Panaspis massaiensis ( Angel, 1924 ) Kilunda & Conradie & Wasonga & Jin & Peng & Murphy & Malonza & Che, 2019

Kilunda, Felista Kasyoka, Conradie, Werner, Wasonga, Domnick Victor, Jin, Jie-Qiong, Peng, Min-Sheng, Murphy, Robert W., Malonza, Patrick Kinyatta & Che, Jing, 2019, Revalidation and resurrection of Panaspis massaiensis (Angel, 1924) and the description of a new species of Panaspis Cope (Squamata: Scincidae) from south-eastern Kenya, Zootaxa 4706 (2), pp. 255-274 : 261-265

Panaspis massaiensis ( Angel, 1924) comb. nov.

Maasai Snake-eyed skink

( Fig. 3–4 )

Chresonymy

Ablepharus wahlbergii: Loveridge (1920: 158, 1923: 963 , 1929: 79, 1933: 324, 1936:72, 1957:219; Barbour & Loveridge (1928: 163).

Panaspis wahlbergii: Broadley & Howell (1991:16) ; Spawls et al. (2002:155).

Afroablepharus wahlbergi: Branch (2005:77) .

Panaspis wahlbergi: Razzetti & Msuya (2002:53) ; Medina et al. (2016 as Tanzania sp. 1 & 2); Spawls et al. (2018: 165).

Angel (1924) described Ablepharus massaiensis from the Maasai plains near Nairobi, and distinguished it from typical P. wahlbergi on the basis that his specimen has 26 midbody scale rows and exhibit two instead of three oculars. But when Barbour & Loveridge (1928) and subsequently Loveridge (1929) examined material from the Nairobi region they concluded that all material exhibit three oculars and that the type of A. massaiensis must be an aberrant with only two oculars. Although they did agree that the Kenyan populations have a higher midbody scale rows (26 versus 24) count than Mozambique material they considered A. massaiensis to be a junior synonym of P. wahlbergi . Our newly collected material from Chyulu Hills in southern Kenya, Nairobi and other similar highland areas conform to the type description in that they exhibit the same dorsal coloration and white ventrolateral stripe and have an average 26 midbody scale rows. Our newly collected genetic material from near the type locality further allowed us to validate the status of Ablepharus massaiensis (see Results above). We therefore confidently assign our material to Ablepharus massaiensis , resurrect this species from the synonym and transfer this species formally to Panaspis .

Holotype. MHNP 1904.306, adult unsexed collected 9 October 1903 by Charles Alluaud from the Maasai plains in Nairobi region, Kenya ( Fig. 3 ). Basic measurements (in mm) of the holotype as provided in the original description: SVL = 42 mm, HL = 6 mm, HW = 4 mm, front limb = 6 mm, back limb = 11 mm.

Comparative additional material examined. NMK-L3857/1 (Field No. KIZ 030203), adult male collected on 18 September 2016 in Chyulu Hills National Park, Makueni County, Kenya (-2.71944°, 37.97389°; 1200 m a.s.l.) by Patrick K. Malonza, Justus Ochong’ and Felista K. Kilunda; NMK-L3857/2 (Field No. KIZ 030202), adult male with same collection details as NMK-L3857/1 ( Fig. 4 ); NMK-L3225 (Field No. CHY04), adult male collected on 13 September 2009 in Chyulu Hills National Park (-2.78811°, 37.96244°; 1759 m a.s.l.) by Patrick K. Malonza and Vincent Muchai; NMK-L2198/2, adult female collected on 6 August 1996 at National Museums of Kenya compound-Nairobi (-1.27370°, 36.81369°; 1658 m a.s.l.) by F. Muhoro, Victor Wasonga and Martha Nzisa; NMK-L2977/3, adult female collected on 18 February, 2002 in Nairobi National Park (-1.3616°, 36.8452°; 1636 m a.s.l.) by Patrick K. Malonza and Vincent Muchai; NMK-L682, adult male collected on 31 July 1970 in Nairobi (-1.27370°, 36.81369°; 1658 m a.s.l.) by Peter Nares; NMK-L2066, adult female collected on 26 August 1993 in Muthwani Village, Mbiuni- Machakos by Luis Malaret; NMK-L2046, adult collected on 22 April 1993 in Mutula Village, Mbiuni- Machakos by Luis Malaret; NMK-L2807 (Field No. GK010), adult female collected on 15 September 2005 in Sagalla Hill-Taita (-3.51368°, 38.57702°; 1100 m a.s.l.) by Greshon Kisombe; NMK-L3319 (Field No. JN0071), adult female collected on 21 March 2010 in Arabuko-Sokoke Forest-Kilifi (-3.27803°, 39.97958°; 31 m a.s.l.) by Joash Nyamache and Peter Mwasi; NMK-L1044, adult female collected on 13 December 1970 at Rapogi Secondary School, Sare-Migori (0.90000°, 34.46667°; 1449 m a.s.l.) by Paul Brother; NMK-L3889, adult male collected in July 1971 in Mandera (3.93333°, 41.86667°; 366 m a.s.l.) by Robert C. Bob Drewes; NMK-L3373 (Field No. PKM0237), adult female collected on 9 June 2010 in Mathews Range (1.2514°, 37.29332°; 1376 m a.s.l.) by Patrick K. Malonza and Justus Ochong; NMK-L3435 (Field No. VM0053), adult female on 13 December 2010 in Mt. Warges Forest-Mathews Range (1.06694°, 37.35225°; 1465 m a.s.l.) by Vincent Muchai, Justus Ochong; NMK-L4094/2 (Field No. W5), collected on 15 February 2018 in Mathews Range (1.0175°, 37.48722°; 1627 m a.s.l.) by Joash Nyamache; NMK-L3059 (Field No. PKM00632), adult male collected on 14 February 2007 in Ngaya forest (0.38869°, 38.02822°; 1334 m a.s.l.) by Stephen Spawls; NMK-L2311 (Field No. 53), adult female collected on 22 May 1999 in Muamba village-Kinondo, Kwale (-4.48333°, 39.41833°; 10 m a.s.l.) by William R. Branch; NMK-L1333 (Field No. PKM0535), adult female collected on 12 December 2012 in Mukurumudzi River- Maumba, Shimba Hills Kwale (-4.39287°, 39.42903°; 55 m a.s.l.) by Patrick K. Malonza, David M. Mulwa and Joash Nyamache; NMK-L2706/1 (Field No. E19), adult male collected on 11 October 2004 in Endau Hill by Patrick K. Malonza and Joash Nyamache; PEM R16769, adult female collected 6 December 2005 from Klein’s Camp, Loliondo Game Controlled Area (Mara); Tanzania (-1.83500°, 35.09731°) by William R. Branch; PEM R17323, adult female collected on 22 January 2007 from Klein’s Camp, Tanzania (-1.83497°, 35.24625°; 1884 m a.s.l.) by William R. Branch; PEM R20799, juvenile collected on 7 December 2005 from Grometi River, Klein’s Camp Concession (Serengeti), Tanzania (-1.83600°, 35.25889°) by William R. Branch; PEM R25125 and -25126 (Field Number. WRB021 and WRB28 respectively), adult males collected 7 December 2008 in Arusha suburb, Tanzania (-3.38614°, 36.68339°) by Joe Berarducci.

Diagnosis. Panaspis massaiensis comb. nov. can be distinguished from other members of the East Africa Panaspis wahlbergi-maculicollis group by the following combination of characteristics: 1) the presence of a white ventrolateral stripe (absent in P. tsavoensis sp. nov. and P. maculicollis ); 2) smaller average size (39.0 mm SVL versus 41.8 mm SVL in P. wahlbergi ); 3) longer tail about 1.5 times SVL (versus 1.1 times SVL in P. wahlbergi ), much shorter tail than P. megalurus (approx. 3 times SVL); 4) fused frontoparietal (divided in P. megalurus ); 5) average 26 midbody scale rows (versus 24 in P. wahlbergi ). Genetic pair-wise difference from its nearest congener P. tsavoensis sp. nov. is 5.87 % (see Table 2 ), from which it can be easily diagnosed with the presence of a white ventrolateral stripe (absent in 5 out of 23 specimens examined); 26 versus 24 midbody scale rows; longer eye-tympanum distance and eye-snout distance ( Table 4 ); higher number of subdigital lamellae under the fourth finger (7–9 versus 5) and higher number of supralabials in front of subocular (6–7 versus 5).

Comparative description of additional material (with specific emphasis on NMK-L3857/1). Adult male. Small slender skink with reduced functional limbs, tail longer than SVL (mostly>60%), rostral scale broader than high, nostril pierced between a single nasal, nasals 2, widely separated behind the rostral, frontonasal wider than long, prefrontals separate, frontal small almost as broad as long, frontoparietal large and concave posteriorly and in contact with frontal, parietal and interparietal, parietals large and in contact behind interparietal, supraoculars 3, supraciliaries 5, loreals 2, preocular 3, upper labials (supralabials) 7 with four anterior–subocular, lower labials (infralabials) 6, mental wider than long, posteriorly curve edged, a large postmental followed by three postpostmentals. Dorsal scales smooth and imbricate, midbody scale rows 26. Ventrals smooth and overlapping, digits clawed, a single row of unicarinate lamellae under the toes, palmer tubercles raised and conical. Tail full. Head short, indistinct from neck; snout conical ( Fig. 3 ). Ear opening oval, clearly visible; body elongated, body scales smooth. Fore and hind limbs relatively long, slender; all five digits moderately long and slender. Tail cylindrical and more than half total body length.

Coloration of NMK-L3857/1. Dorsum brown with six longitudinal black broken stripes along the back. White ventrolateral stripe present from snout to groin. Above the ventrolateral stripe is dark broad black stripe that extends the entire flanks from the snout through the eye to the tail tip. On the back within the brownish vertebral stripe are six longitudinal lines or dashed stripes ( Fig. 3–4 ). Preserved specimen body venter is white ( Fig. 3 ). Lower labials with scattered black spots giving the lips a bared appearance. Tail dorsum brownish with white underside. In life, breeding males have pink-orange underside including tail and limbs ( Fig. 4 ).

Variation of additional material examined. The variation in body measurements and scalation are shown in Table 4 . However, most of the specimens are similar in many characters. There are variations in the absence and presence of the white ventrolateral stripe and the number of dorsal stripes from none to six in the same population. Notably are specimens from the moist coastal strip and the Mathews Range in northern Kenya. It is unclear whether this is a sexual difference or breeding period variation or even cryptic species.

Natural history. Panaspis massaiensis comb. nov. is a terrestrial species found to burrow on loose debris or in holes during the day. They move quickly on the surface within leaf litter or among grasses and can also be found under stones and similar micro-habitats. Loveridge (1920) recorded specimens in the stomachs of Cattle Egret ( Bubulcus ibis ) and Battersby’s Green Snake ( Philothamnus battersbyi ) from Morogoro and Nairobi respectively. The species lays eggs. Loveridge (1923) recorded a large female with six developing eggs measuring 6 x 3 mm.

Distribution, habitat and conservation status. Panaspis massaiensis comb. nov. lives in moist savanna in lowland and highland areas. In Chyulu Hills, the species occurs from about 1000–2200 m a.s.l. Elsewhere the species has been recorded in other areas all within the range of 0–2200 m a.s.l. including Taita Hills (1000–2000 m a.s.l.), Nairobi area (1700 m a.s.l.), Namanga Hills (1500 m a.s.l.), Endau Hill-Kitui East (1000 m a.s.l.), Makuli & Nzaui-Makueni Hills (1000 m a.s.l.), Mitunguu-Meru (1100 m a.s.l.), Kijege Hill-Chiakariga (1100 m a.s.l.), Kiang’ombe Hill (1500 m a.s.l.), Arabuko Sokoke Forest, Kwale (Shimoni, Kinondo, Shimba Hills) 0–400 m a.s.l.) as well as Kajiado, Mukurweni, Thika, Rapogi in Migori, and Mandera. Loveridge (1920:158, 1929:963, 1936:72) documented specimens from Nairobi, Mt Kenya, and Lukenya that can be referred to this species with confidence. Currently the only genetically confirmed records for Tanzania are from Arusha (1400 m a.s.l) and Klein’s camp, Serengeti National Park (approx. 1884 m a.s.l), but it is expected to be much widespread. The following material could be assigned to this species but verification through molecular data and morphological examination is necessary: Longido West, Morogoro, Mkuyuni, Handeni ( Loveridge 1920: 158); Dar es Salam, Mkindo, Mbweni, Kilosa, Mbala, Mpanira-kwa-Sagoi ( Loveridge 1923: 963); Bagamoyo, Mpwapwa, Ugogo, Masiliwa, Turu, Kitungulu, Urungu, Nyamkolo, Lake Tanganyika, Managasini, Kasanga ( Loveridge 1933: 324); Uleia ( Loveridge 1936:72), and Dar es Salaam, Bagamoyo ( Barbour & Loveridge 1928: 163). Caution should be taken as shown by Medina et al. (2016) as more cryptic species might be present between these records and the undescribed northern Mozambique species. For example, specimens from moist coastal Kenya and Tanzania within the Zanzibar Inhambane floristic zone that ranges from Mozambique may need more molecular scrutiny as they may be the same as the undescribed northern Mozambique species. Other potential cryptic species could be those from Mandera and Mathews Range in the Kenyan northern frontier region. Given its widespread distribution and abundance, the species should be categorised as Least Concern under IUCN criteria.

Figures

FIGURE 3. Panaspis massaiensis comb. nov. holotype (MHNP 1904.306) from the Maasai plains Nairobi region, Kenya. Photos: Nicolas Vidal.

FIGURE 4. Life coloration of Panaspis massaiensis comb. nov. from Chyulu Hills. (A) Dorsal view (B) Ventral view. Photos: P.K. Malonza. (C) Comparative Panaspis massaiensis comb. nov. from Mt. Kasigau -Taita Hills. Photo: David Wojnowski.

Tables

TABLE 2: 16S rRNA percentage (%) pairwise distances between groups calculated using the K2P model. The number of base substitutions were obtained from averaging all sequence pairs between groups. Standard error estimate(s) shown above were obtained by a bootstrap procedure (1000 replicates). The Panaspis sp. lineages used here follow Medina et al. 2016.

	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17
1 _ P. sp. 2 Mozambique		1.15	1.11	1.49	1.05	1.25	1.24	1.07	1.20	1.15	0.98	1.31	1.29	1.56	1.09	1.14	1.06
2 _ P. sp. 4 Mozambique	6.03		0.60	1.30	0.87	1.07	1.06	1.01	1.11	0.96	0.63	1.16	0.96	1.32	0.87	0.93	1.01
3 _ P. sp. 3 Mozambique	5.30	1.91		1.36	0.85	1.11	1.08	0.95	1.00	1.04	0.50	1.19	0.94	1.35	0.95	1.01	1.05
4 _ P. sp. 1 Mozambique	9.22	7.35	7.72		1.31	1.32	1.16	1.55	1.59	1.48	1.33	1.23	1.53	0.84	1.43	1.48	1.28
5 _ P. wahlbergi	4.65	4.12	3.65	7.53		1.07	1.11	0.86	0.93	0.87	0.68	1.19	0.98	1.35	0.71	0.83	1.00
6 _P. sp. Ethiopia	6.99	5.41	5.65	7.36	5.54		1.09	1.16	1.22	1.20	1.14	1.20	1.23	1.27	1.14	1.19	1.10
7 _P. sp. Limpopo	6.98	5.89	5.88	5.78	6.12	5.77		1.30	1.39	1.32	1.05	0.44	1.23	1.27	1.26	1.32	1.06
8 _P. sp. 5 Mozambique	5.08	4.83	3.88	9.11	3.67	5.77	7.22		0.97	0.96	0.88	1.29	0.96	1.64	0.89	0.92	1.16
9 _P. sp. 2 Katanga	6.04	5.34	4.49	9.91	4.38	7.02	8.50	4.14		1.02	0.98	1.49	1.11	1.65	1.03	1.04	1.16
10 _P. sp. 2 Tanzania	5.91	4.24	4.46	8.97	3.90	6.38	7.84	4.01	4.48		0.92	1.37	1.07	1.51	0.42	0.25	1.18
11 _P. sp. Malawi	4.12	2.03	1.23	7.35	2.49	5.78	5.53	3.66	4.15	3.76		1.15	0.94	1.37	0.82	0.89	0.96
12 _ P. namibiana	7.72	6.38	6.37	6.39	6.61	6.51	1.12	6.98	9.02	8.10	6.02		1.25	1.35	1.31	1.36	1.17
13 _P. sp. 1 Katanga	7.42	4.49	4.37	9.76	5.09	6.88	7.36	4.73	5.73	5.55	4.27	7.62		1.57	1.04	1.03	1.21
14 _ P. maculicollis	10.10	7.34	7.46	3.18	8.08	7.10	7.11	10.12	10.40	9.21	7.59	7.62	10.25		1.48	1.51	1.41
15 _P. sp. 1 Tanzania	5.43	3.47	3.75	8.34	2.95	5.77	7.22	3.78	4.72	0.90	3.06	7.47	5.31	8.58		0.34	1.12
16 _ P. massaiensis comb. nov	5.85	3.94	4.17	8.91	3.58	6.32	7.78	3.69	4.66	0.39	3.47	8.03	5.25	9.15	0.61		1.18
17 _ P. tsavoensis sp. nov	4.82	4.85	4.95	7.38	5.27	5.79	6.02	5.82	5.84	5.93	4.37	6.52	6.94	8.62	5.32	5.87