Hyperolius marmoratus Rapp, 1842

Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, Zootaxa 5134 (3), pp. 301-354 : 325-328

Hyperolius marmoratus Rapp, 1842 .

Hyperolius marmoratus Rapp, 1842 . Types unknown. “Natal”, KwaZulu-Natal Province, South Africa. Locality obscure. Back dark brown with irregular pale grey markings. Lower limbs flecked.

Hyperolius verrucosus Smith, 1849 . Types unknown. “Kaffirland to the eastward of the Cape Colony” (see comment above), South Africa. Locality obscure. Synonymy by Hewitt (1937). Back greyish brown variegated with spots, stripes or broken bars of pale sienna-yellow or cream-yellow. Extremities pale buff-orange, thighs speckled with dull olive-green. Inner surfaces of thighs reddish-orange.

Hyperolius taeniatus Peters, 1854 . ZMB 4531, Boror, Mozambique. Synonymy by Boulenger (1882). Yellow lateral stripes join at the snout. Legs silver with dark flecks, with a dark lateral band from snout to leg.

Hyperolius sugillatus Cope, 1862 . ANSP 11319, Umvoti, South Africa. Synonymy by Loveridge (1941). Back straw colour, with a yellow band from snout to sacral region, which has a dark purple line forming a lower border to the yellow band on the snout. A purple spot on each eyelid.

Hyperolius citrinus Günther, 1864 . BMNH 1947.2.9.61 and 1947.2.9.63. “Zambesi Expedition” Luabo River, Zambezi Delta, Mozambique. Synonymy by Laurent (1952). Lemon-coloured above and below.

Hyperolius granulosus Peters, 1867 . Syntypes ZMB 4811 , ZMB 75652, “ Capanga am Flüsschen Mutizi östlich von Tette” Capanga, Mozambique. Synonymy by Laurent (1961). Back, sides and upper surfaces of limbs signal red with yellow vermiculation that shows occasional black pigmentation.

Hyperolius variegatus Peters, 1882 . ZMB 10249, 75602, “Cabaceia, Quelimane, Inhambane ”, Mozambique. Synonymy by Laurent (1952). ZMB 75603–75604 were listed as types, but these are Afrixalus sp. ( Tillack et al. 2021). Above dark brown with golden flecks and lines. Lips and ventral surfaces of limbs black with golden dots with red borders. Limbs blood-red, throat marbled in yellow.

Hyperolius marmoratus pondoensis FitzSimons, 1930 . TMP (now DNMNH) 13689. “Port St Johns, Pondoland Coast”, South Africa. Synonymy by Laurent (1952). Olive to chocolate-brown above, with irregular sized grass-green spots, darkly edged. Upper sides irregular and black edged. Ventral surfaces of limbs pink.

Hyperolius breviceps Ahl, 1931 . ZMB 86026, “Chemba”, Mozambique. Synonymy by Laurent (1952). Colour in alcohol: Back, sides and limbs chalk-white to pale grey-brown, with irregular small black or dark brown markings, some of which are rounded. An interrupted dark stripe on the side of the head and snout. Small dark dots on a white throat.

Hyperolius albifrons Ahl, 1931 . ZMB 36095. “Afrika (ohne genauern Fundort) [without precise locality]. Colour in alcohol: Back and sides yellowish-brown. Upper surface of snout silvery-white, extending back to the eyes, continuing as lateral stripes to the level of the shoulder. Limbs the same colour as the back. I follow Tillack et al. (2021) in retaining this taxon in H. marmoratus .

Hyperolius dermatus Ahl, 1931 . ZMB 85999, “Cabayra” (=Cabaceia?), Mozambique. The locality is obscure ( Tillack et al. 2021). Synonymy by Laurent (1985). Colour in alcohol: Above chalk white with a yellow-brown tint that is more pronounced behind the head. Upper lip pale yellow. Yellow lateral band between the limbs.

Hyperolius fuelleborni Ahl, 1931 . ZMB 71184–71186, “Neu-Helgoland” Papya Island, Malawi. Illustrated in Tillack et al. (2021, fig. 7). Other specimens included in the description by Ahl (1931) are placed as synonyms of H. parallelus because the H. marmoratus clade does not occur in the northern Lake Malawi area. Back pale grey with dark brown to black marbling. Sides of head without flecks.

Hyperolius swynnertoni FizSimons, 1941 . TMP 18911 and 18916, “Chirinda Forest”, Zimbabwe. Presently regarded as a good species. I follow the synonymy by Poynton & Broadley (1987). Dark vermiculation on a pale background, sometimes with pale stippling on a dark background.

Hyperolius albofasciatus Hoffman, 1944 . NMB A2622, “Umbe, Nyasaland ” ( Malawi). Implied synonymy by Poynton & Broadley (1987). Dark brown back with three white bands running the length of the body. The middle band starts between the nostrils and joins with the lateral bands just above the vent. Femur pink, tibia above brown with narrow white bands. Arms pale with scattered brown spots.

Hyperolius marmoratus broadleyi Poynton, 1963 . NMZB 5370, “Fern Valley, Umtali” Zimbabwe. Back black with a regular white band from the snout tip to above the vent, and a lateral band from behind the eye to the groin. A thin red line present in the middle of each white band. Limbs with black and red markings.

Hyperolius fumosus Pickersgill, 2007 . LIV 1998.46.67 “Beira, Mozambique ”. It is assigned as a synonym as the colour pattern is part of the known variation. The phase F is silver-white with black speckling with the typical red or pink on the hidden parts of the limbs. The advertisement call (no voucher) has a duration of 64–90 ms, with an emphasised frequency of 2.6–3.8 kHz ( Pickersgill 2007) which matches those along the South African east coast. Only a single clade, H. marmoratus , occurs along the coast including Beira.

Nearly 40 taxa were previously placed in the synonomy of H. marmoratus (Frost 2021) . The type locality is Kwa- Zulu-Natal, which agrees with the clade here assigned to H. marmoratus . Sequences were available from South Africa, Zimbabwe, eSwatini, and Mozambique. The minimum uncorrected p- distances from other clades were 4.0–6.1% ( Table 2 ). Names applied to the sequences in this clade include H. viridiflavus , H. v. broadleyi , H. v. swynnertoni , H. v. marmoratus , H. v. taeniatus , H. v. verrucosus , H. v. albofasciatus , H. marmoratus , H. m. marmoratus , H. m. taeniatus , and H. swynnertoni . This clade occurs from the south coast of South Africa, to eSwatini, Zimbabwe, Malawi and Mozambique ( Fig. 5 ). See Table 3 for synonym localities.

The following extralimital taxa were removed from the synonymy of H. marmoratus and placed as synonyms of H. parallelus : Hyperolius vermiculatus Peters, 1882 (from Angola); Hyperolius plicifera Bocage, 1893 (after Perret, 1976); Rappia rhodoscelis Boulenger, 1901 (from the DRC); Hyperolius nossibeensis Ahl, 1931 (from Angola); Hyperolius decoratus Ahl, 1931 (from Angola); Hyperolius microstictus Ahl, 1931 (from Angola); Hyperolius marungaensis Ahl, 1931 (from Angola); Hyperolius latirostris Laurent, 1943 (from the DRC); Hyperolius melanoleucus kandoensis Laurent, 1943 (from the DRC); Hyperolius fuliginosus Laurent, 1943 (from the DRC); Hyperolius marmoratus epheboides Laurent, 1957 (from the DRC); Hyperolius marmoratus alborufus Laurent, 1964 (from Angola); Hyperolius fuelleborni Ahl, 1931 types from Songwe, Miramba, Rungwe, and Lumbira in Tanzania (types of Hyperolius fuelleborni Ahl, 1931 from Papya Island, Malawi are retained in H. marmoratus ); Hyperolius nyassae Ahl, 1931 (from Tanzania).

The following taxa were removed from the synonymy of H. marmoratus and placed as synonyms of H. glandicolor : Hyperolius asper Ahl, 1931 (after Wieczorek et al. 2000); Hyperolius marmoratus campylogrammus Laurent, 1951 , which occurs north of the range of the H. marmoratus clade.

Hyperolius lestagei Laurent, 1943 (from the DRC) was removed from the synonymy of H. marmoratus , which does not occur in the DRC, and placed as a synonym of H. marginatus .

Hyperolius marmoratus mafianus Laurent, 1961 was removed from the synonymy of H. marmoratus and placed as a synonym of H. mariae , which occurs north of the H. marmoratus clade.

Hyperolius marmoratus Rapp, 1842 , was described from KwaZulu-Natal Province, South Africa. Only one clade extends along the east coast of South Africa, and this clade is assigned to Hyperolius marmoratus ( Fig. 5 ). The localities of types now regarded as junior synonyms of H. marmoratus are listed in Table 3 . The range of this species has expanded southwards to Cape Town and its environs due to multiple introductions ( Bishop 2004, Tolley et al. 2008).

The most common pattern elements in the descriptions of the types included in H. marmoratus are dorsal marbling (25%), and a pale lateral band (25%), followed by small black speckle or spots (19%), and a pale vertebral band (19%). The percentages are not additive, as patterns may be shared. Pickersgill showed how the colour patterns intergrade along the South African coast (2007: 384–385).

An advertisement call from Chimanimani (no voucher) is a brief whistle (0.06 s) with an emphasised frequency of 2608 Hz; a call from Gqeberha (no voucher) is a whistle (0.1 s) showing a slight rise in frequency, at an emphasised frequency of 2608 Hz. Calls from Howick (no vouchers) are long, rising whistles, 0.13 s, at an emphasised frequency of 2583–2928 Hz.

Figures

FIGURE 5. Localities of sequences (black circles), type locality (star) and location of junior synonyms of Hyperolius marmoratus. Locality codes are explained in Table 3.