Hyperolius marginatus Peters, 1854.

Hyperolius marginatus Peters, 1854. ZMB 4806, “Macanga”, Mozambique. Back yellow with dark red spots, lower femur pink. A dark lateral line runs from the top of the snout to the sacrum, with similar dorso-lateral bands starting behind each eye, with short bands running from the sacrum to the vent.

Rappia undulata Boulenger, 1901. MRAC and BMNH 1947.2.9.64, “Pweto et de Lofoi”, DRC. Synonymy by Laurent (1983). Brown above, with a symmetrical dark marking with a pale border. An irregular pale line on the side of the head and body forming an hourglass pattern on the back. Inside of limbs pink.

Rappia rhodoscelis Boulenger, 1901. MRAC and BMNH 1947.2.9.44–45, “Pweto”. It was previously placed as a synonym of H. marmoratus (see Frost 2021). The H. marmoratus clade does not occur in southern DRC. Greybrown above, with a pale lateral band from the shoulder. Very fine dark speckle on the limbs.

Hyperolius argentovittis Ahl, 1931. ZMB 85718, “Ujiji”, (Udjidji) Tanzania. Synonymy by Laurent (1961). Colour in alcohol: Above yellow-brown with three silver-white longitudinal bands. The lateral bands originate behind the eyes. Illustrated in Tillack et al. fig. 5 (2021).

Hyperolius udjidjiensis Ahl, 1931. ZMB 36101, “Udjidji”, Tanzania. Synonymy by Wieczorek et al. (2000). Colour in alcohol: Above pale grey with many small rounded or rectangular markings, often overlapping. Visible parts of limbs the same colour. The markings are a distinct pale brown.

Hyperolius graueri Ahl, 1931 ZMB 85758, “Westliches Russisi-Ufer und Nordwestufer des Tanganyika-see’s” (Rusizi River west bank and NW Lake Tanganyika, Tanzania). Considered a synonym of Hyperolius marmoratus argentovittis by Laurent (1952).

Hyperolius marmoratus pyrrhodictyon Laurent, 1965. MCZ A-46182 “Mazabuka, in the water-grass fringe of the Kafue River” Zambia. Previously placed as a full species H. pyrrhodictyon Laurent, 1965, it falls within the distribution of the H. marginatus clade, and there is no molecular evidence to maintain its species status. The back has no marbling or any spots, but there is a dark red network on the throat and belly.

Hyperolius melanoleucus Laurent, 1941. MRAC 35962–67 Lukafu, DRC. Synonymy by Wieczorek et al. (2000). Colour in alcohol: Back black or brown spotted, with a dark mark from between the eyes to the mid back, and two dark lateral markings, sometimes pale, or with a pattern in white consisting of a medio-dorsal stripe, sometimes interrupted, from snout tip to vent. Two lateral lines starting from the eyelid. Sometimes a dark back with a broad pale vertebral stripe, and an irregular pale lateral line.

Hyperolius melanoleucus kandoensis Laurent, 1943. MRAC 1394, “Kando (près de Tenke, Katanga Province)” DRC. Implied synonymy by Wieczorek et al. (2000). Hyperolius marmoratus does not occur in the DRC. Colouration very variable. Three irregular longitudinal white bands, confluent with other markings, producing a marbling effect.

Hyperolius lestagei Laurent, 1943. MRAC 39679, “Luapula, (près du Lac Bengwelo)” Luapula Province, DRC. Placed as a synonym of H. marmoratus by Schiøtz (1975). However, the clade identified as H. marmoratus does not occur in the DRC. Colour in alcohol: back and visible parts of the limbs uniform grey or blackish, without markings.

Hyperolius angolensis straeleni Laurent, 1943. IRSNB 1.040, “Bukama (Lualaba)”, DRC. Placed as a synonym of H. marginatus by Wieczorek et al. (2000). Two patterns were described. The first consists of a symmetrical, or nearly symmetrical broad vertebral stripe, that may be interrupted, with large irregular markings along the flanks. Tibia marbled. The second pattern consists of a red-brown background with pale vermiculation.

Hyperolius marmoratus rhodesianus Laurent, 1948. BMNH 1937.1.5.1, Matetsi, Zimbabwe. Previously regarded as a good species H. rhodesianus Laurent, 1948, it is placed as a synonym of H. marginatus as it occurs close to the localities of sequences (Fig. 6) forming the H. marginatus clade. Dark lateral area without a pale stripe, and greyish dots on the back, all weakly pigmented.

Hyperolius marmoratus aposematicus Laurent, 1951. MNHNP 33.295, “Lealui s/Zambèze, Rhodesie” (Zambia). The H. marmoratus clade does not extend this far west. The type locality is between sequenced material in the H. marginatus clade. Adult colouration variable, with three wide pale bands and two pale lumbar marks, or black spots, rounded or elongated, separated from each other.

Hyperolius decipiens and H. callichromus (only specimens from north-west Lake Tanganyika), were removed from the synonymy of H. marginatus which does not occur there, and placed in H. parallelus.

Hyperolius albofasciatus Hoffman, 1944 was removed from the synonymy of H. marginatus and placed as a synonym of H. marmoratus based on its type locality (Umbe, Malawi) within the distribution of the H. marmoratus clade. It was previously placed as a subspecies of H. marmoratus by Loveridge (1953).

Hyperolius brieni Laurent, 1943. Schiøtz (1975) placed it as a synonym of H. marginatus, but the type locality is close to sequenced material of the H. parallelus clade, with which it is regarded as a synonym.

Sequences were available from Angola, Zambia, Zimbabwe, Malawi and western Tanzania (Fig. 6). The minimum uncorrected p- distances from other clades were 2.4–5.3% (Table 2). The sequences were reported in GenBank as Hyperolius marginatus, H. viridiflavus aposematicus, H. v. marginatus, H. v. melanoleucus, H. v. pyrrhodictyon, H. v. rhodesianus, H. pyrrhodictyon, and H. parallelus alborufus. Although there are no sequences available from the type locality of Macanga in northern Mozambique, the clade occupies the area where Schitz (1999) mapped H. marginatus. The conservative approach of assigning this clade to H. marginatus thus seems prudent. The type localities of taxa considered junior synonyms of H. marginatus are listed in Table 3.

The most common pattern element in the descriptions of the types included in H. marginatus is a pale lateral band (58%), followed by a pale vertebral stripe (42%) and dorsal marbling (33%). The percentages are not additive, as the elements may be shared.

The advertisement call of a specimen from Lusaka, Zambia consists of a series of clicks (Schiøtz 1975, (no voucher) as H. paralellus melanoleucus) with an emphasised frequency of 2150 Hz. A call from Livingstone, Zambia is similar, with an emphasised frequency of 2000 Hz (Schiøtz 1975, (no voucher) as H. p. aposematicus). A call from Kachalola, Zambia has an emphasised frequency of 2100 Hz (Schiøtz 1975, (no voucher) as H. marginatus). The call is identical to those from Kitwe, Zambia (as H. melanoleucus), Solwezi, Zambia (as H. alborufus) and Nkotakota, Malawi (as H. marginatus) (Pickersgill 2007). Calls in Kasanka National Park (no vouchers) are brief (0.07 s) at an emphasised frequency of 2497 Hz.