & Ineich 2012), D. atra (Gans 1959, 1964; Spawls et al. 2002) and D. abyssina (see below).

Dasypeltis scabra (Linnaeus)

Common Egg-eater

Figs 5A–H & 6

Coluber scaber Linnaeus, 1758: 223. Type locality: ‘in Indiis’ [= “via the Cape … read as South Africa”, Flower 1933: 818; restricted to “probably near Cape Town”, Mertens 1954: 214].

Rachiodon abyssinus (not Duméril, Bibron & Duméril) Duméril, Bibron & Duméril, 1854: Vol. 7: 496 (part, “Nil Blanc”) (MNHN 7227, proposed paralectotype of R. abyssinus, see below).

Dasypeltis scaber, var. abyssinicus (lapsus) (not Duméril, Bibron & Duméril) Peters, 1864: 645 (Sennar, Sudan).

Dasypeltis scabra var. B Boulenger, 1894: 356.

Dasypeltis scabra Günther, 1894: 87 (E. Kikuyu, Kenya); Boulenger, 1896: 20 (between Comia & Mategoi, Somalia), 1897: 229 (Somalia) & 1898: 720 (Lugh, Somalia); Lönnberg & Andersson, 1913: 4 (Kismayu); Loveridge, 1929: 28 (part); Corkill, 1935: 18 (Blue Nile Province, Sudan); Loveridge, 1957: 288 (check-list); Gans, 1959: 141 (part), 1961: 73, 1964: 283, 1971: 461 & 1974: 34; Gans & Laurent, 1965: 67 (Mogadiscio); Lanza, 1972: 177 (Ilala Sala, Awash National Park), 1983: 223 (part) & 1990: 436 (part); Pitman, 1974: 116 (part); Spawls, 1978: 10 (part); Emmrich, 1985: 66, 68 (Oukam, Lake Zwai); Hoevers & Johnson, 1982: 199 (Mareri); Broadley & Howell, 1991: 32; Meirte, 1992: 57; Largen & Rasmussen, 1993: 340 (part); Hughes, 1997: 68 (part); Largen 1997: 89 (part); Behangana & Goodman, 2002: 66 (part, Uganda); Spawls et al., 2002: 418; Branch, 2005: 59 & 2015: 67; Spawls et al., 2006: 130; Ziliani et al., 2006: 193 (Mount Kulal, Kenya); Largen & Spawls, 2010: 490 (part); Geniez, 2015: 211 (Mount Kulal); Spawls, 2017: 31 (Maasai Mara).

Dasypeltis scaber Calabresi, 1927: 54 (Somalia); Scortecci, 1931: 208 (Giohar), 1939: 276 (part, Giohar, Belet Amin) & 1943: 285 (El Dire).

Dasypeltis scabra scabra De Witte, 1962: 139; Bourgeois, 1968: 284.

Dasypeltis scaber scaber Laurent, 1956: 258 (part).

Dasypeltis scabra” Göthel, 2015a: 19 (part) & 2015h: 86 (part).

Dasypeltis cf. scabra Sindaco et al., 2013: 331 (part, Kenya). 

Description. In the study area the dorsal pattern may vary quite considerably (see Gans 1959). Dorsum grey or light brown with darker markings (‘5N’ pattern of Gans 1959); saddles vary from rectangular to squarish, oval, subcircular, weakly hourglass-shaped, weakly transversely enlarged, or irregular, and often a mixture of two or more of these shapes (Figs 5 & 6). Dorsal saddles (46–94, mean 67.4, N = 107) are dark brown or black, and a few may be linked to the dark lateral bars. The centre of the saddle may be paler than the outer rim, as in MHNP 7227 (Sudan) where it is grey with a black edge (Fig. 6). Venter cream-white, usually with grey stipples, or dark blotches, at the edges; sometimes with large bold cones or blotches at the edges and/or mid-ventrally (see Gans 1959, e.g. Pl. 9). Top of head often with 1–2 anterior-pointing (sometimes fragmentary) chevrons and/or other dark markings.

Preoculars usually one on either side of head, but one on one side and two on the other in MCZ 30208 & USNM 62902, and two on either side in BM 01.1.3.4 & MSNG 28808 (N = 104); postoculars usually two on either side of head, but one on one side and two on the other in 6.0% of specimens, one on either side in 6.0% of specimens, and absent on both sides in ZMB 26832 (N = 133); frontal shield heavily pitted (States 3 or 4) in 57.7% of specimens (73.3% of specimens from Tanzania in the south of the study area, N = 30), marginally pitted (States 1 or 2) in 36.1% of specimens, and smooth (State 0) in 6.2% of specimens (N = 97); inter-prefrontal suture weak in 48.9% of specimens, moderately marked in 38.0% of specimens, and deeply sunken in 13.0% of specimens (N = 92); supralabials 11–16, usually 14 (both sides of head), usually 7 [3rd and 4th entering orbit] on either side of head, but variations are: 5[2,3] on right and 6[2,3] on left in ZMUC 6136, 6[2,3] on both sides in BM 68.2.29.101 and BM 1964.1069, 6[3,4] on both sides in ZMB 22518, 6[2,3] on one side and 7[3,4] on the other in FMNH 134939, MSNG 29239 and MZUF 2802, 6[3,4] on left and 7[3,4] on right in BM 68.2.29.100, 7[2,3] on both sides in NMK 3184, and 8[4,5] on both sides in ZMB 51835 (N = 123); anterior temporals usually two on either side of head (88.5%), occasionally 2 left:3 right or 3:2 (3.5%), three (3.5%), 2:1 (3.5%), and one in MRAC 18563 (N = 113); posterior temporals usually three on either side of head (64.6%), occasionally 3:4 or 4:3 (16.8%), 3:2 or 2:3 (7.1%), four (6.2%), two (4.4%), and five in PEM 5626 (N = 113); temporal formula on either side of head usually 2+3, occasionally 2+4 or 2+2; midbody scale rows 22–28 (mean 25.0 ±1.38, N = 102); ventrals 185–226 in males (N = 50), 202–249 in females (N = 76); subcaudals 47-73 in males (N = 45), 43–65 in females (N = 72). Ratio SVL: tail length 3.9–6.2 in males (N = 45), 5.3–8.9 (N = 65) in females; total length: tail length 4.9–7.2 in males (N = 45), 6.3–9.9 in females (N = 65).

INSERT FIGS 5 + 6

Size. Largest male (NMK 2965 – Kora, Kenya) 456 + 88 = 544 mm; LIV 1962.168.14 (Ruzinga, Ankole, Kenya) has a smaller SVL but is the longest male: 453 + 98 = 551 mm; largest female (NMK 3184 – Lake Naivasha, Kenya) 867 + 95 = 962 mm.

Habitat. Found in savannah and grassland from sea level to over 1500 m.

Distribution. From the south-western Cape of South Africa through Namibia, Botswana, Zambia, southern and western Democratic Republic of the Congo, much of south-eastern and eastern Africa as far north as Gojam area (too vague to map) in north-central Ethiopia, with isolated populations along the Nile Valley (see comments under D. bazi) in south-eastern Sudan and adjacent north-eastern South Sudan (see Gans 1959; Broadley 1990; Spawls et al. 2002; Fig. 3), as well as 50 km north of Kutum in western Sudan (see Remarks below).

Sympatry. Sympatric with with D. medici at Belet Amin (Somalia), Kilifi (Kenya), and Amani, Dodoma, Morogoro and Zanzibar (Tanzania); sympatric with D. confusa at Lake Kivu (Rwanda); sympatric with D. atra at Aware Melca and Lake Abiata (Ethiopia), Mulema (Uganda), Lake Naivasha and Mount Kulal (Kenya), Astrida (Rwanda) and Minziro Forest Reserve (Tanzania); sympatric with both D. medici and D. atra at Mt Kilimanjaro (Appendix 1).

Localities.ARABIA’. No specific locality, MNHN 6565. SUDAN. Nil Blanc, MNHN 7227 (Fig. 5, proposed paralectotype of D. abyssina, see below); 40 km E of Rabak, near Kosti [NE 1333A1] RMNH 24399; Sennar State [NE 1334C], BM 09.10.15.35. SOUTH SUDAN. Melut [NE 1032A3] FMNH 134939; ‘Sudan’ (probably East Equatoria state, or vicinity of Wadelai, Uganda; see Remarks), NMW 9964/6. ETHIOPIA. No locality, NMW 15298; Aware Melca [NE 0939B2] BM 16.6.24.11–12; Arba Minch [NE 0637B1] AAU 494, 679, BM 1969.288; Awash National Park [NE 0839D4] (Figs 5G&H); Awash National Park, 2 km NE of entrance to [NE 0839D4] PEM 8602–3; Bulcha [NE 0638A1] ZFMK 17763; Fitu, 135 km ESE of Neghelle [NE 0540B1] ‘5N’/‘2Bx’ dorsal pattern, photographic records; Gojam, BM 1964.1069; Lake Abiata [NE 0738D1] BM 1973.3259; Lake Chamo (west shore) [NE 0537D3] AAU 489; Meki–Dukem area [NE 0838B] ZMB 51835, 77053; Neghelle, 31 km ESE of [NE 0539B2] vertebral stripe over most of back, photographic records; west of Omo River [NE 0436C], BM 00.11.29.3. SOMALIA. environs of Afgoi [NE 0245A1] MZUF 1118, 1133; Afgoi, 6 km E of [NE 0245A1] MZUF 24179; Belet Amin [NE 0042B2] MSNG 42710A; between Comia and Matagoi [NE 0143C] MSNG 29239; Ceelcillan [= El Ellan, NE 0242D1] MZUF 31459; Dinsoor [NE 0242B4] MZUF 24674; Giohar [NE 0245D3] MSNM 785; near Mareri (Labadaah to Bula Yaag) [0042D1] MZUF 27156; Lugh (= Luuq) [NE 0342D1] BM 98.1.28.14, MSNG 28808; Lugh to Ganane [NE 0342D] MZUF 21968; Maheddei Uen [NE 0245D3] MZUF 21968; Mogadishu [NE 0242A2] MZUF 2802; Uagadi [NE 0144D1] MSNM 974. UGANDA. Kaburangiri [SE 0031D3] LIV 1962.106.6; Kakoba [SE 0030C1] LIV 1960.67.6; Kayanja, LIV 1962.330.22; Mbarara, Kasheri [SE 0030D1] BM 51.1.5.79, 1961.1804, LIV 1959.83.1; Mulema [SE 0031D1] BM 03.12.2.21; Namatyobola [SE 0031D1] LIV 1962.106.5; Ruburogoto [SE 0130B1] LICVM 1960.67.4, 1960.67.7, 1962.106.35; Rukoni [SE 0030C4] LIV 1962.106.20; Ruzinga, Isingiro [SE 0030D4] BM 1959.1.5.59, LIV 1962.168.14; Rwensama [SE 0029B4] LIV 1960.67.8; Wadelai [NE 0231C2] BM 07.12.1.19. KENYA. Aberdare Mtns [SE 0036B] USNM 42016; Buna, Wajir [NE 0239D3] NMK 2824; about 45 km W of Ewaro Ngiro, near Narok [SE 0135B3] PEM 5627; Gatab, Mount Kulal [NE 0236D2] MSNPV CR790–2; Gilgil [SE 0036C2] BM 01.1.3.3–4, and Fig. 5E; Hell’s Gate, Naivasha [SE 0036C4] CAS 147836; Isiolo [NE 0037B3] CAS 130784–5, NMK 561; Kilifi [SE 0339D2] BM 51.1.3.70; Kithu-Uri, East Kikuyu [SE 0037A4] BM 93.11.21.72; Kora [NE 0038B1] NMK 2965; Laini, Tana River [SE 0420C2] MCZ 40584; Lake Naivasha [SE 0036C4] NMK 1415, 1472, 2083, 2343, 2853, 3184, 3319, NMP 1260; Loita Hills [SE 0135D2] Fig. 5D; Mombosasa, Witu [SE 0240B3] MCZ 40583; Moshi [0337C2] (Gans 1964); Naivasha [SE 0036C2] CAS 147837–8; Ol Ari Nyiro Ranch [NE 0036C2] NMK 3103; Solio Ranch, Laikipia [SE 0036B4] NMK 3052; Sotik, Uaso Nyiro [SE 0035C1] USNM 41974; Tsavo East National Park [SE 0338B] NMK 3021; Uaso Nyiro Plains, MCZ 11501; Voi [SE 0338B3] NMK 1022; Wajir [NE 0140C3] CAS 130661, NMK 2262, Fig. 5C, and top photograph in Spawls et al. 2004: 418; Wema [SE 0240A1] MCZ 40585. RWANDA. Astrida [SE 0229D1] MRAC 18559; 15 km S of Butare [SE 0229D1] PEM 5624; Gabiro [SE 0130C2] IRSNB 5499, MRAC 9209, 9214; Lake Kivu [SE 0229A1] ZMB 22518; Nyagatare [SE 0130A4] PEM 5625. BURUNDI. Kabuyenge [SE 0329A4] MRAC 18563; Kahiro District [SE 0330C3] MRAC 18565; Mugera [SE 0329B4] MRAC 18564. TANZANIA. Amani [SE 0538A4] ZMB 21062; Chai Camp [SE 0336B4] AMNH 50791; Dar es Salaam [SE 0639C4] NMW 9964/17, ZMB 26832; Dodoma [SE 0635B1] (Fig. 5A); Kibwesa [SE 0629B4] ZMB 22754; Kikombo, Dodoma [0635B2] USNM 72476; Maji Malulu, Usandawi [SE 043B4] MCZ 30205; Mapenduzi, Dodoma [SE 0635B2] PEM 5626; Masiko ya Tembo, vicinity of Sanya Chini [SE 0337A3] (Fig. 5B); Massailand, ZMB 11934a,b, 14097; Minziro Forest Reserve [SE 0131B1] ZMUC 6136–7; Morogoro [SE 0637D3] USNM 62902–3, 62905; Mount Kilimanjaro [SE 0337A2] BM 87.11.3.30; Mpwapwa [SE 0636B3] ZMB 13304, 19669; Ngare, Nanyuki [SE 0235C1] FMNH 27786; Serondera, Serengeti National Park [SE 0234 C4] NMK 1887;Tanga [SE 0539A1] NMW 9964/9; Ukerewe Island [SE 0233A1] MCZ 30207–8; Wembere Steppe [SE 0434A] ZMB 16753; Zengeragusa, Mkalama [SE 0434B1] MCZ 18208.

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it appears to be widespread and abundant and should be regarded as Least Concern.

Remarks. MNHN 6565 is a female from “Arabia” (without precise locality data) with a ‘5N’ dorsal pattern consisting of 55 fairly broad saddles (some ‘diabolo’-shaped), 244 ventrals and 54 subcaudals. Although its provenance is uncertain, we refer this specimen to D. scabra.

According to Gans (1959: 11, 205) specimen BM 07.12.1.19 was collected in ‘Wadelai’, north-western Uganda, by Emin Pasha (also known as Emin Bey, baptised E.C.O.T. Schnitzer [1840-1892]: Chisholm 1911; Anon 2017). A colour photograph of the specimen (sent by P. Campbell, Natural History Museum, London) was made available to the first author, and the ‘5N’ dorsal pattern was confirmed. This appears to represent an isolated population of D. scabra. Specimens of D. scabra (NMW 9964/6, ‘5N’) and D. confusa (NMW 9964/7, ‘5L’) from ‘Sudan’, donated to the Naturhistorisches Museum Wien in 1884 by ‘Emin Bey’, may have been collected nearby in South Sudan’s East Equatoria state, or possibly in the vicinity of Wadelai. Emin Pasha was governor of Equatoria (a region in southern South Sudan that included a portion of north-western Uganda), then a province of Anglo-Egyptian Sudan; he retreated to Wadelai in 1885 in the face of the Madhist advance (Chisholm 1911; Anon 2017).

ZFMK 32479 from 50 km north of Kutum, North Darfur State, western Sudan is a faded male specimen with a damaged head, referable to D. scabra on account of the following characters: ‘5N’ dorsal pattern with dark oval saddles and whitish interspaces, pattern cycles 50, SVL 447 mm, tail length 81 mm, SVL/tail ratio 5.5, ventrals 227, subcaudals 66, midbody scale rows 23. This locality is outside the study area and the only other known record of D. scabra in the north of Africa. It is not associated with the Nile River and represents an extremely isolated population, possibly a relic of a former widespread distribution (see discussion below under D. bazi). The nearest conspecific (RMNH 24399) is from near Kosti about 880 km ESE.

Dasypeltis bazi Saleh & Sarhan

Egyptian Egg-eater

Figs 7A–B, 8, 9A–B, 10–12

Rachiodon abyssinicus lapsus for abyssinus (not Duméril, Bibron & Duméril) Lichtenstein & Von Martens, 1856: 28 (“Oberaegypten” = Upper Egypt).

Rachiodon scaber (not Linnaeus) Jan, 1863: 106 (Cairo); Panceri, 1878: 22 (Egypt).

Dasypeltis scaber (not Linnaeus) Gasco, 1876: 119 (Middle Egypt).

Dasypeltis scabra (not Linnaeus) Anderson, 1896: 114, 1898: 278 (Fayum); Gans, 1959: 141 (part, Egypt, Fayum, Upper Egypt), Pl. IV, Figs 5& 6, Pl. VI, Fig. 1; Gans, 1961: 74 (part), Pl. 1, Figs A & B (Egypt); Marx, 1968: 33 (Egypt); Rehak & Osborn, 1988: 275 (Faiyum); Saleh, 1997: 146 (Faiyum); Baha El Din, 2006: 232 (Fayoum); Geniez, 2015: 210 (Fayoum).

Dasypeltis cf. scabra Sindaco et al. 2013: 93 (part, Egypt).

Dasypeltis scabra” Göthel, 2015a: 19 (part) & 2015h: 86 (part).

Dasypeltis bazi Saleh & Sarhan, 2016: 31. Type locality: Abu-Gandir, Faiyum Governorate, Egypt, holotype AUZC-R09458.

Diagnosis. Assigned to the genus Dasypeltis on account of its slender form, possession of (usually 3–4) rows of reduced, oblique, keeled and serrated lateral scales (little or no serration in D. inornata) and head barely distinct from the neck (moderately distinct in D. fasciata). Distinguished from other congeners by its colour pattern consisting of: large cranio-caudally elongate dark brown saddles, usually bifurcated at each end with a fairly shallow but distinct notch, the saddles about 1.5 to 2 times as long as the pale interspaces between them (similar to D. loveridgei comb. nov., but this species has mostly transversely enlarged and distinctly hourglass-shaped saddles which are mostly about equal in length to the pale interspaces; similar but less elongate saddles in some populations of D. scabra, e.g. Fig. 5G & H); low numbers of pattern cycles 38–49 (lower than all other [patterned] species/populations except D. taylori sp. nov. and a few specimens of D. scabra; D. inornatus always uniform brown; D. gansi uniform or weakly marked); flanks decorated with dark brown spots or blotches (similar in D. loveridgei comb. nov.; and some D. scabra, e.g. 5G & H) rather than vertical bars as in most other (patterned) Dasypeltis species/populations; and venter immaculate white to cream-white apart from a few dark markings at the edges in some specimens (extensive dark markings in D. abyssina and many D. scabra, often extensive grey stippling in D. medici); together with a combination of the following characters: Ventrals 213–229 in males (mostly higher than D. taylori sp. nov. 196–216, lower than D. arabica sp. nov. 236–244), 215–232 in females (lower than D. abyssina 241–271, D. arabica sp. nov. 239–254, D. gansi 235–259, D. latericia 234–262; mostly lower than D. crucifera sp. nov. 231–247); Subcaudals 59–61 in males (lower than D. inornata 81–92, D. fasciata 71–90, D. medici 69–109, D. gansi 68–83, D. palmarum 68–77, D. parascabra 67–74, D. latericia 66–86, D. arabica sp. nov. 63–65), 51–63, 70 in females (mostly lower than D. inornata 69–84, D. fasciata 64–84, D. palmarum 62–77, D. medici 61–90); Tail of moderate length: SVL/tail length 5.0–5.9 in males, 5.1–7.2 in females (in study area, tail long in: D. medici 3.5–4.7 males, 4.5–5.7 females; D. fasciata 4.7–5.2 males, 4.8–6.4 females; see also Gans 1959); Postoculars 2 on either side of head (often one, especially in western populations, of D. atra); Nasal divided below the nostril (undivided in D. sahelensis and D. parascabra); Supralabials 7 (3rd and 4th enter orbit) on either side of head (usually 6[2,3] in D. loveridgei comb. nov.). Half the specimens examined had longitudinally divided anal plates; this plate is usually undivided in other species in the genus.

Description of BM 97.10.28.575 from Fayoum. (at Natural History Museum, London). Adult female (stretched oviducts and three eggs observed) with a total length of 692 mm, snout-vent length 602 mm, tail length 90 mm (Gans 1959) (ratio SVL: tail length 6.7, ratio total length: tail length 7.7) from Fayoum [about 29º18’30”N, 30º51’00”E], collected by A.R. Birdwood (Gans 1959, Pl. VI, Fig. 1; Figs 7 & 8). This is the same specimen discussed and illustrated by Anderson (1898, Pl. 34; see discussion under Remarks below) as D. scabra.

Head (somewhat dehydrated) small and barely distinct from the neck; snout rounded, rostral shield barely visible when viewed from above. The elongate nasal scale (about twice as long as high) is divided below the nostril (which is situated in the anterior half of the nasal); no loreal present. Three internasals between the rostral and paired prefrontals, but the larger, subcircular, central internasal appears to be an aberration. Each prefrontal is larger than an internasal (and the paired prefrontals together are much larger than all three internasals together), but they are of equal length (cranio-caudal orientation). Inter-prefrontal sulcus moderately marked. Frontal large, broadest anteriorly, 1.61 times as long as broad, smooth except for a single row of pits on the lateral borders, and of similar length to a supraocular. Frontal followed by a pair of large parietals, each one similar in size to the frontal. A single preocular and two postoculars present on either side of head. Temporal formula 3+3 (left), 2+3 (right). Seven supralabials on either side, the 3rd and 4th bordering the eye. First pair of infralabials in contact behind the mental, followed by a pair of large and elongate scales on the throat, and another pair of much smaller scales in contact with the first ventral plate. Eyes moderate in size, their greatest diameter subequal to the distance from tip of snout to anterior part of eye socket.

Dorsal scales strongly keeled, unicarinate, in 24 rows at midbody, the third, fourth and fifth rows laterally (counted from the first row in contact with the ventral plates) are denticulate (serrated), reduced in size and obliquely positioned; vertebral scales not enlarged. There are 227 smooth ventrals, anal plate divided, with 55 pairs of smooth subcaudals. After preservation in ethanol the dorsal colouration is pale brown, with 42 large, cranio-caudally elongate, medium-brown saddles (with darker edges, especially on their anterior and posterior ends) from nape to base of tail (i.e. above vent), most of which are about 1.5 to 2 times longer than wide, and bifurcated at each end with a fairly shallow but distinct notch. Inter-saddle region cream-white, distinctly paler than the rest of the dorsal surface. Lateral markings are in the form of medium-sized, medium-brown spots or blotches, with darker edges, on either side between the saddles. Dark vertebral blotches continue along the tail, but lateral markings here are narrow and elongate. There are extensive dark (brown) markings on the top and sides of the head (including the labials), including a pair of elongated stripes on either side of the frontal, a pair of anteriorly-joined teardrop-shaped markings (blunt anteriorly) on the front of the parietals, and a pair of larger, similar markings on the posterior part of the parietals which extend onto the neck. The saddle on the nape has a spike-like anterior extension which terminates bluntly on the medial-posterior part of the parietals where it is in contact with the larger pair of (posterior) parietal markings. The latter are also narrowly fused with the smaller pair of parietal markings. Venter immaculate cream, apart from a few small brown spots and flecks extending from the outer edges of the ventrals to the outer scale rows.

INSERT FIGS 7 & 8

Description of additional Egyptian material. There are six additional specimens (one is a skeleton which we assign to this species) of D. bazi from Egypt, mostly with inexact localities, but potentially all collected at or near Fayoum (see discussion below).

ZMB 1861 (at Museum für Naturkunde, Berlin), adult male (left side inverted hemipenis examined), from “Oberaegypten” (= Upper Egypt, a strip of land on either side of the Nile River from about Memphis just above the Nile Delta southwards to Lake Nasser in Nubia, which may include parts of northern Sudan) collected by Karl Richard Lepsius (born 1810, died 10 July 1844), a Prussian Egyptologist who was leader of an expedition to Egypt conducted between 1842 and 1846 (Figs 9–11). This expedition visited places from Memphis in the north traveling along the White and Blue Nile rivers into central Sudan. The specimen, in remarkably good condition, appears to be the same snake referred to by Lichtenstein & Von Martens (1856).

ZMB 51552, adult female (left side of tail base dissected open, no hemipenis) from “Oberaegypten” (= Upper Egypt), collected by the French physician Dr Alfred Peney (1817–1861), who was together with K.R. Lepsius in Egypt and Sudan; this specimen was apparently later donated to Albert von Barnim and Robert Hartmann who visited Peney on an expedition to Egypt, Nubia and eastern Sudan between 1860 and 1861. Barnim and Hartmann are mentioned on the jar label as donors for this specimen to the “Anatomisch-Zoologisches Museum” in Berlin, and the specimen was incorporated into the Zoologisches Museum Berlin in 1888.

ZMB 5865, adult male (inverted hemipenes present, F. Tillack in litt.) from “Aegypten” (= Egypt), purchased from Mr “Keitel”; specimen catalogued around 1867 by W.C.H. Peters as “Rachiodon abyssinicus Dum. Bib.”.

NMW 26879/1, adult female (no spines in tail, tail distinctly tapered at base) from “Omgebung v. Cairo” (= surroundings of Cairo), donated by Franz Steindachner in January 1874.

NMW 962, mounted adult skeleton with skull (few teeth present), from “Kairo” (= Cairo), donated or collected by Franz Steindachner in 1880, labelled “Dasypeltis scabra”.

MNHN 1847, adult male (left side hemipenis everted) from “Egypt”, collected by Baron Alphonse Marie Delort Gléon Leopard (1843–1899).

Size: ZMB 1861, male, 455 mm SVL + 79 mm tail length = 534 mm total length; ratio SVL: tail length 5.8, ratio total length: tail length 6.8; ZMB 5865, male, 427 + 78 = 505 mm, ratios 5.5 and 6.5; MNHN 1847, male, 410 + 70 = 480 mm, ratios 5.9 and 6.9; ZMB 51552, female, 503 + 70 = 573 mm, ratios 7.2 and 8.2; NMW 26879/1, female, 461 + 61 = 522 mm, ratios 7.6 and 8.6.

Scalation: All have similar scalation and general appearance to BM 97.10.28.575, except: Nostril on right side of NMW 26879/1 situated in about middle of nasal scale (not in anterior part thereof). All have a pair of internasals (not three) between rostral and paired prefrontals, and these are (individually and in combination) smaller than the prefrontals (smaller but of equal length in ZM 51552). Inter-prefrontal sulcus weakly marked in ZMB 5865 and ZMB 51552. Ratio frontal length: frontal width: 1.54 (ZMB 1861), 1.44 (ZMB 5865) and 1.40 (MNHN 1847) in males; 1.38 (ZMB 51552) and 1.43 (NMW 26879/1) in females; frontal smooth in ZMB 51552, covered by small pimple-like protrusions in MNHN 1847. Frontal slightly shorter than each supraocular in NMW 26879/1. In ZMB 1861 the pair of parietals is fused medially, with only the ends separated. Temporals 2+3 on both sides. In NMW 26879/1 and ZMB 1861 the pupils are clearly vertically elliptical. Midbody scale rows 23 or 24 (NMW 26879/1, ZMB 51552). Reduced, oblique, serrated lateral scale rows: 3–5 (MNHN 1847, ZMB 5865) and 3–6 (NMW 26879/1, ZMB 1861, ZMB 51552), but all except NMW 26879/1 and ZMB 51552 also have the scales of the second row weakly serrated although not reduced in size like the other serrated rows, nor obliquely oriented. Ventrals in males 219 (ZMB 5865), 229 (ZMB 1861, MNHN 1847); in females 228 (ZMB 51552), 234 (NMW 26879/1). Anal plate undivided in MNHN 1847, ZMB 1861 and ZMB 51552. Subcaudals in males 59 (MNHN 1847, ZMB 1861), 60 (ZMB 5865); in females 50 (NMW 26879/1), 51 (ZMB 51552).

Colour pattern: Basal colour dorsally in MNHN 1847 is grey, not pale brown. Saddles on back 39 (ZMB 5865), 48 (ZMB 1861, ZMB 51552), 49 (MNHN 1847); about 48 in NMW 26879/1 which is largely faded. Along the tail the lateral markings are in the form of small dark spots/blotches in MNHN 1847, NMW 26879/1 and ZMB 1861. Brown forward-pointing V-shaped markings on back of head and nape in MNHN 1847; distinct, backwards-curving, narrow pale band at back of head, bisected by spike-like extension of a dark and mostly blunt V-shaped marking (almost band-like) on nape in ZMB 5865; ZMB 1861 and ZMB 51552 similar to previous specimen but pale band not interrupted, and the latter with a longer and sharper anterior extension of the nape marking.

INSERT FIGS 9–11

Description of ZFMK 38415 (Dasypeltis cf. bazi). Adult male (600 mm total length) from Erkowit, north-eastern Sudan [18º46’N, 37º07’E], collected dead-on-road (head damaged) in Spring 1982 by Gerhard Nikolaus (Fig. 12).

Ventrals 229, subcaudals 67, midbody scale rows 25, supralabials 7 (3rd and 4th enter orbit) on either side of head, one preocular on either side, two postoculars on either side, 3+4 temporals on either side, 48 pattern cycles. Photographs (e.g. Fig. 12) of the specimen show a dorsal pattern similar to that of BM 97.10.28.575; venter also similar, but yellowish-cream; dorsal and ventral scales, including their arrangement on the throat, similar to holotype; reduced, oblique, serrated lateral scale rows 3–6, but second row (scales similar in size to central dorsals) weakly serrated although not reduced or oblique; frontal similar in shape to that of the holotype, smooth except for 1–2 rows of pits laterally, of similar length to a supraocular, but slightly larger than each of the parietals.

INSERT FIG. 12

Size. Largest male ZMB 1861 (“Upper Egypt”) 455 + 79 = 534 mm [The largest specimen identified as a male by Saleh & Sarhan 2016 measured 482 + 75 = 557 mm, but its sex requires confirmation—see below]. Largest female BM 97.10.28.575 (Fayoum, Egypt) 602 + 90 = 692 mm.

Distribution. Dasypeltis bazi appears to be restricted to an area of about 300 km2 along the El Buts channel (near the village of Ibshwai) in the Fayoum Depression, which is situated about 100 km south-west of Cairo and 40 km west of the Nile River, at elevations of 32–52 m below sea level (Saleh & Sarhan 2016). Apart from sites in the Fayoum, there are a records of this species from only one other specific locality in Egypt, namely Cairo (see discussion below), but its occurrence there was not confirmed by Saleh & Sarhan (2016). Cairo is now a densely populated area, and if these records are accurate, the species may have been extirpated there. Although it has been suggested that it occurs elsewhere in Egypt along the Nile River (e.g. Anderson 1898; Baha El Din 2006), this was also not confirmed by Saleh & Sarhan (2016) who investigated potential habitats in various parts of the country.

Historical records. Based on the literature, D. bazi is a poorly known species. The first record of an egg-eating snake from Egypt is contained in Lichtenstein & Von Martens’ (1856) list of the reptiles and amphibians in the collection of the Royal University of Berlin. He recorded a specimen of Rachiodon abyssinicus (lapsus for abyssinus) from “Oberaegypten” (= Upper Egypt). Jan (1863) later recorded Rachiodon scaber (= D. scabra) from Cairo with reference to a specimen in the Museo Civico di Storia Naturale di Milano. However, the latter specimen was not illustrated, nor were any morphological data provided, and it is no longer to be found at the museum in Milan (S. Scali in litt. 26 September 2014). No other mention has ever been made of the presence of this species in Cairo, a densely populated area, so it may have been extirpated there. In an account of his travels in Egypt, Gasco (1876) recorded eight specimens of Dasypeltis scaber (= D. scabra) from Middle Egypt (= Nile Valley between the Nile Delta and Asyut or Qena Bend), collected in 1873–1874. The title of Gasco’s (1876) paper, the second of two reports on his travels in Egypt, actually refers to the oasis of Fayoum and excursions in the vicinity of Cairo, although it also alludes to hunting on Lake Menzala on the Nile Delta. Anderson (1896: 114) noted that “Count Peracca has been so good as to ascertain from Prof. Costa that two specimens brought back from Egypt by Prof. P. Panceri, the companion of Gasco, one a skeleton and the other in alcohol, are preserved in the Naples Museum. Panceri (1878) did in fact refer to two such specimens (number 3219, alcohol specimen; 3520, skeleton and teeth), as Rachiodon scaber, in his published catalogue of the collection in the Comparative Anatomy section of Università degli Studi di Napoli Federico II. It is not known whether the two specimens from Cairo in the Vienna collection, as mentioned above, were a part of the series alluded to by Gasco and Panceri, but the dates involved do coincide.

Subsequently, in Zoology of Egypt, Anderson (1898: 279) wrote: “After much correspondence with those in Egypt who have been assisting me in my researches, at last, through the efforts of Mr. A. R. Birdwood, after I had almost given up all hope of finding this species, I received the specimen herein recorded alive and am thus enabled to give a figure of it from life.” According to Anderson (1898: 279) the specimen was collected in “the Fayum” (variously spelled as “Faiyum”, “Fayoum”, “Al Fayyum” and “El Faiyūm”), which probably refers to the oasis in general rather than the city of Fayoum itself; the area is also referred to as the Fayoum Depression. Anderson (1898, table) was of the opinion that D. scabra also occurred in the “Valley of White Nile”. In fact, Anderson’s (1898: 279) comment about the “wide distribution” of ‘D. scabra’ and his suggestion that the species occurs “northwards along the valley of the Nile to within about 140 miles [= 225 km] of the Mediterranean [Sea]” is somewhat misleading because apart from the ‘Nil Blanc’ record (a syntype of R. abyssinus; see below) of D. scabra far to the south, there does not appear to have been (nor is there now) any other specific documented evidence to suggest that the Fayoum population was in fact contiguous with other Dasypeltis populations. For his Fayoum specimen Anderson (1898) provided scalation and size data, line drawings of the head and anterior part of the body, and an illustration (painting in colour) of the snake in the process of swallowing a pigeon’s egg.

Anderson’s specimen (BM 97.10.28.575) was later examined by Gans (1959) who remarked on the uniqueness of its dorsal pattern. Gans (1959) also examined three more specimens (also listed and examined here) from Egypt and “Upper Egypt” (ZMB 1861, this is probably the snake mentioned by Lichtenstein & Von Martens 1856). One additional specimen (ZMB 51552) from “Upper Egypt” recorded here can be added to the list of old museum specimens. These non-specific localities could conceivably refer to specimens collected at other localities apart from the Fayoum Depression, perhaps along the Nile River itself or possibly in one of the country’s other oases.

Rehak & Osborn (1988: 275) later recorded a specimen “collected by natives in Faiyum, September 1982” and noted that it “seems extremely rare in Egypt”. Saleh (1997) subsequently provided a photograph of a specimen he collected in Fayoum, again illustrating the much-elongated saddles with spots/blotches on the flanks. Although Saleh (1997) claimed that his specimen from Fayoum was the first to be collected in almost a century, Rehak & Osborn (1988) had in fact documented a specimen collected there about 90 years after Anderson’s (1898) report. Baha El Din (2006) was of the opinion that the only certain locality for D. scabra in Egypt is the Fayoum Depression, but it is also likely to occur in parts of the Nile Valley. He added that Gasco’s (1876) locality of “Middle Egypt” may be in reference to Fayoum, which is located within the northern parts of this region. Baha El Din (2006) also noted that animal collectors from Abu Rawash (30o01’55”N, 31o04’30”E; near the southern edge of the Nile Delta, about 80 km north-east of Fayoum) claimed to find specimens at pigeon breeding houses. If it does in fact occur in the vicinity of Abu Rawash, this would be the most northerly known locality for the species and the most north-easterly extent of the range of the genus Dasypeltis.

The new species was therefore, prior to 2016, known to science from only 16 specimens recorded over a period of about 170 years (first specimen collected by Karl Lepsius in about 1844, see above). Saleh & Sarhan (2016) reported in detail on eight specimens collected at Fayoum in the period 1995–2015 (presumably including Saleh’s [1997] specimen) and also mentioned another 17 specimens collected there since 1995 but released. There are no other records of egg-eaters from Egypt. Venchi & Sindaco’s (2006) checklist of the reptiles of the Mediterranean region did not add any additional records of Dasypeltis. Watkins-Colwell et al. (2010) reported on reptiles recorded during three Yale University expeditions to Egypt in the first half of the 1960s, which included several sites along the Nile River and the Fayoum Oasis, but no Dasypeltis were recorded. Ibrahim (2013) recently reported on the herpetofauna of the Suez Canal Zone, surveyed between 1999 and 2012. Despite the fact that habitat changes in the area at and near the canal (which includes associated freshwater irrigation canals) have allowed ‘invasive’ species to expand their ranges, and that an extensive web of irrigation canals has provided passage for several species to move from the Nile River to the Suez Canal, no specimens of Dasypeltis were recorded. If Dasypeltis occurred in the Nile Delta itself, for which there is no actual evidence, this too would allow easy access to the Suez Canal.

Natural history. Saleh (1997: 146) collected a specimen from Fayoum “in an area with dense wild vegetation, not far from houses at the edge of cultivated land”. Saleh & Sarhan (2016) later provided a detailed description of the habitat of the Fayoum population. All specimens were collected along the El Buts channel, a relict section of the Fayoum ancient natural drainage system. This habitat comprises a fairly deep gorge in the thick and silty Nilotic deposits in the central region of Fayoum Depression. Terraces occupy the sides where small plots of crops are grown (Saleh & Sarhan 2016). Anderson (1898: 280) noted: “On Plate XXXIX. it [his Fayoum snake—perhaps depicted while in captivity] is represented in the act of swallowing a pigeon’s egg, but fowls’ eggs are even within its capacity.” Saleh & Sarhan (2016) later determined that several bird species that produce suitably small eggs occur in sympatry with these snakes at Fayoum. In addition, they noted that traditional pigeon towers and chicken hatcheries also provide a source of eggs. Based on the presence of three large eggs observed in the abdomen of BM 97.10.28.575, this species is, like other congeners for which data is available (e.g. Broadley 1990; Spawls et al. 2002), oviparous.

Mimicry. Dasypeltis bazi has a dorsal colour pattern similar to the saw-scaled viper Echis pyramidum Geoffroy Saint-Hilaire (Schleish et al. 1996; see photographs in Dobiey & Vogel 2007) which it appears to mimic. Like Echis Merrem, it has similar serrated lateral dorsals which it may engage for defensive display by coiling and rubbing together to produce a rasping sound (scale stridulation, Schleish et al. 1996). The type series of Echis pyramidum lucidus Cherlin, 1990 was collected in the Fayoum Oasis (Cherlin 1990). A specimen of the latter subspecies from Biskra in Algeria (www.megasphera.cz/africanvenomoussnakes/, accessed 11 June 2014) has a dorsal pattern almost identical to D. bazi (e.g. Fig. 7A-B). Echis pyramidum was also collected in the Fayoum Oasis during the Yale University Expedition (Watkins-Colwell et al. 2010). It is one of several species of snakes that occur in the same habitat as D. bazi (see Saleh & Sarhan 2016). Anderson (1898) suggested that the similarity of the Fayoum snake to Echis carinatus (Schneider) (= ? E. pyramidum) might explain why this egg-eater is so seldom observed or recognised, but Saleh (1997) noted that the Fayoum Dasypeltis that he obtained was not particularly similar to this viper. Gans (1961, Pl. 1) illustrated the similarity between these two species using ZMB 5865 (‘Egypt’) and a specimen of ‘E. c. carinatus’ (CAS 38722) from Cairo.

Biogeography. Dasypeltis bazi appears to have a relict distribution, the result of a once extensive African savannah being reduced and fragmented following dessication of the region during the Holocene (Kuper & Kröpelin 2006). This is discussed in more detail below (see Discussion).

Conservation. The Fayoum Oasis is a depression or basin linked to the Nile River by a 24-km-long canal called the Bahr Yussef. In prehistory the oasis contained the large freshwater Lake Moeris, which received water when the Nile was in high flood, but it is now a much smaller saltwater lake referred to as Birket Qaran (= Qaroun Lake). In 2300 B.C.E. the natural waterway from the Nile to the lake was widened and deepened to control flooding, regulate water levels during the dry season, and provide the area with irrigation for crops. Unlike other oases in Egypt, which have spring water sources, cultivated land in the Fayoum is formed by fertile mud from the Nile transported there via the Bahr Yussef. The scarcity of Dasypeltis records from the Fayoum is perhaps not too surprising considering the high levels of human activity (e.g. crops and housing) in the region. Saleh & Sarhan (2016: 37) noted: “Through centuries of human occupation and intensive agriculture, the Faiyum natural drainage system was fundamentally modified and almost totally obliterated.” Proposed tourism developments will impose further threats to natural habitat in the area (Khalifa & El-Khateeb 2011).

Despite the benefits provided to the human population of Egypt by the building of the Aswan High Dam, which created Lake Nasser, this dam has permanently altered the ecology of the Nile River region, with some areas now under water and others changed from desert into cultivated fields or wetlands (Watkins-Colwell et al. 2010). Although the establishment of vegetated areas has resulted in some species extending their ranges, e.g. inland from the Nile Delta (Baha El Din 2006; Watkins-Colwell et al. 2010), this does not appear to have benefited Dasypeltis. Since the construction of the dam and the resultant low flow rates, the Nile Delta no longer floods and has been converted into an intensive agricultural area (Van As et al. 2012). In the past specimens may have been introduced downstream—even from as far away as Sudan—during floods, on rafts of floating vegetation, but this no longer occurs. Although the delta is still an important area for a number of animal species, including large numbers of migratory birds (Mockrin & Thieme undated), no egg-eating snakes have been recorded there.

Considering the scarcity of records as indicated above, D. bazi may be a naturally rare species. Anderson (1898: 280) noted: “It seems remarkable that a snake of this kind, so specialized as regards its food-supply, should have been unknown to the ancient Egyptians. There is no evidence, so far as I have been able to discover, that they were acquainted with it, unless the figure, said to occur on the monuments, of a hawk-headed serpent with an egg in its mouth may have been derived from some knowledge of it.” The Egyptian god Cneph is in fact occasionally represented as a serpent with an egg in its mouth (Deane 1833). Anderson (1898) also noted that he had made a special effort to obtain specimens, yet he only eventually secured a single snake, from Fayoum. Rehak & Osborn (1988) also recorded a single specimen and considered Dasypeltis as extremely rare in Egypt. This specimen was apparently collected by a Fayoum villager in September 1982 but was purchased from an animal dealer in Cairo (see Saleh & Sarhan 2016). Saleh (1997) and Baha El Din (2006) classified the Egyptian population of ‘D. scabra’ as regionally Critically Endangered. Baha El Din (2006: 234) noted: “The impact of the large-scale ecological changes that took place in Fayoum since the beginning of last century is unknown, but probably reduced available habitat. The species is threatened by over-collection by professional animal collectors [in Egypt], who target the species.” This suggestion about over-collecting appears to be based on his comment about animal collectors from Abu Rawash. Saleh & Sarhan (2016: 43) concluded that D. bazi was “saved from certain extinction by the development [since the mid-Holocene] of artificial breeding of pigeons and chickens in Faiyum” which provided a source of eggs.

The Erkowit specimen may represent an isolated relict population of D. bazi in north-eastern Sudan. This population may in the past have been part of a more widespread population of D. scabra that became isolated following aridification of the Sahara, resulting in the evolution of a new species, D. bazi. The Egyptian and Erkowit populations themselves may then have become isolated, accounting for their large current-day separation. Habitat at Erkowit has been negatively affected by the impacts of a human population that has grown considerably in the last 60-odd years, with associated removal of wood, pressure from livestock grazing and browsing, and a warming climate (Vetaas et al. 2012). This environment has suffered continuous change from semi-evergreen woodland to open shrubland similar to that of the surrounding desert, and the area can no longer be regarded as a ‘mist oasis’ dominated by Afromontane and Mediterranean plant species (Vetaas et al. 2012). Therefore, there is some uncertainty about the conservation status of the Erkowit population.

Based on a global evaluation of its conservation status using IUCN (2012, 2016) criteria—and considering the Fayoum Oasis population as the only surviving population of this species (see above)—it is evident that D. bazi should be classified globally as Endangered [B1ab(i, ii, iii)] based on its small geographical range (known from an area of 300 km2 [Saleh & Sarhan 2016], extent of occurrence <5000 km2 [the entire oasis is about a third of this size]) which is severely fragmented, with an inferred continuing decline in its extent of occurrence, area of occupancy and quality of habitat (see Baha El Din 2006; Khalifa & El-Khateeb 2011; Saleh & Sarhan 2016).

Remarks. Although Anderson (1898) did not identify his Fayoum snake by a museum accession number, he did mention that reptile and amphibian specimens collected and discussed by him were lodged in the British Museum (now in the Natural History Museum, London), the Museum of Egyptian Government School of Medicine (Cairo) and a few museums on the European mainland. Baha El Din (2006: 233) actually identified Anderson’s specimen as “BMNH 97.10.28.575”. Based on the illustrations, measurements and scalation details provided by Anderson for his Fayoum snake, it is apparent (see below) that his specimen is indeed the same snake preserved at the Natural History Museum in London and labelled as BM 97.10.28.575, which was examined by both Gans (1959, see Pl. 6, Fig. 1) and the first author. Firstly, the body shape, and shape and proportion of dorsal markings in Anderson’s (1898) Pl. 34, Fig. 3 are remarkably similar to the specimen examined (Figs 7 & 8), including the first saddle on the nape with its narrow finger-like forward projection. Secondly, measurements of SVL and tail length by Anderson (602 + 95 mm) and Gans (602 + 90) are similar. Thirdly, there are general similarities with regard to scalation. Anderson’s ventral count of 230 is similar to the count of 227 determined by both Gans and the first author. The slight difference would be accounted for if Anderson included in his count the three pairs of scales between mental and first ventral. Some of Anderson’s scale counts are in exact agreement with those obtained by Gans and this author: 24 midbody scale rows, one preocular and two postoculars. Anderson’s subcaudal count of 49 is slightly lower than the 54 counted by Gans and the 55 counted for the present study. However, because subcaudals in Dasypeltis are paired—and these paired scales are not always clearly aligned—it is possible that Anderson under-counted, thus minimising this count. Also, while Anderson recorded eight supralabials with 4th and 5th entering the orbit, both Gans and the first author recorded only seven supralabials with 3rd and 4th entering the orbit. However, an examination of Anderson’s (1898) Pl. 34, Fig. 3, representing a lateral view of the right side of the head and neck, clearly shows seven supralabials with the 3rd and 4th entering the orbit. It seems that Anderson may have inadvertently or erroneously included the rostral in his count, which would have created the impression that the ‘4th’ and ‘5th’ supralabials entered the orbit. Anderson provided a temporal formula of 3+3, while Gans recorded 2+3, but the first author observed 3+3 on the left side of the head and 2+3 on the right.

A few inaccuracies were noted in Saleh & Sarhan’s (2016) diagnosis of D. bazi. Firstly, these authors state that in D. bazi the frontal shield is longer than wide, but as long as wide in other species of Dasypeltis. In fact the frontal is longer than wide in most if not all congeners (e.g. Gans 1959; Broadley 1990; Trape & Mané 2006; Trape et al. 2012; figures in this paper). Also, the temporal arrangement of 2+3 (two anterior and three posterior temporals on either side of the head) is common in the genus and the norm for several species (Gans 1959; Broadley 1990; this paper); while one preocular on either side of the head is the norm in all species, even though there are occasionally two or three such scales in some species (Gans 1959; this paper). Elsewhere in their paper Saleh & Sarhan (2016) recorded low numbers of longitudinal scale rows at midbody (20–21, one specimen with 22). Egyptian specimens of D. bazi examined for the present study had counts of 23–24. Saleh & Sarhan (2016) also counted ‘anterior’ and ‘posterior’ longitudinal scale rows, and their ‘anterior’ scale row counts (presumably counted halfway between tip of snout and midbody) of 21–25 were closer to the values obtained in the present study. It is possible that they did not use the same method for counting these scale rows as other authors, especially that proposed by Gans (1959) and used in this study, and therefore their comparisons with other species are not feasible. Saleh & Sarhan (2016) added to the confusion when comparing ‘dorsal scale rows’ between species by using the total range of anterior, midbody and posterior scale row counts for D. bazi (i.e. ‘18–24’; but 18–25 in their Table III) and comparing this with what is almost certainly only midbody scale rows recorded by other authors for other species. Lastly, a specimen (AUZC-R12006) identified as a male by Saleh & Sarhan (2016) has a very high ventral count of 250 (and low subcaudal count of 55), and may be a female; the next highest ventral count for a male D. bazi is 229 (see above).

Dasypeltis taylori Bates & Broadley sp. nov.

Taylor’s Egg-eater

Figs 13–14

Dasypeltis scabra var. Medici (not Bianconi) Mocquard, 1888: 131 (‘Comalis’ = Djibouti).

Dasypeltis scabra (not Linnaeus) ? Scortecci, 1930: 213 (Assab, Eritrea) (material not examined); Gans, 1959: 141 (part), Pl. VI, Fig. 6 & Pl. VII, Fig. 1; Balletto, 1968: 268 (part); Lanza, 1983: 223 (part), 1990: 436 (part); Ineich, 1999: 35 (Djibouti).

Dasypeltis scaber (not Linnaeus) Parker, 1949: 66 (Haud, Ogo and Hargeisa, Somalia).

Dasypeltis scabra” Göthel, 2015a: 19 (part) & 2015h: 86 (part).

Holotype. BM 49.2.2.2 (at Natural History Museum, London), a subadult male (Gans 1959) from Haud [former British Somaliland, 08°20’N, 46°00’E at 2100 ft = 640 m] in northern Somalia, collected by Colonel R.H.R. Taylor on 20 April 1932 (Fig. 13).

Paratypes. Seven specimens: SOMALIA (Somaliland). Allotype. BM 49.2.2.3 (at Natural History Museum, London), a subadult female (Gans 1959) with the same data as the holotype, but collected on 1 May 1932 (Gans 1959, Pl. VI, Fig. 6); BM 49.2.2.1 (subadult male) with the same data as the holotype; BM 49.2.2.4, adult female from Hargeisa District [9°35’N, 44°05’E at 4000 ft = 1220 m a.s.l.] collected 29 May 1932; BM 1992.501, male from 2.5 km WSW of Hargeisa [09°34’N, 44°03’E at 1275 m a.s.l.] collected 29 March 1992 by M.R.K. Lambert; BM 49.2.2.5, adult male from Ogo [10°10’N, 42°50’E at 4500 ft = 1370 m a.s.l.] collected 6 August 1933; BM 51.1.7.69, skin of a male from Garadak [09°29’N, 46°18’E at about 1100 m a.s.l.]. DJIBOUTI. MNHN 84–707, adult female from ‘Comalis’ [= Djibouti] (Fig. 14).

Diagnosis. Assigned to the genus Dasypeltis on account of its slender form, possession of (usually 3–4) rows of reduced, oblique, keeled and serrated lateral scales (little or no serration in D. inornata) and head barely distinct from the neck (moderately distinct in D. fasciata). Distinguished from other congeners by a combination of the following characters: Colour pattern usually consists of: large, usually closely-spaced, square to slightly rectangular saddles, separated by narrow, usually sharply defined, white crossbars (a variety of ‘5N’, Gans 1959, Pl. VI, Fig. 6); low numbers of pattern cycles 44–59 (≥62 in patterned species/populations of D. palmarum, D. fasciata, D. abyssina, D. atra, D. arabica sp. nov., D. inornatus always uniform brown; D. gansi uniform or weakly marked); flanks decorated with dark vertical bars (spots or blotches in D. bazi, D. loveridgei comb. nov. and some populations of D. scabra); and venter immaculate apart from a few dark markings at the edges (extensive dark markings in D. abyssina and many D. scabra); Frontal shield extensively pitted, States 3 & 4 (usually marginally pitted in most other species except D. medici, and some populations of D. scabra [especially in the south of its range] and D. loveridgei comb. nov., which are similar); Inter-prefrontal sulcus usually moderately depressed (usually weakly marked in most other species, but moderate to well marked in D. arabica sp. nov., weakly to moderately marked in D. bazi, and variable in D. scabra [often well marked in the south of its range]); Low numbers of ventrals: 196–216 in males (≥219 in: D. arabica sp. nov. 236–244, D. abyssina 226-247, D. crucifera 226, D. gansi 221–240, D. latericia 219–243, mostly lower than D. medici 215–254 and D. bazi 213–229), 208–222 in females (lower than D. abyssina 241–271, D. arabica sp. nov. 239–254, D. gansi 235–259, D. latericia 234–262, D. crucifera sp. nov. 231–247, D. fasciata 225–260, D. parascabra 224–237; mostly lower than D. palmarum 220–248 and D. bazi 215, 219, 224–232); Low numbers of subcaudals: 55–68 in males (lower than D. inornata 81–92, D. fasciata 71–90, D. medici 69–109; mostly lower than D. gansi 68–83, D. palmarum 68–77, D. parascabra 67–74, D. latericia 66–86), 44–52 in females (lower than D. inornata 69–84, D. fasciata 64–84, D. parascabra 57–64, D. palmarum 62–77, D. medici 61–90, D. latericia 59–72, D. gansi 59–73, D. arabica sp. nov. 53–61); Tail of moderate length: SVL/tail length 4.9–5.7 in males, 7.0–8.2 in females (in study area mostly greater than: D. medici 3.5–4.7 males, 4.5–5.7 females; D. fasciata 4.7–5.2 males, 4.8–6.4 females; see also Gans 1959); Nasal divided below the nostril (undivided in D. sahelensis and D. parascabra); Postoculars 2 on either side of head (often one, especially in western populations, of D. atra); Supralabials usually 7 (3rd and 4th enter orbit) on either side of head (usually 6[2,3] in D. loveridgei comb. nov.).

The new species is most similar to D. scabra, which usually has higher numbers of pattern cycles on the back—in surrounding countries/regions only two specimens, from southern Somalia (50–51 cycles) and a single snake from Sudan (54 cycles), have similarly low counts (Appendix 1).

Description of holotype The holotype (BM 49.2.2.2) is a subadult male with a total length of 202 mm, snout-vent length 172 mm, tail length 30 mm; ratio SVL: tail length  5.7, ratio total length: tail length 6.7.

Head small and barely distinct from the neck; snout rounded. Elongate nasal semi-divided (divided below the nostril), no loreal. Paired prefrontals much larger than paired internasals (individually and in combination). Inter-prefrontal sulcus moderately marked. Frontal large, slightly wider anteriorly, 1.51 times longer than broad, and fully pitted (State 4), slightly longer than each supraocular, and followed behind by a pair of large parietals each of which is larger than the frontal. There is a single preocular and two postoculars. Temporals 2+3. Seven supralabials, the 3rd and 4th bordering the eye. First pair of infralabials in contact behind the mental, followed by a pair of large and elongate sublinguals, and another pair of much smaller sublinguals in contact with the first ventral plate. Eyes moderate in size (with vertically elliptical pupils), their length subequal to the distance from tip of snout to anterior part of eye socket.

Dorsal scales strongly keeled, unicarinate, in 24 longitudinal rows at midbody, the third, fourth and fifth rows denticulate and (except for the 3rd row) obliquely placed. Vertebral scale row not enlarged. There are 198 smooth ventrals, the anal is entire and there are 60 pairs of smooth subcaudals. After preservation in ethanol the dorsal colouration is grey with 50 dark dorsal saddles between the nape and the base of the tail, separated by sharply defined white crossbars. Similarly dark, mostly elongate and wavy, lateral bars are linked to these crossbars on either side. Back of head with a distinct forward-pointing chevron; saddle on nape weakly forked posteriorly with a rectangular anterior prolongation. Venter immaculate cream-white apart from a few small brown spots on the outer edges of the ventrals.

INSERT FIG. 13

Variation in paratypes (including allotype, as indicated). The seven paratypes comprise four males and three females. The largest male (BM 49.2.2.5) measures 501 (408 + 83) mm, SVL/tail length ratio 4.9, total length/tail length ratio 5.9; one other male with measurements: BM 49.2.2.1 (188 + 34 mm, ratios 5.5 and 6.5). Largest female (MNHN 84–707) measures 545 (477 + 68 mm, ratios 7.0 and 8.0); other females: BM 49.2.2.4 (530 mm SVL), BM 49.2.2.3 (189 + 23 mm, ratios 8.2 and 9.2; allotype). The allotype and paratypes have similar scalation and general appearance to the holotype, except: Snout less rounded in BM 49.2.2.4. Inter-prefrontal sulcus weakly marked in BM 49.2.2.1 and 49.2.2.5. Frontal distinctly wider anteriorly in BM 49.2.2.4, BM 51.1.7.69, BM 1992.501 and MNHN 84–707; 1.25 to 1.53 times as long as wide; largely (not fully) pitted (State 3) in BM 51.1.7.69 and MNHN 84–707; and about as long as (not longer than) each supraocular in BM 49.2.2.5. Frontal about the same size as each of the parietals in BM 1992.501. One postocular (not two) on both sides of head in BM 49.2.2.4, and one on right side in BM 49.2.2.3 (allotype). Temporals 1+3 in BM 49.2.2.4 and 2+2 in BM 51.1.7.69. Six supralabials, the 2nd and 3rd bordering the eye, in BM 49.2.2.1. In BM 49.2.2.4 the pair of posterior sublinguals is largely separated by a more-or-less quadrangular supranumerary scale which is in contact with the first ventral plate. Length of eye almost equal to distance from tip of snout to anterior part of eye socket in BM 49.2.2.1 and BM 51.1.7.69, distinctly smaller than snout-eye distance in BM 49.2.2.4, 49.2.2.5 and MNHN 84–707. Midbody scale rows 23 in two specimens and 25 in two others (see Appendix 1); about 24 in BM 1992.501 (difficult to count) which has the third and fourth, and in some areas also the fifth, rows denticulate and oblique. Dorsal scales of the anal region distinctly denticulate in BM 49.2.2.4 and MNHN 84–707, feebly denticulate in BM 49.2.2.1, 49.2.2.3 (allotype) and 51.1.7.69. Ventrals 196-216 in males, 208–222 in females (see Appendix 1). Subcaudals 55–68 in males, 44–52 in females (tail truncated in BM 1992.501, male; and BM 49.2.2.4, female) (see Appendix 1). Dorsal pattern cycles 44–59 (see Appendix 1). In a flat skin (BM 51.1.7.69) the two halves of the vertebral saddles are not synchronised, forming an oblique pattern [Gans, 1959, Pl. VII, Fig. 1, erroneously given as ‘BM 51–1–7–59’]; and several dorsal saddles are oblique in BM 49.2.2.1. Dorsal colouration (excluding dark saddles and lateral bars) in preservative mostly rufous brown in BM 49.2.2.3 (allotype), a mixture of rufous brown and grey in BM 49.2.2.1 and 49.2.2.5, and pale brown in BM 49.2.2.4. Several lateral bars non-elongate and irregular in shape in BM 49.2.2.5. Pale crossbars between saddles in BM 49.2.2.4, 49.2.2.5, 51.1.7.69 and 1992.501 are thicker and less sharply defined than in the holotype [and allotype BM 49.2.2.3, as illustrated in Gans 1959, Pl. VI, Fig. 6]. Saddles are mostly rectangular and somewhat elongate in BM 49.2.2.5 and 1992.501, and in BM 49.2.2.4 they are rectangular at midbody but vary considerably in shape elsewhere, from slightly transversely enlarged and oblique, to oval and even ‘5I’-like (see Gans 1959). In BM 1992.501 several lateral bars are in contact with the saddles. Back of head with anterior part of chevron somewhat fragmented in BM 49.2.2.1 and MNHN 84–707, and bulbous in BM 1992.501; top of head somewhat faded in BM 49.2.2.3 (allotype); saddle on nape deeply forked posteriorly in BM 49.2.2.4 & 5, and always with a sharp anterior prolongation. Venter cream with large grey spots laterally, and smaller grey markings elsewhere, in BM 49.2.2.5; and whitish with 1–2 longitudinal rows of brown spots at the edges of the plates in BM 1992.501 and MNHN 84–707. No live specimens or photographic images of living snakes of this species are known to the authors.

INSERT FIG. 14

Etymology. The name of this new species is a patronym honouring Colonel R.H.R. Taylor, who collected about half of the type series while surveying the border between British Somaliland and Ethiopia in 1932–3.

Size. Largest male (BM 49.2.2.5 – Ogo, Somalia) 408 + 83 = 501 mm; largest female (MNHN 84–707 – Djibouti) 477 + 68 = 545 mm.

Habitat. Grasslands to Acacia–Commiphora bushland from 640 m to 1370 m (Figs 15 & 16). BM 1992.501 from 2.5 km WSW of Hargeisa was found on a limestone hillside (1275 m) with Vachellia etbaica Schweinfurth bushes.

INSERT FIGS 15 & 16

Distribution. Dasypeltis taylori sp. nov. has been recorded only from the arid interior of northern Somalia (Somaliland) (Figs 15 & 16) and in adjacent Djibouti (Fig. 3).

Mimicry. This species appears to mimic the saw-scaled viper Echis pyramidum Geoffroy Saint-Hilaire (see photographs in Dobiey & Vogel 2007 and Mazuch 2013). The racer Platyceps taylori (Parker), in the Borama area of Somaliland, has a dorsal pattern (see photograph in Mazuch 2013) remarkably similar to that of D. taylori sp. nov.

Biogeography. Dasypeltis taylori sp. nov. is separated from D. scabra populations in southern Somalia by the arid Ogaden plateau of eastern Ethiopia, an area from which no Dasypeltis have been recorded (e.g. Mazuch 2013). The highlands of east-central Ethiopia are occupied by D. atra, and those of north-western Ethiopia and southern Eritrea by D. abyssina. Dasypeltis taylori sp. nov. may therefore represent a relictual population of a once widespread D. scabra ancestor. Schätti & Ineich (2004) have discussed the radiation of the snake genus Platyceps Blyth, which is even more speciose than Dasypeltis in the Horn of Africa. Similar diversification occurs in the terrapin Pelomedusa subrufa species complex, where P. somalica Petzold et al. 2014 occurs in northern Somalia, P. gehafie (Rüppell) in Eritrea and eastern Sudan, and P. barbata Petzold et al. 2014 in Saudi Arabia and Yemen (Petzold et al. 2014). Typhlops calabressi Gans & Laurent has a very similar distribution to D. taylori sp. nov. (Broadley & Wallach 2009).

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it appears to be widespread and should be regarded as Least Concern.

Remarks. Scortecci (1930) recorded “D. scabra” from Assab on the Red Sea coast of southern Eritrea; this specimen may be D. taylori sp. nov., but as no further details are available and the specimen cannot be found, we desist from assigning it species status and exclude the record from our map.

Dasypeltis atra Sternfeld

Montane Egg-eater

Figs 17A–M & 18

Dasypeltis lineolata Peters, 1878: 206. Type locality: Kitui, Kenya, holotype ZMB 9224.

Dasypeltis palmarum (not Leach) Fischer, 1884a: 12 (Arusha, Tanzania).

Dasypeltis scabra var. F, Boulenger, 1894: 356 (part, Uganda); Angel, 1925: 34 (Nairobi).

Dasypeltis palmarum ? (not Leach) Stejneger, 1894: 733 (Mount Kilimanjaro, Tanzania).

Dasypeltis scabra (not Linnaeus) Boulenger, 1896: 553 (Shoa, Ethiopia); Sternfeld, 1908: 240, 332 (Lake Haramaya, Ethiopia); Roux, 1910: 99 (Bukoba, Tanzania); Boulenger, 1909: 303 (Bugala, Sesse Islands, Uganda); Peracca, 1909: 173 (Fort Portal, Ruwenzori Mtns); Boulenger, 1911: 5 (Bussu, Uganda) & 1912: 332 (Addis Ababa); Loveridge, 1916: 79 (part, Fort Hall); Lönnberg, 1922: 7 (part, east slopes of Mount Elgon, Kenya); Loveridge, 1929: 28 (part); Parker, 1932: 220 (Lake George, Lake Naivasha); Loveridge, 1936a: 34 (part, Harar, Lake Bunyonyi) & 1936b: 256 (Uganda & Kenya); Mertens, 1938: 8 (Ngorogoro, Aberdares); Gans, 1959: 141 (part); Spawls, 1978: 10; Largen & Rasmussen, 1993: 340 (part); Flemming & Bates, 1999: 423 (Ibaya Camp, Mkomazi Game Reserve); Wagner & Böhme 2007: 136 (USNM 49376, Kakamega Forest).

Dasypeltis scabra var. atra Sternfeld, 1912: 272. Type locality: ‘Urwald hinter den Randbergen am Nordwestufer des Tanganyika-Sees’ [= Jungle behind the border mountains on the north-west shore of Lake Tanganyika], Democratic Republic of the Congo; Gans, 1961: 73.

Dasypeltis scabra var. palmarum (not Leach) Angel, 1925: 34 (Nairobi).

Dasypeltis scaber (not Linnaeus) Loveridge, 1929: 28 (part); Parker, 1932: 220 (Lake George, Lake Naivasha); Loveridge, 1936a: 34 (part, Harar, Lake Bunyonyi) & 1936b: 256 (Uganda & Kenya); Uthmöller, 1937: 113 (part, Moshi, Uru).

Dasypeltis scaber scaber (not Linnaeus) Bogert, 1940: 85 (part, Ganame, Ethiopia).

Dasypeltis scaber palmarum (not Leach) Loveridge, 1942: 285 (Uganda, Rwanda) & 1946: 106.

Dasypeltis scabra palmarum (not Leach) Loveridge, 1955: 11 (Gilo, South Sudan).

Dasypeltis scaber ater Laurent, 1956: 261; Curry-Lindahl, 1956: 47 (Rwanda); Hinkel, 1986: 25 (Rwanda).

Dasypeltis scabra atra De Witte, 1962: 139; Bourgeois, 1968: 284.

Dasypeltis atra Gans, 1964: 279, 1971: 461 & 1974: 47; Laurent, 1965: 14; Pitman, 1974: 119; Hinkel & Fischer, 1988: 16 (Rwanda); Hinkel & Fischer, 1990: 137 (Nyungwe Forest, Rwanda); Broadley & Howell, 1991: 32; Fischer & Hinkel, 1992: 144 (Nyungwe and Birunga Forests, Rwanda); Hinkel, 1986: 25 (Rwanda); Meirte, 1992: 57; Largen & Rasmussen, 1993: 340; Hughes, 1997: 68; Broadley, 1998: xxxii; Gravlund et al., 1998: 53 (Mbizi Forest Reserve, Tanzania); Necas & Modry, 1998: 22; Gravlund, 2001: 101; Vonesh, 2001: App. A, Table 3 (Kibale National Park, Uganda); Behangana & Goodman, 2002: 66 (Uganda); Spawls et al., 2002: 414; ? Spawls, 2004: 17 (Keriyo; may be referable to plain phase D. abyssina); Spawls et al., 2006: 129; Largen & Parker, 2004: 25; Wagner & Böhme 2007: 136 (Kakamega Forest); Gartner & Greene, 2008: 368; Bates & Broadley, 2008a,b (abstracts); Broadley & Bates, 2009: 50; Bates, 2009: 3; Largen & Spawls, 2010: 494 (part, ‘Mulu’ = Keriyo, may be plain phase D. abyssina); Bates et al., 2012: 28; Branch, 2015: 67; Göthel, 2015a: 19 & 2015h: 90; Van de Loock & Bates 2016: 16 (Ngangao Forest, Kenya); Mannel & Kucharzewski, 2017: 43; Spawls, 2017: 31 (Maasai Mara).

Dasypeltis scaber var. lineolata (in error) Trape & Mané, 2006: 53.

Description. There are three basic colour morphs: dorsum black (‘2M’ of Gans 1959) with a grey venter (N = 89); dorsum brown (‘2B’ of Gans 1959) with a cream, yellowish or pale orange (e.g. Fig. 17C) venter, usually with grey stippling at the edges (N = 90); dorsum grey or tan brown, decorated with closely-spaced, mostly narrow, transversely enlarged dark saddles, sometimes rhombic, in common or occasional contact with the equally dark lateral bars, or occasionally separated in a more-or-less ‘5N’ pattern similar to that of D. scabra (‘2Bx’ of Gans 1959) and venter cream, usually with grey stippling at the edges (N = 128). In patterned specimens the dorsal saddles (62–130, mean 91.8, N = 51) are dark brown, olive or reddish; and the head may be largely grey to bluish-grey, and/or with few dark markings unlike the boldly dark-light patterned head of D. scabra (see images in Broadley & Bates 2009; Spawls et al. 2002; Göthel 2015e,h; Fig. 17F, I–K). For example, PEM R17336 (Tanzania; Fig. 17K) and BM 03.12.2.22 (Uganda) have grey bodies with reddish saddles/crossbars; but the latter specimen also has a broad red chevron on the top of the head. Saddles and lateral bars may vary in colour between specimens even at the same locality – e.g. at Kitengela in Kenya they may be olive (Fig. 17F) or brown. The ‘lineolata’ phase (Gans, 1959, Pl. VI, Fig. 8; Fig. 17D) shows the extreme reduction of the pattern where most of the back is dark with only narrow, often interrupted or incomplete, pale transverse bars. Some specimens with low numbers of bands are somewhat similar to D. confusa, but lack the large well-defined saddles. Dorsally brown (‘2B’) and black (‘2M’) phase specimens are found together at a few localities such as Bourka in Ethiopia, Bussu, Kampala and Sipi at Mt Elgon in Uganda, and Minziro Forest Reserve in Tanzania; brown and patterned (‘2Bx’) phases are both found at Harar in Ethiopia, Mushongeru in Uganda, and Nairobi and Kacheliba in Kenya; black and patterned specimens both occur at Nakuru and North Nandi Forest in Kenya; and at Muko in Uganda all three colour phases occur together (Appendix 1). All specimens from South Sudan are melanistic (i.e. ‘2M’), except MCZ 53569 which is brown.

Preoculars usually one on either side of head (96.5% of specimens), but one on one side and two on the other in 1.8%, and two on either side in 1.8% (N = 227); postoculars one on either side of head (e.g. Fig. 18) in 58.3% of specimens, one on one side and two on the other in 6.8%, and two on either side in 34.9% (N = 307) (usually one in western populations – especially in Uganda, Rwanda and Burundi, e.g. 93.0% of Rwanda/Burundi snakes [N = 43] had a single postocular on either side; by contrast 77.4% of Ethiopian snakes [N = 31] had two postoculars on either side); frontal shield for the most part smooth, usually with 1–2 rows of shallow pits at the edges (States 1–2; 95.3% of specimens), but occasionally fully smooth (State 0; 3.3% of specimens) or extensively pitted (States 3–4; 1.4% of specimens: NMW 15299, PEM 16778, ZMUC 6122) (N = 213); inter-prefrontal suture usually weakly marked (87.5% of specimens), but moderately marked in 11.5%, and distinctly depressed in 1.0% (BM 1976.2265, MSNG 30412) (N = 200); supralabials 12–16, usually 14 (both sides of head), usually 7 [3rd and 4th entering orbit] on either side (84.7%), but variations are: 6[2,3] on either side of head (e.g. MRAC 18557) in 1.4% of specimens, 6[3,4] on either side (e.g. BM 51.1.3.74) in 5.2% (including MSNG 52410 which had left side of head damaged), 6[3,4] on left and 6[2,3] on right in ZFMK 29731, 6[2,3] on one side (e.g. CAS 122281) in 2.8%, 6[3,4] on one side (e.g. MCZ 40578) in 1.4%, 7[2,3,4] on right in ZFMK 16237, 7[2,3,4] on both sides in NMK 3182, 8[3,4] on one side (e.g. BM 1976.2265) in 1.0%, 8[3,4,5] on one side (e.g. USNM 40897) in 1.7%, and 8[3,4,5] on both sides (e.g. MRAC 15440) in 0.7% (N = 287); anterior temporals usually two on either side of head (93.9%), occasionally one (2.5%), 2 left:1 right or 1:2 (1.2%), 2:3 or 3:2 (1.2%), or three (1.2%) (N = 244); posterior temporals usually three on either side of head (52.7%), often two (18.3%) or 2:3 or 3:2 (14.1%), occasionally 3:4 or 4:3 (8.3%), four (4.1%) or five (0.8%), and exceptionally 4:5 (ZFMK 29733), 3:5 (BM 1976.2265), 5:3 (BM 34.12.15.590) or 2:4 (LIV 1960.67.1) (N = 241); temporal formula on either side of head usually 2+3, often 2+2 or 2+4; midbody scale rows 22–28 (mean 24.1 ±1.25, N = 226) [28 in NMW 26879/3 only]; ventrals 199–232 in males (N = 109), 211–256 in females (N = 167); subcaudals 50–71 in males (N = 103), 45–64 in females (N = 151). One unsexed specimen from Bujumbura, Burundi (MRAC 82–16–R4, probably a male) had 218 ventrals and as many as 75 subcaudals. The brown (‘2B’ of Gans 1959) female (MSNM 7202) from Circao (northern-most record for this species) has 232 ventrals, a much lower count than that of female D. abyssina (241–264) which also occur as brown or uniform plain phase morphs (see below); and brown D. atra in Ethiopia have lower ventral + subcaudal counts than plain phase D. abyssina (both sexes, 269–291 [N = 5] versus 294–323 [N = 7] respectively). Ratio SVL: tail length 4.2–6.7 in males (N = 97), 5.8–8.3 in females (N = 143); total length: tail length 5.2–7.7 in males (N = 97), 6.8–9.3 in females (N = 143).

INSERT FIGS 17 & 18

Size. Largest male (MSNG 30413b – Bussu, Uganda) 611 + 98 = 709 mm; largest female (NMK 2583 – North Nandi Forest, Kenya) 890 + 115 = 1005 mm.

Habitat. Found mainly in high elevation savannah and forest, usually above 1500 m.

Distribution. Kivu Region and vicinity of Lake Albert in eastern Democratic Republic of the Congo (Laurent 1956; Gans 1959), Rwanda, Burundi, south to Mbizi Forest in western Tanzania and east to the extreme northern parts of that country, Uganda, south-western, central and southern Kenya, extreme southern part of South Sudan, south-western and central Ethiopia, with the northern-most record at Circao in north-western Ethiopia (Fig. 3).

Sympatry. Sympatric with D. medici at Nyambene Hills (Kenya) and Moshi (Tanzania); sympatric with D. confusa 10 km S of Juba (South Sudan), at Busingiro and Gulu (Uganda) and Kacheliba (Kenya); sympatric with D. scabra at Aware Melca and Lake Abiata (Ethiopia), Mulema (Uganda), Lake Naivasha and Mount Kulal (Kenya), Astrida (Rwanda) and Minziro Forest Reserve (Tanzania); sympatric with both D. medici and D. scabra at Mount Kilimanjaro (Appendix 1).

Localities. ETHIOPIA. ‘Abessinien’, NMW 9964/2; Addis Ababa [NE 0938B2] MSNG 30412, and Fig. 17E; Aware Melca [NE 0939B2] BM 16.6.24.13; Bedele [NE 0836A4] ZFMK 57537; Bedele, 4.5 km SW of [NE 0836A4] melanistic, photographic records; Bourka [NE 0941A4] MNHN 05.208–10; Circao, Dambria [NE 1237A4] MSNM 7202; El Dire, Tertale [NE 0437C3] MSNM 2975; Fitu, 135 km ESE of Negele [NE 0540B1] ‘2Bx’ pattern, photographic records; Ganame [NE 0841D3] AMNH 20345; Gemu, Gofa [NE 0636B] BM 1964.1068; Genale to Chamo [NE 0837D3] ZFMK 16297; Ghimbi, Wollega [NE 0935B2] BM 1973.3175; Ginir [NE 0740B1] NMW 26879/2; Gololtscha [NE 0840A1] ZFMK 5535; Gumaydo [NE 0537D1] NMW 26879/3; Harar [NE 0942A3] FMNH 4012, MCZ 34923, NMW 9964/1; Jigjiga [NE 0942B4] ZFMK 84993; Kebre Mengist to Negele [NE 0539C3] BM 1975.2144; Keriyo [NE 0938B3] (Fig. 17A); near Kersa Dek, 14 km SE of Neghelle [NE 0539B1] ‘2Bx’ pattern, photographic record; Kounni [NE0940B2] MNHN 05.211; Laga Hardine [NE 0840C4] MNHN 05.205–7; Lake Abiata [NE 0738D1] ZFMK 16236; Lake Alemeya [NE 0942A3] ZMB 27463; Lake Shala [NE 0738B3] ZFMK 16237; Let Marefia [NE 0939D2] ZMB 10286; Neghelle [0539B3] (Fig. 9M); Shek Husen, 4.4 km WSW of [NE0740D1] brown dorsally with cream patches, photographic record; Shoa, MSNG 29047a; Wondo [NE 0638C2] NMW 15299. SOUTH SUDAN. Gilo, Imatong Mtns [NE 0432B2] BM 1978.379–80, FMNH 62215, MCZ 53406–8, 53569, ZFMK 29564, 29719, 34532, 59567; 10 km S of Juba [NE 0431D3] ZFMK 29567; Katire [NE 0432B2] ZFMK 26005; Talanga Forest [NE 0432B1] ZFMK 29731–3. UGANDA. Bihunga [SE 0129B4] LIV 1960.67.1, 1960.50.32; Bitereko [SE 0030B3] LIV 1960.115; Budongo Forest [NE 0131D3] USNM 63481; Bufindi, Kigezi [NE 0129B4] BM 51.1.3.68 & 79, NMZB 217; Bukakata [SE 0031A3] BM 51.1.3.80; Busingiro, Budango [NE 0131D1] BM 1951.1.5.24; Bussu [NE 0033A3] BM 11.7.8.14, MSNG 30413a, b, 52410; Butundiga, Mount Elgon [NE 0134A2] MCZ 40574; Bwamba Forest [NE 0030C3] LACM 38696, 46344; Chagwe, Mabira Forest [NE 0032D2] BM 08.10.20.14–5; Entebbe [NE 0032A2] BM 51.1.3.74; Gulu [NE 0232C4] BM 51.1.3.77; Fort Portal [NE 0030C2] CAS–SU21709–10; Kabingo, Bunyaruguru, BM 1961.1803; Kagabagara [SE 0030D3] LIV 1962.9.28; Kagando [NE 0029B2] LIV 1960.67.5; Kalinzu Forest [NE 0030A3] LACM 60705; Kampala [NE 0032A2] BM 36.7.3.32, 1984.882; Katunguru [SE 0030A1] NMZB 103; Katwe [SE 0029B2] MRAC 9997; Kayonza, Kinkizi [NE 0029D3] LACM 35090, LIV 1960.115.4, 1962.9.27, 1980.150.31, NMZB–UM 2550; Kazinga Channel [SE 0030A1] NMK 404; Kichwamba [SE 0030A1] LIV 1962.9.14; Kinani [SE 0030B4] LIV 1962.9.22; Kyamahunga, Igara [SE 0030A3] BM 51.1.3.75, 1961.1805, NMZB 2549; Kyamugorani [SE 0030B3] LIV 1962.9.4; Lake Bunyonyi [SE 0129B4] FMNH 9913–4, MCZ 34922; Lake Edward [SE 0029B] NMZB 219; Lake George [NE 0030A] BM 32.5.2.99; Lake Nabugabo [SE 0032A3] BM 51.1.3.78; Likiri [SE 0030A2] LIV 1980.150.27; Maozi-Isingiro [SE 0030D4] BM 03.12.2.23; Mihunga, Ruwenzori [NE 0030A3] MCZ 48359; Mityana [NE 0032A3] NMK 3291; Morulinga, 24 km SW of [NE 0234A4] BM 1954.1.12.37; Mubango, Mabira Forest [NE 0033A3] BM 34.12.15.592; Muko, Kigezi [SE 0129B2] BM 34.12.15.585–91, 593–4; Mulema [SE 0031D1] BM 03–12–2–22; Mushongero, Lake Mutanda [SE 0129B1] MCZ 48362–3; Nsika [SE 0030A4] LIV 1980.150.26; Nyabushozi [SE 0030B4] LIV 1962.9.23; Nyakabende [SE 0129B4] MCZ 48360–1, UMMZ 88532/1; Nyamuhumia [SE 0030C1] LIV 1959.81.2; Nyansimbo [SE 0030B1] LIV 1962.50.33, 1980.150.33; Nyarusiza, Bufumbira [SE 0129B3] BM 1976.1849, LIV 1980.150.29–30, NMK 1496, NMZB–UM 2553; Rucho, Ankole, MCZ 54744; Ruhama [SE 0030C4] LIV 1960.67.3; Sango Bay Forest [SE 0031D3] LACM 39242; Sesse Islands, Lake Victoria [SE 0032A3] MSNG 27755a, 40764; Sipi, Mount Elgon [NE 0134A4] MCZ 40564–73. KENYA. No locality, NMK 348, 463, 1962, UMMZ 96274; Athi Plain, NMW 9964/20; Athi-Kapiti Plains, NMK 3129; Athi River [SE 0136B4] NMK 1453, 3085; Butere, Kakamega [NE 0034B1] NMK 2323 (old no. 306); Chemelil, Kisumu [SE 0035A1] CAS 148034–5; Chuka, Meru [SE 0037B3] CAS 122281, and Fig. 17G – ‘2Bx’ pattern; Chyulu Hills [SE 0037D2] NMK 406; Elgonyi, Mount Elgon [NE 0134B1] MCZ 40575; Emakoko, Nairobi National Park [SE 0136B4] photographic record (dorsum grey with brown saddles [similar to Fig. 9F] and two postoculars on left side of head); Embakasi, Nairobi [SE 0136B4] NMK 2849; Hopcroft Ranch, Athi River [SE 0137C1] NMK 3131; Juja Farm [SE 0137A1] USNM 40897; Kabartonjo, Kamasia Mtns [NE 0035D2] CAS 111767; Kabarnet, Baringo [NE 0035D2] BM 1957.1.11.42, NMK 3329; Kabete, Kiambu [SE 0136B3] NMK 479; Kacheliba [NE 0135A3] NMK 486, 488; Kaimosi [NE 0034B2] MCZ 40576–8, 18205, UMMZ 61224, USNM 49078; Kairuni, Nyambeni Mtns [NE 0037B2] LACM 93378–9; Kakamega [NE 0034B4] BM 1959.1.5.58, CAS 142248, NMK 482, 2715, USNM 49376; Kakamega Forest [NE 0034B4] Fig. 16L, ‘2Bx’ pattern; Karen, Nairobi [SE 0136B3] NMK 2503, 3007; Kericho [SE 0035A4] NMK 1260; Kiambu [SE 0136B2] NMK 481; Kiboko Ranch Station [SE 0237B1] NMK 2952 (4); Kijabe [SE 0036D3] NMK 2130, and top photograph of melanistic specimen in Spawls et al. 2002: 415; Kiganjo, Nyeri [SE 0036D4] NMK 2862; Kiserien, Baringo [SE 0136B3] NMK 3123; Kitale [NE 0135A1] NMK 3051, ZMUC 6101; Kitengela [SE 0136D2] Fig. 17F and another photograph, both ‘2Bx’ patterned; Kitengela, Athi River [SE 0136D2] NMK 3413; Kitui [SE 0138A3] ZMB 9224 (holotype of D. lineolata); Lake Baringo [NE 0036C2] NMK 1791, 4174; Lake Naivasha [SE 0036C2] BM 93.11.21.72, 1932.5.2.100; Limuru [SE 0136B1] Fig. 17B, melanistic; Lorgorien, Trans-Mara, NMK 3109; Lumbwa, Pen-y-Bryn [SE 0035A4] CM 2278–9, 2281; 13 km SSW of Maralal [NE 0036D3] LACM 66076; Mbunyi [SE 0137B4] USNM 62907 (Gans’ 1959 locality of ‘Morogoro’ for this specimen is in error, see Loveridge 1929); Mount Elgon, NMK 657–9, 3152; Mount Kenya [SE 0037A2] USNM 40990; Mount Kulal [NE 0236D2] LACM 66395; Muranga [Fort Hall] [SE 0037C1] BM 08.9.17.4; Mweiga [SE 0036B4] LACM 50631; Mwino, Sigor [NE 0135A4] NMK 1965; Nairobi [SE 0136B4] NMK 403, 2486, 3339, 3548; Naitolia (6000 ft = 1829 m a.s.l.) [SE 0036A4] BM 01.1.3.9; Nakuru [SE 0036A3] NMK 484 and 1453, and photographs of ‘2Bx’ patterned and melanistic specimens; Nandi [NE 0035A1] BM 06.7.6.7; Nandi Hills [NE 0035A] NMK 1158; Nanyuki, Laikipia [NE 0037A1] NMK 3490, 3512, 3557; Nbabibi, Naivasha [SE 0036C2] NMK 1960; Ngangao Forest, Taita Hills [SE 0338A4] (photograph of barred/patterned ‘2Bx’ specimen: Van de Loock & Bates 2016, Fig. 2); Ngong [SE 0136B3] BM 1960.1.6.40; North Nandi Forest, Chemesia [NE 0034B4] NMK 2576, 2583, 2590; Nyambene Hills [NE 0037B] NMK 1487, 1489, 1699; South Nandi Forest [NE 0035A1] NMK 4176; Subukia [SE 0036A1] FMNH 58337; Timbilil, Mau Forest [SE 0035A4] NMK 3182; Uganda Railway, BM 96.3.27.18. RWANDA. Astrida [SE 0229D1] MRAC 18554–8; Bugarama [SE 0229C1] MRAC 86–01–R–10 & 11; Bugesera [SE 0230A1] ZFMK 61616; Bulimbi [SE 0130C1] MRAC 18577; Cyamudongo [SE 0228D2] ZFMK 61617; between Gikongoro and Nyungwe Forest Reserve [SE0229C2] PEM 5623; Kamohora District, MRAC 14329; Kibungo [SE 0230B1] ZFMK 55105; Kisenyi [SE 0129C1] MCZ 48364, MRAC 2723, 2725, 2757, 3821, 5871–2, 9773, ZMB 20458, 27156, 27539; Lake Gando [SE 0129C2] MRAC 14333, 14341; Mwogo River near Butare [0230A1] MRAC 74–23–R–22; Mutura, Mount Virunga [SE 0129B2] MRAC 7509; Nyanza [SE 0429B3] MRAC 15477; Ruhengeri [SE 0129D1] IRSNB 2657; Wankeri [SE 0129D1] MRAC 14331, 14340: Rwankuba [SE 0129C2] MRAC 18259; Shangugu [SE 0228B4] MRAC 4074–5; Tare, Astrida [SE 0229D1] MRAC 18560–2. BURUNDI. Bujumbura [SE 0329A4] MRAC 82–16–R3 & 4; Kamagaba Marsh, MRAC 18583; Kibira Forest [SE 0229C4] (Fig. 17C); Makamba [SE 0429B2] MRAC 18584; Rumonge [SE 0329C4] MRAC 18585. TANZANIA. Bukoba [SE 0131B4] ZMB 16749–50; Ibaya Camp, Mkomazi Game Reserve [SE 0337D4] BM 1994.403, NMBO 7375; Kasoge, Mahale Peninsula [SE 0629B2] BM 1970.2177; Klein’s Camp, NE Serengeti National Park [SE 0135C3] PEM 16778, 17329, 17336, 17941; Masiko ya Tembo, vicinity of Sanya Chini [SE 0337A3] (Fig. 17J); Mbizi Forest Reserve [SE 0731D3] ZMUC 6122; Minziro Forest Reserve [SE 0131B1] ZMUC 6138–41; Moru area, Serengeti National Park [SE 0234D1] photograph of ‘2Bx’ marked specimen; Moshi [SE 0337A4] ZSM 81/1936 (3); Mount Kilimanjaro [SE 0337A] ZMB 17039 (2); Oldeani [SE 0335B3] photograph of melanistic ‘2M’ specimen; Sitete, Mahale Peninsula [SE 0629B2] BM 1970.2178.

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it is widespread and abundant and should be regarded as Least Concern.

Remarks. Dasypeltis scabra var. atra was described by Sternfeld (1912) on the basis of three ‘completely black’ specimens, with single postoculars, from the forest behind the border mountains on the north-west shore of Lake Tanganyika, Democratic Republic of the Congo. Photographs of the three syntypes housed at Museum für Naturkunde, Berlin were provided by F. Tillack (pers. comm, 10 July 2017). ZMB 27761 is medium brown above and light brown below (may have faded); ZMB 22516A and ZMB 22516B are dark brown to black above and grey below. It can be confirmed that all three specimens have a single postocular on either side of the head, and the supraocular extends over the posterior corner of the eye.

Since its description by Peters in 1878, Dasypeltis lineolata was used as a valid name by Fischer (1884b) in reference to a snake (247 ventrals, 69 subcaudals) from Ssibange in Gabon, probably referable to D. fasciata or possibly D. palmarum, and later by Stejneger (1894) who concluded that it was a ‘colour variety’ (possibly implying a subspecies) of Dasypeltis abyssina. The only other valid usage of the name was by Saleh & Sarhan (2016). While referring to available names, Trape & Mané (2006) erroneously used the combination Dasypeltis scaber var. lineolata. The samples of ‘D. cf. lineolata’ from Burundi used in the 16S genetic analysis of Trape et al. (2012) are referable to either D. atra or D. scabra. The name D. atra (or D. scaber ater or D. scabra atra, as indicated), the junior synonym, has been used in the following 42 publications (25 required), by 34 authors/sets of authors (10 required), during the preceding 50 years (1968–2017), and for a period covering more than 10 years (see Article 23.9.1.2 of the ICZN 1999): Bourgeois (1968, as D. scaber ater and D. scabra atra), Gans (1971, 1974), Pitman (1974), De Witte (1975), Hinkel (1986, as D. scaber ater), Hinkel & Fischer (1988, 1990), Fischer & Hinkel (1992), Broadley & Howell (1991), Meirte (1992), Largen & Rasmussen (1993), Hughes (1997), Broadley (1998), Gravlund et al. (1998), Neças & Modry (1998a,b), Gravlund (2001), Spawls et al. (2002), Largen & Parker (2004), Spawls (2004), Branch (2015), Spawls et al. (2006), Trape & Mané (2006), Wagner & Böhme (2007), Gartner & Greene (2008), Bates (2009), Broadley & Bates (2009), Largen & Spawls (2010), Bates et al. (2012), Trape et al. (2012), Göthel (2015a–h), Van de Loock & Bates (2016), Saleh & Sarhan (2016), and Mannel & Kucharzewski (2017), thus satisfying the requirements of Article 23.9.1.2 of the ICZN (1999). However, Saleh & Sarwan’s use of the name D. lineolata means that the provisions of Article 23.9.1.1 (i.e. senior synonym not used as a valid name after 1899; ICZN 1999) are not met. In order to maintain nomenclatural stability the ICZN has been requested (Bates 2017, submitted; Case 3750) to suppress the older name D. lineolata by taking action according to Article 23.9.3 by use of its plenary power (Article 81; ICZN 1999). We maintain prevailing usage of the name D. atra according to Article 82.1 (ICZN 1999).

Dasypeltis fasciata A. Smith

Western Forest Egg-eater

Fig. 19A–C

Dasypeltis fasciatus A. Smith, 1849 (footnote to text accompanying Plate 73). Type locality: Sierra Leone.

Dasypeltis scaber fasciatus Loveridge, 1942: 284 (Bundibugyo, Uganda).

Dasypeltis fasciata Loveridge, 1957: 289 (check-list); Gans, 1959: 137 & 1961: 73; Pitman, 1974: 121; Spawls et al., 2002: 416; Spawls et al., 2006: 130; Branch, 2015: 68; Göthel, 2015a: 19 & 2015h: 81. 

Description. Dorsum grey or pale brown with numerous broad dark saddle bands (108–141, N = 5) which are often ill-defined (preserved specimens, e.g. Fig. 19A) and usually arranged in a manner similar to the ‘5N’ pattern, with saddles alternating with broad lateral bars; dorsal body pattern expressed as chequerboard on skin between scales and on lower edges of these (see Gans 1959, Pl. III); top of head vermiculate; venter cream to pale grey, usually with grey stippling at the edges, but sometimes extensively stippled (e.g. TCWC 23839, which also has the throat and anterior part of the belly pale yellow, Fig. 19B).

Preoculars usually one on either side of head (83.3% of specimens), but two on either side in MCZ 48357 and UMMZ 88531 (N = 12); postoculars usually two on either side of head, but two on left side and one on right side of MCZ 48358B, and one on both sides in LACM 46341 (N = 12); frontal shield for the most part smooth, usually with 1–2 rows of shallow pits at the edges (States 1–2), but more extensively pitted (State 3) in LACM 38697 (N = 11); inter-prefrontal suture weakly marked (N = 11); supralabials 13–14, usually 14 (both sides of head), usually 7 [3rd and 4th enter orbit] on either side of head, but 6 [2,3] on right side of head in LACM 46346 and MCZ 48357 (N = 12); anterior temporals usually two on either side of head (75.0%), occasionally 2 left: 3 right or 3:2 (16.7%), and three in LACM 46341 (N = 12); posterior temporals usually three on either side of head (58.3%), occasionally 3:4 or 4:3 (25.0%), two in LACM 39243, and four in LACM 46343 (N = 12); temporal formula on either side of head usually 2+3, occasionally 2+2, 2+4, 3+3 and 3+4; midbody scale rows 22–25 (mean 23.5 ±0.80, N = 12); ventrals 212 (LACM 39243), 229–230 in males (N = 3), 237–248 in females (N = 9); subcaudals 59 (LACM 39243), 71–76 in males (N = 3), 64–76 in females (N = 9). Ratio SVL: tail length 4.7–5.2 in males (N = 3), 4.8–6.4 in females (N = 9); total length: tail length 5.7–6.2 in males (N = 3), 5.8–7.4 in females (N = 9).

INSERT FIG. 19

Size. Largest male (LACM 38697 – Ntandi, Bwamba Forest, Uganda) 525 + 111 = 636 mm; largest female (TCWC 23839 – Ntandi, Bwamba Forest, Uganda) 611+ 108 = 719 mm.

Habitat. Lowland forest at elevations of about 1000 m to 1147 m (Katera).

Distribution. Forested areas of west and central Africa, from Guinea to Togo, and from south-eastern Nigeria south to north-western Angola, and east through southern Central African Republic and much of the Democratic Republic of the Congo (Gans 1959, 1964; Trape & Mané 2012), reaching its eastern limit in western and southern Uganda (Fig. 3).

Localities. UGANDA. Bundibugyo [NE 0030C] MCZ 48537, 48538a,b; UMMZ 88531; Bwamba Forest, Ntandi [NE 0030C3] LACM 38697, 46341–3, 48345–6, TCWC 23839; Katera, Sango Bay Forest [SE 0031D3] LACM 39243.

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it appears to be widespread and abundant and should be regarded as Least Concern.

Dasypeltis abyssina (Duméril, Bibron & Duméril)

Ethiopian Egg-eater

Figs 20A–G

Rachiodon abyssinus Duméril, Bibron & Duméril, 1854: Vol. 7: 496, Pl. 81, Figs 1–2. Type locality: Abyssinia, proposed lectotype MNHN 6567 (see below); Duméril, 1860: 198.

Rachiodon scaber var. abissinicus (lapsus) Jan, 1863: 106 (Ethiopia).

Dasypeltis abyssinicus (lapsus) Blanford, 1870: 457 (Senafe, Eritrea); Zavattari, 1930: 193.

Dasypeltis abyssina Fischer, 1884b: 48; Trape & Mané, 2006: 53; Largen & Spawls, 2010: 490; Göthel, 2015a: 19, 2015h: 92.

Dasypeltis scabra (not Linnaeus) Boulenger, 1893: 354 (var. D, Abyssinia); Calabresi, 1925: 107 (Gondar); Zavattari, 1930: 193 (Senafe); Gans, 1959: 141 (part, Eritrea, Gheleb; Pl. VI, Figs 3 &4); Largen & Rasmussen, 1993: 340 (part, 140 km NE of Gondar on road to Asmara); Spawls, 2004: 17 (Keriyo); Largen & Spawls, 2010: 488 + 490 (part, e.g. ‘Mulu’ = Keriyo).

Dasypeltis scaber (not Linnaeus) Loveridge, 1936: 34 (part, Gondar).

Lectotype. Dumeril et al. (1854) described Rachiodon abyssinus on the basis of two syntypes in the Muséum national d’Histoire naturelle in Paris, one from “Abyssinia” (= Ethiopia) and another collected during an expedition to the sources of the White Nile under direction of M. Arnaud (labelled “Nil Blanc” = White Nile, perhaps above Khartoum in Sudan). Although their description is compound (e.g. ventrals are given as 222–260, clearly referring to MNHN 7227 and MNHN 6567 respectively – see above), the information on colour, which refers to a yellow base colour which exaggerates the darker dorsal markings, appears to be specifically in reference to the illustrated snake (MNHN 6567). Trape & Mané (2006) re-instated D. abyssina which has vivid ‘5N’ dorsal colouration, heavily black-bordered ventrals and a high ventral count of 260. They were of the opinion that the two syntypes were probably referable to different species, and designated MNHN 6567 (female), illustrated in plate 81 in the description (see also Fig. 20A–B), as ‘holotype’ of R. abyssinus. Although we agree that the two specimens do in fact represent different species, the action by these authors is not in accordance with the International Code on Zoological Nomenclature (ICZN 1999) as a syntype cannot be designated as a holotype, only as a lectotype or paralectotype.

MNHN 6567 agrees closely with the heavily-marked belly illustrated in plate 81 of Duméril et al. (1854), but the dorsal pattern is dissimilar (Fig. 20A). The illustration depicts a snake with a pale vertebral stripe, flanked on either side by distinct but disjointed markings. As can be seen in the photograph (Fig. 20A) of this specimen, while the centre of the back has a pale background, this is interrupted frequently by the often oblique dark markings or saddles which are continuous across the centre of the back. There is also a general separation of the saddles from the lateral bars, even though these may be in occasional contact. An indication of the cause of the poor condition of the specimen (head and neck damaged) is provided by plate 81 in which the snake is illustrated, together with a section of its skeleton comprising 13 pairs of ribs and their associated vertebrae, and a section of the vertebrae from the neck region, indicating considerable mutilation of the body. It was therefore not possible to perform a count of the ventral plates, and as an appreciable portion of the tail tip was missing, no subcaudal count was possible either. Nevertheless, there seems to be no doubt that the snake illustrated in plate 81 and described by Duméril et al. (1854) is the specimen represented by MNHN 6567.

We take this opportunity to designate MNHN 6567 (female, no hemipenal spines in tail) as lectotype of Rachiodon abyssinus. In accordance with Recommendation 74C of the 1999 Code we hereby list the following data pertaining to the lectotype, as based on a recent examination of the damaged specimen (Fig. 20A–B) in Paris by the first author: Type locality: “Abyssinia” (= Ethiopia). Snout-vent length about 572 mm; about 86 pattern cycles from nape to above vent. Dorsal colour pattern consists mostly of oblique dark saddles, usually wider than long, and often confluent or forming zigzags on the back and tail; lateral bars (usually extending onto the edges of the ventral plates) somewhat irregular, usually broader above than below, and often in contact with the pale interspaces between saddles, sometimes in narrow contact with the edges of the saddles, but never forming a distinct or broadly linked pattern. Ventral scales with extensive brown markings on their anterior half, consisting of a large rounded cone on either side joined to a smaller cone in the centre. Midbody scale rows 25; lateral dorsal scale rows 3 to 6 are keeled and serrated; nasal scale divided between nostril and first upper labial.

Paralectotype. Duméril et al.’s (1854) other syntype of R. abyssinus, MNHN 7227 (female, no hemipenal spines in tail) from ‘Nil Blanc’, now the paralectotype, is referable to D. scabra. It measures 673 + 90 mm and has a grey dorsum with ‘5N’ pattern (55 pattern cycles) consisting of brown oval saddles with black borders, and lateral bars—situated adjacent to the pale inter-saddle bars—also brown with black borders (Fig. 6); venter cream with dark speckling and two longitudinal rows of large spots at the edges; preoculars 1 on either side of head; postoculars 2 on either side; frontal shield with marginal pitting (State 2); inter-prefrontal suture moderately developed; ventrals 226; subcaudals 52; midbody scale rows 25, rows 3–5 keeled and serrated; supralabials 7 (3rd and 4th enter orbit) on either side; nasal scale divided below the nostril; temporals 2+3.

Diagnosis. Assigned to the genus Dasypeltis on account of its slender form, possession of (usually 3–4) rows of reduced, oblique, keeled and serrated lateral scales (little or no serration in D. inornata) and head barely distinct from the neck (moderately distinct in D. fasciata). Occurs as both patterned and uniform (brown or grey above, immaculate cream below) phases. Patterned individuals are distinguished from other species by their unique dorsal and ventral colour patterns, dorsally consisting mostly of oblique dark saddles, usually wider than long, and often confluent or forming zigzags on the back and tail; lateral bars much-elongated, usually broader above than below, and typically extending onto the edges of the ventral plates, irregular in shape and often in contact with the pale interspaces between saddles, sometimes in narrow contact with the edges of the saddles; pattern cycles 65–98 (≤62 in D. crucifera sp. nov., D. taylori sp. nov. and D. bazi; D. inornatus always uniform brown above; D. gansi uniform or weakly marked above); and ventral scales with extensive black, grey or brown markings on their anterior half, consisting of a large rounded cone on either side joined to a smaller rounded cone in the centre (Gans 1959, Pl. 9, Fig. 7; Fig. 20B; some D. scabra have heavy dark markings on the venter, but never as extensive or in the same pattern as described above); in live specimens the labials and interspaces between saddles are cream-yellow (Fig. 20C–F) as illustrated for one of the two syntypes (MNHN 6567, see discussion below) which, in addition, apparently also had some yellow on the sides of the head (Duméril et al. 1854, Pl. 81): within the genus this combination of dark markings with extensive cream-yellow or yellow also occurs, occasionally, only in southern populations of D. scabra (e.g. Peters 1864; Göthel 2015). In addition to its colour pattern as described above, this species can also be distinguished by the following combination of characters: High ventral scale counts of 226–247 in males (higher or mostly higher than D. scabra 180–226, D. taylori sp. nov. 196–216, D. confusa 199–227, D. sahelensis 207–221, D. inornata 208–225, D. parascabra 210–223), 241–271 in females (higher or mostly higher than D. scabra 198–249, D. taylori sp. nov. 208–222, D. sahelensis 212–237, D. confusa 212–242, D. bazi 215–232, D. inornata 219–237, D. parascabra 224-237); Subcaudals 57–68 in males (mostly higher than D. loveridgei comb. nov. 47–57; lower or mostly lower than D. inornata 81–92, D. gansi 68–83, D. palmarum 68–77, D. parascabra 67–74, D. latericia 66–86, D. fasciata 59, 71–91, D. medici 69–109), 49–65 in females (mostly lower than D. inornata 69–84, D. fasciata 64–84, D. medici 61–90); Inter-prefrontal sulcus usually weakly marked (weakly to moderately marked in D. bazi, usually moderately marked in D. taylori sp. nov., moderate to well marked in D. arabica sp. nov., and variable in D. scabra [often well marked in the south of its range]); Frontal marginally pitted, usually State 1 or 2 (extensively pitted, States 3 & 4, in D. medici, D. taylori sp. nov. and many populations of D. scabra especially in the south of its range); Tail of moderate length: SVL/tail length 5.3–6.1 in males, 5.5–8.1 in females (in study area mostly greater than: D. medici 3.5–4.7 males, 4.5–5.7 females; D. fasciata 4.7–5.2 males, 4.8–6.4 females; see also Gans 1959); Nasal divided below the nostril (undivided in D. sahelensis and D. parascabra); Postoculars 2 on either side of head (often one, especially in western populations, of D. atra); Supralabials usually 7 (3rd and 4th enter orbit) on either side of head (usually 6[2,3] in D. loveridgei comb. nov.).

Ventral counts in D. abyssina are mostly higher (226–247 in males, 241–271 in females) than the nearest populations of D. scabra in south-eastern Sudan and adjacent South Sudan (213; 222–227) and Ethiopia (212–218; 219–243), and much higher than D. taylori sp. nov. which occurs in northern Somalia and Djibouti (196–216; 208–222) (Appendix 1). Dorsally plain brown specimens are similar to the brown/grey phases of D. atra in north-eastern Africa and D. scabra in South Africa (e.g. De Waal 1978), although the venter may be yellowish-cream (MCZ 80984) rather than the usual plain cream. In addition, D. abyssina can often be separated from D. atra on account of its higher ventral counts (199–232 in atra males, 212–256 in females) and possession of two (rather than 1–2) postoculars on either side of the head. Plain phase D. abyssina are always distinguished from plain phase D. atra in Ethiopia (the only area where the two species are sympatric) by having higher ventral + subcaudal counts (both sexes, 294–323 [N =7] versus 269–291 [N = 5] respectively).

Description. Dorsum of patterned specimens with oblique dark saddles (65–98, N = 11), usually wider than long, and often confluent or forming zigzags on the back and tail. Lateral bars dark, long, mostly broader above than below, somewhat irregular and often in contact with the pale interspaces between saddles, sometimes in narrow contact with the edges of the saddles, and may extend onto the ventral plates as in the holotype, but never forming a distinct or broadly linked pattern. Ventral scales cream with extensive black, grey or brown markings on the anterior half, consisting of a large rounded cone on either side joined to a smaller cone in the centre. In live specimens the labials and interspaces between saddles are cream-yellow, and there may also be some cream-yellow on the sides of the head.

Four specimens from Eritrea in the Vienna collection (NMW) were listed under D. scabra by Gans (1959), with the dorsal patterns of three of them recorded as ‘5I’ (i.e. black rectangular saddles and lateral bars forming complete closure around the pale inter-saddle areas; see Gans 1959, Pl. VI, Fig. 4, and Fig. 20G). This material was examined in detail by the first author and all four specimens have at least some indication of the ‘5I’ D. arabica sp. nov.-like dorsal pattern: NMW 15300 has almost the entire dorsum patterned ‘5I’; NMW 15301–3 is similar but has mostly D. abyssina-like markings (as described above) at midbody; NMW 15301–1 has mostly abyssina-like markings anteriorly and mostly ‘5I’ markings posteriorly, although the pattern is not synchronised on either side of the midline just posterior to midbody (Gans 1959, Pl. VI, Fig. 3); while NMW 15301–2 has abyssina-like markings on the first quarter of the body and ‘5I’ markings on the rest of the dorsum. Gans (1959: 125) noted that although some ‘5I’ specimens had an immaculate venter (presumably including ZMB 7631 from Bogos = D. crucifera sp. nov. and BM 03.6.21.41 from El Kubar = D. arabica sp. nov. – see below), NMW 15301–2 (Gans, 1959, Pl. IX, Fig. 7) and NMW 15301–3 both had heavily dark-bordered ventrals like the holotype of D. abyssina (Duméril, Bibron & Duméril 1854, Pl. 81, Fig. 1–2; Fig. 20B). Re-examination of this material indicated that NMW 15301–1 also has the typically well marked abyssina-like ventral pattern, although the markings are pale grey in colour, while the ventral pattern of NMW 15301–3 is actually somewhat different to the others in that it consists of a large bold dark spot at the edges of the ventral plates, usually with 2–3 more-or-less disconnected paler blotches towards the centre. The distinct ventral markings of NMW 15300 are brown, which is perhaps an artifact of preservation.

INSERT FIG. 20

A plain tan brown to greyish phase with a largely unmarked cream to cream-yellow venter is also known, sometimes occurring in sympatry (at Gondar and Keriyo) with the patterned phase. In fact one of the six males and as many as six of the nine females are plain phase (marked by asterisks in ‘Localities’ below). A detailed description of two plain phase specimens follows: MCZ 80984 (Gondar to Asmara): dorsal scales are mostly pale tan, but the edges at the base and sides (anterior proximal half) of each scale is grey; NMB R10257 (Keriyo): dorsal scales pale tan with numerous tiny dark dots, and basal one-third of scale, as well as area around apical pits, grey. In both specimens the lowermost scale row on either side is without dark markings, while the scales of the adjacent row above have few dark markings on the lower edges of the scales, but the upper part of the scale base is dark; dark markings at the base of scales creates the impression of a very weak vertebral stripe and similar lateral bands, especially in NMB R10257. Dorsal scales of the latter specimen (and the plain phased NMZB 17099, also from Keriyo) are more elongate than in patterned conspecifics, and have rounded tips (compared to dorsals less elongate, sharply triangular and acutely-tipped in NMB R10256, a patterned specimen from the same locality as NMB R10257). However, the shape and serration of dorsal scales in MCZ 80984 is similar to that of patterned specimens. Belly and subcaudal region immaculate cream. In addition, these three plain phase specimens have only two rows of weakly serrated, inclined and reduced dorsals on either side of the body.

Preoculars 1 on either side of head (N = 18), postoculars usually 2 on either side of head, but one in BM 1902.12.13.86 (N = 18); frontal shield usually for the most part smooth with marginal pitting (States 1 & 2; with a single row in most specimens examined in detail), but entirely smooth in BM 1902.12.13.85 (State 0) (N = 13); inter-prefrontal suture usually weakly marked, but moderately marked in two specimens (MZUF 783, NMW 15301/2) (N = 13); supralabials 7 (3rd and 4th enter orbit) on either side of head (N = 17), but in BM 1902.12.13.85—although the head is damaged—it is evident that only the 2nd and 3rd supralabials on either side enter the orbit; anterior temporals usually two on either side of head (94.4%), but 1 left and 2 right in NMZB 17099 (N = 18); posterior temporals three on either side of head (38.5%), 3:4 or 4:3 (23.1%), 2:3 or 3:2 (23.1%), 1:2 in BM 1902.12.13.86 and 4 on right (left side damaged) in BM 1902.12.13.85 (N = 13); temporal formula on either side of head usually 2+3, occasionally 2+4 or 2+2; midbody scale rows 21–25 (mean 23.1 ±1.27, N = 19); ventrals 226–247 in males (N = 6), 241–271 in females (N = 10); subcaudals 57–68 in males (N = 4), 49–65 in females (N = 9). Dorsally brown D. abyssina have higher ventral + subcaudal counts than plain phase D. atra in Ethiopia (both sexes, 294–323 [N =7] versus 269–291 [N = 5] respectively). Ratio SVL: tail length 5.3–6.1 in males (N = 4), 5.5–8.1 in females (N = 10); total length: tail length 6.3–7.1 in males (N = 4), 6.5–9.1 in females (N = 10).

Size. Largest male (NMW 15301–3 – Eritrea) has a SVL of 504 mm (tail trunctated); largest female (MNHN 6567 – ‘Abyssina’, proposed lectotype, see below) 665 + 90 = 755 mm (see Duméril et al. 1854).

Habitat. Open savannah in the Ethiopian Highlands at elevations of 1900–2450 m (Figs 21 & 22). A subadult male, BM 69.11.4.38, although having an ‘Arabia’ (e.g. Gans 1959, Appendix 1) label in its museum jar, was in fact collected at Senafe, Eritrea at 2450 m (Blanford 1870), probably in Juniperus coniferous forest. Keriyo, north of Addis Ababa, is the southern-most locality for this species, situated at an altitude of 2320 m (Spawls 2004).

INSERT FIGS 21 & 22

Distribution. Restricted to central and north-western Ethiopia, and central Eritrea (Fig. 3).

Localities. (Plain phase specimens indicated by asterisks.) ERITREA. No locality, NMW 15301–1, 15301–2, 15301–3; Gheleb [NE 1538D4] NMW 15300; Senafe [NE 1439C2] BM 69.11.4.38. ETHIOPIA. No locality, MNHN 6567 (proposed lectotype, see below); Gondar [NE 1237C2] FMNH 12728, MSNM 2699a*,b,c, MZUF 783; Gondar to Asmara [ca. NE1338A = Ras Daschen Terera] MCZ 80984*; Keriyo [NE 0938B3] NMBO 10256–7*, NMZB 17099*; Mai Canetta [NE 1338D4] MSNM 2700a*,b*; Zegie, Lake Tsana [NE 1137C2] BM 1902.12.13.85–6*.

Biogeography. In the terrapin Pelomedusa subrufa Bonnaterre species complex, P. barbata Petzold et al. occurs in Saudi Arabia and Yemen, with closely-related populations of P. gehafie Rüppell in Eritrea and eastern Sudan, and P. somalica Petzold et al. in northern Somalia (Petzold et al. 2014). Similarly, D. arabica sp. nov. inhabits Saudi Arabia and Yemen, D. crucifera sp. nov. is found in northern Eritrea, D. abyssina is restricted to central Eritrea and the northern half of Ethiopia, while D. taylori sp. nov. occurs only in Somaliland and adjacent Djibouti.

Conservation. In terms of the regional (study area) and global distribution of this species, using IUCN (2012, 2016) criteria, it appears to be widespread and should be regarded as Least Concern.

Remarks. Saleh & Sarhan’s (2016: 26) comment that Trape et al. (2012) concluded that D. abyssina is a valid species “based on molecular phylogenetic data” is incorrect. Trape et al. (2012) did not include D. abyssina in their genetic analysis but followed Trape & Mané (2006) who treated this species as valid on morphological grounds.

Dasypeltis crucifera Bates sp. nov.

Cross-marked Egg-eater

Figs 23A–B, 24A–C & 25A–B

Dasypeltis scabra (not Linnaeus) Gans, 1959: 141 (part, Bogos, Pl. V, Fig. 4; Appendix 1, p. 204); Largen & Rasmussen, 1993: 340–341 (part, Agordat, Keren).

Holotype. ZMB 7631 (at Museum für Naturkunde, Berlin), an adult female (right side ovary examined) from Bogos, Eritrea (Gans 1959, Pl. V, Fig. 4; Fig. 23A–B & 24A–C). The specimen was purchased from Edward Gerrard (a London taxidermist and dealer of natural history objects) by the Zoologisches Museum der Königlichen Friedrich-Wilhelms-Universität zu Berlin during the second half of the 19th century (F. Tillack in litt., 15 April 2016). Bogos is a name that may refer to the city of Keren (also known as Senhit or Blin: 15°46’40”N, 38°27’29”E; NE 1538C4), but it may have been used as a more general term referring to the land of the Bogo people (an indigenous ethnic group) in Anseba Region, between the highlands and lowlands of Eritrea.

Paratypes. Two specimens: ERITREA. Allotype. MZUT 3582 (at Museo Regionale di Scienze Naturali di Torino, Turin), a juvenile male from Agordat [15°33’N, 37°53’E; NE 1537D2] (the specimen tag notes ‘Giachetti! cei det. 21.8.88’ [Fig. 25A–B]); MSNG 6738, probably female, from Keren [15°46’40”N, 38°27’29”E; NE 1538C4].

Additional material. One specimen: ERITREA. MZUT 3469, an adult female from Agordat [as above].

Diagnosis. Where data for MZUT 3469 (see “Additional material”) falls outside the ranges recorded for the type series, this is indicated below.

Assigned to the genus Dasypeltis on account of its slender form, possession of (usually 3–4) rows of reduced, oblique, keeled and serrated lateral scales (little or no serration in D. inornata) and head barely distinct from the neck (moderately distinct in D. fasciata). Differs from all other taxa in the genus by its unique dorsal colour pattern, in which dark brown oval saddles and lateral bars almost completely enclose pale cross-markings that form the inter-saddle areas (a variation of the ‘5I’ pattern found in Arabian Dasypeltis; Figs 23A & 25A), together with a combination of the following characters: Pattern cycles 51–62 (higher than D. bazi 38–49; lower or mostly lower than D. fasciata 66-141, D. abyssina 65-98, D. atra 62-130, D. arabica sp. nov. 62-79; D. inornatus always uniform brown above, D. gansi uniform or weakly marked); flanks decorated with dark vertical bars (spots or blotches in D. bazi, D. loveridgei comb. nov. and some populations of D. scabra); and venter immaculate apart from a few dark markings at the edges (extensive dark markings in D. abyssina and in many D. scabra); Ventrals: 226 in male (higher than D. taylori sp. nov. 196–216, D. sahelensis 207–221, D. inornata 208–225, D. parascabra 210–223), 231–243, 247 (MZUT 3469) in females (higher or mostly higher than D. taylori sp. nov. 208–222); Subcaudals 61 in male (higher than D. loveridgei comb. nov. 47–57; lower than D. inornata 81–92, D. fasciata 59, 71–91, D. medici 69–109, D. gansi 68–83, D. palmarum 68–77, D. parascabra 67–74, D. latericia 66–86, D. arabica 63–65), 49–52 in females (lower than D. inornata 69–84, D. fasciata 64–84, D. parascabra 57–64, D. palmarum 62–77, D. medici 61–90, D. gansi 59–73; D. latericia 59–72, D. arabica sp. nov. 53–61); Inter-prefrontal sulcus weakly marked (moderately to deeply marked in D. arabica sp. nov., weakly to moderately marked in D. bazi, usually moderately marked in D. taylori sp. nov., and variable in D. scabra [often well marked in the south of its range]); Frontal shield marginally pitted, State 1 or 2 (extensively pitted, States 3 & 4, in D. medici, D. taylori sp. nov. and many populations of D. scabra especially in the south of its range); Nasal divided below the nostril (undivided in D. sahelensis and D. parascabra); Postoculars 2 on either side of head (often one, especially in western populations, of D. atra); Supralabials usually 7 (3rd and 4th enter orbit) on either side of head (usually 6[2,3] in D. loveridgei comb. nov.).

Description of holotype. The holotype (ZMB 7631) is an adult female with a total length of 506 mm, snout-vent length 442 mm, tail length 64 mm. Ratio SVL: tail length 6.9, total length: tail length 7.9.

Head small and slightly distinct from the neck; snout rounded. Elongate nasal semi-divided (divided below the nostril), nostril pierced in anterior half of nasal, no loreal. Paired prefrontals larger than the paired internasals (individually and in combination). Inter-prefrontal sulcus weakly marked. Frontal large, widest anteriorly, slightly longer than broad, and smooth except for a single row of pits on the lateral borders (State 1). There is a single preocular and two postoculars on either side of head. Temporals 2+3 on either side of head. Seven supralabials, 3rd and 4th bordering the eye, on either side of head. First pair of infralabials in contact behind the mental, followed by a pair of large and elongate sublinguals in broad contact, and another pair of smaller sublinguals in narrow contact with one another and in contact with the first ventral plate, but separated posteriorly and medially by an oval extranumerary scale in broad contact with the first ventral plate. Eyes large (with vertically elliptical pupils), their greatest diameter much less than the distance from tip of snout to anterior part of eye socket.

Dorsal scales strongly keeled, unicarinate, in 25 rows at midbody, the third, fourth and fifth rows denticulate and obliquely placed. Dorsal scales in the anal region denticulate (see Gans 1959: 204). Vertebral scale row not enlarged. There are 231 ventrals, the anal plate is entire and there are 52 pairs of smooth subcaudals. After preservation in ethanol the dorsal colouration is pale brown with 62 ocelli or pattern cycles (markings are darker brown, saddles with paler brown centres) between nape and base of tail. Pale markings on the back are in the form of crosses, largely surrounded by brown, oval or diamond-shaped (sharpest ends anteriorly and posteriorly) dorsal saddles and similarly dark lateral bars, but with the tips of the crosses often separated from the dark markings. The tail has faint dark vertebral blotches anteriorly. Head with a chevron marking, interrupted on the left, with a dark arrowhead marking present on the nape. Venter immaculate apart from a few small brown spots extending from the outer scale row onto the outer edge of a ventral.

INSERT FIGS 23 & 24

Variation in paratypes (including allotype, as indicated). The allotype (MZUT 3582) is a juvenile male (211 + 39 = 250 mm; ratio SVL: tail length 5.4, total length: tail length 6.4); the paratype (MSNG 6738) is a female (368 + 45 = 413 mm, ratios 8.2 and 9.2). Both specimens have similar scalation and general appearance to the holotype, but differ as follows: frontal distinctly longer than broad in MSNG 6738; on the right side of the head of the allotype there are 2+2 temporals; pair of small posterior sublinguals in contact with first ventral plate and not separated by an extranumerary scale (also allotype); longitudinal rows of dorsals at midbody 22 in allotype, 24 in MSNG 6738; ventrals 226 in allotype (male), 243 in MSNG 6738 (female); pairs of subcaudals 61 in allotype (male), 49 in MSNG 6738 (female); pattern cycles on the back 52 in allotype, 51 in MSNG 6738; in allotype, forward-pointing chevron marking at the back of the head is followed by a dark oval marking that extends onto the nape, where it is followed by a backwards-facing chevron; in MSNG 6738 there is a forward-pointing chevron on top of the head, followed by a forward-pointing arrowhead on the nape with a connecting stalk extending onto the neck. The dark (brown) markings (often diamond-shaped) of the allotype are less distinct than in the holotype, and only barely distinguishable from the ground colour on the posterior part of the body and on the tail; in MSNG 6738 the dark oval saddles are often more rounded at the ends such that the pale crosses are less X-like in appearance. Serration of lateral scale rows and scales above the anal region, and condition of the anal plate, not determined. No live specimens are known to the authors, nor are any photographs available.

INSERT FIG. 25

Description of additional material: MZUT 3469 (female). This specimen is not currently available for examination and no photograph of it is available to determine its dorsal pattern (F. Andreone pers comm., 3 April 2016). However, it has 247 ventrals, 50 subcaudals and 23 midbody scale rows.

Size. Largest male (allotype, MZUT 3582 – Agordat) 211 + 39 = 250 mm; largest female (holotype, ZMB 7631 – Bogos) 442 + 64 = 506 mm.

Etymology. The name of the new species refers to the pale cross-marks between the saddles on its back.

Habitat. This species occurs in the hilly area between the lowlands associated with the drainage of the Barka River, the Sahel and the Highlands of Hamasien. Agordat is situated at low altitudes (about 600 m), while the city of Keren is at an altitude of about 1417 m.

INSERT FIG. 26

Distribution. Known from only two areas (Bogos/Keren and Agordat) in northern Eritrea (Fig. 3). Keren and Agordat are about 75 km apart.

Mimicry. Dasypeltis crucifera sp. nov. may mimic Echis megalocephalus Cherlin, 1990 (see photograph of “E. cf. megalocephalus” from Ginda in Dobiey & Vogel 2007; Fig. 30).

Biogeography. In the terrapin Pelomedusa subrufa Bonnaterre species complex, P. barbata Petzold et al. occurs in Saudi Arabia and Yemen, with closely-related populations of P. gehafie Rüppell in Eritrea and eastern Sudan, and P. somalica Petzold et al. in northern Somalia (Petzold et al. 2014). Similarly, D. arabica sp. nov. inhabits Saudi Arabia and Yemen, D. crucifera sp. nov. is found in northern Eritrea, D. abyssina is restricted to central Eritrea and the northern half of Ethiopia, while D. taylori sp. nov. occurs only in Somaliland and adjacent Djibouti.

Conservation. Using IUCN (2012, 2016) criteria and considering that it is known from only two areas, this species should be regarded as Data Deficient.

Remarks. The holotype (ZMB 7631) was listed under D. scabra (‘5I’ dorsal pattern) by Gans (1959) who failed to mention its distinctive cross-mark pattern. He also indicated that while other African specimens with the ‘5I’ dorsal pattern had heavily-marked venters (here referred to D. abyssina), this specimen had ‘immaculate’ underparts.

Dasypeltis arabica Broadley & Bates sp. nov.

Arabian Egg-eater

Figs 27A–C, 28A–B & 29A–B

Dasypeltis scabra (not Linnaeus) Boulenger, 1905: 180 (El Kubar, Yemen); Leviton & Aldrich, 1954, xxiii (checklist); Gans 1959: 141 (part, El Kubar) & 1961: 73 (part, Arabia); ? Corkhill & Cochran, 1965: 485 (Little Aden oil refinery); Spanò, 1972: 132 (El Siyani, Yemen); Gasperetti, 1988: 233, 410, Fig. 39 (1) (Ajibah, Saudi Arabia); Schätti & Desvoignes, 1999: 82; Schätti & Gasperetti, 1994: 386 (Aqabat Raydah & Abha, Saudi Arabia).

Dasypeltis saeizadi Hoser, 2013: 60 (“Saudi Arabia”) (nomen dubium); Göthel, 2015a: 19 & 2015h: 92; Saleh & Sarhan, 2016: 36.

Dasypeltis cf. scabra Sindaco et al. 2013: 93 (part, “SW Arabia and Yemen”).

Holotype. BM 1987.2192 (at Natural History Museum, London), an adult female (oviducts examined) from Sana’a, North Yemen [15º19’N, 44º14’E] collected by M. Al-Safadi (Figs 27 & 28).

Paratypes. Six specimens: YEMEN. Allotype. BM 1903.6.26.41 (at Natural History Museum, London; wrongly listed as ‘xBM 03-6-21-41’ by Gans 1959), an adult male (Gans 1959) from El Kubar, South Yemen [14º25’N, 45º01’E] collected by G.W. Bury; BM 1987.2193, adult female from Taiz, North Yemen [13º35’N, 44º01’E] collected by M. Al-Safadi; MSNG 52221, male from El Siyani, North Yemen, collected by G. Scortecci, 13 August 1965. SAUDI ARABIA. BM 1987.1018, subadult female from Ajibah [19º41’N, 41º57’E] collected by J. Gasperetti (Fig. 29); MHNG 2536.41, male from Aqabat Raydah [NE 1942C]; MHNG 2536.42, female from vicinity of Abha [about 18º13’N, 42º30’S].

Diagnosis. Assigned to the genus Dasypeltis on account of its slender form, possession of (usually 3–4) rows of reduced, oblique, keeled and serrated lateral scales (little or no serration in D. inornata) and head barely distinct from the neck (moderately distinct in D. fasciata). Distinguished from other congeners by its unique dorsal colour pattern in which black rectangular saddles and lateral bars form complete closure around the pale inter-saddle areas (Figs 27A & 29A; this pattern also occurs on parts of the back of a few D. abyssina from Eritrea, see pattern‘5I’ of Gans 1959, Pl. VI, Fig. 4); together with a combination of the following characters: Pattern cycles 62–79 (≤62 in D. crucifera sp. nov., D. taylori sp. nov. and D.bazi; D. inornatus always uniform brown above, D. gansi uniform or weakly marked); flanks decorated with dark vertical bars (spots or blotches in D. bazi, D. loveridgei comb. nov., and some populations of D. scabra—e.g. Fig. 5G); and venter immaculate apart from a few dark markings at the edges (extensive dark markings in D. abyssina and many D. scabra); High numbers of ventrals: 236–244 in males (higher than D. scabra 180–226, D. taylori sp. nov. 196–216, D. confusa 199–227, D. atra 199–232, D. sahelensis 207–221, D. inornata 208–225, D. bazi 213–229, D. parascabra 210–223 and D. crucifera sp. nov. 226), 239–254 in females (higher than D. taylori sp. nov. 208–222, D. sahelensis 212–237, D. bazi 215–232, D. inornata 219-237, D. parascabra 224–237); Subcaudals 63–65 in males (higher than D. loveridgei comb. nov. 47–57, D. bazi 59–61, D. crucifera sp. nov. 61; lower than D. inornata 81–92, D. fasciata 59, 71–91, D. medici 69–109, D. gansi 68–83, D. palmarum 68–77, D. parascabra 67–74, D. latericia 66–86), 53–61 in females (higher than D. loveridgei comb. nov. 39–50, D. taylori sp. nov. 44–52, D. crucifera sp. nov. 49–52; lower than D. inornata 69–84, D. fasciata 64–84, D. palmarum 62–77; mostly lower than D. medici 61–90); Inter-prefrontal sulcus usually moderately, occasionally deeply, marked (usually weakly marked in other species, but weakly to moderately marked in D. bazi, usually moderately marked in D. taylori sp. nov., and variable in D. scabra [often well marked in the south of its range]); Frontal shield marginally pitted, usually State 1 (extensively pitted, States 3 & 4, in D. medici, D. taylori sp. nov. and many populations of D. scabra especially in the south of its range); Nasal divided below the nostril (undivided in D. sahelensis and D. parascabra); Postoculars 2 on either side of head (often one, especially in western populations, of D. atra); Supralabials 7 (3rd and 4th enter orbit) on either side of head (usually 6[2,3] in D. loveridgei comb. nov.).

Description of holotype. The holotype (BM 1987.2192) is an adult female with a total length of 714 mm, snout-vent length 625 mm, tail length 89 mm; ratio SVL: tail length 7.0, total length: tail length 8.0.

Head small and barely distinct from the neck; snout rounded. Elongate nasal semi-divided (divided below the nostril), no loreal. Paired prefrontals slightly larger than the paired internasals (individually and in combination). Inter-prefrontal sulcus moderately marked. Frontal large, widest anteriorly, 1.14 times longer than broad, and smooth except for a single row of pits on the lateral borders (State 1). There is a single preocular and two postoculars on either side of the head. Temporals 3+3 (left), 2+4 (right). Seven supralabials, 3rd and 4th bordering the eye, on either side of the head. First pair of infralabials in contact behind the mental, followed by a pair of large and elongate sublinguals in broad contact which extend backwards to separate a smaller pair of posterior ones, which are followed by a pair of supranumerary shields, the right one being partially fused with the first ventral. Eyes moderate in size (with vertically elliptical pupils), their length much less than the distance from tip of snout to anterior part of eye socket.

Dorsal scales strongly keeled, unicarinate, in 25 rows at midbody, the third, fourth, fifth and sixth rows denticulate and (except for the 3rd row) obliquely placed. Dorsal scales in the anal region very feebly denticulate. Vertebral scale row not enlarged. There are 240 smooth ventrals, the anal plate is entire and there are 53 pairs of smooth subcaudals. After preservation in ethanol the dorsal colour pattern consists of black rectangular saddles and similarly coloured lateral bars (on a pale brown to grey background) that form complete closure around the pale (whitish) square to rectangular inter-saddle areas. There are 67 ocelli or pattern cycles (dark with pale centres) between nape and base of tail. The tail has faint dark vertebral blotches anteriorly. Head with remnants of fragmented chevrons, and a dark forward-pointing arrowhead marking is present on the nape. Venter immaculate cream to white apart from a few dark spots extending from the outer scale rows onto the outer edges of ventrals.

INSERT FIGS 27 & 28

Variation in paratypes (including allotype, as indicated). The six paratypes comprise three males and three females. The largest male (MSNG 52221) measures 563 (486 + 77) mm; BM 03.6.21.41 (allotype) measures 325 + 56 mm. Measured females (BM 1987.2193: 400 + 64 mm, BM 1987.1018: 246 + 60 mm) are smaller than the holotype. The paratypes have similar scalation and general appearance to the holotype, but differ as follows: Midbody scale rows are 23 (3 specimens, including allotype), 24 (1) or 25 (2) (Appendix 1). Ventrals 236–244 in males (244 in allotype), 239–254 in females (Appendix 1). Subcaudals 63–65 in males (65 in allotype), 53–61 in females (Appendix 1). Paired prefrontals much larger than the paired internasals in MHNG 2536.41 & 42; inter-prefrontal sulcus usually moderately developed but deeply marked in MHNG 2536.41; frontal with two rows of pits (State 2) on its outer surface in MHNG 2536.41; temporals in 2+3 arrangement in MHNG 2536.41 & 42; usually third, fourth and fifth (also sixth in two specimens) lateral scale rows reduced in size, serrated and inclined; serration on scales above vent faint or absent. Pupils appear round in BM 1987.2193. In MHNG 2536.41 the dark dorsal markings are joined along only the last one-fifth of the body (from snout to vent), elsewhere being similar to the ‘5N’ markings of D. scabra (see Gans 1959), while in MHNG 2536.42 most of the markings on the body are D. scabra-like, but with several ‘5I’ markings posteriorly. Dorsal pattern cycles 62–79 (Appendix 1), the first neck band divided in BM 03.6.21.41 (allotype) and MHNG 2536.42 (divided by an elongated vertebral band). A narrow, somewhat irregular, chevron is present on the back of the head in BM 1987.1018. No live specimens are known to the authors, nor are any photographs available. (Frontal and eye-snout measurements, and patterning on the top of the head, not recorded for all paratypes and additional material. For MSNG 52221 only size [see above], colour pattern, and numbers of supralabials, preoculars, postoculars, temporals, midbody scale rows, ventrals and subcaudals were recorded.)

INSERT FIG. 29

Size. Largest male (MSNG 52221 – El Siyani, North Yemen) 486 + 77 = 563 mm; largest female (BM 1987.2192, holotype – Sana’a, North Yemen) 625 + 89 = 714 mm.

Etymology. The name of the new species refers to the region of origin (Arabia) of the type series.

Habitat. This species occurs in the south-western highlands of Saudi Arabia and Yemen from 1000 m to 2100 m (annual rainfall exceeds 300 mm).

Distribution. South-western part of Saudi Arabia and western half of Yemen (Fig. 3).

Mimicry. Dasypeltis arabica sp. nov. may mimic Echis coloratus Günther (see photographs from the Sinai Peninsula in Dobiey & Vogel [2007]). Gasperetti (1988) showed that E. coloratus also occurred from Saudia Arabia to the south coast of Yemen, and illustrated a topotype from Jabel Grayyah in Saudia Arabia (his Fig. 123) showing pale vertebral spots (described as ‘pinkish’ in the description of the holotype by Günther [1878]). The new species is also similar to Echis megalocephalus (Fig. 30).

INSERT FIG. 30

Biogeography. Dasypeltis arabica sp. nov. has a similar distribution to Naja arabica Scortecci, except that the latter also occurs further east in the highlands of Dhofor, Oman (Trape et al. 2009). The geological history of the region is summarised by Leviton (1986): “During the entire Mesozoic and early Cenozoic, Arabia, Nubia and Somalia were assembled in one continental block at some times covered by shallow seaways (Bosellini, 1986). The onset of Neogene rifting, accompanied by the ‘slight oblique drifting’ of Africa and Arabia resulted in the opening of the Gulf of Aden.” Boulenger (1905) described, from El Kubar in Yemen, Atractaspis andersoni which was considered closely related to A. leucomelas Boulenger of Somalia, and also Zamenis [= Platyceps] variabilis, closely related to Platyceps brevis Boulenger of Somalia. In the terrapin Pelomedusa subrufa Bonnaterre species complex, P. barbata Petzold et al. occurs in Saudi Arabia and Yemen, with closely-related populations of P. gehafie Rüppell in Eritrea and eastern Sudan, and P. somalica Petzold et al. in northern Somalia (Petzold et al. 2014). Similarly, D. arabica sp. nov. inhabits Saudi Arabia and Yemen, D. crucifera is limited to northern Eritrea, D. abyssina is restricted to central Eritrea and the northern half of Ethiopia, while D. taylori sp. nov. occurs only in Somaliland and adjacent Djibouti. Lavin & Papenpuss (2012) cite several recent molecular studies that show that the Red Sea and Gulf of Aden are barriers to various reptiles between the Arabian Peninsula and Africa.

Conservation. Using IUCN (2012, 2016) criteria this species appears to be widespread and should be regarded as Least Concern.

Remarks. Corkill & Cochran (1965) recorded a specimen (whereabouts unknown, not plotted on map) of “D. scabra” from Little Aden (oil terminal) on the coast in southern Yemen, but its identity remains problematic.

We concur with the opinions, recommendations and proposals of Kaiser et al. (2013) and Kaiser (2014) regarding best taxonomic practices, and therefore reject the unscientific and unethical taxonomy of Hoser (2013, description of Dasypeltis saeizadi). We note specifically that the privately published and personally edited work of Hoser (2013) is in direct violation of the spirit and intention of the Code (ICZN 1999) as indicated by, inter alia, Recommendation 8A which explicitly encourages publication in “appropriate scientific journals or well-known monographic series” (see also comments by Schleip [2014] regarding non-compliance with the ICZN [1999]). It can be added that in their Snakes of the World catalogue, Wallach et al. (2014: ix) state: “The Australasian Journal of Herpetology [in which the Hoser 2013 paper was published], of which Hoser is the editor, reviewer, publisher, and sole author, is not considered a valid publication under the spirit of the Code and therefore all proposed names are nomina illegitima (see Wallach et al., 2009 and Kaiser et al., 2013).”

Hoser (2013) named ‘BMNH – 1987.1018’ as holotype and sole type specimen of what he considered a new species, namely D. saeizadi, from ‘Saudi Arabia’. His diagnosis is based entirely on a vague comparison of colour patterns be tween D. scabra, D. abyssina (presumably based on MNHN 6567, the Ethiopian syntype of R. abyssinus – the only specimen of this taxon known at the time, incorrectly referred to as the ‘holotype’ of D. abyssina by Trape & Mané 2006, see discussion above) and the proposed new species. Hoser (2013: 60) noted that the latter is “readily separated” from both D. scabra and D. abyssina “by the dorsal colouration”. However, he proceeded to make the following erroneous statements in this regard: According to Hoser (2013: 60): D. abyssina has “indistinct markings on the head and body and a complete absence of barring on the upper labials, or if present, it [barring on supralabials] is effectively indistinct.” However, the proposed lectotype (MNHN 6567) of D. abyssina and all other known (patterned) specimens of this species (see species account above) have very distinct dark markings dorsally and laterally on the head, body and tail, including distinct barring on the supralabials (see Duméril et al. 1854, Pl. 81, Fig. 1 in colour & Fig. 20 in the current paper); while even the Nil Blanc ‘syntype’ (proposed paralectotype, MNHN 7227 = D. scabra) has distinct markings as described above. Hoser (2013: 60) then stated that his new species is “more similar to D. scabra in dorsal colouration, but lacks the distinctive whitish patches interspercing the dorsal midline blotches or squares. In turn these blotches or squares tend somewhat towards being pointed at the anterior end of the body”. Again this is incorrect as the dorsal pattern of BM 1987.1018 differs distinctly from D. scabra (Gans’ 1959 ‘5I’ versus ‘5N’ dorsal colour patterns respectively), the pale patches on its back are very distinct, and the saddles are for the most part not at all markedly pointed anteriorly (Fig. 29A). Hoser (2013: 60) added that his proposed new species “In contrast to the other two species [D. scabra and D. abyssina]” has a “darkening on the upper temples” and “Like D. abyssina there is a complete absence of barring on the upper labials”, “or if present, it [barring on the supralabials] is effectively indistinct.” Once again this is incorrect as Fig. 29A–B clearly shows that the upper temples of BM 1987.1018 are not darker than other parts of the head, nor are they noticeably darker than in D. abyssina (Duméril et al. 1854, Pl. 81, Fig. 1; Fig. 20C–G) or D. scabra (e.g. Gans 1959, Pl. IV-V; Broadley 1990, Fig. 152; Bates et al. 2014, p. 411, lower figure; Fig. 5), and the supralabials (of BM 1987.1018) are in fact distinctly barred, as are those of D. abyssina (Duméril et al. 1854, Pl. 81, Fig. 1; Fig. 20C–G). Hoser (2013) did not provide any additional features for distinguishing his proposed new species. It is therefore evident from the afore-mentioned that Hoser’s (2013) description is a mis-representation of reality—his description is not based on the true features of the specimen he names as holotype. His description cannot therefore be considered valid and in accordance with the requirements of the Code (e.g. Article 1, which requires that an actual specimen be used for the description of an extant or extinct species of animal). More specifically, Article 1.3.1 excludes from the provisions of the Code any name proposed for a hypothetical concept, which is what Hoser’s (2013) name must be considered, as it can be shown (see above) that it is not based on the features of the specimen he named as holotype (no paratypes).

According to P. Campbell at the Natural History Museum, London (in litt. 3 September 2015), no record exists of Raymond Hoser having ever borrowed any Dasypeltis from that institution, or of him having spent time at that museum examining such specimens. If Hoser had indeed examined the specimen, or even an image thereof, the characteristic ‘5I’ dorsal pattern (see Gans 1959, and Figs 27A & 29A) of Arabian populations of Dasypeltis would have been obvious immediately. Hoser (2013) also disregarded Recommendation 73C of the Code (1999) by not providing full collection details for the specimen and neglecting to provide any supporting meristic or morphometric data, including ventral, subcaudal and midbody scale row counts—all considered standard fare in modern snake descriptions. The lack of any such data in his description is a further indication that he did not examine the specimen (or an illustration thereof) that he named as holotype. In consideration of the above, Hoser’s (2013) name must be considered a species inquirendum and a nomen dubium and rejected as an available name for D. arabica sp. nov.

Multivariate analyses

For the analysis of all populations, two-way permanova (10 000 permutations) detected that there is no interaction between gender and species (df 9, F = –16.69, p = 0.67). However, it detected significant differentiation at species level (df 9, F = 2.43, p < 0.001) and at gender level (df 1, F = 5.28, p < 0.001) respectively, justifying separate analysis of sexes. For the Horn of Africa analysis, two-way permanova (10 000 permutations) detected that there is no interaction between gender and species (df 9, F = –3.19, p = 0.53). However, it detected significant differentiation at species level (df 6, F = 1.29, p < 0.001) and at gender level (df 1, F = 2.24, p < 0.001) respectively, justifying separate analysis of sexes.

A total of 249 specimens (D. medici: 9 males, 20 females; D. confusa: 6, 13; D. scabra: 24, 40; D. bazi: 3, 2; D. taylori: 3, 2; D. atra: 40, 62; D. fasciata: 2, 3; D. abyssina: 4, 7; D. crucifera: 1, 2; D. arabica: 2, 4) had complete sets of data and were used in the analyses (separately for males and females) of the entire study area. A total of 64 specimens (D. scabra: 6 males, 13 females; D. bazi: 3, 2; D. taylori: 3, 2; D. atra: 7, 9; D. abyssina: 4, 7; D. crucifera: 1, 2; D. arabica: 2, 4) with complete data were used in analyses (separately for sexes) of the Horn of Africa and Egypt. In the analyses for males from the latter area, numbers of preoculars was invariable and was therefore excluded. For pattern cycles, plain coloured specimens (including ‘intermediate’ D. medici with barely distinguishable darker dorsal markings) were scored as ‘0’.

Principal Component Analysis for north-eastern Africa and south-western Arabia

In the analysis of males (Fig. 31) the first two principal components (PC) explained most of the variance (36.2%). Only D. medici formed a distinct cluster distinguishable from all other populations, mostly along PC 2 for which numbers of subcaudals (0.54) and ventrals (0.40) had the highest loadings (Table 3). For females the first two components also explained most of the variance (37.2%), and again only D. medici formed a distinct cluster distinguishable from all other populations (Fig. 31), with the highest loadings (on PC 2) for subcaudals (–0.49) and pattern cycles (0.49) (Table 4). However, both sexes of D. scabra, D. atra and D. confusa also formed clusters and were partly distinguishable along PC 2.

INSERT FIG. 31

INSERT TABLES 3 & 4

Discriminant Function Analysis for north-eastern Africa and south-western Arabia

As all groups (as male or female of a species) are real geographical entities it is appropriate to conduct Discriminant Function Analysis to determine whether or not they are distinguishable. In the analysis of males of the various species (Fig. 32), 83.9% of specimens were correctly classified, but this included only 50% of D. confusa, while one out of three specimens of both D. taylori sp. nov. and D. bazi were mis-classified (Table 5). Dasypeltis medici was easily distinguishable from all other species along axis 1, on which numbers of subcaudals (0.81) and ventrals (0.60) load most heavily; Dasypeltis scabra and D. atra are largely separated along axis 2, on which fontal pitting (0.67) and pattern cycles (0.54) load most heavily; D. abyssina and D. arabica are also separated from D. scabra along axis 2, on which number of ventrals (0.41) is also an important factor (Table 6). For females (Fig. 32), 75.5% of specimens were correctly classified, but this included only 61.5% of D. confusa, 50% of D. taylori sp. nov., D. arabica sp. nov. and D. crucifera sp. nov., and neither of two specimens of D. bazi (Table 7). Dasypeltis medici is again easily distinguishable from all other species along axis 1, on which numbers of subcaudals (0.92) loads most heavily; Dasypeltis scabra and D. abyssina are well separated along axis 2, on which fontal pitting (0.67) and ventrals (0.64) load most heavily; and D. scabra and D. atra are partially separable on this axis (Table 8).

INSERT FIG. 32

INSERT TABLES 5–8

Principal Component Analysis for the Horn of Africa and Egypt

In the analysis of males the first two principal components (PC) explained most of the variance (40.1%); for females these components also explained most of the variance (38.7%). Although some species are largely distinguishable, none of the groups for males or females formed distinct clusters (Fig. 33; Tables 9 & 10).

INSERT FIG. 33

INSERT TABLES 9 & 10

Discriminant Function Analysis for the Horn of Africa and Egypt

In the analysis of males (Fig. 34) all specimens with the exception of one of three D. bazi were correctly classified, suggesting that they are 100% distinguishable in morphometric space (Table 11). All species were fully or partly (D. abyssina and D. arabica sp. nov.) distinguished along axis 1—on which ventrals (2.51) and midbody scale rows (2.09) load most heavily—except D. atra and D. bazi which are nevertheless largely separated along axis 2, for which pattern cycles (0.82) and frontal pitting (–0.68) load most heavily (Table 12). For females (Fig. 34), 81.6% of specimens were correctly classified; this figure was at least 75% for all species except D. bazi for which one of two specimens was incorrectly classified (Table 13). Dasypeltis scabra was distinguished from D. arabica sp. nov. and D. abyssina along axis 1, while the latter two species were distinguished along axis 2; D. taylori sp. nov. and D. bazi were distinguished from D. arabica sp. nov. and D. abyssina along axis 1; D. scabra is also distinguished from D. crucifera sp. nov. along axis 2; D. atra is distinguished from D. arabica sp. nov. along axis 2, and also partly from D. scabra on this axis (Table 14). Ventrals (1.07) load by far the most heavily on axis 1, whereas subcaudals (1.02) and pattern cycles (0.68) load most heavily on axis 2.

INSERT FIG. 34

INSERT TABLES 11–14

SUMMARY OF SPECIES 

A morphological analysis of the genus Dasypeltis in north-eastern Africa and south-western Arabia indicates that as many as 10 species occur there. This includes three new species (D. taylori sp. nov., D.crucifer sp. nov. and D. arabica sp. nov.) previously subsumed under the name D. scabra, as well as two (D. abyssina and D. bazi) that had until recently also been considered referable to D. scabra. The ranges of these five species do not overlap. This taxonomy is partly supported by the multivariate analyses of scale characters and numbers of pattern cycles, and colour pattern is often diagnostic of species.

Dasypeltis medici (Bianconi)

Although it occurs as patterned, weakly patterned and unicolour morphs (dorsally), this species is easily distinguishable by the presence of paired apical pits—on the dorsal scales—which are sharply defined by dark pigment, an extensively pitted frontal shield, and high subcaudal counts (75–109 in males, 61–90 in females). Also, patterned specimens have 1–5 distinct, often narrow, chevrons on the neck (one or two similar chevrons may be present in some other species). In the study area this species is restricted to southern Somalia, south-eastern Kenya, and north-eastern Tanzania; found along the coast but also in some savannah and mountainous areas.

Dasypeltis confusa Trape & Mané

Although often similar in scalation characteristics to D. scabra and D. atra, with which it is sympatric and parapatric, this species is easily distinguished by its colour pattern of large diamond-shaped saddles (often outlined in white) confluent with the lateral bars. In the study area it occurs only in the west (southern South Sudan, Uganda, western Kenya, Rwanda).

Dasypeltis scabra (Linnaeus)

A variable species widely distributed in the study area, from Burundi and Tanzania northwards to southern Somalia and central Ethiopia, with isolated populations along the Nile River in South Sudan and Sudan. It always has the ‘5N’ dorsal pattern, and differs from patterned D. atra by seldom having transversely enlarged saddles on the back, and usually also by having a more boldly marked head. There are usually two postoculars on either side of the head, while D. atra often has only one.

Dasypeltis bazi Saleh & Sarhan

This species has a distinctive colour pattern consisting of large cranio-caudally elongate saddles, usually bifurcated at each end with a fairly shallow but distinct notch, and the flanks are decorated with spots or blotches (not bands or vertical bars). The only locality at which this species is currently known to occur, namely the Fayoum Depression, is separated from other populations of Dasypeltis by over 1 300 km.

Dasypeltis taylori Bates & Broadley sp. nov.

This species, characterised by a combination of features including a strongly pitted frontal scale, is found in Somaliland and Djibouti, separated from populations of D. scabra to the south by the arid Ogaden plateau. The dorsal saddles are often large, closely-spaced, square or slightly rectangular, and separated by narrow, usually sharply defined, white crossbars; there are low numbers of pattern cycles (44–59).

Dasypeltis atra Sternfeld

Occurs as three colour phases: brown dorsally and cream to yellow or orange ventrally; black dorsally (unique in the genus) and grey ventrally; and patterned, sometimes similar to the ‘5N’ pattern of D. scabra but more typically with transversely widened saddles, often in contact with the lateral bars. Typically the frontal scale is only marginally pitted and the inter-prefrontal suture weakly marked; and western populations usually have only a single postocular (rather than two), a condition rare in others of this genus. This species is widespread in the study area, from northern Tanzania to Burundi, Rwanda, Kenya, Uganda, south Sudan and Ethiopia as far north as Circao. In some areas it is sympatric and/or parapatric with D. scabra, D. confusa, D. medici, D. fasciata and D. abyssina.

Dasypeltis fasciata A. Smith

Dorsum patterned, with body pattern always expressed as chequerboard on skin between scales and often also on lower edges of these, rather than on most of the scales themselves as in patterned specimens of other species. Also characterised by high subcaudal counts (59, 71–76 in males, 64–76 in females). In the study area this highly arboreal species is restricted to forests in western Uganda.

Dasypeltis abyssina (Duméril, Bibron & Duméril)

Easily recognisable by the combination of its unique dorsal pattern (including oblique transversely widened saddles which may merge to form zigzags on the back) and heavily marked venter. This species also has the highest ventral counts in the genus (241–271 in females, 226–247 in males). It is restricted to the Ethiopian highlands that extend into Eritrea.

Dasypeltis crucifera Bates sp. nov.

Found near the northern limits of D. abyssina, but at lower altitudes in northern Eritrea below the Ethiopian highlands. Distinguished by its unique dorsal pattern in which dark brown oval saddles and lateral bars almost completely enclose pale cross-markings that form the inter-saddle areas; venter largely unmarked (D. abyssina, its nearest congener, has a markedly different dorsal pattern and a heavily marked venter).

Dasypeltis arabica Broadley & Bates sp. nov.

Easily distinguished by its unique dorsal pattern consisting of black rectangular saddles and lateral bars that form complete closure around the pale inter-saddle areas (pattern ‘5I’ of Gans 1959); the venter is mostly unmarked. It has high ventral counts. This is the only species in the genus restricted to Arabia, where it occurs in the highlands of Saudi Arabia and Yemen.

DISCUSSION

Dasypeltis scabra is by far the most widespread species in the genus, and the most widely distributed of the 10 species found in north-eastern Africa and south-western Arabia. It ranges from the Cape of Good Hope through large parts of southern, central (excluding forests) and eastern Africa (‘5N’ dorsal pattern of Gans 1959; Broadley 1990; this study), with apparently isolated populations along the Nile River (Sudan and South Sudan) and near Kutum (western Sudan). The isolated Egyptian population in the Fayoum Depression is now considered a separate species (D. bazi), as are populations in Somaliland/Djibouti (D. taylori sp. nov.), northern Eritrea (D. crucifera sp. nov. ), the northern Ethiopian highlands (D. abyssina) and western Arabia (D. arabica sp. nov.). Dasypeltis confusa has a large range in western and central Africa (‘5L’ of Gans 1959), extending as far east as South Sudan, Uganda, western Kenya and Rwanda. Dasypeltis fasciata is arboreal and restricted to forests but has a large range in western and central Africa, extending as far east as Uganda. Dasypeltis medici, now considered a monotypic species, occurs in coastal and inland forests from southern Somalia to northern KwaZulu-Natal in South Africa; it also occurs in savannah where it may be sympatric with D. scabra. Dasypeltis atra, at one time considered a montane forest subspecies of D. scabra, is clearly a separate species (e.g. Gans 1964) with an extensive range from eastern Democratic Republic of the Congo to Rwanda, Burundi, Uganda, South Sudan, Kenya and the Ethiopian highlands. The range of D. atra is overlapped almost entirely by that of D. scabra, and although the two species appear to be mostly parapatric, the former also occurs in some savannah areas (e.g. Broadley & Bates 2009) where it is sympatric with the latter.

Gans (1959) considered that evolutionary relationships might be indicated by how dorsal colour patterns were derived. For example, he demonstrated how a generalised ‘5N’ D. scabra pattern could have given rise to the ‘5L’ D. confusa pattern, leading to the D. palmarum pattern (saddles and lateral bars confluent and much narrowed with increased inter-saddle areas), and how the marked D. medici pattern could have been derived from ‘5N’. His diagrammatic scheme of evolutionary history, in which the more derived species are smaller (body length), have better developed lateral body serration, and increased pitting on the head, places D. inornata as the basal species, followed by D. fasciata, “montane” (= D. atra), D. palmarum, and ‘D. scabra’ (including ‘5N’ = D. scabra, ‘5L’ = D. confusa and ‘5I’ = D. arabica sp. nov.), with D. medici as the most derived species. Gans (1959: 172) noted that D. scabra may have been derived “separately from the montane population [= D. atra], representing an invasion of the savannah habitat from the forest, and D. palmarum would be derived from scabra as a form adapted to the coastal dry belt of central West Africa.” However, Gans (1959: 173) warned that in Dasypeltis “…the use of different characters permitted the erection of sharply differing phylogenies.”

It is now generally accepted that the genus Dasypeltis, along with the mainly African genera Crotaphopeltis, Telescopus, Dipsadoboa, Toxycodryas and Boiga, is part of the Boigini clade of the colubrid subfamily Colubrinae (Gravlund 2001; Kelly et al. 2003; Lawson et al. 2005; Figueroa et al. 2016). Relationships within the genus, however, are not well understood, with only three published molecular phylogenies, one (16S only) dealing mainly with West African species (Trape et al. 2012), another using the same data but with two samples of D. bazi from Egypt added (Saleh & Sarhan 2016), and a multigene study that used the same species as Trape et al. (2012) but added additional samples, including three ‘D. atra’ (Figueroa et al. 2016). The phylogeny of Trape et al. (2012) indicated that D. confusa (samples from Togo) was the sister group to all other West African species (including D. fasciata) as well as D. scabra from ‘South Africa” and ‘D. cf. lineolata’ (possibly D. atra) from Burundi. Trape et al.’s (2012) study supported species status for all West African taxa (including D. latericia, previously considered a subspecies of D. gansi). It also suggested a close relationship between D. fasciata, D. gansi and D. latericia, and between D. sahelensis and D. parascabra. With the inclusion of one additional species, D. bazi, Saleh & Sarhan’s (2016) phylogeny suggested different relationships, but most of these were not well supported (e.g. D. scabra was paired with D. latericia, and D. confusa paired with D. cf. lineolata [? D. atra]). The phylogeny of Figuero et al. (2016) identified two main clades, one with D. confusa + D. fasciata versus the rest, but D. fasciata was also included in the larger clade. This latter clade was divided into two well supported subclades, one consisting of D. atra, D. scabra and D. latericia, and the other containing D. atra, D. gansi, D. fasciata and D. sahelensis. The confounding positions of both D. fasciata and D. atra in different clades may suggest the presence of cryptic species, but alternatively it may reflect incorrect identification of specimens. The relationships indicated by these studies may change or be resolved further with greater taxon sampling, particularly of north-eastern, eastern and southern African lineages (A. Barlow et al. in prep.).

Geniez & Guillod (2003) considered the populations of Dasypeltis in Morocco, Egypt and south-western Arabia as being Afrotropical relicts. Dasypeltis bazi appears to have a relict distribution indicative of a former widespread range in northern Africa that was reduced and fragmented following dessication of the region during the Holocene (Kuper & Kröpelin 2006). Oases such as the Fayoum appear to be vestiges of what was once an expansive savannah, probably with riparian-fringed links like the Nile River, and may act as climatic refuges. The geographical ranges of some snakes in Egypt, such as Telescopus dhara obtusus (Reuss), are closely associated with the Nile Valley and this species also occurs in apparent isolation on the Red Sea coast in the south-east of the country. Its distribution range may be similar to the past distribution of D. bazi, a species that is perhaps also represented by an apparently isolated population near the Red Sea coast at Erkowit.

During the Last Glacial Maximum and terminal Pleistocene (21000–8500 B.C.E.) the Sahara desert extended about 400 km further south than it does now, probably incorporating, inter alia, all of modern-day Sudan (Kuper & Kröpelin 2006). Monsoon rains that started 8500 B.C.E. resulted in hyperarid conditions being replaced by savannah-like environments, similar to the current-day Sahel, with formation of lakes and temporary rivers (Baha El Din 2006), perhaps allowing Dasypeltis (and many other animal species) from areas in central Africa to disperse and establish populations as far north as Morocco and Egypt. However, retreating monsoonal rains from 5300 B.C.E. resulted in dessication of the Eastern Sahara, with a return to full desert conditions throughout Egypt by 3500 B.C.E. (Kuper & Kröpelin 2006). Most Afrotropical animal species (or at least most populations) would then have retreated southwards as the only favourable (non-desert) habitat remaining in the region would have been in the Nile Valley and at scattered oases (Baha El Din 2006) such as Fayoum, Siwa, Qara, Bahariya, Farafra, Dakhla and Kharga. Although populations of Dasypeltis in Egypt (Fayoum and possibly in the Nile Valley itself) could be relics of a former widespread species that became established there during favourable conditions, they may alternatively be the descendants of populations that dispersed north along the Nile Valley in later times, using the valley as a ‘green corridor’ (see also Escoriza 2010). According to Baha El Din (2006), the Nile River (and its associated valley) varied considerably in size and impact during the Quaternary, before achieving its current form; and its function as a conduit for African fauna and flora was established as recently as 800 000 BP. Saleh & Sarhan’s (2016) study indicated that divergence times between D. bazi and other African Dasypeltis species predate the formation of the Nile River by about a million years, and this together with the absence of Dasypeltis records in the Nile Valley [in Egypt], suggests that the occurrence of these snakes in Fayoum is the result of vicariant events related to the formation of the Sahara rather than by dispersion across this arid desert.

Geniez & Guillod (2003) were of the opinion that while the Nile Valley represented a northward corridor of the Ethiopian region for some species of sub-Saharan reptiles, the Moroccan population (now known as D. sahelensis, with most of its range in the Sahel; Trape & Mané 2006) is isolated. According to Escoriza (2010), the occurrence of ‘D. scabra’ and ‘Boaedon fuliginosus’ (Boie) in northern Africa may be the result of range expansions during humid paleoclimatic conditions in the late Pleistocene or recent Holocene. Escoriza (2014) suggested that populations of D. sahelensis crossed the Western Sahara desert from the Sahel (or associated savannahs) to northwestern Africa (Western Sahara and Morocco) in humid conditions 6000–9000 years ago during the Holocene. This species, together with populations of the widespread snakes ‘B. fuliginosus’ and Bitis arietans (Merrem), was subsequently isolated in the relatively moist Atlantic coastal region of north-western Africa during the next period of aridification (Schleich et al. 1996; Geniez & Guillod 2003).

The nearest record to D. bazi from the Fayoum Oasis in Egypt is Erkowit in eastern Sudan near the Red Sea coast, but the latter town is situated about 1330 km away to the south-east. The specimen from Erkowit (D. cf. bazi) may represent an isolated relict population of D. bazi. This specimen is similar to Egyptian specimens of D. bazi, both in terms of colour pattern and scalation. This Egyptian/Erkowit population may in the past have been part of a more widespread population of D. scabra that became isolated following aridification of the Sahara during the Holocene, resulting in the evolution of a new species, D. bazi. The two populations may then have become isolated as climatic and environmental conditions changed, with the Fayoum Oasis (D. bazi) and Red Sea Hills (D. cf. bazi =  Erkowit population) functioning as refuges, and accounting for their large current-day separation. Erkowit is a small town situated on a minor plateau (mostly 1000–1200 m a.s.l.) in the Red Sea Hills—a northern extension of the Ethiopian Plateau—at the eastern edge of the arid Sahel. Sea mist is blown in by north-easterly winds during winter to provide a moisture-temperature gradient at Erkowit, which has evergreen Afromontane vegetation only 10 km from the surrounding desert shrub (Vetaas et al. 2012). The Erkowit area was once regarded as a mist oasis in the desert, representing a potential refugium for Dasypeltis. Baha El Din (2006) suggested that some species of Afrotropical reptiles extended their ranges into Saharan Africa using the Nile River (and its associated valley) and Red Sea coast and mountains as conduits, although some populations in these areas are merely relicts. The cobra Naja nubiae Wüster & Broadley, 2003, for example, has a patchy distribution in eastern Chad, Sudan, Eritrea and at sites along the Nile River in Egypt.

Corkhill (1935) noted that in Sudan, D. scabra occurred only in ‘Blue Nile Province’—a region which his map indicates as being situated slightly east and south of Khartoum—but he did not list any specific localities. There are in fact four known Dasypeltis records for Sudan and one for South Sudan (none appear to be referrable to D. bazi), all but one located at or near the banks of the White Nile River. The exception is the newly-reported specimen (ZFMK 32479) from 50 km north of Kutum in Sudan, situated 1750 km from Fayoum and 1395 km WSW of Erkowit, identifiable as D. scabra (see above). One of the two syntypes of Rachiodon abyssinus (= Dasypeltis abyssina), collected in the mid-19th century during an expedition to the sources of the White Nile under the direction of M. d’Arnaud, and labelled “Nil Blanc” (= White Nile, possibly above Khartoum in Sudan), was examined by the author and is in fact a D. scabra (MNHN 7227, female, ‘5N’ dorsal pattern – dark-bordered brown saddles with vertically-elongate lateral bars between them, 54 pattern cycles, 226 ventrals, 53 pairs of subcaudals; Fig. 6). Khartoum, where the White and Blue Niles converge, is about 600 km south-west of Erkowit. A specimen (BM 09.10.15.35) of D. scabra from ‘Sennar State’ (which straddles the White Nile; Fig. 3) in south-eastern Sudan has some ‘5N’ markings dorsally, but also the peculiar ‘5I’ dorsal pattern over parts of its back (photograph examined) as described by Gans (1959) for some populations of Eritrean and Arabian ‘D. scabra’, where the dark saddles and lateral bars are joined, enclosing a pale inter-saddle region. This female specimen was in fact examined by Gans (1959) who recorded 65 pattern cycles, 227 ventrals and 52 pairs of subcaudals. The fourth Sudanese specimen (RMNH 24399, female) was collected 40 km north of Rabak, near Kosti (Fig. 3), on the banks of the White Nile and has the typical ‘5N’ D. scabra dorsal pattern, 50 pattern cycles, 222 ventrals and 55 subcaudals. There is one additional D. scabra record (FMNH 134939: male, ‘5N’ pattern, 64 pattern cycles, 213 ventrals, 62 pairs of subcaudals) on the White Nile (east bank) at Melut in the north-eastern part of South Sudan. Other records of Dasypeltis in adjacent parts of Eritrea and north-western Ethiopia are referable to D. taylori sp. nov., D. abyssina, and D. atra (see below). The four above-mentioned ‘5N’ D. scabra records on the White Nile, possibly representing a single population, appear to be considerably isolated from other populations of this species. The northern-most D. scabra record in the Nile Valley is Sennar State, about 700 km south-west of Erkowit. Apart from Erkowit, the nearest other D. scabra locality is the Addis Abeba area (Largen & Spawls 2010; Fig. 3) about 740 km ESE of Melut. There is one additional (apparently isolated) record of D. scabra from Wadelai, much further upstream, on the Albert Nile in north-western Uganda (Fig. 3).

As discussed in the species account, some Eritrean specimens of D. abyssina have the ‘5I’ dorsal pattern typical of D. arabica sp. nov. over parts of their backs. This, together with other morphological similarities (e.g. high numbers of ventrals and subcaudals) between D. abyssina and D. arabica, together with their geographical proximity, suggests a close evolutionary relationship. A continuous population may have been separated by the opening of the Gulf of Aden (see Leviton 1986), with subsequent divergence of the resultant sub-populations. The dorsal colour pattern of D. crucifera sp. nov. is similar to this ‘5I’ pattern and the close geographical proximity suggests that it may be closely related to the high-elevation D. abyssina. Dasypeltis taylori sp. nov. is separated from the nearest populations of D. scabra—a species it most resembles—by the arid Ogaden plateau, and may represent a relictual population of a once widespread D. scabra.

A molecular phylogeny of the genus, including samples of all species and with reasonable coverage of their ranges, will go a long way towards elucidating evolutionary relationships. Considering the vast areas covered by the ranges of most species, the possibility of additional (possibly cryptic) species cannot be discounted. Sampling of all species may be difficult at this time as some parts of Africa are still relatively inaccessible, not only physically, but in some cases also as a result of armed conflicts (e.g. parts of Somalia, Sudan, South Sudan, Eritrea-Djibouti border). Relationships and levels of divergence for the various isolated populations of D. scabra (e.g. 50 km N of Kutum, Sudan; populations along the Nile River in Sudan/South Sudan; Wadelei, Uganda) and D. cf. bazi (Erkowit, eastern Sudan) should also be investigated.

Key to the genus Dasypeltis in north-eastern Africa and south-western Arabia

1a. Paired apical pits on dorsal scales sharply defined by dark pigment; frontal shield extensively (State 3) to entirely (State 4) pitted; patterned specimens with 1–5 distinct chevrons on neck; subcaudals 75–109 in males, 61–90 in females ………………… D. medici

1b. Paired apical pits on dorsal scales not sharply defined by dark pigment; frontal varies from smooth to extensively pitted (States 0–4); when present, chevrons restricted to nape ………………………………………………………………………………………………………………………….. 2

2a. Dorsum patterned, with body pattern always expressed as chequerboard on skin between scales and often also on lower edges of these; frontal usually marginally pitted (States 1–2); lateral scales always strongly serrated; subcaudals 71–76 (once 59) in males, 64–76 in females ………………………………………………………………………………………………… D. fasciata

2b. Dorsum without pattern or with pattern expressed across the scales; frontal varies from smooth to extensively pitted (States 0–4); lateral serration present in varying degrees; subcaudals 47–73 in males, 43–65 in females ………………………………………………………… 3

3a. Dorsum uniform black, brown, grey or red, or pale brown or grey with a pattern of dense, dark brown, olive or reddish, usually transversely-enlarged, saddles, often joined to the lateral bars; when present 62–130 (mean 92.6) pattern cycles from nape to base of tail; postoculars 1 or 2 on either side of head ……………………………………………………….. D. atra

3b. Dorsum light brown or grey, uniform, or with a pattern of darker dorsal markings linked or alternating with lateral bars; 39–98 pattern cycles from nape to base of tail; usually 2 postoculars on either side of head …………………………………………………………………………. 4

4a. Dark dorsal markings (saddles) usually large and diamond-shaped, linked on either side to the dark lateral bars (or dorsum uniform pale brown); ventrals 199–224 in males, 212–239 in females ………………………………………………………………………………………. D. confusa

4b. Dark dorsal markings (saddles) variable in size but not diamond-shaped, usually not linked on either side to the dark lateral bars (or dorsum uniform pale brown in D. abyssina, but then ventrals 226 in males, 241–263 in females) …………………………………. 5

5a. Dark dorsal markings (saddles) and lateral bars completely or largely enclose a pale saddle patch; venter unmarked except for small dark specks at the lateral edges; frontal marginally pitted (States 1 & 2). …………………………………………………………………………… 6

5b. Dark dorsal markings (saddles) and lateral bars mostly separated, not enclosing a pale saddle patch; markings on venter variable; frontal pitting variable…………………………….. 7

6a. Pale markings on the back more-or-less square and completely surrounded by dark rectangular dorsal saddles and lateral bars; pattern cycles on back 62–79; inter-prefrontal sulcus moderately to deeply marked; ventrals 236–244 in males, 239–254 in females; subcaudals 63–65 in males, 53–61 in females ………………………………. D. arabica sp. nov.

6b. Pale markings on the back in the form of crosses, largely surrounded by dark oval dorsal saddles and lateral bars, but with the tips of the crosses often separated from the dark markings; pattern cycles on back 51–62; inter-prefrontal sulcus faintly marked; ventrals 226 in males, 231–247 in females; subcaudals 61 in males, 49–52 in females ……………………………………………………………………………………………… D. crucifera sp. nov.

7a. Dorsal markings (saddles) often oblique and wider than long, almost confluent or forming zig-zags, usually not linked to lateral bars; parts of the labials and interspaces between saddles are cream-yellow (in life); venter with heavy dark markings, each plate decorated by, anteriorly, a large rounded cone on either side joined to a smaller rounded cone in the centre; uniform phase pale grey-brown above, cream below; inter-prefrontal sulcus usually faintly marked; frontal usually marginally pitted (States 1 or 2); ventrals 226–247 in males, 241–271 in females ………………………………………………………………… D. abyssina

7b. Dorsal markings (saddles) usually not oblique, variable in shape (e.g. round, oval, square, rectangular), usually not confluent or forming zig-zags; no yellowish markings; venter varies from immaculate to heavily-marked, but not as extensively as above; inter-prefrontal sulcus variable; frontal pitting variable; ventrals 185–229 in males, 202–249 in females ……………………………………………………………………………………………………………… 8

8a. Dorsal markings (saddles) variable, often oval to rectangular (antero-posteriorly); pattern cycles 46–94 (mean 67.6); markings on venter variable; ventrals 185–226 in males, 202–249 in females ………………………………………………………………………………………… D. scabra

8b. Dorsal markings (saddles) large, often broad or elongate; pattern cycles 38–59; venter usually unmarked except for a few dark spots laterally ……………………………………………. 9

9a. Dorsal markings (saddles) cranio-caudally elongate, well separated, usually bifurcated at each end with a shallow but distinct notch; flanks with spots or blotches; ventrals 213–229 in males, 215–234 in females ………………………………………………………………… D. bazi

9b. Dorsal markings (saddles) broad, often squarish, usually separated by narrow sharply-defined white crossbars; flanks with elongate, wavy markings; ventrals 196–216 in males, 208–222 in females ………………………………………………………….. D. taylori sp. nov.

Acknowledgements

We thank the following museums and their curators and collection managers for providing access to specimens: Natural History Museum, London (C.J. McCarthy, P. Campbell); Museum National d’Histoire Naturelle, Paris (I. Ineich, N. Vidal); Museum für Naturkunde, Berlin (R. Günther, F. Tillack); Naturhistorisches Museum, Vienna (Silke Schweiger); Musée Royal de l’Afrique Centrale, Tervuren (D. Meirte); Museum of Comparative Zoology, Harvard (J.P. Rosado); Field Museum of Natural History, Chicago (A. Resetar); American Museum of Natural History, New York (C.W. Myers, D. Kizirian, D.A. Dickey); United States National Museum, Washington (G.R. Zug); University of Michigan Museum of Zoology, Ann Arbor (A.G. Kluge); California Academy of Sciences, San Francisco (R.C. Drewes, J.V. Vindum); Addis Ababa University Museum (G. Redda); Makerere University Zoological Museum, Kampala (R. Kityo); National Museums of Kenya, Nairobi (P.K. Malonza); Port Elizabeth Museum/Bayworld (W.R. Branch, W. Conradie); and KwaZulu-Natal Museum, Pietermaritzburg (C. Stoffels).

We thank the curators of the following museums for sending us material on loan: Los Angeles County Museum (R. Bezy); Museum Alexander Koenig, Bonn (W. Böhme); Muséum d’Histoire Naturelle, Geneva (A. Schmitz); National Museum of Kenya, Nairobi (P.K. Malonza); Naturhistorisches Museum, Vienna (H. Grillitsch); and Statens Naturhistoriske Museum, Københavns Universitet, Copenhagen (J.B. Rasmussen); and the curators and other individuals associated with the following museums for providing us with meristic data and/or images of specimens in their care: Museo Civico di Storia Naturale, Genova (G. Doria, M. Petri); Museo Civico di Storia Naturale, Milan (S. Scali); Museo di Storia Naturale, Università di Pavia, Pavia (E. Razzetti); Museo Zoologico “La Specola”, Università di Firenze (A. Nistri); Museo Regionale di Scienze Naturali di Torino, Turin (F. Andreone); Museum Alexander Koenig, Bonn (W. Böhme, M. Flecks, U. Bott); Statens Naturhistoriske Museum, Københavns Universitet, Copenhagen (J.B. Rasmussen); Institut Royal des Sciences Naturelles de Belgique, Brussels (G. Lenglet); National Museum of Kenya, Nairobi (P.K. Malonza); Port Elizabeth Museum/Bayworld (W.R. Branch, W. Conradie); Natural History Museum, London (P. Campbell); Museum National d’Histoire Naturelle, Paris (I. Ineich); Zoologisches Museum der Universität, Berlin (F. Tillack), Museum Alexander Koenig, Bonn (W. Böhme and U. Bott); California Academy of Sciences, San Francisco (J. Vindum); and Texas A&M University, Texas (T. Hibbits).

We also thank B. Hughes for providing scalation data for Ugandan specimens in the collection of the World Museum, Liverpool; the following individuals for photographs of Dasypeltis (see figures for credits): W.R. Branch, F. Finke, E. Greenbaum, T. Mazuch, S. Spawls, C. Tilbury and V. Trailin; V. Trailin for additional photographic records of D. scabra and D. atra in Ethiopia; S. Spawls for additional photographic records of D. atra and D. scabra in Kenya, and D. atra in Tanzania; T. Mazuch for additional photographic records of D. scabra and D. atra in Ethiopia; J. Marais for photographic records of Kenyan D. medici (Malindi) and D. atra (Emakoko); T. Dudu for photographic records of D. atra from Serengeti in Tanzania; C. Doria for a photographic record of a melanistic D. atra from Oldeani in Tanzania; A. Bauer, T. Mazuch, F. Tillack, D. Meirte, C. Kucharzewski, M.-O. Rodel, I. Ineich and R. Charles for access to old literature; S. Spawls for helping to trace various localities; and U. Ziliani for helping to trace records of D. scabra from Turkana County in Kenya. Frank Tillack provided collection details for Egyptian specimens in the collection in Berlin.

The first author’s trip to the natural history museums in London, Paris and Berlin in September and October 2011 was funded by the Reptile Speciation Project (a South African National Research Foundation [NRF] grant to K.A. Tolley, South African National Biodiversity Institute, Cape Town), his visit to the Royal Museum for Central Africa in Tervuren in October 2012 was made possible by an African Biodiversity Information Centre scholarship, and a later trip (September/October 2015) to the museum in Paris was funded by the National Museum (Bloemfontein).

We are indebted to J. Kersten (Bulawayo) and M. van Rensburg (National Museum, Bloemfontein) for preparing the distribution map; Zhongning Zhao (University of the Free State, Bloemfontein) for assistance with multivariate analyses; and W.R. Branch (Port Elizabeth Museum) for commenting on parts of the Discussion. The reviewers, Prof. A.M. Bauer (Villanova University, Philadelphia) and S. Spawls (Norwich, England) are thanked for their valuable comments and suggestions for improving the manuscript.

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Table 1. Numbers of pattern cycles and other pattern morphs in Dasypeltis from north-eastern Africa and south-western Arabia. Codes in parentheses under ‘Range’ and ‘Other morphs’ are those used by Gans (1959) for the various dorsal colour patterns.

  N Range Mean S.D. Other morphs
D. medici 24 57–108 (8) 74.0 11.9 Faintly patterned to uniform (3): 37

Total patterned (8): 35

D. confusa 30 50–79 (5L) 61.7 7.1 Uniform brown: 11
D. scabra 107 46–94 (5N) 67.4 10.0
D. bazi (Egypt) 5 39–49 (5N) 45.2 4.4
D. cf. bazi (Sudan) 1 48
D. taylori sp. nov. 8 44–59 (5N) 52.8 5.1
D. atra 51 62–130 (2Bx) 91.8 14.1 Melanistic (2M): 89, Brown (2B): 91, Total 2Bx: 135
D. fasciata 5 108–141 (4) 117.8 13.3
D. abyssina 11 65–98 (5N, 5I) 80.7 11.4 Uniform brown/tan: 7
D. crucifera sp. nov. 3 51-62 (5I) 55.0
D. arabica sp. nov. 6 62–79 (5I) 70.0 5.7

Table 2. Ventral and subcaudal scale counts in Dasypeltis from north-eastern Africa and south-western Arabia. Values for males are in bold.

                                      VENTRALS                                                                                                     SUBCAUDALS

Males                                                    Females                                               Males                                               Females

N Range Mean S.D. N Range Mean S.D. N Range Mean S.D. N Range Mean S.D.
D. medici 29 220–252 235.8 8.1 49 218–252 234.4 7.3 27 75–109 88.3 8.1 47 61–90 77.5 5.7
D. confusa 14 199–224 209.9 7.1 30 212–239 226.1 7.9 12 51–67 60.2 5.3 24 48–61 54.1 3.0
D. scabra 50 185–226 209.3 8.9 76 202–249 226.9 8.5 45 47–73 62.7 5.6 72 43–65 53.6 4.8
D. bazi (Egypt) 3 219–229 225.7 5.8 3 227–234 229.7 3.8 3 59–60 59.3 0.6 3 50–55 52.0 2.6
D. cf. bazi (Sudan) 1 229     1 67    
D. taylori sp. nov. 5 196–216 202.2 8.0 3 208–222 213.7 7.4 4 55–68 60.5 5.4 2 44–52 48.0 5.7
D. atra 109 199–232 212.7 6.8 167 211–256 225.7 7.9 103 50–71 61.9 3.9 151 45–64 53.5 3.3
D. fasciata 3 212–230 223.7 10.1 9 237248 242.7 3.8 3 59–76 68.7 8.7 9 6476 68.3 3.7
D. abyssina 6 226–247 235.5 8.2 10 241271 256.0 9.3 4 57–68 61.5 4.6 9 4965 56.7 5.0
D. crucifera sp. nov. 1 226     3 231–247 240.3 8.3 1 61     3 49–52 50.3 1.5
D. arabica sp. nov. 3 236–244 239.3 4.2 4 239–254 244.8 6.9 3 63–65 63.7 1.2 4 53–61 56.8 3.5

Table 3. Loading scores for Principal Component Analysis of Dasypeltis males in north-eastern Africa and south-western Arabia.

      PC 1       PC 2
Midbody scale rows 0.2344 –0.3343
Ventrals 0.3292 0.3966
Subcaudals 0.3438 0.5406
Supralabials 0.0471 0.0288
Preoculars 0.0193 0.2555
Postoculars 0.4007 –0.2850
Anterior temporals 0.3447 0.0854
Posterior temporals 0.3890 –0.2412
Pattern cycles 0.3316 –0.2163
Frontal pitting 0.3194 0.2843
Inter-prefrontal sulcus 0.2706 –0.3135

Table 4. Loading scores for Principal Component Analysis of Dasypeltis females in north-eastern Africa and south-western Arabia.

   PC 1       PC 2
Midbody scale rows 0.2960 0.2813
Ventrals 0.2792 –0.0893
Subcaudals 0.3261 –0.4928
Supralabials 0.1168 0.0920
Preoculars 0.1024 0.3277
Postoculars 0.4236 0.1670
Anterior temporals 0.3542 –0.2803
Posterior temporals 0.4146 –0.0816
Pattern cycles 0.2204 0.4892
Frontal pitting 0.3748 –0.1808
Inter-prefrontal sulcus 0.1955 0.4113

Table 5. Observed (rows) and predicted (columns) classifications of male Dasypeltis from north-eastern Africa and south-western Arabia according to the Discriminant Function Analysis.

% correct D.

atra

p = .430

D. scabra

p = .258

D. confusa

p = .065

D. medici

p = .097

D. fasciata

p = .022

D. taylori

p = .032

D. abyssina

p = .043

D. arabica

p = .022

  D.

bazi

p = .032

D. atra 85.0 34 5 0 0 0 0 1 0 0
D. scabra 83.3 2 20 1 0 0 0 0 0 1
D. confusa 50.0 1 2 3 0 0 0 0 0 0
D. medici 100 0 0 0 9 0 0 0 0 0
D. fasciata 100 0 0 0 0 2 0 0 0 0
D. taylori 66.7 0 1 0 0 0 2 0 0 0
D. abyssina 100 0 0 0 0 0 0 4 0 0
D. arabica 100 0 0 0 0 0 0 0 2 0
D. bazi 66.7 1 0 0 0 0 0 0 0 2
Total 83.9 38 28 4 9 2 2 5 2 3

Table 6. Standardised Canonical Discriminant Function Coefficients (Sigma-restricted parameterisation) for the analysis of male Dasypeltis in north-eastern Africa and south-western Arabia.

Function 1 Function 2
Midbody scale rows –0.001839 0.229281
Ventrals 0.604129 –0.405545
Subcaudals 0.812478 0.007712
Supralabials 0.137091 –0.034382
Preoculars –0.006490 –0.090852
Postoculars 0.099070 0.304857
Anterior temporal 0.007210 –0.104399
Posterior temporal –0.075543 0.170241
Pattern cycles –0.085729 0.541770
Frontal pitting 0.190038 0.669032
Inter-prefrontal sulcus 0.020179 0.300686
Eigenvalue 6.032452 2.042220
Cumulative Proportion 0.591676 0.791982

Table 7. Observed (rows) and predicted (columns) classifications of female Dasypeltis from north-eastern Africa and south-western Arabia according to the Discriminant Function Analysis.

% correct D.

atra

p = .400

D. scabra

p = .258

D. confusa

p = .084

D. medici

p = .129

D. fasciata

p = .019

D. taylori

p = .013

D. abyssina

p = .045

D. arabica

p = .026

  D.

bazi

p = .013

D. crucifera

p = .013

D. atra 74.2 46 6 6 0 0 0 3 1 0 0
D. scabra 75.0 2 30 3 0 0 2 0 2 1 0
D. confusa 61.5 3 2 8 0 0 0 0 0 0 0
D. medici 95.0 1 0 0 19 0 0 0 0 0 0
D. fasciata 100 0 0 0 0 3 0 0 0 0 0
D. taylori 50.0 0 1 0 0 0 1 0 0 0 0
D. abyssina 100 0 0 0 0 0 0 7 0 0 0
D. arabica 50.0 1 1 0 0 0 0 0 2 0 0
D. bazi 0 1 1 0 0 0 0 0 0 0 0
D. crucifera 50.0 1 0 0 0 0 0 0 0 0 1
Total 75.5 55 41 17 19 3 3 10 5 1 1

Table 8. Standardised Canonical Discriminant Function Coefficients (Sigma-restricted parameterisation) for the analysis of female Dasypeltis in north-eastern Africa and south-western Arabia.

Function 1 Function 2
Midbody scale rows 0.155036 –0.307790
Ventrals 0.174019 0.644477
Subcaudals 0.919766 0.182047
Supralabials 0.000409 0.009540
Preoculars –0.005440 0.252592
Postoculars 0.203638 –0.274530
Anterior temporal 0.420286 0.032552
Posterior temporal –0.077620 0.051138
Pattern cycles –0.041610 –0.269700
Frontal pitting 0.256583 –0.673860
Inter-prefrontal sulcus –0.110640 –0.282740
Eigenvalue 5.033508 1.395425
Cumulative Proportion 0.627753 0.801783

Table 9. Loading scores for Principal Component Analysis of Dasypeltis males in the Horn of Africa and Egypt.

    PC 1      PC 2
Midbody scale rows 0.3324 –0.0988
Ventrals 0.1386 –0.4114
Subcaudals 0.0951 0.1427
Supralabials 0.1531 –0.3130
Postoculars 0.5229 –0.1184
Anterior temporal 0.2389 0.0896
Posterior temporal 0.3864 0.1763
Pattern cycles 0.5052 –0.1264
Frontal pitting 0.1129 0.6546
Inter-prefrontal sulcus 0.3001 0.4529

Table 10. Loading scores for Principal Component Analysis of Dasypeltis females in the Horn of Africa and Egypt.

       PC 1        PC 2
Midbody scale rows 0.4654 –0.2276
Ventrals –0.3110 0.4003
Subcaudals 0.0721 0.2901
Supralabials –0.2623 0.3800
Preoculars 0.1817 0.3029
Postoculars 0.3175 0.3540
Anterior temporal 0.3243 0.0786
Posterior temporal 0.3693 0.2800
Pattern cycles 0.4324 –0.0104
Frontal pitting 0.0859 –0.4338
Inter-prefrontal sulcus 0.2073 0.2637

Table 11. Observed (rows) and predicted (columns) classifications of male Dasypeltis from the Horn of Africa and Egypt according to the Discriminant Function Analysis.

% correct D.

atra

D. scabra D. taylori D. abyssina D. arabica   D.

bazi

p = .280 p = .240 p = .120 p = .160 p = .080 p = .120
D. atra 100 7 0 0 0 0 0
D. scabra 100 0 6 0 0 0 0
D. taylori 100 0 0 3 0 0 0
D. abyssina 100 0 0 0 4 0 0
D. arabica 100 0 0 0 0 2 0
D. bazi 66.7 1 0 0 0 0 2
Total 96.0 8 6 3 4 2 2

Table 12. Standardised Canonical Discriminant Function Coefficients (Sigma-restricted parameterisation) for the analysis of male Dasypeltis in the Horn of Africa and Egypt.

Function 1 Function 2
Midbody scale rows 2.08508 –0.334990
Ventrals 2.51335 –0.269100
Subcaudals 0.08589 –0.155450
Supralabials 0.05782 –0.447450
Postoculars –1.36866 0.590286
Anterior temporal 1.34341 0.105930
Posterior temporal 0.11259 0.006628
Pattern cycles –0.70219 –0.824390
Frontal pitting 0.13149 –0.68330003
Inter-prefrontal sulcus –0.36369 –0.534840
Eigenvalue 42.25546 2.395576
Cumulative Proportion 0.88362 0.933713

Table 13. Observed (rows) and predicted (columns) classifications of female Dasypeltis from the Horn of Africa and Egypt according to the Discriminant Function Analysis.

% correct D.

atra

D. scabra D. taylori D. abyssina D. arabica   D.

bazi

D. crucifera
p = .237 p = .316 p = .053 p = .184 p = .105 p = .053 p = .053
D. atra 77.8 7 1 0 1 0 0 0
D. scabra 75.0 1 9 0 0 1 1 0
D. taylori 100 0 0 2 0 0 0 0
D. abyssina 100 0 0 0 7 0 0 0
D. arabica 75.0 0 1 0 0 3 0 0
D. bazi 50.0 0 1 0 0 0 1 0
D. crucifera 100 0 0 0 0 0 0 2
Total 81.6 8 12 2 8 4 2 2

Table 14. Standardised Canonical Discriminant Function Coefficients (Sigma-restricted parameterisation) for the analysis of female Dasypeltis in the Horn of Africa and Egypt.

Function 1 Function 2
Midbody scale rows –0.187479 0.117839
Ventrals 1.074553 0.262493
Subcaudals –0.216661 1.018205
Supralabials –0.492329 –0.208635
Preoculars 0.097186 0.420874
Postoculars 0.178724 –0.409874
Anterior temporal –0.063169 0.143802
Posterior temporal 0.379009 0.083376
Pattern cycles –0.460762 0.680089
Frontal pitting –0.298385 –0.084354
Inter-prefrontal sulcus –0.280103 0.377127
Eigenvalue 3.706452 2.549360
Cumulative Proportion 0.462892 0.781277

Appendix 1. Morphological data for specimens of Dasypeltis used in this study. Museum catalogue numbers with asterisks denote specimens with full sets of data used in the multivariate analyses; a few specimens with full data, obtained subsequent to these analyses, are marked by +. Under ‘Sex’: M = male, F = female, U = undetermined. Counts for supralabials, preoculars, postoculars, anterior temporals and posterior temporals include both sides of the head. Under ‘Cycles’: B denotes plain brown or greyish specimens with no discernible dorsal pattern; for D. medici, patterned specimens for which cycle counts are not available are indicated as P, while dorsally uniform or weakly marked specimens are coded as L. For D. atra, patterned specimens for which cycle counts are not available are indicated as ‘2Bx’, while specimens of the ‘lineolata’ morph (a variety of ‘2Bx’) are indicated as ‘lin’ (see Gans 1959). Frontal pitting: 0 = smooth, 1 = marginally pitted (States 1 & 2), 2 = extensively pitted (States 3 & 4); Inter-prefrontal sulcus: 0 = weakly marked, 1 = moderately marked, 2 = deeply sunken.

Species Catalogue No. Country Locality Sex SVL Tail SVL/Tail Midbody scale rows Ventrals Subcaudals Supralabials Preoculars Postoculars AnterTemps PosterTemps Pattern cycles Frontal  pitting Inter-prefrontal sulcus
D. abyssina NMW 15301/2* ERITREA ? M 412 74 5.568 23 239 60 14 2 4 4 5 77 1 1
D. abyssina NMW 15301/3 ERITREA ? M 504 24 247 14 2 4 4 6 93 1 0
D. abyssina BM 69.11.4.38* ERITREA Senafe M 270 45 6 24 239 61 14 2 4 4 5 67 1 0
D. abyssina FMNH 12728* ETHIOPIA Gondar M 348 57 6.105 23 236 57 14 2 4 4 5 95 1 0
D. abyssina BM 1902.12.13.85 ETHIOPIA Zegie, Lake Tsana M 23 226 2 4 4 8 65 0 0
D. abyssina BM 1902.12.13.86* ETHIOPIA Zegie, Lake Tsana M 235 44 5.341 23 226 68 14 2 2 4 3 B 1 0
D. abyssina NMW 15301/1* ERITREA ? F 539 80 6.738 23 251 54 14 2 4 4 7 73 1 0
D. abyssina NMW 15300* ERITREA Gheleb F 540 72 7.5 24 271 54 14 2 4 4 7 81 1 0
D. abyssina MNHN 6567 ETHIOPIA ? F 665 90 7.389 25 260
D. abyssina MSNM 2699A ETHIOPIA Gondar F 655 91 7.198 22 248 59 14 2 4 4 B
D. abyssina MZUF 783* ETHIOPIA Gondar F 416 52 8 25 254 49 14 2 4 4 6 98 1 1
D. abyssina NMBO 10256* ETHIOPIA Keriyo F 422 52 8.115 25 264 51 14 2 4 4 6 89 1 0
D. abyssina NMBO 10257* ETHIOPIA Keriyo F 483 87 5.552 22 241 65 14 2 4 4 7 B 1 0
D. abyssina NMZB 17099* ETHIOPIA Keriyo F 475 70 6.786 22 247 59 14 2 4 3 6 B 1 0
D. abyssina MSNM 2700A ETHIOPIA Mai Canetta F 663 84 7.893 21 263 60 14 2 4 4 B
D. abyssina MCZ 80984* ETHIOPIA Ras Dashen Terara F 511 74 6.905 24 261 59 14 2 4 4 6 B 1 0
D. abyssina MSNM 2699B ETHIOPIA Gondar U 364 59 6.169 22 222 61 14 2 4 4 73
D. abyssina MSNM 2699C ETHIOPIA Gondar U 240 22 228 14 2 4 4 77
D. abyssina MSNM 2700B ETHIOPIA Mai Canetta U 220 28 7.857 21 257 58 14 2 4 4 B
D. arabica MHNG 2536.42* SAUDI ARABIA vicinity of Abha M 24 238 63 14 2 4 4 6 69 1 1
D. arabica BM 03.6.26.41* YEMEN El Kubar M 325 56 5.804 23 244 65 14 2 4 4 8 72 1 1
D. arabica MSNG 52221 YEMEN El Siyani M 486 77 6.312 23 236 63 14 2 4 4 6
D. arabica BM 1987.1018* SAUDI ARABIA Ajibah F 246 60 4.1 23 246 55 14 2 2 4 6 79 1 1
D. arabica MHNG 2536.41* SAUDI ARABIA Aqabat Raydah F 43 25 254 61 14 2 4 4 6 71 1 2
D. arabica BM 1987.2192* YEMEN Sana’a F 625 89 7.022 25 240 53 14 4 4 5 7 67 1 2
D. arabica BM 1987.2193* YEMEN Taiz F 400 64 6.25 25 239 58 14 2 4 4 7 62 1 1
D. atra MRAC 18583 BURUNDI Kamagaba Marsh M 457 86 5.314 24 208 57 14 2 2 4 5 2Bx 1 0
D. atra MRAC 18585 BURUNDI Rumonge M 457 24 215 12 2 2 4 6 2Bx 1 0
D. atra NMW 9964/2+ ETHIOPIA ‘Abessinien’ M 350 65 5.385 24 209 57 14 2 4 4 7 76 1 0
D. atra MSNG 30412 ETHIOPIA Addis Ababa M 227 50 14 2 4 2 4 0 2
D. atra ZFMK 57537* ETHIOPIA Bedele, Ullubabor M 505 88 5.739 22 220 57 14 2 2 4 7 M 1 0
D. atra MNHN 05-209 ETHIOPIA Bourka M 478 23 230 14 2 4 4 6 B 1 0
D. atra MNHN 05-210* ETHIOPIA Bourka M 293 45 6.511 22 231 58 14 2 4 4 4 M 1 0
D. atra BM 1964.1068* ETHIOPIA Gemu Gofa District M 435 75 5.8 25 224 58 14 2 4 4 6 85 0 0
D. atra ZFMK 16297* ETHIOPIA Genale to Chamo M 504 87 5.793 25 214 55 14 2 4 4 6 B 0 0
D. atra BM 1973.3175 ETHIOPIA Ghimbi, Wollega M 390 76 5.132 61 14 2 4 4 6 M 1 0
D. atra MCZ 34923 ETHIOPIA Harar M 259 46 5.63 23 225 60 14 2 4 4 6 2Bx 1 1
D. atra NMW 9964-1* ETHIOPIA Harar M 352 58 6.069 22 232 58 14 2 4 4 6 B 1 1
D. atra MNHN 05-205 ETHIOPIA Laga Hardine M 328 58 5.656 25 211 61 14 2Bx 1 0
D. atra MNHN 05-206* ETHIOPIA Laga Hardine M 325 60 5.417 25 219 58 14 2 4 4 6 74 1 0
D. atra ZMB 27463 ETHIOPIA Lake Alemaya M 245 40 6.125 23 231 60 14 2 4 4 5 2Bx 1 1
D. atra ZFMK 16237* ETHIOPIA Lake Shala M 414 72 5.75 23 220 58 14 2 4 6 8 79 0 0
D. atra MSNG 29047A ETHIOPIA Shoa M 22 216 54 14 2 2 4 1 1
D. atra BM 96.5.19.57 ETHIOPIA Shoa (Ragazzi) M 315 49 6.429 22 221 59 14 2 4 4 7 M 0
D. atra NMK 483 KENYA ? M 453 96 4.719 216 67 14 2 B
D. atra CAS 148035* KENYA Chemelil, Kisumu M 420 83 5.06 23 209 62 14 2 3 4 6 M 1 0
D. atra CAS 122281* KENYA Chuka, Meru M 505 110 4.591 23 201 67 13 2 4 4 6 B 1 0
D. atra BM 04.12.23.10* KENYA Fort Hall (= Murang’a) M 183 33 5.545 26 221 67 14 2 4 4 6 106 1 0
D. atra NMK 3329 KENYA Kabarnet, Baringo M 454 199 14 2 B
D. atra NMK 479 KENYA Kabete, Kiambu M 520 97 5.361 215 57 14 2 B
D. atra MCZ 40577* KENYA Kaimosi M 346 64 5.406 24 211 65 14 2 3 4 6 M 1 0
D. atra CAS 142248* KENYA Kakamega Forest Stn. M 515 97 5.309 23 209 59 14 2 4 4 6 M 1 0
D. atra NMK 3007 KENYA Karen, Nairobi M 464 93 4.989 206 61 14 4 B
D. atra NMK 2952/1 KENYA Kiboko Ranch Station M 198 37 5.351 206 67 14 2 2Bx
D. atra NMK 2862 KENYA Kiganjo, Nyeri M 428 81 5.284 200 55 14 2 M
D. atra CG 2278 KENYA Lumbwa, Pen-y-Bryn M 508 99 5.131 23 213 62 2 3 4 6 106 1 0
D. atra CG 2279 KENYA Lumbwa, Pen-y-Bryn M 285 47 6.064 24 213 58 2 4 6 7 62 1 0
D. atra NMK 659 KENYA Mount Elgon M 434 94 4.617 211 60 B
D. atra LACM 50631* KENYA Mweiga M 476 96 4.958 23 210 59 15 2 2 4 6 B 1 0
D. atra NMK 3339 KENYA Nairobi M 483 90 5.367 222 61 14 2 76
D. atra NMK 3548 KENYA Nairobi M 421 85 4.953 212 62 14 2 95
D. atra NMK 403 KENYA Nairobi M 384 66 5.818 207 59 14 2 B
D. atra BM 1960.1.6.40* KENYA Ngong M 420 83 5.06 23 209 65 14 2 4 4 6 88 1 1
D. atra NMK 2576 KENYA North Nandi Forest, Chemisia M 456 98 4.653 208 59 14 2 M
D. atra NMK 2590 KENYA North Nandi Forest, Chemisia M 466 99 4.707 208 60 14 2
D. atra NMK 1497 KENYA Nyambene Hills, Meru M 421 87 4.839 207 61 12 2 B
D. atra NMK 4506 KENYA Ol Ari Nyiro Ranch M 365 26 210 14 2 4 4 6 74 1 0
D. atra NMK 4176 KENYA South Nandi Forest M 465 71 6.549 208 50 14 2 M
D. atra FMNH 58337 KENYA Subukia M 546 110 4.964 23 212 60 14 2 2 4 4 2Bx 1
D. atra MRAC 18554 RWANDA Astrida M 297 54 5.5 23 210 61 14 2 2 4 6 2Bx 1 0
D. atra PEM R5623 RWANDA Btwn Gikongoro & Nyungwe For. R. M 429 81 5.296 23 211 59 14 3 3 4 6 2Bx 1 0
D. atra ZFMK 61616* RWANDA Bugesera M 426 84 5.071 24 216 62 14 2 2 4 6 M 1 0
D. atra MRAC 2723 RWANDA Kisenyi M 70 22 207 66 14 3 2 4 5 2Bx 1 0
D. atra MRAC 2725 RWANDA Kisenyi M 404 96 4.208 22 207 66 14 2 2 2 4 2Bx 1 0
D. atra MRAC 2727* RWANDA Kisenyi M 414 88 4.705 23 218 63 13 2 2 4 5 B 1 0
D. atra MRAC 3821* RWANDA Kisenyi M 383 83 4.614 24 210 66 14 2 2 4 4 B 1 0
D. atra MRAC 5871 RWANDA Kisenyi M 413 94 4.394 24 208 66 14 2 2 4 7 2Bx 1 0
D. atra MRAC 15477 RWANDA Nyanza M 226 40 5.65 24 212 63 14 2 3 5 6 2Bx 1 0
D. atra MRAC 14331 RWANDA Rwankeri District M 352 61 5.77 23 212 60 14 2 2 4 6 2Bx 1 0
D. atra MRAC 18562 RWANDA Tare, Astrida M 24 2 4 5 B 0
D. atra BM 1978.380 SOUTH SUDAN Gilo, Imatong Mtns M 530 104 5.1 23 14 2 2 4 4 M 1 0
D. atra FMNH 62215 SOUTH SUDAN Gilo, Imatong Mtns M 501 102 4.912 23 219 63 14 2 3 4 6 M 1
D. atra MCZ 53406* SOUTH SUDAN Gilo, Imatong Mtns M 536 116 4.621 24 210 66 14 2 4 4 6 M 1 0
D. atra MCZ 53407 SOUTH SUDAN Gilo, Imatong Mtns M 459 94 4.883 23 212 60 14 M 1 0
D. atra MCZ 53408* SOUTH SUDAN Gilo, Imatong Mtns M 518 116 4.466 23 217 65 14 2 2 4 6 M 1 0
D. atra ZFMK 29564 SOUTH SUDAN Gilo, Imatong Mtns M 24 221 56 15 2 2 4 8 M
D. atra ZFMK 29719 SOUTH SUDAN Gilo, Imatong Mtns M 24 218 63 14 2 2 4 6 M
D. atra ZMUC 6116 SOUTH SUDAN Gilo, Imatong Mtns M 493 23 210 14 2 2 4 4 M 1 0
D. atra ZFMK 26005 SOUTH SUDAN Katire M 22 219 65 14 2 2 4 6 M 1
D. atra ZFMK 29731 SOUTH SUDAN Talanga Forest M 23 214 66 12 2 2 4 4 M
D. atra ZMB 16749* TANZANIA Bukoba M 398 79 5.038 23 213 67 14 2 2 4 5 100 1 0
D. atra ZMB 16750 TANZANIA Bukoba M 278 48 5.792 23 214 59 14 4 2 4 6 2Bx 1 1
D. atra BM 1994.403* TANZANIA Ibaya Camp, Mkomazi G.R. M 453 82 5.524 26 224 59 14 2 4 4 6 90 1 0
D. atra NMBO 7375 TANZANIA Ibaya Camp, Mkomazi G.R. M 211 38 5.553 26 220 65 4 4 6 92 1 1
D. atra BM 1970.2177 TANZANIA Kasoge, Mahale Peninsula M 405 91 4.451 25 14 2 2 2 7 109 1 0
D. atra ZMUC 6122 TANZANIA Mbizi Forest Reserve M 398 23 200 67 14 2 3 4 6 B 2 1
D. atra ZMUC 6139* TANZANIA Minziro Forest Reserve M 455 91 5 23 211 66 14 2 2 4 6 M 1 0
D. atra ZSM 81/1936a TANZANIA Moshi M 355 23 208 14 2Bx
D. atra ZSM 81/1936b TANZANIA Moshi M 370 75 4.933 23 206 58 14 2Bx
D. atra LIV 1980.150.30 UGANDA ? M 445 85 5.235 205 58 2 4 4 B
D. atra LIV 1960.67.1 UGANDA Bihunga, Bukwezi M 440 96 4.583 215 69 2 6 6 M
D. atra USNM 63481 UGANDA Budongo Forest M 560 115 4.87 23 221 63 14 2 3 4 5 M 1
D. atra LIV 1962.50.32 UGANDA Buhunga, Buweju M 494 98 5.041 62 2Bx
D. atra BM 11.7.8.14* UGANDA Bussu, Jinja M 480 95 5.053 24 212 63 14 2 4 4 6 M 1 1
D. atra MSNG 30413B* UGANDA Bussu, Jinja M 611 98 6.235 23 209 65 14 2 4 4 4 B 1 0
D. atra BM 51.1.3.74* UGANDA Entebbe M 183 35 5.229 25 202 62 12 2 4 4 6 B 1 0
D. atra CAS-SU 21710* UGANDA Fort Portal, Kingami M 438 92 4.761 23 212 63 14 2 3 4 6 B 1 0
D. atra LIV 1962.9.28 UGANDA Kagabagara, Isingiro M 508 100 5.08 210 62 2 4 5 B
D. atra BM 36.7.3.32 UGANDA Kampala M 463 101 4.584 23 208 64 14 2 2 4 6 2Bx 1 0
D. atra LIV 1980.150.31 UGANDA Kayonza, Kinkizi, Kigezi M 292 53 5.509 208 61 2 4 6 B
D. atra NMZB-UM 2550* UGANDA Kayonza, Kinkizi, Kigezi M 450 92 4.891 22 210 63 14 2 3 4 6 M 1 0
D. atra NMK 404 UGANDA Kazinga Channel M 380 87 4.368 221 66 14 2 B
D. atra LIV 1962.9.14 UGANDA Kichwamba, Bunyaruguru M 348 60 5.8 210 57 4 4 6 2Bx
D. atra ZMB 27539 UGANDA Kisenyi M 224 43 5.209 23 217 67 14 2 2 4 6 2Bx 1 0
D. atra BM 1961.1805* UGANDA Kyamuhunga, Igara, Ankole M 405 83 4.88 23 208 63 14 2 2 4 5 M 1 0
D. atra FMNH 9914 UGANDA Lake Bunyonyi M 446 86 5.186 24 211 61 14 2 2 4 8 2Bx 1
D. atra BM 32.5.2.99* UGANDA Lake George M 450 104 4.327 24 208 71 14 2 3 4 6 B 1 0
D. atra BM 51.1.3.78* UGANDA Lake Nabugabo M 468 92 5.087 23 220 69 14 2 2 4 5 B 1 0
D. atra LIV 1980.150.27 UGANDA Lukiri, Mitoma M 447 85 5.259 215 61 2 4 6 M
D. atra BM 08.10.20.14* UGANDA Mabira Forest, Chagwe M 179 30 5.967 25 218 61 14 2 2 4 6 B 1 0
D. atra BM 03.12.2.23* UGANDA Msozi – Isingiro M 455 98 4.643 24 214 67 14 2 2 3 5 M 1 0
D. atra BM 34.12.15.585* UGANDA Muko, Kigezi M 355 65 5.462 23 205 61 14 2 2 4 6 M 0 0
D. atra BM 34.12.15.586* UGANDA Muko, Kigezi M 410 82 5 24 208 64 14 4 2 4 4 85 1 0
D. atra BM 34.12.15.587* UGANDA Muko, Kigezi M 201 38 5.289 23 206 65 14 2 2 4 6 107 1 0
D. atra BM 34.12.15.589 UGANDA Muko, Kigezi M 215 43 5 24 204 68 14 2 2 4 5 2Bx 1 0
D. atra BM 34.12.15.590 UGANDA Muko, Kigezi M 420 23 211 14 2 3 4 8 M 1 0
D. atra BM 34.12.15.591* UGANDA Muko, Kigezi M 455 88 5.17 23 207 64 14 2 2 4 5 B 1 0
D. atra BM 34.12.15.593 UGANDA Muko, Kigezi M 327 59 5.542 23 218 63 14 2 2 4 7 2Bx 0
D. atra MCZ 48363 UGANDA Mushongero, Lake Mutanda M 320 63 5.079 24 206 63 14 2 2 4 6 2Bx 1 0
D. atra LIV 1980.150.26 UGANDA Nsika, Buhwezi M 320 212 3 4 6 M
D. atra LIV 1962.9.23 UGANDA Nyabushozi M 400 83 4.819 213 65 2 4 6 M
D. atra UMMZ 88532/1 UGANDA Nyakabende M 236 35 6.743 24 205 63 14 2 2 4 4 2Bx 1
D. atra LIV 1962.50.33 UGANDA Nyansimbo, Mitoma M 328 62 5.29 206 62 4 4 5 B
D. atra NMZB-UM 2549 UGANDA Nyanuhunga, Igara, Ankole M 430 96 4.479 24 204 63 14 2 2 4 6 2Bx 1 0
D. atra LIV 1980.150.29 UGANDA Nyarusiza, Bufumbira, Kigezi M 460 99 4.646 220 65 2 4 4 2Bx
D. atra NMZB-UM 2553* UGANDA Nyarusiza, Bufumbira, Kigezi M 320 58 5.517 22 208 62 14 2 2 4 5 103 1 0
D. atra MSNG 40764* UGANDA Sesse Islands, Lake Victoria M 472 77 6.13 23 214 67 14 2 4 4 7 B 1 0
D. atra MCZ 40567* UGANDA Sipi, Mount Elgon M 212 39 5.436 23 208 62 14 2 2 4 4 B 1 0
D. atra MCZ 40568 UGANDA Sipi, Mount Elgon M 473 90 5.256 23 219 64 14 2 2 4 6 2Bx 0
D. atra MCZ 40570* UGANDA Sipi, Mount Elgon M 518 97 5.34 23 211 59 14 2 3 4 4 B 1 0
D. atra MRAC 82-16-R-3* BURUNDI Bujumbura F 615 100 6.15 25 215 57 14 2 2 4 6 130 0 0
D. atra MRAC 18584* BURUNDI Makamba F 184 30 6.133 24 219 56 14 2 2 4 6 B 1 0
D. atra BM 16.6.24.13* ETHIOPIA Aware Melca F 247 31 7.968 26 239 52 14 2 2 4 7 B 1 0
D. atra MNHN 05-208 ETHIOPIA Bourka F 712 96 7.417 22 243 52 13 2 3 3 4 2Bx 1 0
D. atra MSNM 2702 ETHIOPIA Circao, Dambria F 530 22 232 14 2 4 4 10 B
D. atra AMNH 20345* ETHIOPIA Ganame F 660 93 7.097 22 256 56 15 2 4 4 6 M 1 0
D. atra NMW 26879/2+ ETHIOPIA Ginir F 346 46 7.522 26 231 50 13 2 4 4 7 82 1 1
D. atra ZFMK 5535* ETHIOPIA Gololtscha F 666 83 8.024 23 250 50 15 2 4 4 6 M 1 1
D. atra NMW 26879/3 ETHIOPIA Gumaydo F 609 28 234 14 2 4 4 4 81 1 0
D. atra FMNH 4012* ETHIOPIA Harar F 248 35 7.086 26 231 53 14 2 4 4 5 85 1 0
D. atra ZFMK 84993* ETHIOPIA Jigjiga F 490 64 7.656 27 222 45 14 2 4 4 8 72 1 0
D. atra MNHN 05-211 ETHIOPIA Kounni F 178 24 7.417 27 219 48 14 2 4 4 4 2Bx 1 0
D. atra MNHN 05-207* ETHIOPIA Laga Hardine F 264 34 7.765 26 228 51 14 2 4 4 6 B 1 0
D. atra ZFMK 16236* ETHIOPIA Lake Abiata F 532 67 7.94 25 244 50 14 2 4 4 8 72 1 0
D. atra ZMB 10286* ETHIOPIA Let Marefia F 266 32 8.313 22 243 48 14 2 2 4 4 M 1 0
D. atra NMW 15299* ETHIOPIA Wondo F 708 85 8.329 23 239 47 14 2 2 4 4 M 2 0
D. atra UMMZ 96274 KENYA ? F 610 93 6.559 24 221 55 12 2 4 2 7 M 1
D. atra LACM 66076* KENYA 13 km SSW of Maralal F 244 34 7.176 25 225 53 14 4 4 4 6 81 1 0
D. atra NMW 9964/20+ KENYA Athi Plain F 478 64 7.469 27 223 47 14 2 2 4 6 B 1 0
D. atra NMK 3085 KENYA Athi River F 416 59 7.051 245 54 14 4 B
D. atra NMK 3129 KENYA Athi-Kapiti Plains F 461 71 6.493 228 58 14 4 B
D. atra NMK 4174 KENYA Baringo F 198 32 6.188 231 49 14 4 B
D. atra NMK 2323 KENYA Butere, Kakamega F 314 49 6.408 234 53 14 4 79
D. atra CAS 148034 KENYA Chemelil, Kisumu F 605 90 6.722 25 225 14 2 4 2 4 M 1 1
D. atra NMK 406 KENYA Chyulu Hills, Makueni F 571 82 6.963 228 51 14 2 B
D. atra MCZ 40575 KENYA Elgonyi F 310 43 7.209 24 223 51 14 2 4 4 6 2Bx 1 0
D. atra NMK 2849 KENYA Embakasi, Nairobi F 232 58 14 4 B
D. atra BM 08.9.17.4 KENYA Fort Hall (= Murang’a) F 500 27 230 14 2 4 4 6 lin 1 0
D. atra NMK 3131 KENYA Hopcroft Ranch, Athi River F 512 75 6.827 244 57 12 2 B
D. atra USNM 40897 KENYA Juja Farm F 575 72 7.986 26 241 55 15 2 4 4 6 lin 1
D. atra BM 1957.1.11.42* KENYA Kabarnet, Baringo F 595 92 6.467 23 236 56 12 2 2 4 6 M 1 0
D. atra CAS 111767 KENYA Kabartonjo, Kamasia Mtns F 645 102 6.324 25 229 55 13 2 2 4 6 B 1
D. atra NMK 486 KENYA Kacheliba F 440 71 6.197 228 58 14 4 2Bx
D. atra MCZ 40576* KENYA Kaimosi F 700 101 6.931 24 220 51 12 2 2 4 6 M 1 0
D. atra MCZ 40578* KENYA Kaimosi F 707 98 7.214 24 225 55 13 2 4 4 7 M 1 0
D. atra USNM 49078 KENYA Kaimosi F 724 97 7.464 24 217 50 14 2 2 4 4 M 1
D. atra MCZ 18205 KENYA Kaimosi (Yala River) F 610 24 221 14 2 3 4 4 2Bx 1 0
D. atra UMMZ 61224 KENYA Kaimosi (Yala River) F 620 88 7.045 23 223 50 12 2 2 4 6 M 1
D. atra LACM 93378* KENYA Kairuni, Nyambeni Mts F 662 114 5.807 26 222 56 14 3 4 4 6 102 1 1
D. atra LACM 93379* KENYA Kairuni, Nyambeni Mts F 203 35 5.8 26 226 57 14 2 4 4 7 B 1 0
D. atra BM 1959.1.5.58 KENYA Kakamega F 665 105 6.333 221 51 14 2 4 4 8 M 0
D. atra USNM 49376 KENYA Kakamega F 448 74 6.054 22 213 55 13 2 4 2 4 M 1
D. atra NMK 2503 KENYA Karen, Nairobi F 550 92 5.978 223 56 12 2 B
D. atra NMK 2952/3 KENYA Kiboko Ranch Station F 175 27 6.481 216 51 14 2 2Bx
D. atra NMK 2952/4 KENYA Kiboko Ranch Station F 230 247 14 2 2Bx
D. atra NMK 2130 KENYA Kijabe F 584 95 6.147 219 51 12 2 B
D. atra ZMUC 6101* KENYA Kitale F 647 91 7.11 25 225 51 14 2 2 4 4 M 1 0
D. atra NMK 3413 KENYA Kitengela, Athi River F 581 77 7.545 232 58 14 2 B
D. atra ZMB 9224 KENYA Kitui F 555 67.5 8.162 27 233 49 14 2 4 4 6 lin 1 0
D. atra BM 1932.5.2.100* KENYA Lake Naivasha F 640 94 6.809 230 50 14 2 4 4 5 M 1 0
D. atra BM 93.11.21.72* KENYA Lake Naivasha F 672 93 7.226 25 223 52 14 2 4 4 7 M 1 0
D. atra NMK 3109 KENYA Lorgorien, Trans-Mara F 655 101 6.485 225 52 14 4 B
D. atra USNM 62907 KENYA Mbunyi F 765 23 227 13 2 2 4 M 1 0
D. atra NMK 657 KENYA Mount Elgon F 539 85 6.341 217 53 14 2 M
D. atra USNM 40990 KENYA Mount Kenya F 622 95 6.547 26 231 54 14 2 4 4 6 M 1
D. atra LACM 66395* KENYA Mount Kulal F 464 68 6.824 24 233 54 14 2 4 4 8 102 1 0
D. atra NMK 1965 KENYA Mwino, Sigor F 359 55 6.527 235 52 14 2 B
D. atra PEM 873* KENYA Nairobi F 650 96 6.771 23 219 54 13 2 2 4 4 B 1 0
D. atra MCZ 54108 KENYA Nairobi (?) F 650 86 7.558 233 53 14 2 4 4 8 lin 1 0
D. atra NMK 2486 KENYA Nairobi (Westlands) F 524 89 5.888 236 54 14 2 105
D. atra BM 01-1-3-9 KENYA Naitolia (6000 feet) F 495 27 238 14 2 4 4 7 81 1 0
D. atra NMK 1453 KENYA Nakuru F 604 97 6.227 218 53 14 2 B
D. atra NMK 484 KENYA Nakuru F 554 89 6.225 220 55 14 4 B
D. atra BM 06.7.6.7* KENYA Nandi F 384 53 7.245 23 218 50 14 2 2 3 5 M 1 0
D. atra NMK 1158 KENYA Nandi Hills F 700 105 6.667 224 50 14 4 M
D. atra NMK 3490 KENYA Nanyuki River, Laikipia F 531 92 5.772 224 53 14 2 M
D. atra NMK 3512 KENYA Nanyuki River, Laikipia F 611 223 14 4 M
D. atra NMK 1960 KENYA Nbabibi, Naivasha F 469 58 8.086 231 48 14 4 B
D. atra NMK 2583 KENYA North Nandi Forest, Chemisia F 890 115 7.739 218 53 14 4 M
D. atra CG 2281 KENYA Pen-y-Bryn Farm , Lumbwa F 705 97 7.268 26 231 49 2 4 4 6 M 1
D. atra BM 96.3.27.18* KENYA Uganda Railway F 232 35 6.629 24 212 51 14 2 2 4 5 104 1 0
D. atra MRAC 18555 RWANDA Astrida F 613 83 7.386 24 224 58 14 2 2 4 4 2Bx 1 0
D. atra MRAC 18556 RWANDA Astrida F 40 25 50 14 2 2 4 6 2Bx 1 0
D. atra MRAC 18557 RWANDA Astrida F 372 56 6.643 24 214 51 12 2 2 4 4 2Bx 1 1
D. atra MRAC 18558 RWANDA Astrida F 207 33 6.273 24 222 54 14 2 2 4 5 2Bx 1 0
D. atra MRAC 86-01-R-10 RWANDA Bugarama F 567 90 6.3 25 219 54 14 3 2 4 7 2Bx 1 0
D. atra MRAC 86-01-R-11 RWANDA Bugarama F 642 95 6.758 24 218 55 14 2 2 4 6 2Bx 1 0
D. atra MRAC 18577 RWANDA Bulimbi F 24 221 14 2 2 4 6 2Bx 1 0
D. atra ZFMK 61617 RWANDA Cyamudongo F 656 100 6.56 24 230 58 12 2 2 4 5 2Bx 1 0
D. atra MRAC 14329 RWANDA Kamohorora District F 293 46 6.37 24 234 58 14 2 3 4 6 2Bx 1 0
D. atra ZFMK 55105* RWANDA Kibungo, Rusumo F 432 68 6.353 24 224 55 14 2 2 4 5 B 1 1
D. atra MCZ 48364* RWANDA Kisenyi F 572 91 6.286 25 227 59 14 2 2 4 5 115 1 0
D. atra MRAC 5872 RWANDA Kisenyi F 424 65 6.523 24 222 50 14 2 2 4 6 2Bx 1 0
D. atra MRAC 9773 RWANDA Kisenyi F 755 115 6.565 24 236 53 14 2 2 4 6 2Bx 1 0
D. atra ZMB 20458 RWANDA Kisenyi F 517 75 6.893 24 222 51 14 2 2 4 5 2Bx 1 0
D. atra ZMB 27156 RWANDA Kisenyi F 409 67 6.104 25 221 54 14 2 2 4 6 2Bx 1 0
D. atra MRAC 14333 RWANDA Lake Gando F 663 103 6.437 24 226 54 14 2 2 4 5 2Bx 1 0
D. atra MRAC 14341 RWANDA Lake Gando F 65 25 218 54 14 2 2 4 4 2Bx 1 0
D. atra MRAC 7509 RWANDA Mutura, Mount Virunga F 587 90 6.522 24 229 59 14 2 2 4 4 2Bx 1 0
D. atra MRAC 74-23-R-22 RWANDA Mwogo near Butare F 24 217 14 2 2 4 4 2Bx 1 0
D. atra MRAC 14340 RWANDA Rwankeri District F 377 58 6.5 24 219 52 14 2 2 4 5 2Bx 1 0
D. atra MRAC 15440 RWANDA Rwankeri District F 673 109 6.174 24 225 60 16 2 2 4 7 113 1
D. atra MRAC 18259* RWANDA Rwankuba F 632 100 6.32 25 228 54 14 2 2 4 7 B 1 0
D. atra MRAC 4074* RWANDA Shangugu F 525 85 6.176 24 225 59 14 2 2 4 8 93 1 0
D. atra MRAC 4075* RWANDA Shangugu F 373 58 6.431 26 229 54 14 2 2 4 5 108 1 0
D. atra MRAC 18560 RWANDA Tare, Astrida F 719 99 7.263 25 225 51 13 2 2 4 6 B 0
D. atra MRAC 18561 RWANDA Tare, Astrida F 542 24 224 14 2 2 4 6 B 1 0
D. atra ZFMK 29567 SOUTH SUDAN 10 km S of Juba F 760 24 236 57 13 2 2 4 5 M
D. atra ZFMK 29565 SOUTH SUDAN Aweil F 760 23 225 51 14 2 2 4 4 M
D. atra BM 1978.379 SOUTH SUDAN Gilo, Imatong Mts F 625 97 6.443 24 229 13 2 4 4 6 M 1 0
D. atra MCZ 53569 SOUTH SUDAN Gilo, Imatong Mts F 372 26 228 14 2 4 4 6 B 1 1
D. atra ZFMK 34532 SOUTH SUDAN Gilo, Imatong Mts F 24 228 63 2 2 4 6 M
D. atra ZFMK 29732 SOUTH SUDAN Talanga Forest F 500 24 227 54 14 2 2 4 6 M
D. atra ZFMK 29733 SOUTH SUDAN Talanga Forest F 430 24 59 14 2 2 4 9 M
D. atra PEM 17329 TANZANIA Klein’s Gate, N.E Serengeti F 317 45 7.044 27 240 56 14 2 4 92 1 0
D. atra PEM 17336 TANZANIA Klein’s Gate, N.E Serengeti F 434 25 227 14 2 4 100 1 0
D. atra PEM 17941 TANZANIA Klein’s Gate, N.E Serengeti F 345 25 225 14 2 4 4 6 80 1 1
D. atra ZMUC 6140* TANZANIA Minziro Forest Reserve F 23 227 54 14 2 2 4 6 M 1 0
D. atra ZSM 81/1936c TANZANIA Moshi (Uru) F 475 60 7.917 23 235 47 14 2Bx
D. atra ZMB 17039A* TANZANIA Mount Kilimanjaro F 197 30 6.567 25 240 55 14 2 4 4 6 93 1 0
D. atra ZMB 17039B TANZANIA Mount Kilimanjaro F 408 62 6.581 26 233 56 14 2 4 4 6 2Bx 1 0
D. atra LIV 1960.115.3 UGANDA Bitereko, Igara F 638 96 6.646 223 54 2 4 5 M
D. atra BM 51.1.3.68 UGANDA Bufundi, Kigezi F 501 75 6.68 24 222 55 14 2 2 108 1 0
D. atra BM 51.1.3.79 UGANDA Bufundi, Kigezi F 665 96 6.927 24 216 50 14 2 2 4 6 2Bx 1 0
D. atra NMZB 217 = SRM 3.2* UGANDA Bufundi, Lake Bunyonyi F 580 88 6.591 25 221 52 14 2 2 4 6 B 1 0
D. atra BM 51.1.3.80* UGANDA Bukakata F 645 101 6.386 24 227 56 14 2 2 4 5 B 1 0
D. atra BM 1951.1.5.24* UGANDA Busingiro, Budango F 760 106 7.17 23 230 54 14 2 4 4 6 M 1 0
D. atra MCZ 40574 UGANDA Butandiga,  Mount Elgon F 650 97 6.701 24 224 52 14 2 4 4 6 2Bx 1 0
D. atra LACM 38696* UGANDA Bwamba Forest (Ntandi) F 567 92 6.163 25 226 55 14 2 2 4 6 B 1 0
D. atra LACM 46344* UGANDA Bwamba Forest (Ntandi) F 24 217 64 14 2 3 4 6 B 1 1
D. atra CAS-SU 21709 UGANDA Fort Portal, Kingami F 410 70 5.857 25 228 60 14 2 2 4 6 2Bx 1 1
D. atra MCZ  54738* UGANDA Fort Portal, Kingami F 700 108 6.481 27 216 54 14 2 2 4 7 M 1 0
D. atra BM 51-1-3-77* UGANDA Gulu F 475 63 7.54 26 232 48 14 2 4 4 6 88 1 1
D. atra BM 1976.2265 UGANDA Ibanda F 390 63 6.19 22 52 15 2 2 5 8 M 1 2
D. atra BM 12.10.12.2* UGANDA Jinja F 422 57 7.404 26 230 51 14 2 4 4 6 B 1 0
D. atra BM 1961.1803* UGANDA Kabingo, Bunyaruguru F 445 67 6.642 25 211 53 14 2 4 4 6 B 1 0
D. atra LIV 1960.67.5 UGANDA Kagando, Kigagati F 205 30 6.833 219 49 4 4 6 2Bx
D. atra LACM 60705* UGANDA Kalinzu Forest F 392 59 6.644 25 213 53 15 2 2 4 6 M 1 0
D. atra BM 1984.882* UGANDA Kampala F 710 119 5.966 23 223 57 14 2 2 4 6 B 1 0
D. atra MUZM UGANDA Kampala F 475 73 6.507 25 224 56 14 3 B
D. atra MUZM UGANDA Kampala F 310 49 6.327 25 220 54 14 2 2 4 6 M
D. atra LACM 39242* UGANDA Katera, Sango Bay Forest F 668 100 6.68 25 222 56 14 2 2 4 6 B 1 0
D. atra NMZB 103 = SRM 3.1* UGANDA Katunguru F 325 54 6.019 25 216 56 14 2 2 4 8 M 1 0
D. atra MRAC 9997* UGANDA Katwe F 25 217 60 14 2 4 4 6 M 1 0
D. atra LACM 35090* UGANDA Kayonza, Impenetrable F. F 530 87 6.092 24 224 57 14 2 2 4 6 B 1 0
D. atra LIV 1962.9.27 UGANDA Kayonza, Kinkizi F 657 105 6.257 216 53 2 4 6 M
D. atra LIV 1962.9.22 UGANDA Kinani, Nyabushozi F 222 2 4 4 M
D. atra BM 51.1.3.75 UGANDA Kyamahunga F 630 94 6.702 24 225 55 14 2 2 4 6 2Bx 1 0
D. atra LIV 1962.9.4 UGANDA Kyamogorani, Kashari F 648 100 6.48 226 55 2 4 6 B
D. atra FMNH 9913 UGANDA Lake Bunyonyi F 527 76 6.934 24 224 52 14 2 2 4 4 2Bx 1
D. atra MCZ 34922 UGANDA Lake Bunyonyi F 628 100 6.28 24 223 51 15 2 2 4 4 2Bx 1 0
D. atra NMZB 219 = SRM 3.4* UGANDA Lake Edward F 620 101 6.139 26 218 54 14 2 2 4 6 B 1 0
D. atra BM 08.10.20.15* UGANDA Mabira Forest, Chagwe F 391 59 6.627 26 219 54 14 2 4 4 6 B 1 0
D. atra BM 51.1.3.76* UGANDA Masese, Jinja F 754 121 6.231 24 225 56 14 2 2 4 6 B 1 0
D. atra MCZ 48359 UGANDA Mihunga, Ruwenzori F 230 24 222 14 2 2 4 6 M 1 0
D. atra NMK 3291 UGANDA Mityana F 530 78 6.795 232 14 4 M
D. atra BM 1954.1.12.37* UGANDA Morulinga, 24 km S of Moroto F 300 43 6.977 24 224 54 14 2 4 4 6 82 1 1
D. atra CAS 85300* UGANDA Mount Elgon F 620 90 6.889 23 220 52 14 2 2 4 4 M 1 0
D. atra BM 34.12.15.592* UGANDA Mubango, Mabira Forest F 830 133 6.241 23 230 59 14 2 2 4 6 M 1 0
D. atra BM 34.12.15.588 UGANDA Muko, Kigezi F 625 102 6.127 24 214 55 14 2 2 4 6 2Bx 1 0
D. atra BM 34.12.15.594* UGANDA Muko, Kigezi F 365 45 8.111 24 222 52 14 2 2 4 6 M 1 0
D. atra BM 03-12-2-22* UGANDA Mulema F 500 68 7.353 25 226 51 14 2 4 4 6 92 1 0
D. atra MCZ 48362* UGANDA Mushongero, Lake Mutanda F 438 64 6.844 24 230 52 14 2 2 4 6 B 1 0
D. atra MCZ 48360 UGANDA Nyakabande F 677 98 6.908 25 226 53 14 2 2 5 10 2Bx 1 0
D. atra MCZ 48361 UGANDA Nyakabande F 538 74 7.27 24 223 51 14 2 2 4 6 2Bx 1 0
D. atra UMMZ 88532/2 UGANDA Nyakabende F 325 45 7.222 25 216 51 14 4 2 4 6 2Bx 1
D. atra LIV 1959.83.2 UGANDA Nyamuhumia, Igara F 365 49 7.449 218 47 4 4 4 2Bx
D. atra LIV 1980.150.33 UGANDA Nyansimbo, Mitoma F 222 38 5.842 223 55 2 4 5 B
D. atra BM 1976.1849* UGANDA Nyarusiza, Bufumbira, Kigezi F 270 42 6.429 24 214 53 14 2 2 4 6 95 1 0
D. atra NMZB-UM 2552* UGANDA Nyarusiza, Bufumbira, Kigezi F 310 50 6.2 24 216 54 14 2 2 4 6 96 1 0
D. atra MCZ 54744 UGANDA Rucho, Ankole F 592 102 5.804 25 225 59 14 2 2 4 6 2Bx 1 0
D. atra LIV 1960.67.3 UGANDA Ruhama, Rwampara F 455 65 7 220 52 4 4 6 2Bx
D. atra MSNG 27755A* UGANDA Sesse Islands, Lake Victoria F 616 76 8.105 22 229 55 14 2 4 4 4 B 1 0
D. atra MCZ 40564* UGANDA Sipi, Mount Elgon F 725 100 7.25 23 224 55 14 2 2 4 4 M 1 0
D. atra MCZ 40565* UGANDA Sipi, Mount Elgon F 621 90 6.9 24 218 52 14 2 2 4 4 M 1 0
D. atra MCZ 40566 UGANDA Sipi, Mount Elgon F 515 77 6.688 24 218 52 14 2 2 4 4 2Bx 1 0
D. atra MCZ 40569* UGANDA Sipi, Mount Elgon F 710 106 6.698 24 232 53 14 2 3 4 6 M 1 0
D. atra MCZ 40571* UGANDA Sipi, Mount Elgon F 620 84 7.381 24 220 52 14 2 3 4 7 M 1 0
D. atra MCZ 40572 UGANDA Sipi, Mount Elgon F 342 50 6.84 218 50 14 2 4 4 4 B
D. atra MCZ 40573 UGANDA Sipi, Mount Elgon F 225 35 6.429 223 54 14 2 4 4 4 B
D. atra MRAC 82-16-R4 BURUNDI Bujumbura U 23 218 75 14 2 2 4 5 B 1 0
D. atra MSNM 2975 ETHIOPIA El Dire, Tertale U 277 42 6.595 24 219 49 14 2 4 4 6 69
D. atra BM 1973.3175 ETHIOPIA Ghimbi U M
D. atra BM 1975.2144 ETHIOPIA Kebre Mengist to Negele U 170 14 2 2 4 6 2Bx 1 1
D. atra NMK 1962 KENYA ? U 377 72 5.236 203 59 14 4 2Bx
D. atra NMK 348 KENYA ? U 192 27 7.111 212 53 12 4 B
D. atra NMK 1348 KENYA Athi River U 636 93 6.839 221 50 14 4 B
D. atra NMK 488 KENYA Kacheliba U 536 87 6.161 229 54 14 2 98
D. atra NMK 482 KENYA Kakamega U 257 39 6.59 216 48 14 2 B
D. atra NMK  2715 KENYA Kakamega U 411 65 6.323 221 54 14 2 2Bx
D. atra NMK 1260 KENYA Kericho U 712 101 7.05 216 51 16 4 B
D. atra NMK 481 KENYA Kiambu U 199 32 6.219 226 69 12 4 B
D. atra NMK 2952/2 KENYA Kiboko Ranch Station U 182 29 6.276 251 64 14 4 2Bx
D. atra NMK 3123 KENYA Kiserian, Baringo U 235 33 7.121 227 51 14 4 B
D. atra NMK 3051 KENYA Kitale U 317 69 4.594 214 52 14 2 2Bx
D. atra NMK 1791 KENYA Lake Baringo U 619 107 5.785 228 57 14 2 104
D. atra NMK 3152 KENYA Mount Elgon U 208 40 5.2 214 59 12 4 2Bx
D. atra NMK 658 KENYA Mount Elgon U 565 90 6.278 221 52 12 2 B
D. atra NMK 3557 KENYA Nanyuki River, Laikipia U 539 91 5.923 221 54 14 4 M
D. atra NMK 1489 KENYA Nyambene Hills, Meru U 594 97 6.124 225 52 14 2 92 1 1
D. atra NMK 3182 KENYA Timbilil, SW Mau Forest U 195 232 14 4 B
D. atra IRSNB 2657 RWANDA Ruhengeri U 245 46 5.326 23 209 69 14 2 2 4 5 106 1 0
D. atra PEM 16778 TANZANIA Klein’s Camp, N.E. Serengeti U 251 27 244 14 2 4 75 2 0
D. atra ZMUC 6138 TANZANIA Minziro Forest Reserve U 212 38 5.579 22 216 61 14 2 2 4 6 M 1 0
D. atra ZMUC 6141 TANZANIA Minziro Forest Reserve U 23 214 68 12 2 2 4 6 B 1 0
D. atra BM 1970.2178 TANZANIA Sitete, Mahale Peninsula U 435 43 64 14 2 2 4 4 B 1 0
D. atra MSNG 30413A UGANDA Bussu, Jinja U 786 102 7.706 25 230 60 14 2 4 4 6 M 1 0
D. atra MSNG 52410 UGANDA Bussu, Jinja U 586 71 8.254 23 226 54 12 2 4 4 B
D. atra LIV 1960.115.4 UGANDA Kayonza, Kinkizi U 2 4 5 B
D. atra NMK 1495 UGANDA Nyarusiza U 664 93 7.14 226 50 14 2
D. atra NMK 1496 UGANDA Nyarusiza U 591 94 6.287 221 59 14 2 B
D. bazi NMW 26879/1 EGYPT Cairo, surroundings of F 461 61 7.557 24 234 50 14 2 4 4 6 1 1
D. bazi ZMB 1861* EGYPT Oberaegypten M 455 79 5.759 23 229 59 14 2 4 4 6 48 1 1
D. bazi MNHN 1847* EGYPT M 410 70 5.857 23 229 59 14 2 4 4 6 49 2 1
D. bazi ZMB 5865* EGYPT M 427 78 5.474 23 219 60 14 2 4 6 6 39 2 2
D. bazi BM 97.10.28.575* EGYPT Fayoum F 602 90 6.689 24 227 55 14 2 4 5 6 42 1 1
D. bazi ZMB 51552* EGYPT Uberegypte F 503 70 7.186 24 228 51 14 2 4 4 6 48 0 0
D.  cf. bazi ZFMK 38415 SUDAN Erkowit M 25 229 67 14 2 4 6 8 48 1
D. confusa NMW 9964/2* ETHIOPIA ? M 362 67 5.403 24 208 57 14 2 4 4 7 75 1 0
D. confusa ZMB 56436* ETHIOPIA ‘Addis Ababa’ M 475 87 5.46 24 213 57 14 2 4 4 6 79 0 1
D. confusa NMK 485 KENYA Kacheliba M 319 55 5.8 212 51 14 4 67
D. confusa NMK 489 KENYA Kacheliba M 424 74 5.73 216 54 14 4 65
D. confusa MCZ 53403* SOUTH SUDAN Nimule M 232 42 5.524 26 208 63 14 2 4 4 6 55 1 1
D. confusa FMNH 58361 SOUTH SUDAN Torit M 396 79 5.013 25 224 67 14 2 4 4 6 B 1
D. confusa FMNH 58362 SOUTH SUDAN Torit M 330 69 4.783 23 210 63 B 1
D. confusa FMNH 62209 SOUTH SUDAN Torit M 551 25 203 14 2 4 4 6 B 1
D. confusa FMNH 62214 SOUTH SUDAN Torit M 394 78 5.051 24 220 64 14 2 4 4 5 54 1
D. confusa FMNH 62217 SOUTH SUDAN Torit M 235 45 5.222 25 212 64 14 2 4 4 6 B 1
D. confusa MUZM 10/R1* UGANDA Ajai Game Reserve M 395 84 4.702 25 200 66 14 2 4 4 8 50 1 0
D. confusa MCZ 47815* UGANDA Busingiro, Budongo Forest M 358 64 5.594 26 199 54 14 2 4 4 10 54 1 1
D. confusa BM 51-1-5-39* UGANDA Lira, Lango M 325 62 5.242 25 205 62 14 2 4 4 6 62 1 0
D. confusa UMMZ 88529 UGANDA Lira, Lango M 45 25 14 2 4 4 7 61 0
D. confusa LIV 1960.51.2 UGANDA Nakosa, Bunyaruguru M 390 208 4 4 6
D. confusa CAS 141784* KENYA Mumias F 203 27 7.519 26 226 48 14 2 4 4 6 63 1 0
D. confusa ZMB 77441 RWANDA Lac Kivu F 620 24 224 11 2 4 4 6 71 1 0
D. confusa ZFMK 29694 SOUTH SUDAN 10km S of Juba F 25 55 14 2 4 4 6 50
D. confusa FMNH 58496 SOUTH SUDAN Juba F 575 92 6.25 26 228 54 14 2 4 4 8 54 1
D. confusa MCZ  53401* SOUTH SUDAN Logutoic F 77 26 221 55 14 2 4 4 6 60 1 0
D. confusa FMNH 62216 SOUTH SUDAN Lokwi F 165 25 6.6 25 229 49 14 2 3 4 6 69 1
D. confusa MCZ 53402 SOUTH SUDAN Mongalla F 524 23 225 14 2 4 4 6 58 1 0
D. confusa NMW 9964/7+ SOUTH SUDAN ‘Sudan’ (taken as East Equatoria) F 465 67 6.940 25 231 55 14 2 4 4 4 62 1 0
D. confusa FMNH 58363 SOUTH SUDAN Torit F 492 72 6.833 25 223 54 14 2 4 4 6
D. confusa FMNH 62207 SOUTH SUDAN Torit F 369 25 239 14 2 4 4 6 61 1
D. confusa FMNH 62208 SOUTH SUDAN Torit F 428 25 236 14 2 4 4 6 B 1
D. confusa FMNH 62210 SOUTH SUDAN Torit F 417 64 6.516 24 230 55 14 2 4 4 6 1
D. confusa FMNH 62211 SOUTH SUDAN Torit F 187 32 5.844 25 233 56 14 2 4 4 6 B 1
D. confusa FMNH 62212 SOUTH SUDAN Torit F 435 62 7.016 25 238 53 14 2 4 4 6 68 1
D. confusa FMNH 62213 SOUTH SUDAN Torit F 378 25 237 14 2 4 4 5 64 1
D. confusa MCZ  53405* SOUTH SUDAN Torit F 324 51 6.353 25 236 57 14 2 4 4 6 73 1 1
D. confusa MCZ 53404 SOUTH SUDAN Torit F 546 24 236 14 2 4 4 6 64 1 1
D. confusa FMNH 58336 SOUTH SUDAN Yei F 252 51 4.941 27 226 58 13 2 4 4 7 58
D. confusa BM 51.1.3.69* UGANDA Bisu, Kigezi F 518 79 6.557 26 222 56 14 2 4 4 6 61 1 0
D. confusa LACM 39043* UGANDA Bugoma Forest F 166 31.5 5.188 23 213 61 14 2 4 4 6 61 1 0
D. confusa MCZ 47830* UGANDA Gulu, Acholi F 301 52 5.788 23 212 56 14 2 4 4 6 B 1 0
D. confusa NMZB 14333* UGANDA Kyambura Game Reserve F 520 73 7.123 25 229 52 14 2 4 4 8 63 1 0
D. confusa NMZB 15411* UGANDA Kyambura Game Reserve F 485 73 6.644 26 215 52 14 2 4 4 5 B 1 0
D. confusa MCZ 47801* UGANDA Lira, Lango F 444 59 7.525 25 228 50 14 2 4 4 8 B 1 0
D. confusa UMMZ 88530 UGANDA Lira, Lango F 56 27 218 50 14 2 4 4 8 B 1
D. confusa BM 51.1.3.84* UGANDA Moyo, West Madi F 245 37 6.622 26 229 57 14 2 4 4 6 55 1 0
D. confusa LIV 1962.106.19 UGANDA Nyanshimbo, Ibanda F 501 226 4 4 7
D. confusa LIV 1960.67.2 UGANDA Sanga, Nyabushozi F 481 73 6.589 213 53 4 4 7
D. confusa BM 1954 1.12.8* UGANDA Semliki F 503 72 6.986 24 218 52 12 2 4 3 6 59 1 0
D. confusa BM 1954 1.12.9* UGANDA Semliki F 77 26 215 55 14 2 4 3 6 55 1 2
D. confusa BM 1976.848* UGANDA Serere, Teso F 567 81 7 26 227 55 14 2 4 4 5 B 1 0
D. crucifera MZUT 3582* ERITREA Agordat M 211 39 5.41 22 226 61 14 2 4 4 5 52 1 0
D. crucifera ZMB 7631* ERITREA Bogos F 442 64 6.906 25 231 52 14 2 4 4 6 62 1 0
D. crucifera MZUT 3469 ERITREA Agordat F 23 247 50
D. crucifera MSNG 6738* ERITREA Keren F 368 45 8.178 24 243 49 14 2 4 4 6 51 1 0
D. fasciata LACM 38697 UGANDA Ntandi, Bwamba Forest M 525 111 4.73 23 230 71 14 2 4 4 7 2 0
D. fasciata LACM 46342* UGANDA Ntandi, Bwamba Forest M 509 116 4.388 23 229 76 14 2 4 4 6 141 1 0
D. fasciata LACM 39243* UGANDA Sango Bay Forest, Katera M 472 91 5.187 24 212 59 14 2 4 4 4 108 1 0
D. fasciata MCZ  48357 UGANDA Bundibugyo, Bwamba F 667 119 5.605 24 241 68 13 4 4 4 6 1 0
D. fasciata MCZ  48358A UGANDA Bundibugyo, Bwamba F 730 132 5.53 23 239 67 14 2 4 4 6 1 0
D. fasciata MCZ  48358B UGANDA Bundibugyo, Bwamba F 635 100 6.35 25 248 66 14 2 3 4 6 0
D. fasciata UMMZ 88531 UGANDA Bundibugyo, Bwamba F 627 107 5.86 23 248 64 14 4 4 4 6 1
D. fasciata LACM 46341 UGANDA Ntandi, Bwamba Forest F 238 50 4.76 22 237 76 14 2 2 6 7 1 0
D. fasciata LACM 46343* UGANDA Ntandi, Bwamba Forest F 586 113 5.186 24 241 71 14 2 4 5 8 116 1 0
D. fasciata LACM 46345* UGANDA Ntandi, Bwamba Forest F 312 57 5.474 24 242 71 14 2 4 4 6 111 1 0
D. fasciata LACM 46346 UGANDA Ntandi, Bwamba Forest F 598 103 5.806 24 245 65 13 2 4 5 6 1 0
D. fasciata TCWC 23839* UGANDA Ntandi, Bwamba Forest F 611 108 5.657 23 243 67 14 2 4 4 7 113 1 0
D. medici NMK 2631 KENYA Gede M 679 146 4.651 220 78 14 4 L
D. medici NMK 3200 KENYA Gede M 384 91 4.22 235 80 14 2 L
D. medici BM 51.1.3.73* KENYA Kilifi M 465 116 4.009 24 229 87 14 2 4 4 6 L 2 0
D. medici MCZ 40581* KENYA Lamu Island M 502 132 3.803 23 226 94 14 2 4 4 6 L 2 0
D. medici NMK 1561 KENYA Malindi M 487 127 3.835 230 84 14 4 L
D. medici NMK 2535 KENYA Malindi M 560 125 4.48 220 75 12 4 L
D. medici NMK 2558 KENYA Malindi M 241 79 14 2 L
D. medici NMK 3292 KENYA Malindi M 616 133 4.632 241 80 14 4 L
D. medici ZMB 50191 KENYA Malindi M 461 115 4.009 24 239 91 14 2 4 4 5 P 1 0
D. medici NMK 3956 KENYA Mount Warges, Wamba M 466 106 4.396 24 239 83 14 2 4 4 6 P 2 0
D. medici NMK 1699 KENYA Nyambene Hills, Meru M 424 104 4.077 233 14 2 4 4 6 P 2 0
D. medici CAS 135751 KENYA Tiwi, Kwale District M 370 85 4.353 24 224 84 14 2 4 4 6 P 2 1
D. medici MZUF 2028 SOMALIA Alessandra Island, Gelib M 422 107 3.944 24 233 93 12 2 4 4 6 1 1
D. medici CAS 148077* SOMALIA Mareri, Lower Juba River M 375 95 3.947 24 230 89 14 2 4 4 6 L 2 1
D. medici MZUF 27155 SOMALIA Mareri, Lower Juba River M 496 137 3.62 24 231 98 13 2 4 4 6 2
D. medici NMW 9964/18+ TANZANIA Dar-es-Salaam M 430 104 4.135 24 246 94 14 3 4 4 6 66 2 0
D. medici FMNH 251325 TANZANIA East Usambara Mts M 400 95 4.211 24 249 89 14 2 3 4 5 P 2 0
D. medici ZMUC 6144* TANZANIA Longido M 522 129 4.047 24 239 89 14 2 4 4 6 79 2 1
D. medici SMF 19460* TANZANIA Mafia Island M 471 119 3.958 23 252 98 14 2 4 4 7 68 2 0
D. medici MCZ 18206* TANZANIA Makindu River M 525 130 4.038 24 240 89 14 2 4 4 6 67 2 0
D. medici NMZB 17318 TANZANIA Mlinga Forest Res. M 460 124 3.71 23 240 103 14 2 4 4 6 P 2 0
D. medici USNM 62904 TANZANIA Morogoro M 475 115 4.13 24 237 83 14 2 4 5 7 67 2
D. medici USNM 62906 TANZANIA Morogoro M 470 105 4.476 24 248 86 14 2 4 4 6 P 2
D. medici SMNS 4216E TANZANIA Moshi M 26 230 14 2 4 4 6 72 2 0
D. medici SMNS 4216F TANZANIA Moshi M 315 69 4.565 26 232 80 15 2 4 4 7 92 1
D. medici ZMUC 6121* TANZANIA Msolwa, Ruhebo Mtns M 444 117 3.795 25 234 81 14 2 2 4 6 70 2 0
D. medici ZMB 17453* TANZANIA Usambara M 575 158 3.639 25 242 109 14 2 4 6 10 64 2 0
D. medici ZMB 8597* TANZANIA Zanzibar Coast M 480 117 4.103 24 243 94 14 3 4 5 6 69 2 0
D. medici NMK 3249 TANZANIA Zanzibar Island M 451 129 3.496 235 93 12 2
D. medici NMK 402 KENYA Chyulu Hills F 616 120 5.133 239 66 14 4
D. medici AMNH 88631 KENYA Gede F 641 130 4.931 25 235 80 14 2 3 4 6 2 0
D. medici NMK 2597 KENYA Gede F 600 141 4.255 229 82 14 2 L
D. medici NMK 2619* KENYA Gede F 580 123 4.715 26 227 81 14 2 4 6 7 L 2 0
D. medici NMK 480 KENYA Giriama, Kilifi F 466 103 4.524 226 72 14 4
D. medici ZMB 37600 KENYA Kibognoto – Kilimanjaro F 496 103 4.816 27 218 74 14 2 4 6 8 1 0
D. medici NMK 1050 KENYA Kikambala, Kilifi F 612 112 5.464 231 79 14 4
D. medici BM 51.1.3.72* KENYA Kilifi F 600 119 5.042 25 228 80 14 2 3 4 6 L 2 0
D. medici MCZ 40582* KENYA Lamu Island F 575 130 4.423 24 226 84 14 2 4 4 6 L 2 0
D. medici NMK 2522 KENYA Malindi F 567 141 4.021 229 80 14 2 L
D. medici NMK 2526 KENYA Malindi F 707 131 5.397 235 77 14 4 L
D. medici NMK 2527 KENYA Malindi F 545 124 4.395 247 85 14 4 L
D. medici NMK 2529 KENYA Malindi F 654 127 5.15 232 76 14 4 L
D. medici NMK 2530 KENYA Malindi F 616 148 4.162 234 79 14 4 L
D. medici NMK 2531 KENYA Malindi F 597 134 4.455 225 82 14 4 L
D. medici NMK 2532 KENYA Malindi F 494 116 4.259 227 79 14 4 L
D. medici NMK 2533 KENYA Malindi F 535 117 4.573 220 76 14 2 L
D. medici NMK 2587 KENYA Malindi F 566 128 4.422 239 75 14 4 L
D. medici NMK 3290 KENYA Malindi F 597 121 4.934 234 79 14 2 L
D. medici BM 98.1.8.15* KENYA Maungu F 452 92 4.913 25 228 81 14 2 4 5 8 L 2 0
D. medici NMW 26926+ KENYA Mombasa, 30 km S of F 289 66 4.379 26 233 81 14 2 4 4 6 86 2 0
D. medici BM 87.11.3.31* KENYA Mount Kilimanjaro F 500 104 4.808 24 230 80 14 2 4 5 6 L 2 0
D. medici USNM 16755* KENYA Mount Kilimanjaro F 702 25 232 67 13 2 4 7 6 L 2 0
D. medici MCZ 40580 KENYA Mount Mbololo F 645 120 5.375 26 231 72 14 2 4 6 9 L
D. medici ZFMK 63582 KENYA near Mombasa F 613 119 5.151 25 231 74 14 2 4 4 6 2 0
D. medici NMK 2377 KENYA Nguni, Mwingi F 481 84 5.726 223 82 14 4
D. medici ZMB 9244* KENYA Taita F 105 25 232 73 14 2 3 4 7 L 2 0
D. medici ZMB 17496 KENYA Takaungu F 405 77 5.26 25 238 74 14 2 4 6 8 2 1
D. medici FMNH 142635* KENYA ‘Vicinity of Nairobi’ F 912 149 6.121 25 235 71 14 2 4 6 8 L 2 1
D. medici BM 98.1.8.14 KENYA Voi to Ndi F 274 52 5.269 24 241 75 14 2 4 4 6 P 2 0
D. medici MSNG 42710b SOMALIA Belet Amin F 845 22 232 13 2 4 4 6 L 0 0
D. medici MCZ 23274* TANZANIA Amani, Usambara Mts F 640 145 4.414 26 241 87 14 2 4 4 6 108 2 0
D. medici ZMB 37598* TANZANIA Amani, Usambara Mts F 546 117 4.667 26 241 82 14 2 4 4 7 77 2 0
D. medici ZMUC 6123 TANZANIA Bombani, E. Usambara Mts F 649 124 5.234 25 239 73 14 2 4 4 6 P 2 0
D. medici ZMUC 6133* TANZANIA Bombani, E. Usambara Mts F 599 124 4.831 25 245 78 14 2 4 4 6 76 2 0
D. medici NMZB 11265* TANZANIA Mafia Island F 410 87 4.713 26 252 88 14 2 4 4 6 L 2 0
D. medici AMNH 143717 TANZANIA Morogoro F 262 54 4.852 24 243 83 14 2 4 4 6 64 2 2
D. medici USNM 62901 TANZANIA Morogoro F 595 118 5.042 25 240 81 14 2 4 4 6 59 2
D. medici ZMB 27616* TANZANIA Morogoro F 630 120 5.25 26 237 72 14 2 4 4 6 70 2 0
D. medici SMNS 4216A* TANZANIA Moshi F 680 135 5.037 24 237 75 15 2 4 5 7 L 2 1
D. medici SMNS 4216B* TANZANIA Moshi F 117 25 233 69 14 2 4 5 8 L 2 0
D. medici SMNS 4216C* TANZANIA Moshi F 500 93 5.376 26 233 61 14 2 4 5 7 L 1 0
D. medici SMNS 4216D* TANZANIA Moshi F 419 82 5.11 25 232 75 14 2 4 6 8 L 2 0
D. medici ZFMK 82342 TANZANIA Mount Kwaraha F 287 25 235 14 2 4 4 6 78 1 0
D. medici MCZ 23273* TANZANIA Nyange, Uluguru Mts F 463 94 4.926 25 244 82 14 2 4 4 6 57 2 0
D. medici CM 36586 TANZANIA Usa River, Arusha F 460 85 5.412 26 232 75 14 2 4 4 6 L
D. medici BM 50.1.5.40 TANZANIA Zanzibar F 650 131 4.962 25 247 77 14 2 4 4 6 P 2 0
D. medici ZMB 17219* TANZANIA Zanzibar F 568 112 5.071 25 242 77 14 2 4 6 8 82 2 0
D. medici ZMB 5737* TANZANIA Zanzibar F 595 133 4.474 25 247 90 14 2 4 5 8 80 2 0
D. medici NMK 4121 KENYA Kitobo Forest, Taveta U 672 106 6.34 233 66 14 4
D. medici NMK 2538 KENYA Malindi U 597 134 4.455 231 70 2 L
D. medici NMK 2540 KENYA Malindi U 422 123 3.431 229 79 4 P
D. medici NMK 4131 KENYA River Tana, Mchelelo Forest U 484 83 5.831 232 84 12 4
D. medici ZMUC 6124 TANZANIA Amani, E. Usambara Mts U 330 73 4.521 26 246 84 14 2 4 4 8 91 2
D. medici ZMUC 6143 TANZANIA East Usambara Mts U 182 46 3.957 230 92 14 4 4 4 5 59 1 1
D. medici ZMB 48150 TANZANIA Kimboza Forest U 241 14 2 4 4 4 75
D. medici MCZ 54827 TANZANIA Mafia Island U 14 2 4 6 7 P 2 1
D. medici ZMUC 6111 TANZANIA Zanzibar Island U 382 78 4.897 25 251 83 14 2 4 4 6 P 2 0
D. scabra PEM 8602* ETHIOPIA Entrance to Awash Nat. Park M 385 71 5.423 23 212 57 14 2 4 5 6 51 2 2
D. scabra BM 1973.3259* ETHIOPIA Lake Abiata M 435 80 5.438 24 215 56 14 2 4 4 6 75 1 1
D. scabra ZMB 51835* ETHIOPIA Meki M 420 70 6 23 214 57 16 2 4 4 7 77 1 0
D. scabra BM 00.11.29.3 ETHIOPIA west of Omo River M 445 24 212 14 2 4 4 6 60 2 0
D. scabra AAU 489 ETHIOPIA west shore of Lake Chamo M 313 52 6.019 25 218 62 14 2 4 4 4
D. scabra MSNPV CR790* KENYA Gatab, Mount Kulal M 384 81 4.741 24 220 63 14 2 4 4 6 92 1 1
D. scabra MSNPV CR792* KENYA Gatab, Mount Kulal M 312 64 4.875 24 219 66 14 2 4 4 6 84 1 2
D. scabra BM 01.1.3.3* KENYA Gilgil M 398 79 5.038 26 208 65 14 2 2 4 7 76 2 0
D. scabra CAS 130785* KENYA Isiolo M 385 75 5.133 25 203 60 14 2 4 4 6 79 1 0
D. scabra NMK 561 KENYA Isiolo M 386 71 5.437 220 59 14 4 81
D. scabra BM 93.11.21.72* KENYA Kithu-Uri, E. Kikuyu M 350 59 5.932 25 206 54 14 2 4 4 6 66 1 0
D. scabra NMK 2965 KENYA Kora M 456 88 5.182 221 62 14 4 65
D. scabra NMK 1472 KENYA Lake Naivasha M 367 63 5.825 213 61 14 4 80
D. scabra NMK 2343 KENYA Lake Naivasha M 412 84 4.905 206 57 14 4 69
D. scabra NMK 3052 KENYA Solio Ranch, Laikipia M 288 54 5.333 204 50 14 4 66
D. scabra NMK 3021 KENYA Tsavo East N.P. M 236 41 5.756 208 47 14 4 57
D. scabra NMK 1022 KENYA Voi M 298 52 5.731 208 52 14 4 73
D. scabra PEM 5624 RWANDA 15 km S of Butare M 351 76 4.618 25 197 67 14 2 4 4 7 1 1
D. scabra MRAC 18559 RWANDA Astrida M 136 25 192 14 2 4 4 5 61 1
D. scabra MRAC 9209* RWANDA Gabiro M 192 35 5.486 26 201 63 14 2 4 4 7 58 1 2
D. scabra ZMB 22518* RWANDA Lake Kivu M 420 108 3.889 24 201 73 12 2 3 4 7 64 1 0
D. scabra MZUF 24179 SOMALIA 6 km E of Afgoi M 376 78 4.821 23 208 63 14 2 4 4 8 2
D. scabra MSNG 29239 SOMALIA Btwn Comia and Matagoi M 462 74 6.243 25 214 62 13 2 4 4 6 2 1
D. scabra MZUF 31459* SOMALIA Ceelcillan = El Ellan M 389 81 4.802 25 204 62 14 2 4 4 6 74 2 0
D. scabra MZUF 24674* SOMALIA Dinsoor M 285 58 4.914 24 209 64 14 2 4 4 6 70 2 2
D. scabra MSNM  2071 SOMALIA Giohar M 397 22 203 14 2 4 4 4 50
D. scabra MZUF 21968* SOMALIA Lugh – Ganame M 391 78 5.013 24 210 62 14 2 4 4 6 76 2 1
D. scabra MZUF 1134 SOMALIA Maheddei Uen M 461 89 5.18 22 209 63 14 2 4 4 6 2 2
D. scabra FMNH 134939* SOUTH SUDAN Melut M 370 72 5.139 25 213 62 13 2 4 4 6 64 1 0
D. scabra NMW 9964/6+ SOUTH SUDAN ‘Sudan’ (taken as East Equatoria) M 449 88 5.102 23 215 67 14 2 4 4 4 59 1 0
D. scabra AMNH 50791 TANZANIA Chai Camp M 241 45 5.356 27 208 69 14 2 4 4 8 2
D. scabra NMW 9964/17 TANZANIA Dar es Salaam M 23 205 14 2 4 4 6 60 1
D. scabra ZMB 26832* TANZANIA Dar es Salaam M 369 82 4.5 24 201 68 14 2 0 4 5 69 2 0
D. scabra BM 1954.1.2.154* TANZANIA Kange Estate M 400 85 4.706 23 206 67 14 2 2 4 6 71 2 1
D. scabra ZMB 22754 TANZANIA Kibwesa M 432 77 5.61 25 221 61 14 2 4 4 7 1 0
D. scabra MCZ 30205 TANZANIA Maji Malulu, Usandawi M 181 27 225 14 2 4 4 6 54 2 1
D. scabra ZMB 14097* TANZANIA Massailand M 387 78 4.962 27 223 69 14 2 3 4 6 67 1 0
D. scabra USNM 62902* TANZANIA Morogoro M 335 64 5.234 24 218 62 14 3 4 4 5 69 2 0
D. scabra ZMB 19669* TANZANIA Mpwapwa M 398 87 4.575 26 214 69 14 2 4 4 6 54 2 0
D. scabra NMK 1887 TANZANIA Seronera, Serengeti N.P. M 344 73 4.712 211 72 14 4
D. scabra ZMB 16753* TANZANIA Wembere Steppe M 418 83 5.036 26 215 63 14 2 4 4 7 56 2 1
D. scabra BM 68.2.29.101* TANZANIA Zanzibar M 370 74 5 24 214 72 12 2 4 4 6 47 2 0
D. scabra MCZ 18208* TANZANIA Zengeragusa, Mkalama M 460 78 5.897 25 226 58 14 2 4 4 6 79 1 1
D. scabra LIV 1962.330.22 UGANDA Kayanja, Nyabusonji M 360 76 4.737 200 66
D. scabra BM 51.1.5.79* UGANDA Mbarara, Ankole M 375 84 4.464 25 207 68 14 2 4 4 7 78 2 0
D. scabra LIV 1960.67.4 UGANDA Ruborogoto, Ishingiro M 198 38 5.211 188 63 4 4 6
D. scabra LIV 1960.67.7 UGANDA Ruborogoto, Ishingiro M 329 72 4.569 200 65 4 4 8
D. scabra LIV 1962.168.14 UGANDA Ruzinga, Ankole M 453 98 4.622 199 63 4 4 5
D. scabra LIV 1960.67.8 UGANDA Rwensama, Kashali M 352 83 4.241 185 64 4 4 6
D. scabra BM 07.12.1.19* UGANDA Wadelai M 333 67 4.97 23 214 67 14 2 4 4 6 65 1 0
D. scabra MNHN 6565* ‘ARABIA’ ? F 252 39 6.462 24 244 54 14 2 4 4 6 55 2 1
D. scabra MRAC 18563* BURUNDI Kabuyenge F 702 103 6.816 26 234 60 14 2 2 2 4 75 2 1
D. scabra MRAC 18565* BURUNDI Kahiro Dist. F 547 77 7.104 25 227 53 14 2 4 4 6 52 2 1
D. scabra MRAC 18564* BURUNDI Mugera F 495 72 6.875 25 220 52 14 2 4 4 6 60 2 1
D. scabra NMW 15298* ETHIOPIA ? F 590 87 6.782 28 219 53 14 2 4 6 7 56 2 1
D. scabra AAU 494 ETHIOPIA Arba Minch F 229 48 14 2 4 4 6
D. scabra AAU 679 ETHIOPIA Arba Minch F 668 92 7.261 22 243 54 14 2 4 4 5
D. scabra BM 16.6.24.12* ETHIOPIA Aware Melca F 415 71 5.845 24 221 59 14 2 4 4 6 56 1 0
D. scabra ZFMK 17763* ETHIOPIA Bulcha F 522 80 6.525 27 241 50 14 2 4 4 6 79 1 0
D. scabra PEM 8603* ETHIOPIA Entrance to Awash Nat. Park F 217 29 7.483 24 234 57 14 2 4 4 6 54 2 1
D. scabra BM 1964.1069 ETHIOPIA Gojam District F 525 76 6.908 23 219 47 12 2 4 4 7 0 0
D. scabra ZMB 77053* ETHIOPIA Meki F 825 121 6.818 25 231 57 14 2 4 4 6 85 0 2
D. scabra BM 00.11.29.2* ETHIOPIA W of Omo River F 593 87 6.816 25 226 53 14 2 4 4 6 55 2 1
D. scabra PEM 5627 KENYA 45 km W of Ewaro Ngiro (Narok) F 26 236 49 14 2 4 4 7 1 0
D. scabra USNM 42016 KENYA Aberdare Mtns F 305 38 8.026 26 220 47 14 2 4 4 6 69
D. scabra NMK 2824 KENYA Buna, Wajir Dist. F 411 73 5.63 222 55 14 4 67
D. scabra MSNPV CR791* KENYA Gatab, Mount Kulal F 324 61 5.311 22 219 64 14 2 4 4 6 94 1 1
D. scabra BM 01.1.3.4* KENYA Gilgil F 532 60 8.867 28 229 43 14 4 4 5 6 73 2 0
D. scabra CAS 147836 KENYA Hell’s Gate, Naivasha F 350 42 8.333 26 233 46 14 2 4 4 6 2 1
D. scabra CAS 130784* KENYA Isiolo F 550 76 7.237 26 234 51 14 2 4 4 7 88 1 0
D. scabra BM 51-1-3-70 KENYA Kilifi F 480 25 227 14 2 2 4 6 70 2 0
D. scabra MCZ 40584* KENYA Laini, Tana River F 427 62 6.887 25 225 53 14 2 4 4 6 83 1 0
D. scabra NMK 1415 KENYA Lake Naivasha F 230 50 14 4 72 2 1
D. scabra NMK 2083 KENYA Lake Naivasha F 229 52 14 4 72 1 2
D. scabra NMK 2855 KENYA Lake Naivasha F 247 240 14 4 74
D. scabra NMK 3184 KENYA Lake Naivasha F 867 230 14 4
D. scabra NMK 3319 KENYA Lake Naivasha F 657 98 6.704 215 50 14 4 72
D. scabra NMP 1260* KENYA Lake Naivasha F 509 76 6.697 27 228 52 14 2 4 4 8 72 1 1
D. scabra NMK 2819 KENYA Lolgorien, Trans-Mara F 372 58 6.414 232 55 14 6 69
D. scabra NMK 3005 KENYA Lolgorien, Trans-Mara F 281 42 6.69 233 53 14 4 73
D. scabra NMK 930 KENYA Manyani, Tsavo West N.P. F 368 57 6.456 231 52 14 4 60
D. scabra MCZ 40583 KENYA Mombosasa, near Witu F 648 97 6.68 25 228 55 14 2 4 4 6 76
D. scabra CAS 147837* KENYA Naivasha F 300 34 8.824 26 238 45 14 2 4 4 6 67 1 1
D. scabra CAS 147838* KENYA Naivasha F 325 40 8.125 25 226 45 14 2 4 4 6 75 2 1
D. scabra NMK 3103 KENYA Ol Ari Nyiro Ranch F 390 53 7.358 228 49 14 4 75
D. scabra USNM 41974 KENYA Sotik, Uaso Nyiro F 343 42 8.167 26 230 45 14 2 4 4 6 71 2
D. scabra MCZ 11501* KENYA Uaso Nyiro Plains F 362 46 7.87 26 221 47 14 2 4 3 6 75 1 1
D. scabra CAS 130661 KENYA Wajir F 565 95 5.947 26 65 14 2 4 4 6 56 2 2
D. scabra NMK 2262 KENYA Wajir F 232 50 14 2 63 2 0
D. scabra MCZ 40585* KENYA Wema (Ngatana) F 268 40 6.7 25 215 55 14 2 4 4 6 73 0 0
D. scabra MRAC 9214* RWANDA Gabiro F 637 83 7.675 25 235 48 14 2 4 5 6 83 2 0
D. scabra MZUF 1118* SOMALIA Afgoi F 180 31 5.806 25 218 56 14 2 4 6 7 55 0 0
D. scabra MZUF 1133* SOMALIA Afgoi F 344 57 6.035 24 228 64 14 2 4 4 7 74 2 1
D. scabra MSNG 42710A SOMALIA Belet Amin F 587 78 7.526 23 216 51 14 2 4 4 6 1 0
D. scabra MZUF 24996* SOMALIA Dinsoor F 557 93 5.989 28 222 57 14 2 4 4 7 68 2 0
D. scabra MSNG 28808 SOMALIA Lugh F 519 66 7.864 24 222 54 14 4 4 6 4 2 1
D. scabra BM 98.1.28.14* SOMALIA Lugh F 218 32 6.813 26 227 58 14 2 4 4 6 58 2 0
D. scabra MSNM  2148 SOMALIA Magadi (= Uagadi) F 496 80 6.2 22 202 57 14 2 4 4 6 51
D. scabra MZUF 2802* SOMALIA Mogadishu F 210 36 5.833 25 224 59 13 2 4 4 6 76 2 0
D. scabra MZUF 27156* SOMALIA near Mareri F 367 57 6.439 24 215 54 14 2 4 4 7 79 1 0
D. scabra RMNH 24399 SUDAN 40 km E of Rabak F 222 55 50
D. scabra MNHN 7227* SUDAN Nil Blanc F 673 90 7.478 25 226 52 14 2 4 4 6 55 1 1
D. scabra BM 09.10.15.35 SUDAN Sennar Province F 240 23 227 14 2 4 4 6 65 1 1
D. scabra ZMB 21062* TANZANIA Amani, Usambara Mtns F 740 124 5.968 27 234 65 14 2 4 4 7 63 2 1
D. scabra PEM 5626* TANZANIA Mapenduzi, Dodoma F 265 42 6.31 26 249 60 14 2 3 6 10 60 2 2
D. scabra ZMB 11934A* TANZANIA Massailand F 283 41 6.902 26 231 61 14 2 3 3 6 59 2 0
D. scabra ZMB 11934B* TANZANIA Massailand F 793 110 7.209 27 242 54 14 2 4 4 6 69 2 0
D. scabra USNM 62903 TANZANIA Morogoro F 500 85 5.882 26 232 57 14 2 4 5 8 67 0
D. scabra USNM 62905 TANZANIA Morogoro F 447 72 6.208 26 223 59 14 2 4 4 6 79 2
D. scabra BM 87.11.3.30* TANZANIA Mount Kilimanjaro F 346 52 6.654 25 230 58 14 2 2 3 6 67 1 0
D. scabra ZMB 13304* TANZANIA Mpwapwa F 403 63 6.397 27 225 57 14 2 2 4 6 62 2 0
D. scabra FMNH 27786 TANZANIA Ngare, Nanyuki F 539 26 229 55 14 2 4 4 8 69 2
D. scabra NMZB 1681* TANZANIA Tumba, Rukwa Valley F 710 103 6.893 26 238 54 14 2 3 4 6 60 2 2
D. scabra NMZB 659* TANZANIA Tumba, Rukwa Valley F 760 109 6.972 26 236 53 14 2 4 4 6 65 2 2
D. scabra MCZ 30207* TANZANIA Ukerewe Island F 524 70 7.486 26 223 49 14 2 4 4 8 62 2 1
D. scabra MCZ 30208 TANZANIA Ukerewe Island F 708 25 231 14 3 4 4 6 64 2 1
D. scabra BM 68.2.29.100* TANZANIA Zanzibar F 476 78 6.103 25 226 59 13 2 4 4 6 46 2 0
D. scabra BM 68.2.29.102* TANZANIA Zanzibar F 335 51 6.569 27 233 58 14 2 4 4 7 53 2 0
D. scabra LIV 1962.106.6 UGANDA Kaburangiri, Kashali F 270 32 8.438 216 52 4 4 5
D. scabra LIV 1960.67.6 UGANDA Kakoba, Kashari F 612 85 7.2 216 49 3 4 6
D. scabra LIV 1959.83.1 UGANDA Mbarara, Ankole F 640 90 7.111 217 51 2
D. scabra BM 1961.1804* UGANDA Mbarara, Kashari F 690 105 6.571 25 216 53 14 2 4 4 6 71 2 2
D. scabra BM 03.12.2.21* UGANDA Mulema F 550 84 6.548 27 219 54 14 2 4 4 6 72 2 0
D. scabra LIV 1962.106.5 UGANDA Namatyobola, Kashali F 354 53 6.679 206 51 3 4 6
D. scabra LIV 1962.106.35 UGANDA Ruborogoto, Ishingiro F 810 113 7.168 222 53 4 4 6
D. scabra LIV 1962.106.20 UGANDA Rukoni, Kwampala F 480 63 7.619 220 50 4 4 6
D. scabra BM 1959.1.5.59 UGANDA Ruzinga, Isingiro F 650 92 7.065 230 57 14 2 4 4 6 78 1 1
D. scabra BM 1969.288 ETHIOPIA Arba Minch U 155 27 5.741 26 213 59 14 2 4 4 7 82 0 0
D. scabra BM 16.6.24.11 ETHIOPIA Aware Melca U 210 28 7.5 25 235 55 14 2 4 4 6 66 1 1
D. scabra IRSNB 5499 RWANDA Gabiro U 443 66 6.712 26 226 54 14 2 4 4 5 70
D. scabra PEM 5625 RWANDA Nyagatare U 163 31.5 5.094 25 194 59 14 2 4 4 6 55 2 0
D. scabra USNM 72476 TANZANIA Kikombo, Dodoma U 200 28 7.143 26 253 67 14 2 4 4 6 68 1
D. scabra ZMUC 6136 TANZANIA Minziro Forest Reserve U 176 27 6.519 25 216 52 11 2 4 3 5 63 2 1
D. scabra ZMUC 6137 TANZANIA Minziro Forest Reserve U 188 31 6.065 26 226 56 14 2 4 4 6 1 0
D. scabra NMW 9964/9+ TANZANIA Tanga U 195 37 5.270 25 204 63 14 2 3 4 6 65 1 0
D. taylori BM 1992.501 SOMALIA 2.5 km SW of Hargeisa M 305 199 14 2 4 4 6 44 2 1
D. taylori BM 51.1.7.69* SOMALIA Garadak M 23 202 59 14 2 4 4 4 49 2 1
D. taylori BM 49.2.2.1* SOMALIA Haud M 188 34 5.529 23 196 55 12 2 3 4 6 55 2 0
D. taylori BM 49.2.2.2* SOMALIA Haud M 172 30 5.733 24 198 60 14 2 4 4 6 50 2 1
D. taylori BM 49.2.2.5 SOMALIA Ogo M 408 83 4.916 25 216 68 14 2 4 4 6 53 0
D. taylori MNHN 84-707* DJIBOUTI ? F 477 68 7 25 222 52 14 2 4 4 6 59 2 1
D. taylori BM 49.2.2.4 SOMALIA Hargeisa Dist. F 530 24 208 14 2 2 2 6 53 2 1
D. taylori BM 49.2.2.3* SOMALIA Haud F 189 23 8.217 24 211 44 14 2 3 4 6 59 2 1

FIGURE CAPTIONS

 

Figure 1. Tail length relative to snout-vent vent in males and females of Dasypeltis atra, D. confusa, D. fasciata, D. medici, D. scabra and D. abyssina in north-eastern Africa.

2_Fig.-2A_Dasypeltis-medici_vicinity-of-Dar-es-Salaam,-coastal-Tanzania_W.R

Figure 2. Dasypeltis medici. A: Vicinity of Dar-es-Salaam, Tanzania (Photo: W.R. Branch); B: Juvenile from East Usambara Mountains, Tanzania (Photo: W.R. Branch); C: Watamu, Kenya (Photo: S. Spawls); D: Tanga, Tanzania (Photo: S. Spawls); E & F: Diani Beach, Kenya (Photos: F. Finke). [A & B are colour pattern ‘8’ (‘medici’), C & D are pattern ‘3’ (‘lamuensis’, mostly unmarked), and E & F—same individual—are intermediate (weakly marked)]

3_Fig.-2B_Dasypeltis-medici_juv_E-Usambara-Mountains,-Tanzania_W.R

Figure 3. Geographical distribution of the genus Dasypeltis in north-eastern Africa and south-western Arabia.

4_Fig.-2C_Dasypeltis-medici_Watamu,-Kenya_S

Figure 4. Dasypeltis confusa. [‘5L’ colour pattern of Gans 1959] A: Bisu (Kigezi), Uganda. (BM 1951.1.3.69, Photo: M.F. Bates); B: Kacheliba, Kenya (NMK 489, Photo: P. Malonza); C: Mumias, Kenya (CAS 141784, Photo: J. Vindum); D: Lake Kivu, Rwanda (ZMB 77441, Photo: F. Tillack).

5_Fig.-2D_Dasypeltis-medici_Tanga,-N-Tanzania_S

Figure 5. Dasypeltis scabra. [‘5N’ colour pattern of Gans 1959] A: Dodoma, Tanzania (Photo: W.R. Branch); B: Masiko Tembo, Tanzania (Photo: W.R. Branch), C: Wajir, Kenya (Photo: S. Spawls); D: Juvenile from Loita Hills, Kenya (Photo: T. Mazuch); E: Gilgil, Kenya (Photo: S. Spawls); F: MSNM 974, Magadi, Somalia (Photo: Museo Civico di Storia Naturale, Milan); G: Awash National Park, Ethiopia (Photo: S. Spawls); H: Awash National Park, Ethiopia (Photo: S. Spawls).

Figure 6. Dasypeltis scabra. Close-up dorsal view (in alcohol) of MNHN 7227 (syntype of Rachiodon abyssinus) from the White Nile River (possibly near Khartoum, Sudan) showing a ‘5N’ pattern (see Gans 1959) consisting of dark oval saddles and dark dorso-ventrally elongated lateral bars, with paler centres. (Photo: M.F. Bates)

Figure 7. Dasypeltis bazi. A: Dorsal and lateral views of the largest known female (BM 97.10.28.575) from Fayoum, Egypt (Photo: M.F. Bates); B: Ventral view of the same specimen (Photo: Natural History Museum, London).

 

Figure 8. Dasypeltis bazi. Dorsal (left), lateral (middle) and ventral (right) views of the head of the largest known female (BM 97.10.28.575) from Fayoum, Egypt. (Photo: Natural History Museum, London)

 9_Fig.-4A_Dasypeltis-confusa_NHMUK-1951.1.3.69_M.F

Figure 9. Dasypeltis bazi. Dorsal (A) and ventral (B) views of a male (ZMB 1861) from ‘Upper Egypt’. (Photo: F. Tillack)

 10_Fig.-4B_Dasypeltis-confusa_NMK-S489_Male_P

Figure 10. Dasypeltis bazi. Dorsal (left), lateral (middle) and ventral (right) views of the head of a male (ZMB 1861) from ‘Upper Egypt’. (Photo: F. Tillack)

 11_Fig.-4C_Dasypeltis-confusa_Mumias-2_cropped_CAS-141784_J

Figure 11. Dorsal views of Dasypeltis loveridgei comb. nov. [‘6N’ colour pattern of Gans 1959; specimen also has 6 supralabials on either side of head, with 2nd and 3rd in contact with orbital] (female, ZMB 33670a, top) from Otjimbingwe, Namibia, and Dasypeltis bazi (male, ZMB 1861, below) from ‘Upper Egypt’ showing the general similarity in colour pattern. (Photo: M.F. Bates)

 12_Fig.-4D_Dasypeltis-confusa_Lake-Kivu,-Rwanda_F

Figure 12. Dasypeltis cf. bazi. Dorsal view of the anterior part of the body of a specimen (ZFMK 38415) from Erkowit, Sudan. (Photo: W. Böhme)

13_Fig.-5A_Dasypeltis-scabra_Dodoma,-Tanzania_W.R

Figure 13. Dasypeltis taylori sp. nov. Body and head views of the holotype (BM 1949.2.2.2) from Haud, Somaliland. (Photos: Natural History Museum, London)

14_Fig.-5B_Dasypeltis-scabra_Masiko-Tembo,-Tanzania_W.R

Figure 14. Dasypeltis taylori sp. nov. Paratype (MNHN 1884.0707) from Djibouti. (Photo: I. Ineich)

15_Fig.-5C_Dasypeltis-scabra_Wajir,-Kenya_S

Figure 15. Haud, Somaliland—type locality of Dasypeltis taylori sp. nov. (Photo: T. Mazuch)

16_Fig.-5D_Dasypeltis-scabra_juv_Loita-Hills,-Kenya_T

Figure 16. Vicinity of Hargeisa, Somaliland—habitat of Dasypeltis taylori sp. nov. (Photo: T. Mazuch)

17_Fig.-5E_Dasypeltis-scabra_Gilgil,-Kenya_S

Figure 17. Dasypeltis atra. Black morph (‘2M’ of Gans 1959)—A: Keriyo near Mulu Cattle Farm, Ethiopia (Photo: S. Spawls); B: Limuru, Kenya (Photo: S. Spawls); Brown morph (‘2B’ of Gans 1959)—C: ELI 1264 (field number), Kibira Forest, Burundi (Photo: E. Greenbaum); Patterned morph (‘9’ of Gans 1959)—D: Holotype (ZMB 9224) of Dasypeltis lineolata Peters from Kitui, Kenya (Photo: M.F. Bates); Patterned morph (‘2Bx’ of Gans 1959)—E: Addis Ababa, Ethiopia (Photo: S. Spawls); F: Kitengela, Kenya (Photo: S. Spawls); G: Chuka Mountain, Kenya (Photo: S. Spawls); H: ELI 2760 (field number), Bwindi Impenetrable National Park, Uganda (Photo: E. Greenbaum); I: Addis Ababa, Ethiopia (Photo: S. Spawls); J: Masiko Tembo, Tanzania (Photo: W.R. Branch); K: PEM R17336, Klein’s Camp, Mkomazi Game Reserve, Tanzania (Photo: W.R. Branch); L: Kakamega Forest, Kenya (Photo: T. Mazuch); M: Neghelle, Ethiopia (Photo: V. Trailin).

Figure 18. Dasypeltis atra from Kakamega Forest, Kenya, with a single postocular. (Photo: T. Mazuch)

19_Fig.-5G_Dasypeltis-scabra_Awash-National-Park,-Ethiopia_S

Figure 19. Dasypeltis fasciata. [Colour pattern ‘4’ (checker board) of Gans 1959] A & B: Dorsal and ventral views, respectively, of TCWC 23839, Nandi, Bwamba Forest, Uganda (Photos: T. Hibbits); C: Makeni, Sierra Leone (Photo: W.R. Branch).

20_Fig.-5H_Dasypeltis-scabra_Awash-Nat-Park,-Ethiopia_-Echis-mimic_S

Figure 20. Dasypeltis abyssina. Dorsal (A) and ventral (B) aspects of the lectotype (MNHN 6567) from Ethiopia (Photos: M.F. Bates); C: Mulu Cattle farm, Ethiopia (Photo: S. Spawls); D: Ethiopia (Photo: C. Tilbury); E: Mulu Cattle farm, Ethiopia (Photo: S. Spawls); F: Keriyo near Mulu, Ethiopia (Photo: S. Spawls); G: Eritrea (NMW 15301–3, Photo: M.F. Bates).

 

Figure 21. Habitat of Dasypeltis abyssina and D. atra near Lake Tana, north of Addis Zemen, Ethiopia. (Photo: T. Mazuch)

22_Fig.-7A_D

Figure 22. Habitat of Dasypeltis abyssina south of Gondar, northern Ethiopia. (Photo: T. Mazuch)

23_Fig.-7B_Dasypeltis-scabra-97.10.28

Figure 23. Dasypeltis crucifera sp. nov. Dorsal (A) and ventral (B) views of the holotype (ZMB 7631) from Bogos, Eritrea (Photo: F. Tillack).

24_Fig.-8A_Dasypeltis-aegyptia_BM-97.10.28

Figure 24. Dasypeltis crucifera sp. nov. Dorsal (A), lateral (B) and ventral (C) aspects of the head of the holotype (ZMB 7631) from Bogos, Eritrea (Photo: F. Tillack).

25_Fig.-8B_Dasypeltis-aegyptia_BM-97.10.28

Figure 25. Dasypeltis crucifera sp. nov. Dorsal (A) and ventral (B) views of the allotype (MZUT 3582) from Agordat, Eritrea (Photo: F. Andreone).

26_Fig.-8C_Dasypeltis-aegyptia_BM-97.10.28

Figure 26. Keren, Eritrea—type locality of Dasypeltis crucifera sp. nov. (Photo: T. Mazuch)

Figure 27. Dasypeltis arabica sp. nov. [‘5I’ colour pattern of Gans 1959] Dorsal (A), ventral (B) and lateral (C) views of the holotype (BM 1987.2192) from Sana’a, northern Yemen. (Photos: M.F. Bates)

Figure 28. Dasypeltis arabica sp. nov. [‘5I’ colour pattern of Gans 1959] A: Dorsal (left), lateral (middle) and ventral (right) aspects of the head of the holotype (BM 1987.2192) from Sana’a, northern Yemen (Photo: Natural History Museum, London); B: Dorso-lateral aspect of the head of the holotype (in water, Photo: M.F. Bates).

Figure 29. Dasypeltis arabica sp. nov. [‘5I’ colour pattern of Gans 1959] A: Dorsal view of a paratype (BM 1987.1018) from Ajibah, Saudi Arabia. (Photo: M.F. Bates); B: Dorsal (left), lateral (middle) and ventral (right) aspects of the head of the paratype (Natural History Museum, London).

Figure 30. Echis megalocephalus (BM 1987.3217) from Nocra Island, Dahlak Archipelago, Eritrea—a possible model species for mimicry by Dasypeltis arabica sp. nov. and Dasypeltis crucifera sp. nov. (Photo: T. Mazuch)

Figure 31. Principal Component Analysis of the genus Dasypeltis in north-eastern Africa and south-western Arabia. (MBSR = midbody scale rows, V = ventrals, SC = subcaudals, SL = supralabials, Pre Oc = preoculars, Pos Oc = postoculars, Ant T = anterior temporals, Pos T = posterior temporals, Cycles = pattern cycles, Frontal = frontal pitting, Sulcus = Inter-prefrontal sulcus)

Figure 32. Discriminant Function Analysis of the genus Dasypeltis in north-eastern Africa and south-western Arabia.

33_Fig.-12_Erkowit_ZFMK-38415-cropped

Figure 33. Principal Component Analysis of the genus Dasypeltis in the Horn of Africa and Egypt. (MBSR = midbody scale rows, V = ventrals, SC = subcaudals, SL = supralabials, Pre Oc = preoculars, Pos Oc = postoculars, Ant T = anterior temporals, Pos T = posterior temporals, Cycles = pattern cycles, Frontal = frontal pitting, Sulcus = Inter-prefrontal sulcus)

34_Fig.-13_Daypeltis-taylori_Haud,-Somaliland_NHMUK-1949.2.2.2-(old-no

Figure 34. Discriminant Function Analysis of the genus Dasypeltis in the Horn of Africa and Egypt.

 

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