A new sand frog from Namaqualand, South Africa (Pyxicephalidae: Tomopterna) Author Wilson, Lyle Author Channing, Alan text Zootaxa 2019 2019-05-23 4609 2 225 246 journal article 26713 10.11646/zootaxa.4609.2.2 ad74907d-57fc-4ce0-9aab-dc7b6adaf63d 1175-5326 3187583 BE4E293B-FFF3-445B-A5BB-3625BF8E01ED Tomopterna branchi sp. nov. Namaqua Sand Frog Figs. 4 , 5 . Holotype . A male, ZMB 87233, field number AC3383 ( Fig. 4 ), collected in the dry bed of the Buffels River where it is crossed by the N7 highway, Namaqualand , South Africa , 8 May 2015 , by N. Telford ( WGS84 : 29.67791°S , 17.05214°E ). Paratypes . ZMB 87234, a female from Kamieskroon, and ZMB 87235 a male from Springbok . Diagnosis. The new species has been confused with Tomopterna delalandii , and is similar to all other species of sand frogs in morphology and burrowing behaviour. It has an advertisement call of a rapidly-repeated series of high-pitched notes, the most common type of call in sand frogs. We assign it to the genus Tomopterna based on the presence of teeth on the maxilla, the inner metatarsal tubercle strongly flanged, the outer metatarsals bound into the sole, and the presence of vomerine teeth, all characters that distinguish the genus Tomopterna ( Poynton 1964 ) . The following comparisons are based largely on data in Wasonga & Channing (2013) . The new species has single subarticular tubercles, distinguishing it from those with divided tubercles: T. gallmanni , T. krugerensis and T. wambensis . It has more than three phalanges of the fourth toe free of web, distinguishing it from those with more webbing: T. elegans , T. gallmanni , T. luganga , T, marmorata , T. tandyi and T. wambensis . There is a row of isolated glands below the tympanum, distinguishing it from those species with a distinct continuous or interrupted ridge: T. cryptotis , T. damarensis , T. elegans , T. gallmanni , T. kachowskii , T. krugerensis , T. luganga , T. natalensis , T. tandyi , T. tuberculosa and T. wambensis . The outer metatarsal tubercle is absent or present as a small white bump, which distinguishes it from those species with prominent outer metatarsal tubercles: T. delalandii , T. elegans , T. kachowskii , and T. tandyi . There is a transverse dark interocular bar present, which distinguishes it from those species without this pattern element: T. damarensis , T. luganga and T. natalensis . This bar is variable in T. elegans , T. kachowskii , and T. tandyi . The back is smooth or has small warts, distinguishing it from T. gallmanni and T. tuberculosa , which have prominent dorsal tubercles. FIGURE 4. Photo of the live holotype (ZMB 87233) of Tomopterna branchi , from the Buffels River, Namaqualand, South Africa. Photo: A. Channing. The advertisement call consists of a rapidly-repeated series of high-pitched notes, distinguishing it from the knocking call of T. krugerensis and T. gallmanni . The first two harmonics are emphasised (933–995 Hz; 1866–1990 Hz) at a note repetition rate of 3 s-1 . Tomopterna natalensis also has two emphasised harmonics, but at higher frequencies ( 1272–1488 Hz; 2544–2976 Hz). It is further distinguished from those species with only a single harmonic emphasised, at a higher frequency ( T. wambensis , T. tandyi , T. damarensis , T. sp. 3 and T. cryptotis ), and T. luganga and T. marmorata with a single emphasised harmonic at a lower frequency. The call of Tomopterna delalandii has the second harmonic emphasised, at a similar frequency to T. branchi , but the note repetition rate is approximately twice that of T. branchi . The call of T. elegans is unknown. The new species may occur with T. delalandii and T. tandyi . It can be distinguished from T. delalandii which has a distinct ridge below the tympanum (isolated glands), a prominent outer metatarsal tubercle (weak or absent), and an advertisement call with a note repetition rate of about 6 s-1 (about 3 s-1 ). It differs from T. tandyi which has more than three phalanges of the fourth toe free of webbing (three free), a ridge below the tympanum (isolated glands), a prominent outer metatarsal tubercle (weak or absent), and an advertisement call with a single harmonic emphasised at 1.9–2.1 kHz (two harmonics emphasised). The uncorrected p -distances (as percentages) of T. branchi compared to all other species of sand frogs based on 16S rRNA vary from 2.3–6.0%. The uncorrected p -distances between T. branchi and T. delalandii are 2.2–3.4% for 16S ( Table 2 ) and 2.5–4.3% for tyrosinase exon 1. FIGURE 5 . Dorsal, ventral and lateral views of the holotype of Tomopterna branchi ZMB 87233. TABLE 2 . List of sequenced material and sources. Rows in bold are multiple sequences from the same voucher.

Species (GenBank identification if different)	Voucher	Locality	Co-ordinates	16S Accession number	tyr Accession number	Reference
T. branchi sp. nov .	ZMB 87233	Buffels River, South Africa	29.6779°S, 17.0521°E	MK289551	MK300847	This study
T. branchi sp. nov .	AC3246	Buffels River, South Africa	29.6779°S, 17.0521°E	MG450788	This study
T. branchi sp. nov .	AC3247	Buffels River, South Africa	29.6779°S, 17.0521°E	MG450789	MK300848	This study
T. branchi sp. nov .	AC3248	Buffels River, South Africa	29.6779°S, 17.0521°E	MG450790	This study
T. branchi sp. nov .	AC3250	Buffels River, South Africa	29.6779°S, 17.0521°E	MG450791	This study
T. branchi sp. nov .	AC3251	Buffels River, South Africa	29.6779°S, 17.0521°E	MG450792	This study
T. cryptotis	No voucher	Bredell, South Africa	26.0871°S, 28.3039°E	AF215506	Vences et al. 2000
T. tandyi	AC3063	Khorab, Namiba	19.6631°S, 17.3283°E	MK335431	This study
T. cryptotis	AC1154	Bloemfontein, South Africa	30.3300°S, 26.8003°E	AY255099	Dawood & Uqubay 2004
T.cryptotis	AC1100	Bubye, Zimbabwe	21.7081°S, 30.4986°E	AY255090	Dawood & Uqubay 2004
T. damarensis	AC2653	Naukluft, Namibia	24.2661°S, 16.2411°E	MK335418	This study
T. damarensis	AC3114	Farm Oas, Namibia	27.4900°S, 19.2288°E	MK335419	This study
T. damarensis	AC3130	Weavers Rock, Namibia	20.6983°S, 16.8417°E	MK335420	This study
T. damarensis (“ T . sp.)	CAS 254855	Pediva, Angola	16.2833°S, 12.5666°E	KU662310	Ceríaco et al . 2016
T. damarensis	MCZ A-148577	Epupa Falls, Namibia	17.0000°S, 13.2333°E	KX869908	Heinicke et al. 2017
T. damarensis	MCZ A-149503	Farm Omandumba, Namibia	21 29 51.1 / 15 37 47.9	KX869909	Heinicke et al. 2017
T. damarensis	TM83916	Khorixas, Namibia	20.3666°S, 14.9666°E	AY255091	Dawood & Uqubay 2004
T. damarensis ( “ T. cf. tandyi ” )	ZFMK 66403	Khorixas, Namibia	20.3666°S, 14.9666°E	AF215419	Vences et al. 2000
T. damarensis	ZFMK 66403	Khorixas, Namibia	20.3666°S, 14.9666°E	DQ019610	Van der Meijden et al. 2005
T. damarensis	ZFMK 66403	“Namibia”	20.3666°S, 14.9666°E	DQ347301	Bossuyt et al. 2006
T. delalandii	AC1134	Stellenbosch, South Africa	33.9823°S, 18.8282°E	AY454372	Lehtinen et al . 2004
T. delalandii	AC1144	Stellenbosch, South Africa	33.9823°S, 18.8282°E	AY255085	Dawood & Uqubay 2004
T. delalandii	AC2832	Riet River, South Africa	32.6746°S, 21.4648°E	MK289552	This study
T. delalandii	AC3222	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289553	This study
T. delalandii	AC3223	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289554	MK300849	This study
T. delalandii	AC3224	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289555	MK300850	This study
T. delalandii	AC3225	Rondevlei, South Africa	34.0584°S, 18.5402°E	MK289556	MK300851	This study
T. delalandii	AC3226	Bellville, South Africa	33.9316°S, 18.6235°E	MK289557	MK300852	This study
T. delalandii	AC3227	Bellville, South Africa	33.9316°S, 18.6235°E	MK289558	MK300853	This study

......continued on the next page ......continued on the next page Description of holotype . A male ( Figs. 4 , 5 ) SVL 42.4; the body is robust; head short (HL/SVL 0.37, HW/SVL 0.44), not wider than trunk, not longer than wide (HL/HW 0.83); snout short (SL/HL 0.44), rounded in dorsal view, truncated in profile, slightly projecting beyond lower jaw, narrow (SL/IND 1.97); canthus rostralis rounded; loreal region slightly concave; nostrils situated on slight projections, closer to the eye than snout tip (ENL/SNL 0.86); eyes directed anterolaterally, slightly protruding, relatively small (ED/HL 0.40); eye diameter less than snout length (ED/SL 0.91); interorbital distance is less than upper eyelid (IOS/ELD 0.46), and nearly equal to internarial distance (IOS/INS 0.94); Internarial distance just more than half eye diameter (IOS/ED 0.56); vomerine teeth inconspicuous; tympanum visible, smaller than eye diameter (TD/ED 0.46); a row of isolated glands below tympanum; upper jaw without dentition; choanae small, round, vomerine teeth absent; tongue notched posteriorly, 7.0 at widest part; median lingual processes present; vocal sac single, darkly pigmented anteriorly; dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs, gular and abdomen smooth. Arms slender; hand moderately large (HND/SVL 0.26); tips of fingers not enlarged into discs; relative length of fingers: IV <II <I <III; subarticular tubercles single and distinct, with one on fingers I, II and IV, and two on finger III; fingers without webbing; thenar tubercle distinct; metacarpals with supernumerary tubercles; pale nuptial pads present on upper surface of fingers I and II. TABLE 2. (Continued)

Species (GenBank identification if different)	Voucher	Locality	Co-ordinates	16S Accession number	tyr Accession number	Reference
T. delalandii	AC3228	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289559	MK300854	This study
T. delalandii	AC3229	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289560	MK300855	This study
T. delalandii	AC3230	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289561	MK300856	This study
T. delalandii	AC3231	Stellenbosch, South Africa	33.9823°S, 18.8202°E	MK289562	MK300857	This study
T. delalandii	AC3232	Noordhoek, South Africa	32.7181°S, 19.0668°E	MK289563	MK300858	This study
T. delalandii	AC3233	Noordhoek, South Africa	34.0974°S, 18.3960°E	MK289564	MK300859	This study
T. delalandii	AC3234	Noordhoek, South Africa	34.0969°S, 18.3734°E	MK289565	This study
T. delalandii	AC3236	Port Elizabeth, South Africa	34.0016°S, 25.5277°E	MG450762	MK300860	This study
T. delalandii	AC3237	Port Elizabeth, South Africa	34.0016°S, 25.5277°E	MG450763	This study
T. delalandii	AC3238	Port Elizabeth, South Africa	34.0016°S, 25.5277°E	MG450764	MK300861	This study
T. delalandii	AC3239	Cape St Francis, South Africa	34.2077°S, 24.8300°E	MG450765	MK300862	This study
T. delalandii	AC3240	Cape St Francis, South Africa	34.2077°S, 24.8300°E	MG450766	MK300863	This study
T. delalandii	AC942	Stellenbosch, South Africa	33.9823°S, 18.8282°E	AY255086	Dawood & Uqubay 2004
T. delalandii	AC942	Stellenbosch, South Africa	33.9823°S, 18.8282°E	DQ283403	Frost et al . 2006
T. delalandii	KTH076	Anysberg Nature Reserve, South Africa	33.5192°S, 20.6492°E	EF136565	Tolley et al . 2010
T. delalandii	No voucher	Stellenbosch, South Africa	33.9823°S, 18.8282°E	HQ014421	Van der Meijden et al . 2011
T. delalandii	SAIAB 185255-1	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450784	This study
T. delalandii	SAIAB 185255-2	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450785	This study
T. delalandii	SAIAB 185255-3	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450786	This study
T. delalandii	SAIAB 185255-4	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450787	This study
T. delalandii	SAIAB 186279-1	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450771	MK300865	This study
T. delalandii	SAIAB 186279-10	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450780	This study
T. delalandii	SAIAB 186279-11	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450781	This study
T. delalandii	SAIAB 186279-12	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450782	This study
T. delalandii	SAIAB 186279-13	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450783	This study
T. delalandii	SAIAB 186279-2	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450772	This study
T. delalandii	SAIAB 186279-3	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450773	MK300866	This study
T. delalandii	SAIAB 186279-4	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450774	This study

......continued on the next page TABLE 2. (Continued)

Species (GenBank identification if different)	Voucher	Locality	Co-ordinates	16S Accession number	tyr Accession number	Reference
T. delalandii	SAIAB 186279-5	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450775	This study
T. delalandii	SAIAB 186279-6	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450776	This study
T. delalandii	SAIAB 186279-7	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450777	This study
T. delalandii	SAIAB 186279-8	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450778	This study
T. delalandii	SAIAB 186279-9	Port Alfred, South Africa	33.6131°S, 26.8881°E	MG450779	This study
T. delalandii	WC-0102	Port Elizabeth, South Africa	34.2077°S, 24.8300°E	MG450767	This study
T. delalandii	WC-0105	Cape St Francis, South Africa	34.2077°S, 24.8300°E	MG450768	This study
T. delalandii	WC-10-125	Cape St Francis, South Africa	34.2077°S, 24.8300°E	MG450769	This study
T. delalandii	WC-10-150	Cape St Francis, South Africa	34.2077°S, 24.8300°E	MG450770	MK300864	This study
T. elegans	MVZ:herp:242739	Buq, Somalia	10.6227°N, 47.1779°E	HQ700692	Zimkus & Larson 2011
T. elegans (“ T. cryptotis ”)	MVZ:herp:242739	Buq, Somalia	10.6227°N, 47.1779°E	JX564898	Zhang et al. 2013
T. gallmanni	NMK A/5145/1	Mogwooni Ranch, Kenya	0.2416°N, 36.9666°E	JX088641	Wasonga & Channing 2013
T. gallmanni	NMK A/5045	Lewa Wildlife Conservancy	0.1527°N, 37.4377°E	JX088642	Wasonga & Channing 2013
T. gallmanni	NMK A/5159	Road from Baragoi to South Horr, Kenya	1.3504°N, 36.8850°E	JX088645	Wasonga & Channing 2013
T. gallmanni	NMK A/5039/2	Laikipia Nature Conservancy, Kenya	0.5666°N, 36.4000°E	JX088643	Wasonga & Channing 2013
T. gallmanni	NMK A/5151/1	Baawa Village, Kenya	0.0458°N, 36.8152°E	JX088644	Wasonga & Channing 2013
T. kachowskii	MVZ:Herp:241321	Borama, Somalia	9.9705°N, 43.1460°E	HQ700690	Zimkus & Larson 2011
T. kachowskii	MVZ:Herp:241323	Borama, Somalia	9.9705°N, 43.1460°E	HQ700691	Zimkus & Larson 2011
T. kachowskii	AAU<ETH>: A:2009-310	Addis Ababa, Ethiopia	9.0243°S, 38.7492°E	HQ700687	Zimkus & Larson 2011
T. kachowskii	AAU<ETH>: A:2008-317	Addis Ababa, Ethiopia	9.0243°S, 38.7492°E	HQ700689	Zimkus & Larson 2011
T. kachowskii	AAU<ETH>: A:2008-321	Addis Ababa, Ethiopia	9.0243°S, 38.7492°E	HQ700688	Zimkus & Larson 2011
T. kachowskii	USNM:589360	Day village, Djibouti	11.7800°N, 42.6300°E	MG700266	Gotte et al. in press
T. kachowskii	USNM:589356	Day village, Djibouti	11.7800°N, 42.6300°E	MG700268	Gotte et al. in press
T. kachowskii	USNM:589358	Day village, Djibouti	11.7800°N, 42.6300°E	MG700264	Gotte et al. in press
T. kachowskii	USNM:589359	Day village, Djibouti	11.7800°N, 42.6300°E	MG700265	Gotte et al. in press
T. kachowskii	USNM:589357	Day village, Djibouti	11.7800°N, 42.6300°E	MG700267	Gotte et al. in press

TABLE 2. (Continued)

Species (GenBank identification if different)	Voucher	Locality	Co-ordinates	16S Accession number	tyr Accession number	Reference
T. krugerensis	AC 1506	Tembe Elephant Park, South Africa	27.0472°S, 32.4147°E	AY255094	Dawood & Uqubay 2004
T. krugerensis	AC1508	Tembe Elephant Park, South Africa	27.0472°S, 32.4147°E	AY255098	Dawood & Uqubay 2004
T. luganga	TM85107	Dodoma, Tanzania	6.1833°S, 37.6666°E	AY547276	Channing et al. 2004
T. luganga	AD310	Iringa, Tanzania	7.7666°S, 35.7000°E	DQ017056	Haas et al. 2006
T. luganga	AD309	Iringa, Tanzania	7.7666°S, 35.7000°E	AY751302	Haas et al. 2006
T. luganga	MCZ:A:HM1331	Iringa, Tanzania	7.7666°S, 35.7000°E	HQ700695	Zimkus & Larson 2011
T. luganga	TM 85104	Kigwembimbe, Tanzania	7.8195°S, 35.7810°E	AY547275
T. marmorata	AC1534	Livingstone, Zambia	17.8419°S, 25.8542°E	AY255084	Dawood & Uqubay 2004
T. marmorata	AC1387	Legalamectse NR, South Africa	24.4411°S, 30.2947°E	AF371233	Dawood & Uqubay 2004
T. milletihorsini	ZFMK 76391	Chelcha, Mauritania	16.6614°N, 9.6149°E	AY014383	Kosuch et al. 2001
T. milletihorsini	ZFMK 76391	Ayoun el-Atrous, Mauritania	16.6614°N, 9.6149°E	AY255101	Dawood & Uqubay 2004
T. natalensis	No voucher	Ithala, South Africa	27.5380°S, 31.2885°E	AF215508	Vences, 2000
T. natalensis	AC1391	Mafefe Road, South Africa	24.1833°S, 30.0666°E	AY255088	Dawood & Uqubay 2004
T. natalensis	AC1451	Andries Vosloo Kudu Reserve, South Africa	33.0376°S, 26.7880°E	AY255089	Dawood & Uqubay 2004
T. natalensis	ES 255	Kyalami, South Africa	26.0178°S, 28.0687°E	AY547277	Channing et al. 2004
T. natalensis	TM84342	Tygerskloof Plantation, South Africa	27.8757°S, 31.3180°E	AY205286	Dawood & Stam 2006
T. natalensis	ZFMK 68815	Port St Johns, South Africa	31.5101·S, 29.4419°E	AF215509	Vences, 2000
T. natalensis	ZFMK 68815	Port St Johns, South Africa	31.5101·S, 29.4419°E	DQ347300	Bossuyt et al. 2006
T. tandyi	AC1556	Grootfontein, Namibia	19.5666°S, 18.1166°E	AF371234	Dawood et al . (2002)
T. tandyi	AC1567	Hardap, Namibia	24.5500°S, 17.9333°E	AF436073	Dawood & Uqubay 2004
T. tandyi	AC2006	Adelaide, South Africa	32.7195°S, 26.2948°E	AF436071	Dawood et al . (2002)
T. tandyi	AC3410	Okonjima, Namibia	20.8628°S, 16.6410°E	MK335421	This study
T. tandyi	AC3412	Queenstown, South Africa	31.8975°S, 26.8753°E	MK335422	This study
T. tandyi	AD14	Blouberg NR, South Africa	23.0500°S, 29.1333°E	AF436072	Dawood et al . (2002)
T. tandyi	AD231	Blouberg NR, South Africa	23.0307°S, 29.1253°E	AY838891	Dawood & Stam 2006
T. tandyi (“ T. marmorata ”)	No voucher	Mt Meru, Tanzania (“South Africa”)	3.4411°S, 36.6323°E	AF215507	Vences et al. 2000

......continued on the next page TABLE 2. (Continued)

Species (GenBank identification if different)	Voucher	Locality	Co-ordinates	16S Accession number	tyr Accession number	Reference
T. tuberculosa	AC1670	Ongongo , Namibia	19.1401°S , 13.8178°E	AY255087	Dawood & Uqubay 2004
T. tuberculosa	AC1672	Ongongo , Namibia	19.1401°S , 13.8178°E	AY255100	Dawood & Uqubay 2004
T. tuberculosa	AC3106	Zootecnica , Humpata , Angola	14.9162°S , 13.2562°E	MK 335423	This study
T. tuberculosa	AC3064	Zootecnica , Humpata , Angola	14.9816°S , 13.4335°E	MK 335424	This study
T. tuberculosa	AC3075	Humpata , Angola	14.9658°S , 13.3445°E	MK 335425	This study
T. wambensis	NMK A/5057/1	Ikave , Kenya	1.5888°S , 37.9000°E	JX088646	Wasonga & Channing 2013
T. wambensis	NMK A/5149/1	Mpala Research Center , Kenya	0.3533°N , 36.8722°E	JX088647	Wasonga & Channing 2013
T. wambensis	NMK A/5149/2	Mpala Research Center , Kenya	0.3533°N , 36.8722°E	JX088648	Wasonga & Channing 2013
T. wambensis	NMK A/4323	Bura Hasuma River , Kenya	3.8000°N 36.2833°E	JX088656	Wasonga & Channing 2013
T. wambensis	MCNPV CA253	Lugga Daudi River , Kenya	3.8166°N , 36.3500°E	JX088655	Wasonga & Channing 2013
T. wambensis	MCNPV CA250	Illeret , Kenya	4.3000°N , 36.2166°E	JX088653	Wasonga & Channing 2013
T. wambensis	MCNPV CA337	Kasa Well , Kenya	3.6000°N , 36.3000°E	JX088654	Wasonga & Channing 2013
T. wambensis	MCNPV CA334	17 km S Allia Bay , Kenya	3.7000°N , 36.3333°E	JX088650	Wasonga & Channing 2013
T. wambensis	NMK A/5152/1	Sordor Pan , Kenya	0.9833°N , 37.3333°E	JX088651	Wasonga & Channing 2013
T. wambensis	NMK A/5149/3	Mpala Research Center , Kenya	0.3533°N , 36.8722°E	JX088649	Wasonga & Channing 2013
T. wambensis	NMK A/5149/2	Mpala Research Center , Kenya	0.3533°N , 36.8722°E	JX088648	Wasonga & Channing 2013
T. wambensis	NMK A/5145/1	Kitobi Forest , Kenya	3 26 29 S , 37 13 E	JX088652	Wasonga & Channing 2013
T. wambensis	AC2062	Arusha , Tanzania	3.3563°S , 36.7081°E	AY547278	Wasonga & Channing 2013
T. wambensis	AC2159	Usangu Flats , Tanzania	8.5801°S , 34.2264°E	MK 335426	This study
T. wambensis (“ T. sp.”)	MCZ:A:HM1329	Iringa , Tanzania	7.7786°S , 35.6861°E	HQ700694	Zimkus & Larson 2011
T. wambensis (“ T. sp.”)	MCZ:A:HM1257	Iringa , Tanzania	7.7786°S , 35.6861°E	HQ700693	Zimkus & Larson 2011
T. sp. 1 “Beira”	AC1115	Beira, Mozambique	19.8123°S , 34.8459°E	AY255092	Dawood & Uqubay 2004
T. sp. 1 “Beira”	AC1116	Beira, Mozambique	19.8123°S , 34.8459°E	AY255093	Dawood & Uqubay 2004
T. sp. 2 “Mutinondo”	AC3376	Mutinondo, Zambia	12.4560°S , 31.2950°E	MK335427	This study
T. sp. 2 “Mutinondo”	AC3377	Mutinondo, Zambia	12.4560°S , 31.2950°E	MK335428	This study
T. sp. 2 “Mutinondo” (“ T. tuberculosa ”)	RdS 880	Tatanda, Tanzania	8.5166°S , 31.5000°E	KC179967	De Sa et al . 2012
T. sp. 3 “Shankara”	AC1549	Shankara, Namibia	17.9650°S , 20.5025°E	AY255095	Dawood & Uqubay 2004
T. sp. 3 “Shankara”	AC2649	Mbambi, Namibia	17.9833°S , 20.9734°E	MK335430	This study
T. sp. 3 “Shankara”	ZMB 83072	Ondjiva, Angola	17.0678°S , 15.7084°E	MK335429	This study

Hind limbs stout, tarsal tubercle absent, represented by a small raised area; tibia short (TL/SVL 0.40); heels not reaching each other when knees are flexed and thighs are held at right angle to body; foot shorter than tibia (TL/ FOT 0.84); relative length of toes: I<II<V<III<IV; toes without expanded discs; subarticular tubercles: one on toe I and II, two on toe III, three on toe IV and two on toe V; pedal webbing formula I 1– 2 II 1– 2 III 1– 3 IV 3.5– 2 V ; thin margin of webbing extending to tips; inner metatarsal tubercle prominent and shovel-shaped, continuing as a tarsal ridge, smaller than eye diameter (IMT/ED 0.63); outer metatarsal tubercle present as a small white bump. Measurements of the holotype and paratypes are presented in Table 3 . TABLE 3. Measurements of the types of Tomopterna branchi . For definitions of measurements see text.

Voucher	Status	Sex	SVL	HW	HL	IOS	DAE	IND	END SL
ZMB87233	Holotype	♂	42.4	18.8	15.6	3.2	7.3	3.5	3.2	6.9
ZMB87234	Paratype	♂	43.1	18.0	15.2	3.1	7.4	3.6	2.7	6.1
ZMB87335	Paratype	♀	32.6	18.2	15.0	3.2	7.7	3.8	3.0	6.5
continued.
Voucher	Status	Sex	TYM	ED	ETD	EL	HND	FOT	TIB	IMT
ZMB87233	Holotype	♂	2.9	6.3	1.6	7.0	11.1	20.1	16.8	4.0
ZMB87234	Paratype	♂	2.8	5.9	1.2	7.1	10.4	19.0	15.7	4.3
ZMB87335	Paratype	♀	2.6	6.2	2.8	6.9	11.2	22.3	18.2	4.2

Colour in life . The dorsal pattern of the holotype consists of green patches with black warts on a reddish-brown background ( Fig. 4 ). The transverse bar between the eyes is green with black flecks. The iris is pale golden with dark veins. Small pale tubercles are densely scattered over the back and sides. The flanks have irregular brown and white marbling. The upper surfaces of the limbs are brown, with green or darker brown transverse bars. The venter is white with a black throat. Colour in preservative . The pattern is clearly visible ( Fig. 5 ) as a darker brown on a paler brown background. Paratype variation. The male from Nababeep ZMB 87234 ( SVL 43.1 ) and the female from Kamieskroon ZMB 87235 ( SVL 43.6 ) have similar morphology to the holotype, and practically identical body proportions. A male from Springbok (no voucher) had pale dorsolateral stripes and a thin vertebral stripe. In life, it had a greybrown background with black and brown warts, and a pale occipital patch. Advertisement call. Calls were recorded at the town dam in Springbok, 28 January 1985 , air temperature 22°C. No voucher was collected. The call consists of a number of rapidly repeated notes, which have the first two harmonics emphasised 933–997 Hz and 1866–1990 Hz. The note repetition rate is 3 s-1 . In contrast, the call of T. delalandii from Elands Bay (air temp 14° C) shows a much faster note repetition rate, and has only one frequency, the first harmonic, emphasised ( Fig. 6 ). Eggs and tadpoles. Eggs are unknown. Tadpoles were found in shallow pools in the riverbed 23 October 2012 and in a small cup-sized depression against a rock in the otherwise dry sandy bed of the Buffels River, 7 May 2015 . The tadpoles are practically identical in proportions and pigmentation to those of Tomopterna cryptotis illustrated by Channing et al. (2012) . A brief description of AC3247 from the Buffels River is given here. Gosner stage 36, TL 31.8; oral disc 0.8 width of head at level of disc; a single row of marginal papillae present; anterior and posterior jaw sheaths 80% pigmented; LTRF 4(2–4)/3(1); nostrils elliptical; internarial ratio NW/ID 0.25; orbitonasal line not visible; pineal spot a small white dot; extra-ocular proportion (HWE-EO)/EO 0.64; spiracle situated 0.6 along body; opening constricted, visible laterally; vent ratio TUB/VW 2.8; height of caudal muscles at base of tail/height of trunk 0.66; small brown spots around eyes to nostril and snout tip, then over the dorsal surface of the body where it is most dense; sides and top of tail muscle lightly marbled; ventral fin unpigmented, dorsal fin transparent with some fine black lines and blotches. Distribution and habitat. The species is presently known from the Buffels River drainage of Namaqualand ( Fig. 7 ), but may be more widely distributed once further investigations have been undertaken. This area includes the Namaqualand Hardeveld vegetation units of the Succulent Karoo Biome ( Mucina et al . 2006 ). A photo of the type locality is shown in Fig. 8 . Tadpoles occupy shallow pools together with tadpoles of Amietia poyntoni . Etymology. The specific epithet refers to Dr W.R (Bill) Branch ( 1946–2018 ), who passed away while this paper was being prepared. We suggest that the English common name, Namaqua Sand Frog, be used.