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The geographically widespread species Afrixalus laevis (Anura: Hyperoliidae) currently has a disjunct distribution in western Central Africa (Cameroon, Equatorial Guinea, Gabon, and possibly adjacent countries) and the area in and near the Albertine Rift in eastern Democratic Republic of the Congo and neighboring countries. At least two herpetologists have previously suggested that these disjunct populations represent distinct species, and herein, we utilize an integrative taxonomic approach with molecular and morphological data to reconcile the taxonomy of these spiny reed frogs. We sequenced 1554 base pairs of the 16S and RAG1 genes from 34 samples of A. laevis and one sample of A. orophilus (sympatric with eastern populations of A. laevis), and combined these data with previously sequenced GenBank Afrixalus samples via the bioinformatics toolkit SuperCRUNCH. Phylogenetic trees, dated phylogenetic analyses, and species-delimitation analyses were generated with RAxML, BEAST, and BPP, respectively. Eleven mensural characters were taken from multiple specimens of A. laevis and A. orophilus, and compared with paired t-tests and analyses of covariance. These combined results suggested populations of A. laevis in western Central Africa (Cameroon and Bioko Island, Equatorial Guinea) represent one species, whereas populations from the Albertine Rift and nearby forests represent two undescribed taxa that are sister to A. dorsimaculatus. The two new species (A. lacustris sp. nov. and A. phantasma sp. nov.) are distinguished by our phylogenetic and species-delimitation analyses, significant differences in several mensural characters, qualitative morphological differences, and by their non-overlapping elevational distribution.

African reed frogs are widespread and highly polymorphic for colour pattern. The Hyperolius viridiflavus complex includes 121 described taxa, including subspecies, which are presently synonymised as 16 species. However, the taxonomy is confused as colour patterns have been used to identify species, despite an inadequate understanding of pattern variation and overlap. This study examined 267 sequences of the 16S rRNA fragment from individuals across the range of the group, from Senegal to Ethiopia, and to the southern tip of Africa. Maximum likelihood analysis showed that there were 14 species that only partially agreed with existing taxonomy. This study recognised the following species: Hyperolius viridiflavus, H. marmoratus, H. marginatus, H. parallelus, H. nitidulus, H. glandicolor, H. tuberculatus, H. mariae, H. noblei, H. goetzei, H. spatzi, H. bangwae, H. dintelmanni and an undescribed species from southern Cameroon. Hyperolius swynnertoni was found to be embedded within H. marmoratus, H. hutsebauti was embedded within H. parallelus, H. rhodesianus and H. pyrrhodictyon were embedded within H. marginatus and H. sheldricki was embedded within H. glandicolor. There was no sequence information to determine the status of Hyperolius nimbae. The ranges of the species are presented, although these are expected to change as more sequences become available. The use of DNA sequences to identify animals is recommended in this group, as the variation in advertisement calls and morphology is not well understood.

Snakes are usually solitary animals except when sheltering under cover, during male-to- male combat, mating and where species use communal dens for hibernation. The usual predation methods used are discussed including a case of cooperative hunting. Observations of possible cases of cooperative foraging by the South African Brown Water Snake,Lycodonomorphus rufulus(Lichtenstein, 1823), are provided along with a suggested explanation of the advantages of such cooperation.

We present a commented catalogue of the type specimens of the Afro-Malagasy frog family Hyperoliidae at the herpetological collection of the Museum für Naturkunde Berlin (ZMB). In current publications and databases, many names based on ZMB primary types are listed as synonyms of other species, the types often declared as lost. Consequently, the respective names are often no longer considered in current taxonomic work. We traced 146 nominal taxa of the family Hyperoliidae in the ZMB collection of which currently 130 are presented by primary types (88 holotypes, 10 lectotypes and 32 taxa based on syntype series); 50 of these taxa are currently considered as valid. Primary types of nine taxa could not be located during our inventory of the collection holdings. Seven taxa are exclusively represented by secondary types (paratypes). Many of these types comprise taxa where types have been thought to be lost. As a further service to the community, we provide important details about collectors and their travel routes, as well as respective documents stored in the collection of the Department of Historical Research at ZMB. This should make it easier to potentially compare the ZMB types in future taxonomic revisions.

Afrixalus lacustris Greenbaum, Dehling, Kusamba & Portik sp. nov. Great Lakes Spiny Reed Frog urn:lsid:zoobank.org:act: 41D3C41A-4501-4BEC-BFB6-B9C6C1518DC3 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1950: 24; Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. UTEP 20805 (field no. ELI 605), adult male, from the vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC, collected on 21 December 2010 by Chifundera Kusamba and Félix I. Alonda (Figs. 8D–F, 12A–B). Paratopotype. UTEP 20806 (field no. ELI 606), adult male, collected with the holotype. Referred specimens. Specimens with an asterisk were not included in morphological analyses due to poor quality of preservation, lack of a voucher, or unavailability of specimen(s): UTEP 20807–08 (field nos. ELI 644, 649), vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC; UTEP 20809, 22422 (field nos. ELI 669–70), Baraka, near shore of Lake Tanganyika (04.10832° S, 29.09705° E, 777 m), South Kivu Province, DRC; UTEP 20810–11 (field nos. EBG 1316, 1319), vicinity of Irangi (01.8873° S, 28.4495° E, 820 m), South Kivu Province, DRC; UTEP 22423 (field no. MUSE 10192), Kahuzi-Biega National Park, Nanwa (02.362138° S, 28.192589° E, 1566 m), South Kivu Province, DRC; UTEP 22424 (field no. MUSE 10137), Itombwe Plateau, Mbandakila (03.47937° S, 28.413049° E, 1135 m), South Kivu Province, DRC; RMCA 77-020 -B-133–136* (four specimens), Itombwe Plateau, Tubutubu (2300 m), South Kivu Province, DRC; RMCA 77-020 -B-89–92 (field nos. MF 14–15 [two specimens]), Kitutu (ca. 03.28° S, 28.11° E, 700 m), South Kivu Province, DRC; RMCA 77-020 - B-0069–71 (1 specimen), Itula (ca. 03.27° S, 28.12° E, 650 m), South Kivu Province, DRC; RMCA 77-020 -B-0081–83, 77-20-B-0086, Kamituga (ca. 03.54° S, 27.69° E, 1050 m), South Kivu Province, DRC; RMCA 77-020 - B-0072–73 (1 specimen), Mwana (ca. 03.15° S, 28.47° E, 1650 m), South Kivu Province, DRC; UTEP 22417 (field no. CRSN 2773), Toyokana (02.02734° N, 030.06653° E, 1294 m), Ituri Province, DRC; (field no. EPLU 395 *, photo and tissue only), ca. 0.5 km E of Epulu, near Mt. Mbiya (01.39594° N, 28.58093° E, 755 m), Ituri Province, DRC; UTEP 22416 (field no. DFH 247), Bwindi Impenetrable National Park, Buhoma (00.99045° S, 29.61884° E, 1523 m), Western Region, Uganda; (field nos. DFH 1102–03 * photos and tissues only), Kibale Forest National Park, Ngogo Research Center (00.49795° N, 30.42262° E, 1363 m), Western Region, Uganda; CAS 202032–36, Bwindi Impenetrable National Park, Buhoma rd., 2 km S (by rd.) of Bizenga River (01.00975° S, 29.620694° E), Western Region, Uganda; CAS 202133, Bwindi Impenetrable National Park, small tributary to Ishasha River (00.905° S, 29.704972° E, 1280 m), Western Region, Uganda; CAS 202109, Bwindi Impenetrable National Park, Bizenga River at Buhoma rd. (00.99275° S, 29.615722° E), Western Region, Uganda; CAS 256035 *, Bwindi Impenetrable National Park, rd. north of Ruhija, Western Region, Uganda; CAS 256128–31 *, Mabira Forest Reserve, marshy area adjacent to unpolluted stream (0.4508° N, 32.9493° E, 1121 m), Mukono District, Central Region, Uganda. Diagnosis. The species is referred to the genus Afrixalus for exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 18.9–22.2 mm, in females 20.8–25.7 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. fulvovittatus (males 23–27 mm, females 25–28 mm), A. lacteus (males 22–27 mm, females 25–29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), A. vittiger (males 22–25 mm, females 25–28 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger size than A. delicatus (males 15–19 mm, females 17–22 mm) and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species showing a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). It is most similar to A. phantasma sp. nov. but differs from this species in being smaller (Table 1) and showing an overall darker coloration and more contrasting dorsal pattern that is obvious even in preserved specimens (Figs. 7–8, 12). It is readily distinguished from A. phantasma sp. nov. by the more extensive hand and toe webbing (Fig. 8E–F). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8D–F); head small (HL/SVL 0.31, HW/SVL 0.31), about as long as wide (HW/HL 0.99); snout relatively long, rounded in dorsal view and in lateral profile, slightly wider than long; canthus rostralis hardly discernible, slightly concave between eye and nostril in dorsal view; loreal region oblique; nostrils rounded, directed anterolaterally and slightly dorsally; situated much closer to tip of snout and to eye, separated from each other by distance about equal to distance between eye and nostril (IN/EN 1.02); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.39), its diameter shorter than snout (ED/SL 0.90); interorbital distance much wider than upper eyelid (IO/EW 1.61) and wider than internarial distance (IO/IN 1.31); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue heartshaped, longer than broad, bilobed for about one-fourth of its length, free distally for about two-thirds its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; tiny pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; few tubercles at rear end of mandible without pointed tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8E); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender; hand large (HaL/SVL 0.28); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2-/2 II 2-/3- III 2.25/2 IV; thenar tubercle oval, small and very low, about one-fourth length of metacarpal of Finger I; inner palmar tubercle and outer palmar tubercle very low, small, rounded, almost indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.44), slightly shorter than thigh; heels not touching each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.94); relative length of toes: I I 1.5/2+ II 1+/2+ III 1+/2+ IV 2/1+ V (Fig. 8E); inner metatarsal tubercle moderately prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light brown with more or less regular, locally more densely clustered dark brown speckling; distinct dark brown transverse stripe between upper eyelids, another less distinct one in scapula region, and another narrow one at level of anterior end of pelvis; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; distinct dark brown band along canthus, continuing behind eye on both sides of trunk to level of pelvis; tubercles below eye and in tympanic region at rear end of jaw white; dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; skin of ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 12). During day, basic dorsal coloration light brown; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible. Coloration in preservative. Dorsal basic coloration largely faded to light brown; darker dorsal pattern elements dark brown, clearly visible; color of gular gland, fingers and toes faded to white (Fig. 8D–F). Variation. The paratypes match the holotype in general appearance and proportions. Coloration and color pattern varies from light-cream to yellowish-orange with a few relatively large and darker blotches and vermiculation, light speckling and indistinct dorsolateral stripe to an almost reticulated pattern of contrasting bright and dark elements with a broad and conspicuous brown dorsolateral band (Fig. 12). Pedal webbing variation is I 1.5[100]/2+[100] II 1+[100]/2+[100] III 1 +[50],1.25[50]/2+[100] IV 2 [100]/1+[100] V. Bioacoustics. The advertisement call and other vocalizations are unknown. Ecology and natural history. We collected males and females in swamps in forest openings and near forest edges (Fig. 11B), from vegetation at the edge of streams (or on vegetation overhanging them) in forest, and from non-forest vegetation near the shore of Lake Tanganyika. Egg deposition has not been observed and tadpoles are unknown. An adult male (CAS 256035) was found in Bwindi Impenetrable National Park (Uganda) ca. 2 m above ground in a tree fern “either calling or moving to a calling site (similar to ‘running’ of Kassina)” (D. Blackburn, in litt., 22 July 2021). Laurent (1955) noted the species from transitional forest. In his paper on amphibians of Virunga National Park, Laurent (1972) noted the species is most commonly found in swamps with herbaceous vegetation in secondary forest, and more rarely, in primary forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it from “une mare sombre” (i.e., a dark pond) in syntopy with Chiromantis at Kitutu (DRC), marshes where it reproduces by sticking its eggs under the leaves of fountain grass (Pennisetum, Shabunda, DRC), a shaded bank of a pond in syntopy with Hyperolius “ tuberculatus ” (likely H. hutsebauti sensu Bell et al. 2017) and H. frontalis; the center of this pond had reeds harboring H. kivuensis and Afrixalus orophilus (Shabunda, DRC). Laurent (1983:352) listed the species (along with Phrynobatrachus petropedetoides and Hyperolius frontalis) from “shadowy puddles of the thick transition forest.” Etymology. The species epithet is the Latin adjective “ lacustris,” meaning belonging to or dwelling in lakes; in allusion to the distribution of the new species in the region of the African Great Lakes. Distribution and conservation. The species is distributed from lowland rainforest of the Congo Basin at 460 m (Omaniundu, DRC, Laurent 1982) to transitional forests of the Albertine Rift at 1650 m (Mwana, DRC) and east as far as Mabira Forest, Uganda (1121 m). As shown in Figure 5, the species has a relatively large distribution, including several national parks and protected areas (e.g., Virunga National Park, Bwindi Impenetrable National Park), and thus is not likely to be threatened based on a limited distribution. Using GeoCAT, its extent of occurrence is estimated as 409,238 km ² and its area of occupancy as 160 km ² (Bachman et al. 2011). We therefore classify this species as Least Concern according to the IUCN Red List criteria (IUCN 2021). Four specimens (RMCA 77-020-B-133–136) from Tubutubu, Itombwe Plateau (2300 m) are problematic because they are morphologically consistent with A. lacustris, but they were collected from a substantially higher elevation than all other known specimens. Coordinates for Tubutubu provided by RMCA (4° S, 28.933° E) suggest the locality is east of the highest point of the plateau at an elevation of about 1400 m on Google Earth. Although it is possible that the specimens were collected at the latter elevation, Laurent (1982) mentions bamboo forest as the habitat, which usually occurs at the highest elevations of the Itombwe Plateau (Doumenge 1998). No clarity is offered by Laurent (1964), because he does not mention any Afrixalus in his study of the ecology and distribution of amphibians of the Itombwe Plateau. Given this conflicting information, we do not include the Tubutubu specimens in our current understanding of the elevational distribution of A. lacustris (see gray triangle in Fig. 5)., Published as part of Greenbaum, Eli, Portik, Daniel M., Allen, Kaitlin E., Vaughan, Eugene R., Badjedjea, Gabriel, Barej, Michael F., Behangana, Mathias, Conkey, Nancy, Dumbo, Bonny, Gonwouo, Legrand N., Hirschfeld, Mareike, Hughes, Daniel F., Igunzi, Félix, Kusamba, Chifundera, Lukwago, Wilber, Masudi, Franck M., Penner, Johannes, Reyes, Jesús M., Rödel, Mark-Oliver, Roelke, Corey E., Romero, Soraya & Dehling, J. Maximilian, 2022, Systematics of the Central African Spiny Reed Frog Afrixalus laevis (Anura Hyperoliidae), with the description of two new species from the Albertine Rift, pp. 201-232 in Zootaxa 5174 (3) on pages 218-223, DOI: 10.11646/zootaxa.5174.3.1, http://zenodo.org/record/6986207, {"references":["Ahl, E. (1930) Ueber die afrikanischen Arten der Baumfroschgattung Megalixalus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1930, 89 - 102.","Laurent, R. (1950) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National Albert, Mission G. F. De Witte (1933 - 1935), 64, 5 - 120, 7 pls.","Laurent, R. F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxieme Serie, 22, 1 - 125, 11 pls.","Laurent, R. F. (1982) Le genre Afrixalus Laurent (Hyperoliidae) en Afrique Centrale. Annales du Musee Royal de l'Afrique Centrale, Serie in Octavo, Science Zoologique, 235, 1 - 93.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, New York, xi + 418 pp.","Spawls, S., Howell, K. M. & Drewes, R. C. (2006) Pocket Guide to the Reptiles and Amphibians of East Africa. A & C Black Publishers, London, 240 pp.","Channing, A. & Rodel, M. - O. (2019) Field Guide to the Frogs & Other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.","Laurent, R. F. (1955) Apercu de la biogeographie des batraciens et des reptiles de la region des grand lacs. Bulletin de la Societe Zoologique de France, 79, 290 - 310.","Bell, R. C., Parra, J. L., Badjedjea, G., Barej, M. F., Blackburn, D. C., Burger, M., Channing, A., Dehling, J. M., Greenbaum, E., Gvozdik, V., Kielgast, J., Kusamba, C., Lotters, S., McLaughlin, P. J., Nagy, Z. T., Rodel, M. - O., Portik, D. M., Stuart, B. L., VanDerWal, J. & Zamudio, K. R. (2017) Idiosyncratic responses to climate-driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands. Molecular Ecology, 26 (19), 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260","Laurent, R. F. (1983) About the herpetofauna of Central African montane forest. In: Rhodin, A. G. J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Cambridge, Massachusetts, pp. 350 - 358.","Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial Conservation Assessment Tool. ZooKeys, 150, 117 - 126. https: // doi. org / 10.3897 / zookeys. 150.2109","IUCN (2021) IUCN Red List Categories and Criteria: Version 2021 - 3. Available from: https: // www. iucnredlist. org / (accessed 25 May 2022)","Doumenge, C. (1998) Forest diversity, distribution, and dynamique in the Itombwe Mountains, South-Kivu, Congo Democratic Republic. Mountain Research and Development, 18 (3), 249 - 264. https: // doi. org / 10.2307 / 3674036","Laurent, R. F. (1964) Adaptive modifications in frogs of an isolated highland fauna in Central Africa. Evolution, 18 (3), 458 - 467. https: // doi. org / 10.1111 / j. 1558 - 5646.1964. tb 01622. x"]}

Afrixalus phantasma Dehling, Greenbaum, Kusamba & Portik sp. nov. Ghost Spiny Reed Frog urn:lsid:zoobank.org:act: 1B2DBF26-5B0A-4DA8-B65E-0BC01704C1B4 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. ZFMK 103454 (field no. JMD 723), adult male, from Gishwati Forest (01.823745° S, 29.360373° E, 2084 m), now part of Gishwati-Mukura National Park, Western Province, Rwanda, collected on 5 April 2011 by J. Maximilian Dehling and Bonny Dumbo (Figs. 7A, 8A–C). Paratypes. ZFMK 103455 (field no. JMD 722), adult female, collected with the holotype; ZFMK 103456 (field no. JMD 2015-30), adult male, ZFMK 103457 (field no. JMD 2015-31), adult female, from the type locality in Gishwati Forest, both collected on 27 September 2015 by J. Maximilian Dehling and Bonny Dumbo; ZFMK 103458–60 (field nos. JMD 677–679), three adult males, collected on 19 March 2011 by J. Maximilian Dehling, ZFMK 103461–62 (field nos. JMD 954–955), two adult males, collected on 16 February 2013 by J. Maximilian Dehling, all from Kamiranzovu Swamp (02.477165° S, 29.158243° E, 1962 m), Nyungwe National Park, Western Province, Rwanda; UTEP 20791–20792 (field nos. EBG 2838, 2843), two adult males, collected on 21 December 2009 by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Maurice Luhumyo from Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC. Referred specimens. UTEP 20802 (field no. EBG 1198), forest ca. 4 km NW of Lwiro (02.2226° S, 28.7754° E, 2077 m), South Kivu Province, DRC; UTEP 20803, 22418–20 (field nos. EBG 1232, 1238–40), Kahuzi-Biega National Park, Mugaba (02.2750° S, 28.6631° E, 2298 m), South Kivu Province, DRC; UTEP 20804 (field no. EBG 1283), Kahuzi-Biega National Park, Chinya (02.2671° S, 28.6455° E, 2267 m), South Kivu Province, DRC; UTEP 20793–94 (field nos. EBG 2844–45), Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC; UTEP 20795–20800, 22421 (field nos. ELI 414–20), Nyakasanza Swamp near Tshibati (02.22829° S, 28.77972° E, 1979 m), South Kivu Province, DRC; UTEP 20801 (field no. ELI 425), Chanjoka (02.21261° S, 28.77644° E, 2115 m), South Kivu Province, DRC. Diagnosis. The species is referred to the genus Afrixalus by exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 20.1–24.6 mm, in females 22.6–26.4 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. lacteus (males 22–27 mm, females 25– 29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31 mm, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger than A. brachycnemis (males 18–21 mm, females 20–22 mm), A. delicatus (males 15–19 mm, females 17–22 mm), A. spinifrons (males to 20 mm, females to 25 mm), and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species with a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). The advertisement calls of the following species of Afrixalus consist of a long series of clicks, often initiated by a long buzzing note, and thus differ from the call of A. phantasma: A. aureus, A. brachycnemis, A. clarkei, A. crotalus, A. delicatus, A. dorsalis, A. fornasini, A. fulvovittatus, A. knysnae, A. nigeriensis, A. morerei, A. osorioi, A. quadrivittatus, A. septentrionalis, A. spinifrons, A. vibekensis, A. vittiger, A. weidholzi, and A. wittei. The advertisement calls of the remaining species for which information is available differ from the call of A. phantasma in the following characteristics (in parentheses): A. dorsimaculatus (soft, short buzzing rattle, usually repeated three times), A. equatorialis (series of five notes, repeated at 15–20/s and initiated by a long buzz, energy maximum at 2000–2500 Hz), A. lacteus (8–9 notes, 14–15 pulses,>300 pulses/s), A. paradorsalis (2–3 notes, energy maximum at 2700 Hz), A. stuhlmanni (2–9 notes, energy maximum at 4200–5100 Hz), and A. sylvaticus (2–5 notes, energy maximum at 4000–4500 Hz). The new species is most similar to A. lacustris sp. nov. from which it differs by a number of characteristics (see below). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8A–C); head small (HL/SVL 0.31, HW/SVL 0.30), about as long as wide (HW/HL 0.98); snout relatively long, rounded in dorsal view and in lateral profile, slightly longer than wide; canthus rostralis hardly distinct, straight between eye and nostril; loreal region oblique; nostrils rounded, directed anterodorsally and slightly laterally; situated much closer to tip of snout and to eye, separated from each other by distance larger than distance between eye and nostril (IN/EN 1.15); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.41), its diameter shorter than snout (ED/SL 0.84); interorbital distance much wider than upper eyelid and wider than internarial distance (IO/IN 1.21); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue short and moderately broad, bilobed for about one-sixth of its length, free distally for about half its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; minute pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; small, subcircular, elevated glandular area in tympanic region posterior to eye and rear end of mandible, bearing large tubercles with pointed keratinous tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8B); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender;hand large (HND/SVL 0.32); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2+/2.25 II 2/3+ III 2.75/2.25 IV; thenar tubercle oval, small and low, about one-fourth length of metacarpal of Finger I; palmar tubercles indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.48), about as long as thigh; heels meeting each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.96); relative length of toes: I I 2/2.25 II 1.5/3- III 1.75/3 IV 3-/1.75 V (Fig. 8C); inner metatarsal tubercle moderately prominent, elongated, about twothirds length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light yellowishbrown with more or less regular dark brown speckling; indistinct darker brown transverse stripe between upper eyelids; large dark brown, irregularly shaped spot in scapula region, two less distinct, somewhat lighter brown, irregularly shaped spots at level of anterior end of pelvis, separated from each other by half their width; cloacal region with dark brown mottling; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; weak brown stripe along canthus, continuing behind eye on anterior half of trunk on both sides; tubercles below eye, and in tympanic region at rear end of jaw white; indistinct dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 7A). During day, basic dorsal coloration brighter, very light brown to bright cream-colored; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible (Fig. 7B). Coloration in preservative. Dorsal basic coloration largely faded to yellowish white; darker dorsal pattern elements light to dark brown, clearly visible; yellow color of gular gland, fingers and toes faded to white (Fig. 8A–C). Variation. The paratypes match the holotype in general appearance, proportions, coloration and color pattern. Mensural variation within the species is shown in Table 1. Pedal webbing variation is I 2+[100]/2.5[90],2.25[10] II 2[50],2+[40],2.25[10]/3[10],3+[40],3.5[50] III 1.75 [80],2-[20]/3[100] IV 3 -[40],3[60]/1.75[100] V. Bioacoustics. Series of advertisement calls of six different males were recorded at the following ambient temperatures: 10.9°C (N = 1), 13.6°C (N = 3), and 16.2°C (N = 2). The advertisement call consisted of five to six, rarely four pulse groups (notes) (Fig. 9). Depending on the ambient temperature, notes were repeated at a rate of 7.1–10.0/s (10.9°C), 8.2–11.7/s (13.6°C), and 10.2–11.4/s (16.2°C). The highest repetition rate was always between the first three notes of a series, the lowest at the end of the series. Each note consisted of 10–11 pulses (Fig. 9). Probably due to echo effects, pulsation was often veiled in the waveforms, especially towards the end of the note. Pulse repetition rate varied from 190–210/s at 10.9°C, 222–250/s at 13.6°C, and 277–292/s at 16.2°C, resulting in a note length of 72–94 ms, 55–70 ms, and 36–50 ms, respectively. Amplitude modulation was prominent within individual notes (Fig. 9). Total call length showed linear temperature dependence and varied from 388–397 ms at 16.2°C, 472–499 ms at 13.6°C, and 572–620 ms at 10.9°C for five-note calls (Fig. 10). Energy maximum showed linear temperature dependence, with 3020–3150 Hz at 10.9°C, 3370–3550 Hz at 13.6°C, and 3660–3810 Hz at 16.2°C (Figs. 9–10). There was no marked frequency modulation. Prominent harmonics were at about 6000–7000 Hz and 9000–11000 Hz (Fig. 9). Ecology and natural history. We collected males and females in swamps in forest openings, or at forest edges (Fig. 11A). Males were found calling above standing bodies of water with thick lower vegetation cover. In Gishwati Forest and Kamiranzovu Swamp in Nyungwe Forest, the species was found calling in syntopy with Hyperolius castaneus and H. discodactylus. One adult male (UTEP 20802) from Kahuzi-Biega National Park (DRC) was found 3.5 meters above ground in montane forest. Tadpoles are unknown. Laurent (1955, 1983) listed the species from bushes in montane forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it in banana trees in a deep valley (Nyungwe, Rwanda) and “des trous de prospection” (i.e., prospecting holes likely for gold mining, Upper Lubitshako, Kabobo Plateau, DRC). Etymology. The species epithet derives from the Greek noun φάντασμα (phántasma), meaning ghost or phantom, in allusion to the coloration and general appearance of the new species. The epithet is used as an invariable noun in apposition. Distribution and conservation. The occurrence of the species has been confirmed for several locations in western Rwanda (Nyungwe and Gishwati Forest) and eastern DRC (in and near Kahuzi-Biega National Park, Itombwe Plateau and Kabobo Plateau [Laurent 1982], Fig. 5). We expect the species to be eventually found in the AR of southwestern Uganda and northwestern Burundi. So far, the species has been recorded from only a narrow elevation range between 1962 m (Kamiranzovu Swamp, Rwanda) and ca. 2400 m (May ya Moto [and possibly Luemba], Itombwe Plateau, Laurent 1982). Given the relatively limited overall geographic distribution of the species with an estimated extent of occurrence of 19,088 km 2 (Fig. 5), an estimated area of occupancy of 52 km 2 (both calculated with GeoCAT; Bachman et al. 2011), the detection of the amphibian chytrid fungus in one adult male (UTEP 20791, Greenbaum et al. 2015), and the conservation challenges facing natural areas of the AR (Greenbaum 2017; Ayebare et al. 2018), we categorize this species as Vulnerable under the IUCN Red List criteria (IUCN 2021).

Hyperolius tuberculatus (Mocquard, 1897) Rappia tuberculata Mocquard, 1897. MNHNP 96.570 “Lambaréné”, Gabon (Table 3). Back and exposed parts of limbs light reddish-brown back with many fine tubercles. A brown stripe is sometimes present from the eye to the snout tip. Another mark of the same colour reaches the eye from the top of the head, while another starts at the lower edge of the eye, running obliquely to the angle of the jaw. This clade consists of 30 sequences from Gabon, Bioko Island (Equatorial Guinea) and Cameroon (Fig. 9B). All sequences were reported in GenBank as Hyperolius tuberculatus. The sequence reported in Veith et al. (2009) as Hyperolius tuberculatus Complex A forms part of this species. The uncorrected p- distances from other clades were 2.1–6.2% (Table 2). The pattern element in the description of the type of H. tuberculatus consists of various stripes on the head. The advertisement call (no vouchers) is a high-pitched long series of notes (Bosch et al. 2000, from Río Muni, Equatorial Guinea), with an emphasised frequency of 3228 Hz, and a mean note rate of 11.6 s- 1. A call from Adjap, Cameroon (no voucher) is a series of double pulsed notes with a note rate of 6 s-1, and an emphasised frequency of 3000 Hz (Amiet & Goutte 2017)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 338, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Veith, M., Kosuch, J., Rodel, M. - O., Hillers, A., Schmitz, A., Burger, M. & Lotters, S. (2009) Multiple evolution of sexual dichromatism in African reed frogs. Molecular Phylogenetics and Evolution, 51, 388 - 393. https: // doi. org / 10.1016 / j. ympev. 2008.12.022","Bosch, J., De la Riva, I. & Marquez, R. (2000) Advertisement calls of seven species of hyperoliid frogs, from Equatorial Guinea. Amphibia-Reptilia, 21, 246 - 255.","Amiet, J. - L. & Goutte, S. (2017) Chants d'Amphibiens du Cameroun. J. - L. Amiet & Editions Petit Genies, Nyons, 280 pp."]}

Hyperolius parallelus Günther, 1858. Hyperolius parallelus Günther, 1858. Syntypes BMNH 1947.2.9.41–43 “Süd Afrika”. Boulenger (1882) corrected the locality for the northern specimens to Ambriz, Angola. Above dark brown with three white parallel bands. Upper lip yellowish. Hyperolius marmoratus var. angolensis Steindachner, 1867 type in NHMW, “ Angola ”. Synonymy by Broadley 1971. Back and upper surfaces of limbs brown. Back with irregular yellow flecks or reticulation, or with a pair of yellow lateral bands and irregular yellow flecks in the middle of the back. Limbs with yellow spots and lines. Hyperolius insignis Bocage, 1868. Syntypes MBL (now MUHNAC) T. 21–164, “Benguella”, type localities corrected to São Salvador and Novo Redondo, and ZMB 6462 to Benguela, Angola by Perret 1976. Synonymy by Boettger (1888). Colour in alcohol: Overall yellow tinted with red. Two wide blue-black bands with bright red edges run from the head to the vent. A similar colour band runs from behind the eye to the shoulder, continuing interrupted to the vent. The upper lip has small blue spots with red borders, with similar markings on the upper limbs. Hyperolius toulsonii Bocage, 1868. MBL (now MUHNAC) T.27–275, “Loanda” (Luanda, Angola). Type destroyed in 1978 fire. Synonymy by Perret, 1976. Colour in alcohol: back overall grey, with three wide longitudinal white bands. Thighs uniform grey, with forearms and thighs with large rounded white markings. Hyperolius huillensis Bocage, 1873. MBL (now MUHNAC) 2-166 destroyed in 1978 fire, “Huilla”, Angola. Placed in the synonomy of H. parallelus as sequences from Huila are embedded in the H. parallelus clade. Back and upper surfaces of limbs, excluding the thighs are red with yellow spots, most with black borders. Ventral surfaces and thighs a uniform pale yellow. Hyperolius vermiculatus Peters, 1882. Syntypes ZMB 10050, “ Malanje ”, Angola. It was initially placed as a synonym of H. marmoratus by Laurent (1952), but H. marmoratus only occurs on the east coast of southern Africa. Back and upper surfaces of limbs signal red with yellow vermiculation, with black pigment sometimes visible. Rappia plicifera Bocage, 1893. Syntypes MBL (now MUHNAC) T.22-209 destroyed in 1978 fire. Syntypes NHMW 22895, Caconda, Duque de Bragança, Angola. Synonymy by Perret (1976). Colour in alcohol: Above marbled and spotted in brown-violet or red on a grey base. Lower surfaces and thighs a uniform bright yellow. Rappia marmorata var. taeniolata Bocage, 1895. Types in MBL (now MUHNAC) considered destroyed. “ Duque de Bragança ”, “Huilla”, “Caconda” and “Cahata”, Angola. Synonymy by Laurent (1952). Back and flanks burgundy with various irregular yellow lines and spots, with black borders. Rappia marmorata var insignis Bocage, 1869. Syntypes MBL (now MUHNAC) T 21–164 destroyed by fire, surviving syntype ZMB 6462 (Tillack et al. 2021). St. Salvador du Congo, Angola north of Quanza, Novo Redondo, Dombe, Angola. Synonymy by Perret (1976). Back black with three longitudinal white or yellow bands with red edges. Rappia burgeoni De Witte, 1921. MRAC 585, “Madyu (Uelé)” DRC. See the section on obscure localities, above. Watsa, Upper Uelé, DRC. This taxon from the northern DRC was regarded as a synonym of H. viridiflavus pachydermus by Laurent (1951a). Dehling (2015) demonstrated that Rappia pachyderma Werner, 1908 was a junior synonym of Hyperolius papyri (Werner, 1908), a species of the H. nasutus group (Channing et al. 2013) and that the synonymization of several species of the H. viridiflavus group with this species by Laurent (1951) were thus unwarranted. However, the clade identified as a subspecies of H. viridiflavus by Laurent (1951) does not occur in the northern DRC, while the clade identified as H. parallelus is well represented in the northern DRC. Pale yellow above, with a dark brown lateral band from the snout tip through the eye to the vent. A band of similar colour on the tibia. A darker vertebral band present. Rappia rossii Calabresi, 1925. Syntypes MSNM 584 “Regione dell’Alto Uellè (Congo Belga)” Watsa, DRC. The H. viridiflavus clade is not known as far west as Watsa. See also Dehling (2015) and image of syntype in Blackburn & Scali (2014). Above blue-grey with white tubercles with red centres. Thighs and fingers with pink speckle. Hyperolius bitaeniatus Ahl, 1931. ZMB 39004, “Konde-Nyika, Deutsch-Ost-Afrika” (northern tip of Lake Malawi), Tanzania. Placed as a synonym of H. marmoratus mariae by Laurent (1952) but here placed in H. parallelus as the H. mariae clade does not occur in the northern Lake Malawi area. Colour in alcohol: Above light brown, with two yellow or white longitudinal stripes. A dark lateral band runs between the arms and leg. Hyperolius nyassae Ahl, 1931. ZMB 39006, “Langenburg” Lumbira, Tanzania. Placed as a synonym of H. marmoratus nyassae (Laurent 1952), but here placed in H. parallelus as the H. marmoratus clade does not occur in the northern Lake Malawi area. Colour in alcohol: Back and upper surfaces of limbs white to yellow or brown. Some individuals have a large triangular mark between the eyes, pointing backwards. A thin dark line runs from the nostril to the eye, where it widens to become a lateral band that ends in the middle of the body. A pair of large markings between the flank and the sacrum, with another pair over the vent. The slightly darker markings are thinly edged in white or yellow. The tibia has a similar transverse marking. Hyperolius fuelleborni Ahl, 1931. ZMB 86128 –86131, “S’ongwe” Songwe, Tanzania, ZMB 85972 –85973, “Miramba bei Langenburg”, Miramba near Lumbira, Tanzania, ZMB 77465 –77468, “Langenburg” Lumbira, Tanzania, ZMB 85929 –85963, 85974–85988, 86017, 86132–86137, 90929–90948, “Rugwe” (Rungwe village, Mbeya Region, Tanzania). Placed as a synonym of H. marmoratus nyassae by Laurent (1952), but here moved to H. parallelus as the H. marmoratus clade does not occur in the northern Lake Malawi area. The specimens ZMB 71184 –71186, “Neu-Helgoland” Papya Island, Malawi, are retained in H. marmoratus. Colour in alcohol: Pale above with more or less distinct dark brown to black markings that overlap to form a marbled pattern. Hyperolius marungaensis Ahl, 1931. ZMB 10736, “Marunga, Angola ”, corrected to northern Marunga, DRC by Tillack et al. (2021). It was considered a synonym of H. marmoratus by Laurent (1961). The clade identified as H. marmoratus is restricted to the east coast of southern Africa. The colour pattern (Ahl 1931: fig. 226) consists of a dark back with three large round markings on the head, five large circular markings arranged in a vertebral line, and another five longer white markings behind each eye running to the flank. Upper lip with a broad white band. Hyperolius decoratus Ahl, 1931. Lectotype ZMB 36112, “Longa” (=Cuito Cuanavale), Angola. Regarded as a synonym of H. marmoratus (Perret 1962), but that clade is restricted to the east coast. Colour in alcohol: above wine-red, with many more or less large round, white markings with black borders. Three broad longitudinal stripes, the first running from snout tip to vent, the other two run from the side of the snout to the flanks. Similar patterns are found on the visible surfaces of the limbs. Hyperolius microstictus Ahl, 1931. ZMB 36100, ZMB 77762, “Longa, oberhalb Minnesera” (=Cuito Cuanavale), Angola. Synonymised with H. marmoratus decoratus by Perret (1962) but that clade is restricted to the east coast. Colour in alcohol: Above and on the visible parts of the limbs when sitting, bright wine-red with many irregular, pointed white markings. Hyperolius decipiens Ahl, 1931. ZMB 39003, “Westliches Russisi-Ufer und Nordwestufer des Tanganyika-see’s” (Rusizi River west bank and NW Lake Tanganyika, Tanzania). Synonymy with H. marmoratus argentovittis by Laurent (1952) but here moved to H. parallelus as the H. marmoratus clade does not occur in the northern Lake Malawi area. Colour in alcohol: Above brown to blue-grey without markings. Upper lip with white border. Hyperolius nossibeensis Ahl, 1930. ZMB 50098 –50100, “Nossi-Bé” Madagascar. Corrected by Tillack et al. (2021) to Lunda, Angola. Originally placed as a synonym of H. marmoratus, which is restricted to the east coast. The Angolan coastal locality places it within the distribution of the H. parallelus clade. Colour in alcohol: Above red-brown with a number of silver-white flecks and lines, that flow together to form three longitudinal bands, that may be interrupted and indistinct. The side of the head and body similarly marbled. Hyperolius callichromus Ahl, 1931. ZMB 78576, “Westliches Russisi-Ufer und nordwestufer des Tanganyika-see’s” (Rusizi River west bank and NW Lake Tanganyika, Tanzania). Synonymy with H. marginatus by Wieczorek et al. (2000) but here moved to H. parallelus as the H. marginatus clade does not occur in the northern Lake Malawi area. Colour in alcohol: Above deep red-brown, with a wide white vertebral band with black border, that runs from level of nostrils to vent. A similar band runs laterally from upper eyelid to vent. The longitudinal bands may be broken into irregular shapes. Upper lip white, sides with a wide red-brown stripe. Illustrated by Tillack et al. (2021, fig. 6). Paratypes from Dar es Salaam ZMB 85869 –85872 (Tillack et al. 2021), are assigned as synonyms of H. mariae. Hyperolius erythromelanus Monard, 1937. Syntypes in LCFM. Sangev, Angola. Synonymy with H. marmoratus huillensis by Laurent (1952). The H. marmoratus clade does not occur in Angola. Upper surfaces dark red, almost black with many small white spots, arranged in lines edged with black. Upper surfaces of limbs the same colour as the back. Hyperolius angolensis quarrei Laurent, 1943. MRAC 37366, “Lusuku (Lomami)”, DRC. Synonymy with H. marmoratus by Laurent (1952), but that clade is restricted to the east coast. Colour in alcohol: Dorsal surfaces yellow to grey, with rust or red-brown irregular marbled markings. Upper surfaces of limbs with similar markings. Hyperolius latirostris Laurent, 1943. MRAC 1394, “Kasai”, DRC. Considered a subspecies of H. marmoratus by Laurent (1952), but that clade does not extend as far west as the DRC. Back and parts of the limbs visible at rest, red-brown, dappled with pale brown and grey, with a violet tinge to the head. Tibia with a trace of brown. Hyperolius brieni Laurent, 1943. MRAC 5445, “Nyonga (s/Lac Upemba)” DRC. 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. Hyperolius fuliginosus Laurent, 1943. MRAC 35166, “Kasenga (Katanga)” DRC. Synonymy with H. marmoratus by Schiøtz (1975). Hyperolius marmoratus does not occur in the DRC, but the locality is close to sequenced material of H. parallelus. Colour in alcohol: Grey-brown above, on the forearm and tibia, with dark brown marbling with indistinct edges. A pale vertebral band and two indistinct lateral bands are present. Limbs finely dotted or with small black spots. Hyperolius hutsebauti Laurent, 1956. MRAC 52495, “Ibembo, Uelé” DRC. Sequences of specimens identified as H. hutsebauti are embedded in a clade identified as H. parallelus. Earlier studies that recognised H. hutsebauti (e.g. Bell et al. 2017) did not compare material with H. parallelus. Female colouration consists of a beige background with dark marbling edged with red. Hyperolius marmoratus epheboides Laurent, 1957. IRSNB 1.158, “Mukana, marais près de Lusinga 1.810 m ” (Mukana swamp near Lusinga) DRC. The H. marmoratus clade is restricted to the east coast, while the type locality is close to sequenced material in the H. parallelus clade. Colour in alcohol: A beige background, with black markings on the flanks, the side of the head, and on the upper surfaces of the limbs. These are intertwined with longitudinal bright red markings. In life the ventral surface has brilliant red markings. Hyperolius marmoratus alborufus Laurent, 1964. MD 5769, “Cazombo, Alto Zambeze, Moxico, Angola ”. The holotype is figured in Ceríaco et al. (2020). The H. marmoratus clade is restricted to the east coast, while the type locality is close to sequenced material in the H. parallelus clade. Back with red marbling, covering also the upper arm and tibia. The sides of the tibia and the toes with small black spots. A total of 57 sequences were available from Angola, Burundi, Rwanda, DRC, northern Zambia, and northern Namibia. The minimum uncorrected p- distances from other clades were 2.2–6.5% (Table 2). The sequences were reported in GenBank as H. angolensis, H. parallelus, H. huillensis, H. viridiflavus and H. v. argentovittis. The sample is distributed from northern Namibia, through Angola, across to southern Lake Tanganyika, Burundi, Rwanda and northern DRC (Fig. 7). The type localities of taxa now regarded as junior synonyms of H. parallelus are listed in Table 3. Hyperolius parallelus Günther, 1858 was described from “Süd-Afrika”. Boulenger (1882) corrected the locality for the northern specimens to Ambriz, on the northern coast of Angola. Only one clade has a distribution that includes coastal Angola, and it is assigned to Hyperolius parallelus. The most common pattern elements in the descriptions of the types included in H. parallelus are dorsal marbling (39%), or a pale lateral band (30%), or a pale vertebral stripe (30%), followed by small yellow spots sometimes with red centres or black edges (17%). The percentages are not additive, as the pattern elements may be shared. The underside may be red, sometimes with red speckles on the throat. Calls from Humpata, Angola, are brief (0.03 s) at an emphasised frequency of 1189–2153 Hz., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 330-334, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London, 256 pp., 16 plates.","Broadley, D. G. (1971) The reptiles and amphibians of Zambia. Puku, 6, 1 - 143.","Perret, J. - L. (1976) Revision des amphibiens africains et principalement des types, conserves au Musee Bocage de Lisbonne. Arquivos do Museu Bocage, Lisboa, Serie 2 6, 15 - 34.","Boettger, O. (1888) Materialien zur Fauna des unteren Congo II. Reptilien und Batrachier. Bericht uber die Senckenbergische Naturforschende Gesellschaft in Frankfurt am Main, 1888, 3 - 108.","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Dehling, J. M. (2015) On the identities of two enigmatic reed frog taxa from South Sudan, Rappia papyri Werner, 1908 and Rappia pachyderma Werner, 1908. Herpetozoa, 28, 39 - 47.","Channing, A., Hillers, A., Lotters, S., Rodel, M. - O., Schick, S., Conradie, W., Rodder, D., Mercurio, V., Wagner, P., Dehling, J. M., du Preez, L. H., Kielgast, J. & Burger., M. (2013) Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species. Zootaxa, 3620 (3), 301 - 350. https: // doi. org / 10.11646 / zootaxa. 3620.3.1","Blackburn, D. C. & Scali, S. (2014) An annotated catalog of the type specimens of Amphibia in the collection of the Museo Civico di Storia naturale, Milan, Italy. Herpetological Monographs, 28, 24 - 45. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00008","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Bell, R. C., Parra, J. L., Badjedjea, G., Barej, M. F., Blackburn, D. C., Burger, M., Channing, A., Dehling, J. M., Greenbaum, E., Gvozdik, V., Kielgast, J., Kusamba, C., Lotters, S., McLaughlin, P. J., Nagy, Z. T., Rodel, M. - O., Portik, D. M., Stuart, B. L., VanDerWal, J., Zassi-Boulou, A. G. & Zamudio, K. R. (2017) Idiosyncratic responses to climate-driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands. Molecular Ecology, 26, 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260","Laurent, R. F. (1957) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National de l'Upemba. Mission G. F. de Witte (1946 - 1949), 42, 3 - 47.","Ceriaco, L. M. P., Marques, M. P., Andr, I., Afonso, E., Blackburn, D. C. & Bauer, A. M. (2020) Illustrated type catalogue of the \" lost \" herpetological collections of Museum do Dundo, Angola. Bulletin of the Museum of Comparative Zoology, 167, 379 - 440. https: // doi. org / 10.3099 / 0027 - 4100 - 162.7.379"]}

Hyperolius nitidulus Peters, 1875 Hyperolius nitidulus Peters, 1875. ZMB 7729, Lagos, Nigeria. Colour in alcohol: Above violet-grey, a dark brown stripe from the nostrils, over the eye, to the flank, where it becomes many flecks against a pale background. Below this band, white with many small black flecks. Hyperolius pallidus Mertens, 1940. NHMW 16862, “Poli bei Garua, Nordkamerun” (Poli near Garua, Cameroon). Synonymy by Rödel (2000). Above pale grey with fine black speckling on the head and sides of females. Juveniles uniform pale, or with two to four pale longitudinal stripes. Hyperolius viridiflavus angeli Laurent, 1951. MNHNP 1921.34b, “Gribingui, Oubang-Chari” Central African Republic. The H. viridiflavus clade does not extend far west into the Central African Republic, and the type is here assigned to H. nitidulus. Dorsal colour uniform, with fine black speckling on side of head, tibia and lower leg, which may not be visible in some specimens. Hyperolius nitidulus aureus Perret, 1966. MHNG 1038.39 “Ngaoundér”, Cameroon. Golden-yellow above, with a dark brown stripe from the side of the snout over the eye to the vent with a black border, and a similar dark line along the tibia. Table 3 lists the localities of the synonyms. Sequences were available from Côte d’Ivoire, Benin, Ghana, and Cameroon (Fig. 8A). The minimum uncorrected p- distances from other clades were 3.3–7.2% (Table 2). Hyperolius nitidulus bangwae Perret, 1966 was removed from the synonymy of H. nitidulus and elevated to a full species. Rödel et al. (2010) showed that H. nitidulus was genetically different from H. spatzi, and confirmed its status, following earlier proposals (Wieczorek et al. 2001, Schiøtz 1967, Drewes 1984, and Rödel 2000). The 16S rRNA sequences from Rödel et al. (2010) were used to identify this clade. The localities of taxa regarded as junior synonyms of H. nitidulus are listed in Table 8. The type locality of Hyperolius nitidulus Peters, 1875 is Lagos, Nigeria. Although there were no sequences available from Lagos or anywhere else in Nigeria, this clade consists of sequences that were identified in the field as H. nitidulus or H. viridiflavus nitidulus, and that occurs west and east of Nigeria. Only two clades (H. nitidulus and H. spatzi) are known from West Africa. Portik et al. (2019) recognised two sub-groups, one in Cameroon and the other in Ghana. Although the Cameroon sample is monophyletic, and has about a 2% difference from the Ghana samples, it is not reciprocally monophyletic from the samples from Benin, Ghana, Nigeria and Côte d’Ivoire. The most common pattern elements in the descriptions of the types included in H. nitidulus are a dark lateral band (50%), and a uniform back (50%). The percentages are not additive, as the pattern elements may be shared. The advertisement call (no voucher) is described as ‘a single, pure, metallic and very loud tone’. Mean call length is 0.08 s, at a mean frequency of 2638 Hz (Rödel et al. 2010)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 334-335, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Rodel, M. - O. (2000) Herpetofauna of West Africa. Vol 1 Amphibians of the West African Savanna. Edition Chimaira, Frankfurt am Main, 332 pp.","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie, 93, 289 - 464.","Rodel, M. - O., Sandberger, L., Penner, J., Man, Y. & Hillers, A. (2010) The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae). Bonn zoological Bulletin, 57, 177 - 188.","Schiotz, A. (1967) The treefrogs (Rhocophoridae) of West Africa. Spolia zoologica Musei hauniensis, 25, 1 - 346.","Drewes, R. C. (1984) A phylogenetic analysis of the Hyperoliidae (Anura): Treefrogs of Africa, Madagascar, and the Seychelles Islands. Occasional Papers of the California Academy of Sciences, 139, 1 - 70.","Portik D. M., Bell, R. C., Blackburn, D. C., Bauer, A. M., Barratt, C. D., Branch, W. R., Burger, M., Channing, A., Colston, T. J., Conradie, W., Dehling, J. M., Drewes, R. C., Ernst, R. Greenbaum, E., Gvozdik, V., Harvey, J., Hillers, A., Hirschfeld, M., Jongsma, G. F. M., Kielgast, J., Kouete, M. T., Lawson, L. P., Leach, A. D., Loadcer, S. P., Lotters, S., Van der Meijden, A., Menegon, M., Muller, S., Nagy, Z. T., Ofori-Boateng, C., Ohler, A., Papenfuss, T. J., Rossler, D., Sinsch, U., Rodel, M. - O., Veith, M., Vindum, J., Zassi-Boilou, A. - G. & McQuire, J. A. (2019) Sexual dichromatism drives diversification within a major radiation of African amphibians. Systematic Biology, 68, 859 - 875. https: // doi. org / 10.1093 / sysbio / syz 023"]}

Hyperolius mariae Barbour & Loveridge, 1928 Rappia vermiculata Pfeffer, 1893. Syntype ZMB 10988 (other ZMB syntypes destroyed) “Sansibar (= Zanzibar, Tanzania) The name is preoccupied by Hyperolius vermiculatus Peters, 1882, and so is not available, although it appears to be the oldest name for this taxon. Above brown with many small yellow flecks. A dark band runs from eye to snout tip. Eyelid very dark with thin yellow margin. Hyperolius mariae Barbour & Loveridge, 1928. MCZ 13267, “near the mill dam at Derema, Usambara Mtns., Tanganyika Territory ” Tanzania. Colour in life: Above uniform greyish-white, with a black speck on snout, nostrils ringed with black, edge of upper eyelid black. A broad cream-coloured stripe laterally with a broad black one below it. Lower lip with red marks, belly blood-red. Hyperolius callichromus paratypes from Dar es Salaam ZMB 85869–85872 (Tillack et al. 2021), are assigned as synonyms of H. mariae as this is the only clade known from Dar es Salaam. Hyperolius vermicularis Ahl, 1931. ZMB 10988, Zanzibar, Tanzania. (Replacement name for Rappia vermiculata Pfeffer, 1893). Only the H. mariae clade is known from Zanzibar. Sequences from the adjacent mainland form part of the clade. Above brown with very many small yellow flecks, forming vermiculation. A dark band from snout tip to eye, eyelid with a thin yellow margin, tibia with small dark speckles. Hyperolius melanopthalmus Ahl, 1931. ZMB 85670 –85672, “ Sansibar ”, (Zanzibar, Tanzania). Synonymy by Loveridge (1936). Colour in alcohol: Above pale grey, nostrils ringed with black, upper eyelid with dark fleck. A broad lateral dark or violet band between the limbs, sometimes marbled. Hyperolius rubripes Ahl, 1931. ZMB 36110 and 57530, “Kililana”, Kenya. Synonymy by Wieczorek et al. (2000). Colour in alcohol: Above uniform pale yellow, pink or brownish. Hyperolius renschi Ahl, 1931. ZMB type lost (Tillack et al. 2021) Zanzibar, Tanzania. Synonymy by Loveridge (1936). Colour in alcohol: Above pale yellow, with a lateral violet band. Hyperolius marmoratus mafianus Laurent, 1961. BMNH 1937.7.1.42–44, This is a new name for specimens of H. albifrons Ahl, 1931, from Mafia Island, reported on by Parker (1937). Only one clade is known from northern coastal Tanzania. Back and sides yellowish-brown. Upper surface of snout silvery-white or yellowish-green, extending back to the eyes, continuing as lateral stripes to the level of the shoulder. The lateral stripes may branch to form a dorsal marbling, or may unite over the coccyx.A dark subdermal lateral band is present. Limbs the same colour as the back. Hyperolius udjidjiensis Ahl, 1931 was removed from the synonymy of H. mariae and placed in the synonymy of Hyperolius parallelus Günther, 1858 by Wieczorek et al. (2000). Sequences were available from Mafia Island (as H. mariae and H. viridiflavus mariae) and Mtai Forest, Tanzania; Arabuko Sokoke, Kenya (as H. viridiflavus mariae and H. v. rubripes). The minimum uncorrected p- distances from other clades were 4.0–6.5% (Table 2). The sample occurs in coastal Tanzania and Kenya (Fig. 9C). The localities of taxa now regarded as junior synonyms of H. mariae are listed in Table 3. The most common pattern elements in the descriptions of the types included in H. mariae are various markings on the head such as black-ringed nostrils (50%), followed by a dark lateral band (33%). The percentages are not additive, as the pattern elements may be shared. Advertisement calls from Mikumi, Tanzania (no vouchers), show a series of brief notes, with an emphasised frequency of about 3200 Hz, and a note rate of 5 s-1 (Schiøtz 1975, 1999 as H. v. mariae). This is similar to another call (no voucher) from the same locality (Pickersgill 2007, as H. mariae). A call from Kingupira, Tanzania (no voucher) has an emphasised frequency of 2750 Hz (Schiøtz 1975, as H. marginatus), similar to calls from Karonga, Malawi (no vouchers) (Pickersgill, 2007, as H. nyassae) and Bububu, Zanzibar (no vouchers) (Pickersgill 2007, as H. renschi)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 339-340, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Loveridge, A. (1936) Scientific results of an expedition to rain forest regions in eastern Africa. VII. Amphibians. Bulletin of the Museum of Comparative Zoology, 79, 369 - 430.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Parker, H. W. (1937) Reptiles and amphibians from the Mafia Archipelago. Annals and Magazine of Natural History, Series 10, 20, 629 - 632. https: // doi. org / 10.1080 / 00222933708655396","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp."]}

Hyperolius dintelmanni Lötters & Schmitz, 2004 Hyperolius dintelmanni Lötters & Schmitz, 2004. ZFMK 67871 (Table 3), “ From the Edib Hills (ca. 1,200 m above sea level), Bakossi Mountains, South-West Cameroon (4° 57′ N, 9° 39′ E).” Colour in life, phase J: back olive to translucent dark green extending to limbs, with a pale green hourglass pattern. Phase F: back dark brown to black with greenish spots and bright red areas on upper legs, feet and hands. The dorsal surface has a goldenyellow dense marbling. The females may be brown with reddish speckles (Portik et al. 2016). Sequences were available from Edib Village, the type locality, Manjo, Mt Kupe, Mantem, Southwest, Littoral, West, all in Cameroon. The minimum uncorrected p- distances from other clades were 1.9–6.0% (Table 2). The clade was identified following Bell et al. (2017). It is known from eastern Cameroon (Fig. 10D). The most common pattern element in the description of the type of H. dintelmanni is dorsal marbling (100%), followed by green spots (50%) or a red speckle (50%). The percentages are not additive, as the pattern elements may be shared. The advertisement call is unknown., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 343, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Lotters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west- Cameroon. Bonner zoologische Beitrage, 52, 149 - 154.","Portik, D. M., Jongsma, G. F. M., Kouete, M. T., Scheinberg, L. A., Freiermuth, B., Tapondjou, W. P. & Blackburn, D. C. (2016) A survey of amphibians and reptiles in the foothills of Mount Kupe, Cameroon. Amphibian & Reptile Conservation, 10 [Special Section], e 131, 37 - 67.","Bell, R. C., Parra, J. L., Badjedjea, G., Barej, M. F., Blackburn, D. C., Burger, M., Channing, A., Dehling, J. M., Greenbaum, E., Gvozdik, V., Kielgast, J., Kusamba, C., Lotters, S., McLaughlin, P. J., Nagy, Z. T., Rodel, M. - O., Portik, D. M., Stuart, B. L., VanDerWal, J., Zassi-Boulou, A. G. & Zamudio, K. R. (2017) Idiosyncratic responses to climate-driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands. Molecular Ecology, 26, 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260"]}

Hyperolius spatzi Ahl, 1931 Hyperolius spatzi Ahl, 1931. Lectotype ZMB 32602, “Bakel-Kidira (oberes Senegalgebiet)” Senegal (Table 3). Colour in alcohol: Above chalk-white or sometimes with fine dark brown stippling. Three sequences were available, all identified as H. spatzi. The minimum uncorrected p- distances from other clades were 3.4–6.5% (Table 2). The sequences all come from Senegal (Fig. 10C). Rödel et al. (2010) showed that H. spatzi was distinct from H. nitidulus using genetics, morphology and advertisement calls. The clade is identified as H. spatzi as the sequences derive from material collected by Rödel et al. (2010). This species is known from Gambia and Senegal, and its distribution was discussed by Rödel et al. (2010). The pattern elements in the descriptions of the type of H. spatzi are a uniform back (100%), or fine brown stipple (100%). The percentages are not additive, as the pattern elements may be shared. The advertisement call (no voucher) is described as ‘a single, pure, metallic and very loud tone’. Mean call length is 0.08 s, at a mean emphasised frequency of 2638 Hz (Rödel et al. 2010)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 342-343, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Rodel, M. - O., Sandberger, L., Penner, J., Man, Y. & Hillers, A. (2010) The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae). Bonn zoological Bulletin, 57, 177 - 188."]}

Hyperolius bangwae Perret, 1966 Hyperolius nitidulus bangwae Perret, 1966. MHNG 1039.2 “Bangwa”, Cameroon (Table 3). Back beige or brown sometimes with a dark lateral line, or with dark speckles. This clade consists of 50 sequences from Gabon, Equatorial Guinea, Republic of Congo and Cameroon (Fig. 9A), all reported in GenBank as Hyperolus tuberculatus. The sequence reported in Veith et al. (2009) as Hyperolius tuberculatus Complex B forms part of this clade. The uncorrected p- distances from other clades were 1.9–6.3% (Table 2). Two species occur at Bangoua in western Cameroon, H. nitidulus and a taxon previously identified as H. tuberculatus. Perret (1966) described Hyperolius nitidulus bangwae from Bangoua, and I remove it from the synonomy of H. nitidulus and regard it as a full species. The most common pattern elements in the descriptions of the type of H. bangwae are a dark lateral line (100%), and dark dorsal speckling (100%). An advertisement call from Monts Cristal (ZFMK 73143) consists of a series of brief notes at a mean note rate of 9.5 s- 1 with a mean emphasised frequency of 2271 Hz (Lötters et al. 2004, as H. tuberculatus). A specimen from the same locality was sequenced (AY329321) and identified as H. bangwae. The advertisement call attributed to H. tuberculatus from Binguéla, Cameroon (Schiøtz 1999) is within the range of H. bangwae as here envisaged, and is regarded as a call of H. bangwae. The call consists of a series of brief notes, about 13 s-1, with an emphasised frequency of 1900 Hz. This is quite different from the call of H. tuberculatus from Río Muni, Equatorial Guinea (no vouchers), as shown by Bosch et al. (2000). A call from Olemb, Cameroon (as H. tuberculatus) (no voucher) that consists of a series of simple notes with a note rate of 14 s-1, and an emphasised frequency of 3000 Hz (Amiet & Goutte 2017) is provisionally regarded as that of H. bangwae, although the call of H. sp. 1 is unknown., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 338, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie, 93, 289 - 464.","Veith, M., Kosuch, J., Rodel, M. - O., Hillers, A., Schmitz, A., Burger, M. & Lotters, S. (2009) Multiple evolution of sexual dichromatism in African reed frogs. Molecular Phylogenetics and Evolution, 51, 388 - 393. https: // doi. org / 10.1016 / j. ympev. 2008.12.022","Lotters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west- Cameroon. Bonner zoologische Beitrage, 52, 149 - 154.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Bosch, J., De la Riva, I. & Marquez, R. (2000) Advertisement calls of seven species of hyperoliid frogs, from Equatorial Guinea. Amphibia-Reptilia, 21, 246 - 255.","Amiet, J. - L. & Goutte, S. (2017) Chants d'Amphibiens du Cameroun. J. - L. Amiet & Editions Petit Genies, Nyons, 280 pp."]}

Hyperolius noblei Ahl, 1931 Hyperolius noblei Ahl, 1931. ZMB 85765, “Kilwa, Deutsch-Ostafrika ” Tanzania (Table 3). Colour in alcohol: Uniform yellow-brown. A silver-white stripe runs from the side of the snout, over the upper eyelid to form a larger mark behind the eye. Sequences were available from Nhica and Palma, Mozambique (as Hyperolius cf. marmoratus), and Nhica do Rovuma (as Hyperolius marmoratus taeniatus), Mozambique. The two sequences from Nhica (MG829620 and MK509512) were derived from the same specimen (MNHNP 2010.185). The minimum uncorrected p- distances from other clades were 4.2–6.9% (Table 2). Presently this species is only known from southern coastal Tanzania and northern coastal Mozambique (Fig. 10A). The nearest described taxon to this clade is Hyperolius noblei Ahl, 1931. The description matches the specimens collected and sequenced from northern Mozambique (Table 1). The pattern elements in the description of the type of H. noblei are a uniform back, with a transverse silver stripe forming a large mark behind the eye. The advertisement call (no voucher) was recorded on the Afungi Peninsula at Palma, Mozambique (10.8456°S, 40.4732°E). The call is a long (0.08 s) whistle, frequency modulated, with the emphasised frequency at 3500 Hz. A typical call starts at 3000 Hz, and rises steeply in pitch to 3595 Hz in 0.03 s, then maintains that pitch for 0.05 s. The emphasised frequency varies in the population from 3500 Hz to 4565 Hz., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 340, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132."]}

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., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 325-328, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Hewitt, J. (1937) A Guide to the Vertebrate Fauna of the Eastern Cape Province, South Africa. Part II. Reptiles, Amphibians, and Freshwater Fishes. Albany Museum, Grahamstown, 141 pp., 34 pls.","Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London, 256 pp., 16 plates.","Loveridge, A. (1941) South African frogs of the genus Hyperolius in the Museum of Comparative Zoology, Cambridge, Massachusetts. Annals of the Transvaal Museum, 20, 283 - 291.","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. F. (1985) Hyperoliidae. In: Frost, D. R. (Ed.), Amphibian Species of the World: A Taxonomic and Geographical Reference. Association of Systematics Collections and Allen Press, Lawrence, Kansas, pp. 203 - 233.","Poynton, J. C. & Broadley, D. G. (1987) Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae, Annals of the Natal Museum, 28, 161 - 229.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp.","Perret, J. - L. (1976) Revision des amphibiens africains et principalement des types, conserves au Musee Bocage de Lisbonne. Arquivos do Museu Bocage, Lisboa, Serie 2 6, 15 - 34.","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Laurent, R. F. (1957) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National de l'Upemba. Mission G. F. de Witte (1946 - 1949), 42, 3 - 47.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Bishop, P. J. (2004) Hyperolius marmoratus Rapp, 1842. In: Minter, L. R., Burger, M., Harrison, J. A., Braack, H. H., Bishop, P. J. & Kloepfer, D. (Eds.), Atlas and Red Data Book of the Frogs of South Africa, Lesotho and Swaziland. Smithsonian Institution - Man and Biosphere Biodiversity Program, Washington, D. C., pp. 141 - 143.","Tolley, K. A., Davies, S. J. & Chown, S. L. (2008) Deconstructing a controversial local range expansion: conservation biogeography of the painted reed frog (Hyperolius marmoratus) in South Africa. Diversity and Distributions, 14, 400 - 411. https: // doi. org / 10.1111 / j. 1472 - 4642.2007.00428. x"]}

Hyperolius goetzei Ahl, 1931 Hyperolius goetzei Ahl, 1931. ZMB 53181, “ Uhehe, Massai-Steppe” (= Iringa), Tanzania. Colour in alcohol: Above bright grey-red, with large dark brown marbling on the side of the head and body which has darker margins. Hyperolius viridiflavus reesi Schiøtz, 1982. ZMUC R77205, Magombero Forest, Tanzania. The type was collected close to the localities of the sequenced material, and this form is known to be widely distributed on the central Tanzania highlands (Schiøtz, 1999). Phase J colour in life: back silvery-brown with a darker pattern of spots and an irregular hourglass pattern. Sides of head and body dark, without a subdermal dark lateral streak. Phase F: Back yellow to light green, flanks with a conspicuous subdermal lateral streak, overlain by 2–3 yellow vertical stripes. Sequences were available from Kibasira Swamp and Mang’ula, (close to Magombero Forest) Tanzania (Fig. 10B). The minimum uncorrected p- distances from other clades were 4.0–7.0% (Table 2). Poynton (1977) recognised Hyperolius sp. from central Tanzania with lateral black markings that show through the skin, which Schiøtz (1982) described as H. v. reesi. Poynton later (1991) recognised Hyperolius reesi as a good species. The dark lateral band is common in other species, as are yellow transverse stripes in some animals. However, Ahl (1931) described Hyperolius goetzei from Iringa in central Tanzania, which Schiøtz (1999) illustrated in colour from Dabaga near Iringa. The back is uniform with mottled sides. This species is known from central Tanzania (Schiøtz 1999). The localities of the type and taxon regarded as a junior synonym are listed in Table 3. The most common pattern elements in the descriptions of the types included in H. goetzei are dorsal marbling (67%), followed by lateral marbling (33%), small dark spots (33%), or a dark lateral band (33%). The percentages are not additive, as the pattern elements may be shared. The advertisement call (no voucher) is a series of frequency modulated whistles or clicks with a mean length of 0.04 s and an emphasised frequency of 2239 Hz (Iringa, Tanzania)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 340-341, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Schiotz, A. (1982) Two new Hyperolius (Anura) from Tanzania. Steenstrupia, 8, 269 - 276.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Poynton, J. C. (1977) A new Bufo and associated Amphibia from southern Tanzania. Annals of the Natal Museum, 23, 37 - 41."]}

Hyperolius glandicolor Peters, 1878 Hyperolius glandicolor Peters, 1878. ZMB 9299, ZMB 77768, Taita, Kenya. Brown to yellow background, covered densely with very small dark dots. Hyperolius striolatus Peters, 1882. ZMB 9300, Taita, Kenya. Synonymy by Laurent (1961). Above rust coloured with black dots and stripes Megalixalus pantherinus Steindachner, 1891. NHMW 3734, “Leikipia” (Laikipia), Kenya. Synonymy by Wieczorek et al. (2000). Yellow with panther-like black markings on the back and behind the head. Rappia Ferniquei Mocquard, 1902. MNHNP 1902.184, “de la rivière Atchi, l’altitude de 1,450 mètres” Athi River, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Beige-grey above with a slight brown tint from the nostril to the arm, many dark speckles over the dorsal surface and limbs. Rappia symetrica Mocquard, 1902. MNHNP 1924.30–34. “de la rivière Atchi, l’altitude de 1,450 mètres” Athi River, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Above a very pale brown, with a black T shaped mark between the eyes. Many speckles along the midline. A dark band from the nostril, over the eye to the arm. A second mark, sometimes paired, extends obliquely towards the vent. These marks become gradually pink towards the ventral surface. A third irregular mark over the vent, edged with white. Rappia platyrhinus Procter, 1920. BMNH 1947.2.9.46, “Nairobi”, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). “Uniformly brownish-grey above, finely speckled with black pigment cells which are slightly concentrated along canthus rostralis” (Proctor 1920). Hyperolius albolabris Ahl, 1931. ZMB 58748, “Kwa Buosch in Süd Kavirondo”, south-western Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Above pale pink, with numerous dark pink or violet-brown flecks and stripes, usually arranged longitudinally, and with indistinct margins. A wide white stripe above the upper lip, extending back to the flank, and separating the colour of the upper and lower surfaces. Hyperolius asper Ahl, 1931. ZMB 4807, Nairobi, Kenya. Placed in the synonymy of H. glandicolor by Wieczorek et al. (2000). Colour in alcohol: Pale grey above and on the sides of the head, covered with many very small spots resembling a spray of ink. Hyperolius bergeri Ahl, 1931. ZMB but type not found (Tillack et al. 2021). “Guaso Narok (Englisch-Ostafrika)” Uaso Narok, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above pale yellow-green, with distinct blue marbling on the back and tibia. Upper thigh dark brown (red in life?) with a thin yellow-green longitudinal stripe. Hyperolius coeruleopunctatus Ahl, 1931. ZMB 36115, “Nairobi”, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above pale grey, with many small, round blue-black dots (bright blue in life), resembling sprayed ink. Hyperolius pulchromarmoratus Ahl, 1931. ZMB 77751, “ Britisch-Ostafrika ” (Kenya). Locality obscure. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above pale yellow on back, head, and upper surfaces of limbs, with thin sulfur-yellow and blue marbling. In between irregular large, dark brown markings of different sizes. Hyperolius scheffleri Ahl, 1931. ZMB 85759, “Kibwezi, Britisch-Ostafrika ” Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above uniform pale yellow. Thigh dark, probably red in life. Hyperolius marmoratus ommatostictus Laurent, 1951. MNHNP 1912.383 “Kibonoto, Kilimandjaro ” (Kibongoto), Tanzania. Synonymy by Wieczorek et al. (2000). Colour in alcohol: The back pattern consists of round dark brown eye-shaped markings, with a red-brown centre surrounded by a pale zone. Hyperolius marmoratus campylogrammus Laurent, 1951. MNHNP 1924.28, “Bura, alt 1050 m ” (Taita Hills). The H. marmoratus clade does not occur as far north as Kenya. The type occurs close to sequenced material of H. glandicolor from Taita. A transverse band between the eyes, with three longitudinal bands, the lateral bands curved inwards. Often lumbar spots present. Males with uniform backs. Hyperolius viridiflavus pitmani Laurent, 1952. Type unknown, “Lac Bunyoni”, Uganda. Synonymy by Wieczorek et al. (2000). Back with many small yellow spots. Hyperolius sheldricki Duff-MacKay and Schiøtz, 1971. NMK A/366/3, “north of Aruba Dam, Tsavo National Park (East) 3° 19’ S, 30° 54’ E ”, Kenya. It is a typical member of the H. viridiflavus group (Duff-MacKay and Schiøtz 1971). Sequences from material collected in the field as H. sheldricki are embedded in the clade identified as H. glandicolor. The back is light brown, with two dark brown stripes along the midline, joined over the snout. Other thin dark brown lines run from the eye back to the flank, and a pair of interrupted dorso-lateral lines, in places with a pale centre. Hyperolius viridiflavus ngorongoriensis Schiøtz, 1975. ZMUC R079324, Ngorongoro Crater Rim, Tanzania. Synonymy by Wieczorek et al. (2000). Dark brown stripes on a light brown background. Dark spots scattered evenly on the back and upper surfaces of the limbs. Ventral surface of femur and tibia maroon in life, pink in alcohol. Hyperolius orkarkarri Drewes, 1997. CAS 186087, “ Tanzania: Mara Region: Serengeti Dist: Serengeti National Park: Moru Kopjes, on Gong Rock. 02° 43.05 S; 34° 47.6 E at an elevation of 1585 m.” Synonymy by Channing & Howell (2006). Back grey, with a broad white vertebral band. This is interrupted by a dark triangle between the eyes, pointing backwards, and three pairs of lateral extensions, one from the scapular region to the eye, a second medio-dorsal saddle, and a posterior pair from the pelvic region. Hyperolius goetzei Ahl, 1931 was removed from the synonymy of H. glandicolor and elevated to a full species below. This clade consists of 20 sequences from Kenya, Tanzania, Uganda and Rwanda. The minimum uncorrected pdistances from other clades were 2.8–5.9% (Table 2). The sequences were reported in GenBank as Hyperolius glandicolor, H. goetzei, H. viridiflavus, H. v. ferniquei, H. v. glandicolor, H. v. ngorongoriensis, H. v. pantherinus, and H. v. pitmani. This clade occurs in Rwanda, Uganda, northern Tanzania and southern Kenya (Fig. 8B). The clade distribution closely matches the distribution of types that are regarded as junior synonyms, and sequences occur at the type locality. The localities of the type and taxa now regarded as junior synonyms of H. glandicolor are listed in Table 3. The most common pattern element in the descriptions of the types included in H. glandicolor is dorsal marbling (39%), followed by speckle or small spots that may be black or blue (33%). The percentages are not additive, as the pattern elements may be shared. An advertisement call from Kakoneni, Kenya (no voucher) shows individual notes with a length of about 0.1 s, and an emphasised frequency of 2800–3150 Hz (Schiøtz 1975, 1999, as H. sheldricki). A call from Arusha National Park in Tanzania (no voucher) shows brief whistles with frequency modulation, with a mean emphasised frequency of 2200 Hz (Schiøtz 1975). This is similar to calls from Kisessa, Tanzania (no vouchers), as H. goetzei; Arusha, Tanzania, as H. ommatostictus and Lake Naivasha, Kenya, as H. ferniquei; Nyahuru, Kenya, as H. pantherinus (Pickersgill, 2007). An advertisement call consisting of a brief note with an emphasised frequency of 1950 Hz was recorded at Lake Bunyoni, Uganda (no voucher), as H. v. pitmani and at Kabale, Uganda (no voucher) as H. pitmani (Pickersgill 2007). This is similar to calls from Mwanza (no vouchers) in Tanzania that have an emphasised frequency of 2800 Hz (Schiøtz 1975, 1999, Pickersgill 2007, as H. mwanzae) and Kisiizi, Uganda (no vouchers) (Pickersgill 2007, as H. variabilis)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 336-337, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Duff-Mackay, A. & Schiotz, A. (1971) A new Hyperolius (Amphibia Anura) from Kenya. Journal of the East Africa Natural History Society and National Museum, 29, 1 - 3.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, 418 pp.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp."]}

Hyperolius viridiflavus (Duméril & Bibron, 1841) Eucnemis viridi-flavus Duméril & Bibron, 1841. Syntypes MNHNP 412, 4599, NHMW 22896, “Abyssinie”. Adwa, Ethiopia. Back green with yellow spots, limbs dark pink, ventral surfaces yellow with small red spots. Rappia bayoni Boulenger, 1911. Syntypes MSNG and BMNH 1947.2.9.32–36, Entebbe, Busso, Bululu, Jinja and Kabulmuliro, Uganda. Synonymy by Laurent (1950). Pale brown back, with a dark lateral band edged with white, sometimes with a similar colour vertebral band, and a similar band along the outer limb surfaces, which are usually brown or dotted with brown. Hyperolius kwidjwiensis Ahl, 1931. ZMB 52449, “ Insel Kwidjwi im Kivu-See” Idjwi Island, Lake Kivu, DRC. Synonymy by Laurent (1952). Colour in alcohol: Back pale yellow with a thin brown band running from the nostrils to the groin. This band becomes the colour of the back along the body, leaving two thin dark margins running backwards from the level of the eye. Tibia the same colour as the body, with a darker brown stripe. Hyperolius variabilis Ahl, 1931. ZMB 36122, “Deutsch-Ost-Afrika”. Bukoba, Tanzania. Synonymy by Schiøtz (1975). Colour in alcohol: Back silvery grey to white, with highly variable red-brown or black-brown markings. These are reproduced here as part of Fig. 3, after Ahl (1931, fig. 173), drawn from Tornier (1896, pl. 4) (Tillack et al. 2021). Ventral surface pale with reddish spots. Hyperolius punctatissimus Ahl, 1931. ZMB 39013, “Bukoba”, Tanzania. Synonymy by Laurent (1952). Colour in alcohol: Back pale grey with many small dark spots or flecks, that may cover the back to produce a uniform dark grey colour. A pale line runs from the upper lip along the side of the body to the upper leg, demarcating the dark back. Limbs similarly patterned. Hyperolius kandti Ahl, 1931. ZMB 46526, “Kivu-See”. Lake Kivu, DRC. Synonymy by Laurent (1947). Colour in alcohol: Uniform beige back and sides. a thin dark line from nostril to eye that broadens into a lateral brown band to the flank. Tibia with dark flecks. Hyperolius schubotzi Ahl, 1931. Type unknown. “Kissenji”, Rwanda. Synonymy by Laurent (1947). Colour in alcohol: Back grey-blue, covered densely with small dark spots or vermiculations. Upper surfaces of limbs as for the back. Throat pale with small round spots, belly yellow, hidden surfaces of legs red. Hyperolius wettsteini Ahl, 1931. ZMB 36103, “Bukoba”, Tanzania. Synonymy by Laurent (1952). Colour in alcohol: Back and sides violet-grey, with large dark violet flecks arranged in longitudinal rows. Ventral surfaces pale yellow, with brown stipples on the throat. Hyperolius phrynoderma Ahl, 1931. Syntypes ZMB 39000 and ZMB 77734 –36, “Zentrales Deutsch-Ostafrika ”, corrected to north-eastern Tanzania (Tillack et al. 2021). Provisionally placed as a synonym of H. viridiflavus (Tillack et al. 2021). Colour in alcohol: Pale yellow all over. Hyperolius karissimbiensis Ahl, 1931. ZMB 46525, “Bambusurwald und Waldwiesen ca. 2400 m hoch, beim Dorf des Mhcabu Gahama am Karissimbi”, Mhcabu Gahama village on Mt Karissimbi, Rwanda. Synonymy by Laurent (1952). Colour in alcohol: Back pale grey with fine blue stipple. A thin dark line runs from snout tip, below the canthus rostralis to under the eye. It starts again on the flank and runs to the vent. The dark line is bordered by white. A dark vertebral stripe. Dark longitudinal line run from between the eyes, interrupted near the armpit, continuing to the vent. These lines have pale borders. Upper surfaces of limbs yellowish with a dark longitudinal line. Hyperolius mohasicus Ahl, 1931. ZMB 36094, “Mohasi-See, Ruanda ” Muhasi Lake, Rwanda. Synonymy by Schiøtz (1975). Colour in alcohol: Back, sides and upper surfaces of limbs white with many moderately large, rounded or long reddish marks, arranged in longitudinal rows. Small bright red-brown warts present. Hyperolius multicolor Ahl, 1931. ZMB 39002, 39005, 74953–74956. “Bambusurwald und Waldwiesen ca. 2400 m hoch, beim Dorf des Mhcabu Gahama am Karissimbi” Mhcabu Gahama village on Mt Karisimbi, Rwanda. Synonymy with Hyperolius karissimbiensis by Laurent (1950). Colour in alcohol: Head and shoulder yellowish to reddish, back red-brown to brown. A dark brown quadratic mark between the eyes, a thin blue vertebral stripe from the shoulder blades to the sacrum. A large dark brown lateral marking between the limbs reaches up to the level of the sacral vertebrae, with a wide black border. A white to yellow regular stripe runs from the level of the sacrum to the vent. Hyperolius macrodactylus Ahl, 1931. ZMB 39100, “Kivu-See” Lake Kivu, DRC. Synonymy by Laurent (1947). Colour in alcohol: Back and sides bright yellow, with many irregular longitudinal dark brown markings forming a marbling. The upper surfaces of the limbs similarly coloured. Hyperolius flavoguttatus Ahl, 1931. ZMB 39011, “Bukoba”, Tanzania. Synonymy by Laurent (1952). Colour in alcohol: Dark brown back with large numbers of round spots more or less in lines. These were bright yellow in life. Upper surfaces of limbs similarly coloured. Hyperolius monticola Ahl, 1931. ZMB 39010, “Niansa, Ruanda, 1500 m hoch” Nyabisundu, Nyanza District, Rwanda. Synonymy with Hyperolius variabilis by Laurent (1943). Colour in alcohol: White above, with four longitudinal lines of small brown markings. Upper surfaces of limbs similar. Hyperolius oculatus Ahl, 1931. ZMB 58570, Balaibo, DRC. Synonymy with H. viridiflavus pachydermus by Laurent (1952). Colour in alcohol: Grey-brown above, with many irregularly spaced pale brown spots with red-brown centres. Hidden surfaces of limbs bright red. In life the back is green with red spots with yellow borders. Hyperolius stuhlmanni Ahl, 1931. ZMB 13008, “Vitschumbi, Südende des Albert-Eduard-See’s” Witschumbi, Lake Edward, DRC. Synonymy by Schiøtz (1975). Colour in alcohol: Pale above, on the sides and upper surfaces of limbs, with many irregular short dark wormlike lines that divide and intertwine. There are also some paler lines in between the darker markings. Hyperolius irregularis Ahl, 1931. ZMB 36105, ZMB 75606, “Mohasi-See, Ruanda ” Muhasi Lake, Rwanda. Synonymy by Schiøtz (1975). Colour in alcohol: Silver-grey above, on the sides and on upper surfaces of limbs, with more or less large, rounded, or elongated dark to red brown markings. Hyperolius koehli Ahl, 1931. ZMB 26089, “Kissenji, Deutsch-Ost-Afrika”, Kissenji, Rwanda. Synonymy by Laurent (1950). Colour in alcohol: Uniform dark brown above. Hyperolius taeniolatus Laurent, 1940. MRAC 39471, “Lacs Mokoto” Mokoto Lake, DRC. Preoccupied by Hyperolius taeniolatus Bocage, 1895. Synonymy with H. viridiflavus bayoni by Laurent (1950). Colour in alcohol: Upper surfaces dark, with hidden surfaces pink. Head and tibia with pink vermiculation, oriented longitudinally. Tarsus and toes pink with black flecks. Ventral surfaces pale with black stipple. Hyperolius ornatus Laurent, 1940. MRAC 3836, Rutshuru, DRC. Synonymy by Laurent (1972). Colour in alcohol: Back dark with pale reticulation and red dots along the flanks. Upper limbs with large dark spots. Hyperolius schubotzi ruandensis Laurent, 1943. MRAC 38371, “Nyakatare, Ruanda ” (Rwanda). Synonymy by Laurent (1983). Colour patterns very diverse, with dark irregular marbling arranged longitudinally. Hyperolius destefanii Scortecci, 1943. Type lost (Blackburn & Scali 2014). “Nargi”, Ethiopia. Synonymy by Schiøtz (1999). Colour description from Schiøtz (1999): “Dorsum green to bluish green, unmarked or with small dark blue or black dots and freckles; posterior face of thigh iridescent pale blue to purple with blackish speckling”. Hyperolius karissimbiensis xanthogrammus Laurent, 1950. MRAC types unknown. Paratypes in LACM. “Cyumbe, Marais de la Mulindi (alt. 1.950 m.), Territoir de Biumba, Ruanda.” Mulindi Swamp, Rwanda. Synonymy by Laurent (1952). Back pale beige to red-brown, with fine yellow to orange dots and vermiculation arranged in three longitudinal lines. Hyperolius karissimbiensis françoisi Laurent, 1950. Type in MRAC, paratypes in LACM, “ Bururi alt 1900–1950m, arrondissement de Bururi ”, Bururi, Burundi. Synonymy by Laurent (1952). Back with a pale vertebral stripe, two lateral-dorsal parallel lines and another between the eyes. An oblique line on the tibia. Hyperolius karissimbiensis intermedius Laurent, 1950. MRAC B 80021.3242 “Marais Rugushu, Kayanza, alt 1900m, terr. de Ngozi ”, Rugushu Swamp, Burundi. Synonymy by Laurent (1952). The pattern is intermediate between H. karissimbiensis xanthogrammus and H. k. françoisi. A pale-yellow interorbital bar and a transverse yellow bar over the scapula, in addition to the patterns described above. Hyperolius viridiflavus hybridus Laurent, 1951. MRAC B 80021.3865 – 3883, “Etang de la Mukura, alt. 1700 m. près d’Astrida” Mukura pond (= Huye), Rwanda. Colouration variable, with a red thigh. Hyperolius viridiflavus mwanzae Schiøtz, 1975. Swamp north of Mwanza, Tanzania. Phase F translucent green with or without bluish spots. No red pigmentation. Hyperolius burgeoni De Witte, 1921 and Rappia rossii Calabresi, 1925 were provisionally removed from the synonymy of H. viridiflavus as both occur in northern DRC where the H. viridiflavus clade has not been recorded, and placed in the synonymy of H. parallelus. Hyperolius viridiflavus angeli Laurent, 1951 from the Central African Republic was removed from the synonymy of H. viridiflavus which does not occur north of the Congo Basin and placed in the synonymy of Hyperolius nitidulus. Altogether 28 sequences were available from Uganda, northern Tanzania, the Kenyan highlands, Burundi, Rwanda, and Ethiopia. The minimum uncorrected p- distances from other clades were 4.2–7.6% (Table 2). The sequences were reported in GenBank as Hyperolius viridiflavus, H. v. viridiflavus, H. v. variabilis, H. v. destefanii, and H. v. bayoni. The earliest species description in this complex is Hyperolius viridiflavus Duméril & Bibron, 1841, from Adwa in Ethiopia. Only one clade has records from Ethiopia, and this clade is assigned to H. viridiflavus (Fig. 4). See Table 3 for localities of synonyms. The map reflects the distribution in Ethiopia reported in Largen & Spawls (2010). The most common pattern elements in the descriptions of the types included in H. viridiflavus are small blue, black or red dark spots (26%), with the next most common elements being marbling on the back (22%), a dark lateral band (19%), and various markings arranged in longitudinal rows (19%). The percentages are not additive, as some individuals share pattern elements. Many of the descriptions were based on preserved material, and a large number of colour patterns is expected when fresh material is examined. Figure 3 illustrates some of the patterns recognised by Laurent (1983) and Ahl (1931) for material they described, here recognised as H. viridiflavus. This sample indicates the problems of identification based on colour patterns. Lötters et al. (2004) illustrate a call (as H. viridiflavus) from Kakamega Forest, Kenya with a distinct pulsed structure and an emphasised frequency of 2971 Hz. A specimen from the same locality was sequenced (AY323920) (Lötters et al. 2004) and found within the H. viridiflavus clade. An advertisement call from Kitale, Kenya consists of a brief whistle with an emphasised frequency of 2600–3000 Hz (Schiøtz 1975 as H. v. viridiflavus). The advertisement call of a specimen from Sango Bay, Uganda, consists of a very brief note with an emphasised frequency of 2400 Hz (Schiøtz 1999)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 320-325, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Laurent, R. (1950) Exploration du Parc National Albert. Mission G. F. de Witte (1933 - 1935). Genres Afrixalus et Hyperolius. Institut des Parcs Nationaux du Congo Belge, Brussels, pp. 14 - 116, 7 pls.","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Laurent, R. F. (1947) Sur quelques rhacophorides nouveaux de Parc National Albert appartenant aux genres Megalixalus et Hyperolius. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 23, 1 - 8.","Laurent, R. F. (1983) La superespece Hyperolius viridiflavus (Dumeril & Bibron, 1841) (Anura Hyperoliidae) en Afrique Centrale. Monitore Zoologico Italiano, New Series, Supplement 18, 1 - 93.","Tornier, G. (1897) Die Kriechthiere Deutsch-Ost-Afrikas. Dietrich Reimer, Berlin, 206 pp.","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Laurent, R. F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxieme Serie, 22, 1 - 125.","Blackburn, D. C. & Scali, S. (2014) An annotated catalog of the type specimens of Amphibia in the collection of the Museo Civico di Storia naturale, Milan, Italy. Herpetological Monographs, 28, 24 - 45. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00008","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Largen, M. J. & Spawls, S. (2010) The Amphibians and Reptiles of Ethiopia and Eritrea. Edition Chimaria, Frankfurt am Main, 693 pp.","Lotters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west- Cameroon. Bonner zoologische Beitrage, 52, 149 - 154."]}

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., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 328-330, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Laurent, R. F. (1983) La superespece Hyperolius viridiflavus (Dumeril & Bibron, 1841) (Anura Hyperoliidae) en Afrique Centrale. Monitore Zoologico Italiano, New Series, Supplement 18, 1 - 93.","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Laurent, R. F. (1965) The geographical variation of the frog Hyperolius marmoratus (family Hyperoliidae) in Rhodesia, Nyasaland and Tanganyika. Breviora, 219, 1 - 9.","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Laurent, R. (1948 \" 1947 \") Two new forms of the genus Hyperolius. Annals and Magazine of Natural History, Series 2, 14, 294 - 296.","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp."]}

The taxonomic validity of the East African reed frogs Hyperolius mitchelli Loveridge, 1953 and Hyperolius rubrovermiculatus Schiotz, 1975 has been questioned in the literature, with the latter havi...

The identification of biotic and abiotic factors that promote the diversification of clades across Africa and the Arabian Peninsula remains a difficult challenge. A variety of ecological and evolutionary processes can be driving such patterns, and clade-specific traits may also play a role in the evolution of these groups. Comparative evolutionary studies of particular clades, relying on a phylogenetic framework, can be used to investigate many of these topics. Beyond these mechanisms there are abiotic factors, such as geological events, that can drive vicariance and dispersal events for large sets of taxa. The investigation of historical biogeography in a comparative phylogenetic framework can be used to detect such patterns. My dissertation explores these topics using reptiles and amphibians as study systems, and I rely on the generation of molecular sequence data, phylogenetics, and the use of comparative phylogenetic methods to address a variety of questions. I provide the abstract for each chapter below.Chapter 1: The reproductive modes of anurans (frogs and toads) are the most diverse among all the terrestrial vertebrates, and a major challenge is identifying selective factors that promote the evolution or retention of reproductive modes across clades. Terrestrialized anuran breeding strategies, in which breeding is partially or completely independent of water, have evolved repeatedly from the plesiomorphic fully aquatic reproductive mode, a process which is thought to occur through a series of intermediate reproductive stages. A number of selective forces have been proposed for the evolution of terrestrialized reproductive traits, but factors such as water systems and co-evolution with particular ecomorphologies have not been investigated in a comparative phylogenetic framework. We examined these topics and the evolution of reproductive mode in Afrobatrachian frogs, an ecologically and reproductively diverse clade representing more than half of the total frog diversity found in Africa (~400 species). We find that direct development evolved twice independently from terrestrialized reproductive modes involving protected eggs or larvae, supporting the intermediate stages hypothesis. We detected correlated evolution in specific derived ecomorphologies and reproductive traits, including arboreality and arboreal oviposition, and fossoriality and subterranean oviposition. There is support for a link between lotic water systems and terrestrial oviposition, and between arboreal ecomorphology and the use of lentic water systems for reproduction. Our findings indicate clade-specific processes driving the evolution of reproductive modes in the Afrobatrachia, and suggest that other processes such as habitat filtering and co-evolution with ecomorphology may generate global patterns of anuran reproductive modes.Chapter 2: Custom sequence capture experiments are becoming an efficient approach for gathering large sets of orthologous markers with targeted levels of informativeness in non-model organisms. Transcriptome-based exon capture utilizes transcript sequences to design capture probes, often with the aid of a reference genome to identify intron-exon boundaries and exclude shorter exons (< 200 bp). Here, we test an alternative approach that directly uses transcript sequences for probe design, which are often composed of multiple exons of varying lengths. Based on a selection of 1,260 orthologous transcripts, we conducted sequence captures across multiple phylogenetic scales for frogs, including species up to ~100 million years divergent from the focal group. After several conservative filtering steps, we recovered a large phylogenomic data set consisting of sequence alignments for 1,047 of the 1,260 transcriptome-based loci (~630,000 bp) and a large quantity of highly variable regions flanking the exons in transcripts (~70,000 bp). We recovered high numbers of both shorter (< 100 bp) and longer exons (> 200 bp), with no major reduction in coverage towards the ends of exons. We observed significant differences in the performance of blocking oligos for target enrichment and non-target depletion during captures, and observed differences in PCR duplication rates that can be attributed to the number of individuals pooled for capture reactions. We explicitly tested the effects of phylogenetic distance on capture sensitivity, specificity, and missing data, and provide a baseline estimate of expectations for these metrics based on nuclear pairwise differences among samples. We provide recommendations for transcriptome-based exon capture design based on our results, and describe multiple pipelines for data assembly and analysis.Chapter 3: The anuran family Hyperoliidae is Africa’s most diverse group of frogs, with a distribution across sub-Saharan Africa (16 genera, over 200 species), Madagascar (1 genus, 11 species), and the Seychelles Islands (1 monotypic genus). Hyperoliids are found in a variety of habitats and altitudes, but exhibit the highest species richness and co-occurring species in forested regions, including lowland rainforests and montane forests. As many as 14 hyperoliid species have been recorded in near sympatry, and species show high turnover based on both elevational and habitat gradients. Hyperoliids exhibit extensive morphological diversity, ranging from semi-terrestrial forms, to walker-climbers, to largely arboreal forms, and display considerable variation in body size across species and between the sexes. A striking and unique feature of many hyperoliid species is their bright coloration and patterning, which can be highly variable and driven by the presence of color polymorphism or sexual dichromatism. Using phylogenomic data, I reconstructed the evolutionary relationships of the family and investigated the evolution of sexual size dimorphism and sexual dichromatism. My examination of the body sizes of hyperoliid species revealed this family largely exhibits female-biased SSD, and I find evidence for an isometric scaling relationship between body sizes of the sexes. Within the genus Hyperolius I found significant differences in the evolutionary trend of male and female body sizes, namely selection for a single optimal female body size and strong directional selection for males occurring in different subclades. I propose the functional relationship between female body size and arboreal oviposition may be opposing the selection for increased fecundity associated with larger female sizes. The body sizes of males may be the result of a combination of interspecific resource partitioning, division of acoustic space, and reproductive character displacement. Sexual dichromatism has evolved independently in two main clades of hyperoliids: 1) the common ancestor of the Malagasy and Seychelles Islands species (Heterixalus, Tachycnemis), and 2) the common ancestor of Hyperolius, Morerella, and Cryptothylax. A number of subsequent reversals to monochromatism occur in the genus Hyperolius and Heterixalus, with some cases of secondarily derived dichromatism. Chapter 4: The Red Sea has had a profound biogeographic effect on organisms with Afro-Asian distributions, resulting in complex patterns of admixture on the Arabian Peninsula. We investigate the phylogenetic affinities of a monitor lizard (Varanus yemenensis) restricted to the southwestern Arabian Peninsula by sequencing all African monitor species and several Asian monitor species for the mitochondrial gene ND2 and the nuclear marker RAG-1. We find evidence that V. yemenensis is of African origin, being most closely related to the white-throat monitor, V. albigularis, an African species complex distributed from the Horn of Africa to southern Africa. Using divergence-dating analyses, we investigate several biogeographic hypotheses to infer the likely mechanism of colonization of the Arabian Peninsula by this species. Our results reveal that both dispersal across a southern landbridge and overwater dispersal are potential explanations. The patterns observed in V. yemenensis are contrasted with other taxa having similar Afro-Arabian disjunct distributions to better understand the complex biogeographic history of this region.Chapter 5: The Arabian Peninsula is home to a unique fauna that has assembled and evolved throughout the course of major geophysical events, including the separation of the Arabian Plate from Africa and subsequent collision with Eurasia. Opportunities for faunal exchanges with particular continents occurred in temporally distinct periods, and the presence of African, Western Eurasian, and South Asian derived taxa on the Arabian Peninsula signifies the complexity of these historical biogeographic events. The six true toad species (family Bufonidae) endemic to Arabian Peninsula present a considerable taxonomic and biogeographic challenge because they are part of a global bufonid radiation, including several genera surrounding the Arabian Peninsula, and difficult to discriminate morphologically. As they could be derived from African, Western Eurasian, or South Asian toad groups, elucidating their evolutionary relationships has important implications for historical biogeography. Here, we analyze a global molecular data set of 243 bufonid lineages, with an emphasis on new sampling from the Horn of Africa, Western Eurasia, South Asia, and the Arabian Peninsula, to reconstruct the evolutionary relationships of the Arabian species. We produce a robust time-calibrated phylogeny to infer the biogeographic history of this group on and around the Arabian Peninsula. Our phylogenetic analyses indicate two of the endemic Arabian toad species, “Bufo” tihamicus and “Bufo” arabicus, evolved independently within the African genus Amietophrynus. We confirm the Arabian species Duttaphrynus dhufarensis is of South Asian origin, but do not find evidence for the Asian genus Duttaphrynus being present in the Horn of Africa, discrediting a previously proposed Asian bufonid dispersal event to Africa. We also do not find evidence of the African genus Amietophrynus occurring in South Asia, suggesting that unlike many other vertebrate taxa, toads have not used the Arabian Peninsula as a stepping-stone for trans-continental dispersal. Our divergence dating estimates strongly suggest the formation of the Red Sea drove simultaneous divergences between two of the Arabian species (A. tihamicus comb. nov. and A. arabicus comb. nov.) and their closest mainland African relatives in the Early Miocene. We estimate the divergence of D. dhufarensis with its closest South Asian relatives occurred in the mid to Late Miocene, suggesting the temporary or permanent land connections between the Arabian plate and Eurasia facilitated dispersal of this lineage to the Arabian Peninsula. The Arabian bufonid assemblage, despite being comparatively depauperate with respect to surrounding continents, exemplifies the faunal pattern of the Arabian Peninsula, namely being a complex admixture of African, Western Eurasian, and South Asian elements. The historical biogeographic patterns exhibited by Arabian toads and their allies are concordant with studies of other vertebrate taxa, building support for the role of major geological events in driving simultaneous vicariance and dispersal events around the Arabian Peninsula. Although many taxa or groups exhibiting disjunct Afro-Arabian distributions appear to have dispersed more recently from the Horn of Africa via a southern land bridge or overwater dispersal, both Amietophrynus tihamicus and A. arabicus likely represent true African relicts resulting from vicariance associated with the Red Sea formation, a pattern that so far is rare among the vertebrate species investigated.Chapter 6: We summarize the various geologic and climatic-based hypotheses influencing the diversification of Arabian taxa and highlight the role of an understudied biodiversity hotspot, the Arabian Hotspot Area (AHA), in driving patterns of endemism. We evaluate these biogeographic hypotheses and patterns utilizing Arabian amphibians as a study system, and contrast these results with a comprehensive literature review of other Arabian taxa. We investigated biogeographic hypotheses using a temporally explicit phylogenetic framework and by compiling divergence-dating estimates for all available extant Arabian taxa. To examine the effects of climate change, we generated distribution models for Arabian amphibians under current and historical climate regimes to identify historically stable refugial areas for montane and lowland species. We find a striking concordance of divergence times for taxa with similar biogeographic histories, providing strong support for hypotheses derived from geologic events. Most in-situ speciation events occurring on the Arabian Peninsula are concentrated in the late Miocene, with intraspecific divergences of species occurring mainly in the Plio-Pleistocene. Changes in connectivity between the Arabian Peninsula and surrounding continents throughout geologic time allows explicit temporal predictions to be made about biogeographic events for Arabian taxa derived from African, Asian, and Palearctic lineages. We find the AHA is in many ways similar to the montane systems of East Africa, and is characterized by habitat stability and the long-term persistence of localized lineages and the diversification of cyclically fragmented lineages. Historically stable refugial regions for amphibian species mainly occur within the AHA, demonstrating the importance of this region for the persistence of both montane and lowland species.

Afrixalus osorioi (Ferreira, 1906) Material (three specimens): One adult male, one adult female and one juvenile specimen (MHNCUP/ ANF 212–214, formerly UP-MHNFCP-017307; Fig. S6), collected from Quilombo [-9.33333º, 14.90000º, 744 m a.s.l.] Kwanza Norte Province, date unknown. Comments: MHNCUP/ANF 212–214 are the syntypes used by Ferreira (1906) in the description of Rappia osorioi, a species named in homage to Baltasar Osório (1855-1926), a naturalist from the Museu Bocage. Ceríaco et al. (2014b) analyzed these specimens in a review of the type specimens published by Ferreira (1904, 1906), confirming the species to be valid and found in northwestern Angola. Until very recently, the species had only been recorded from its type locality in Angola (Marques et al. 2018), but field surveys in Serra do Pingano, Uíge Province, uncovered additional records of the species for the country (Ernst et al. 2020; LMPC pers. obs. (Fig. 3))., Published as part of Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton's Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, pp. 1-80 in Zootaxa 5028 (1) on pages 11-12, DOI: 10.11646/zootaxa.5028.1.1, http://zenodo.org/record/5453967, {"references":["Ferreira, J. B. (1906) Algumas especies novas ou pouco conhecidas de amphibios e reptis de Angola (Coleccao Newton - 1903 - 1904). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (26), 159 - 171.","Ceriaco, L. M. P., Blackburn, D. C. Marques, M. P. & Calado, F. M. (2014 b) Catalogue of the amphibian and reptile type specimens of the Museu de Historia Natural da Universidade do Porto in Portugal, with some comments on problematic taxa. Alytes, 31, 13 - 36.","Ferreira, J. B. (1904) Reptis e amphibios de Angola da regiao ao norte do Quanza (Colleccao Newton 1903). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (25), 111 - 117.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501."]}

Francisco Newton’s zoological expedition to Angola undertaken between 1903 and 1906 is one of the least studied of the naturalist’s life. Only three major papers regarding the herpetofauna collected in this expedition have been published, and a significant part of the specimens remains unstudied since the 1900’s. Here we review the extant herpetological specimens of this expedition, present an updated taxonomic revision, and provide new insights on their taxonomic status. The extant collection is constituted by 329 specimens (155 amphibians and 174 reptiles), corresponding to 73 species, 39 genera and 22 families, and it is presently housed in the Museu de História Natural e da Ciência da Universidade do Porto (MHNC-UP).

Hyperolius bocagei Steindachner, 1867 Material (five specimens): One adult female and one adult male specimen (MHNCUP/ ANF 178–179, formerly UP-MHNFCP-017306), collected from Cacuaco [-8.78333º, 13.36667º, 6 m a.s.l.] Luanda Province, in 1903; one adult specimen (MHNCUP/ ANF 219, formerly UP-MHNFCP-017318; Fig. S16), collected from Bahia dos Tigres [= Baía dos Tigres] (in error) [-16.58559º, 11.82709º, 98 m a.s.l.] Namibe Province, in 1905; two adult specimens (MHNCUP/ ANF 338–339), collected from unknown locality, date unknown. Comments: Hyperolius bocagei is a species found in northeast and western Angola (Marques et al. 2018). Ferreira (1904) cites male and female specimens (no precise number) of “ Rappia bocagei (Stenid.) ” from “Cacuaco”, corresponding to specimens MHNCUP/ANF 178–179.Another specimen of this species was used by Ferreira (1906) to describe a new variety – R. bocagei var. maculata – from “Golungo Alto”. MHNCUP/ANF 219 might correspond to the cited specimen (see Ceríaco et al. 2014b). The label for specimen MHNCUP/ANF 219 indicates the locality “Bahia dos Tigres”, a desert area on the southern coast of Namibe Province. This is certainly a labelling error, as the habitat is not suitable for Hyperolius species due to its extreme aridity., Published as part of Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton's Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, pp. 1-80 in Zootaxa 5028 (1) on page 14, DOI: 10.11646/zootaxa.5028.1.1, http://zenodo.org/record/5453967, {"references":["Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501.","Ferreira, J. B. (1904) Reptis e amphibios de Angola da regiao ao norte do Quanza (Colleccao Newton 1903). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (25), 111 - 117.","Ferreira, J. B. (1906) Algumas especies novas ou pouco conhecidas de amphibios e reptis de Angola (Coleccao Newton - 1903 - 1904). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (26), 159 - 171.","Ceriaco, L. M. P., Blackburn, D. C. Marques, M. P. & Calado, F. M. (2014 b) Catalogue of the amphibian and reptile type specimens of the Museu de Historia Natural da Universidade do Porto in Portugal, with some comments on problematic taxa. Alytes, 31, 13 - 36."]}

Hyperolius adspersus Peters, 1877 Material (two specimens): Two adult female specimens (MHNCUP/ ANF 210–211, formerly UP-MHNFCP-017298 and UP-MHNFCP-017292; Fig. S7), collected from Cabiri [-8.92125º, 13.66654º, 31 m a.s.l.], Bengo Province, in 1903. Comments: Hyperolius adspersus was originally described from “Chinchoxo”, Cabinda Province (Peters 1877), and has been recorded from a few other localities in Bengo, Lunda-Sul and Moxico provinces (Marques et al. 2018), but its distribution extends northwest to southern Cameroon (Channing & Rödel 2019). The species has remained understudied and is a part of the Hyperolius nasutus Günther, 1865 group (Channing et al. 2013). The extant specimens were used by Ferreira (1904) as the type specimens for what he considered a new taxon, Rappia nobrei, named in homage to Augusto Nobre. However, after revision of the type series, Ceríaco et al. (2014b) suggested that Rappia [= Hyperolius] nobrei is a junior synonym H. adspersus., Published as part of Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton's Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, pp. 1-80 in Zootaxa 5028 (1) on page 12, DOI: 10.11646/zootaxa.5028.1.1, http://zenodo.org/record/5453967, {"references":["Peters, W. C. H. (1877) Ubersicht der Amphibien aus Chinchoxo (Westafrika), welche von der Afrikanischen Gesellschaft dem Berliner zoologischen Museum ubergeben sind. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1877, 611 - 621, 1 pl.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501.","Channing, A. & Rodel, M. - O. (2019) Field Guide to the Frogs & Other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.","Ferreira, J. B. (1904) Reptis e amphibios de Angola da regiao ao norte do Quanza (Colleccao Newton 1903). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (25), 111 - 117.","Ceriaco, L. M. P., Blackburn, D. C. Marques, M. P. & Calado, F. M. (2014 b) Catalogue of the amphibian and reptile type specimens of the Museu de Historia Natural da Universidade do Porto in Portugal, with some comments on problematic taxa. Alytes, 31, 13 - 36."]}

Hyperolius cinnamomeoventris Bocage, 1866 Material (six specimens): One adult specimen (MHNCUP/ ANF 176, formerly UP-MHNFCP-017313; Fig. S17), collected from Quilombo [-9.33333º, 14.90000º, 744 m a.s.l.] Kwanza Norte Province, in 1904; two adult male specimens (MHNCUP/ ANF 197–198, formerly UP-MHNFCP-094965), collected from unknown locality, date unknown; one adult specimen (MHNCUP/ ANF 220, formerly UP-MHNFCP-017314), collected from Bahia dos Tigres [= Baía dos Tigres] (in error) [-16.58559º, 11.82709º, 98 m a.s.l.] Namibe Province, in 1905; one adult specimen (MHNCUP/ ANF 294, formerly UP-MHNFCP-017309), collected from unknown location, date unknown; one adult specimen (MHNCUP/ ANF 344), collected from unknown locality, date unknown. Comments: Hyperolius cinnamomeoventris is a species found along the western coast of Angola and in the northern part of the country (Marques et al. 2018). Ferreira (1906) cites several specimens (without giving the precise number) from “Quilombo”, “Mupèpe” and “Rio Luinha”. Six specimens are extant in the collections, one of which matches the Quilombo locality (MHNCUP/ANF 176). The remaining most likely correspond to the “Mupepe” and “Rio Luinha” specimens, as the adult specimen (MHNCUP/ANF 220) is certainly not from “Bahia dos Tigres” as noted on the label. Specimen MHNCUP/ANF 294 was previously identified as “ Rappia cinctiventris ”. The cited localities are the only published records for the species in Kwanza Norte province (Marques et al. 2018)., Published as part of Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton's Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, pp. 1-80 in Zootaxa 5028 (1) on page 14, DOI: 10.11646/zootaxa.5028.1.1, http://zenodo.org/record/5453967, {"references":["Bocage, J. V. B. (1866) Reptiles nouveaux ou peu connus recueillis dans les possessions portugaises de l'Afrique occidentale, qui se trouvent au Museum de Lisbonne. Jornal de Sciencias Mathematicas, Physicas e Naturaes, I, 57 - 78.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501.","Ferreira, J. B. (1906) Algumas especies novas ou pouco conhecidas de amphibios e reptis de Angola (Coleccao Newton - 1903 - 1904). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (26), 159 - 171."]}

The African reedfrog taxon Alexteroon consists of only three described species with rather restricted geographical ranges. Although the assignment to a distinct genus is supported by multiple evidence, its position within the larger African hyperoliid radiation has been disputed. Available molecular data are scarce and the geographic records are few and scattered. The partially formalin fixed type series were previously not accessible to molecular analyses. This changed only very recently with the advancement of Next Generation Sequencing and ancient DNA techniques. Here we provide a reassessment of the current distribution and identity of all known species in this taxon based on (a) historical and new records, (b) morphological reanalyses of the type material and newly collected specimens from Angola and Gabon, and (c) newly established, nearly complete mitochondrial genome data from historical type and modern non-type material. We also present a molecular phylogeny (five mitochondrial loci 12S, 16S, ND1, ND2, COI, Cytb) for 78 sequences from 75 different species of Hyperolius retrieved from GenBank and 14 newly established Alexteroon sequences. We demonstrate that Alexteroon is more widely distributed than previously thought with records from northern Angola representing major southern range extensions. Results of the quantitative morphometric analyses show that the group has a rather conserved general body plan. Therefore qualitative phenotypic features observable in live specimens appear to be more useful for ad hoc species delimitation. We found Alexteroon to be nested within Hyperolius, corroborating previous findings. However, the combination of molecular data and consistent differences observed in morphology and ecology provide strong support for the distinctiveness of this evolutionary lineage within Hyperolius sensu lato. We therefore treat Alexteroon as a subgenus of Hyperolius and argue that the large and diverse genus Hyperolius is in need of revision that may result in new generic arrangements.

Reed frogs of the genus Hyperolius inhabit a wide range of habitats from open savannah to montane forests in Sub-Saharan Africa with sympatric species frequently exploiting the same localities at tadpole and adult stage. Niche partitioning with respect to morphological traits related to diet and locomotion is expected to shape local Hyperolius communities. In a case study on Albertine Rift Hyperolius in Rwanda and Burundi, we analysed the partitioning of larval and adult morphospace (three-dimensional volume describing size and shape variation) as affected by phylogenetic relationships and environmental constraints. Morphological features and molecular barcoding demonstrated that the Albertine Rift Hyperolius resolved into 12 terminal groups (=species), which represent five phylogenetic lineages. The major morphological source of variation among species was isometric size variation at tadpole and adult stage. Among-species variation of tadpole size (maximum size reached before metamorphosis) predicted the range of snout-vent length variation in adult males indicating a strong coupling between the two stages. The adaptive decoupling hypothesis is not supported at the taxonomic scale of the Hyperolius genus. Size-independent shape traits were assessed using a principal component analysis. The two traits which explained most of the morphological variance at the two ontogenetic stages were related to feeding (mouth architecture in tadpoles, head shape in adults) and to locomotion (body/tail height in tadpoles, leg morphology in adults). Partitioning of the shape axes indicated that savannah-dwelling and forest-dwelling species exploit different volumes of the available morphospace. Transition from open savannah habitats to closed-canopy forests seems to induce modifications of both tadpole and adult morphology.

The reed frog genus Hyperolius (Afrobatrachia, Hyperoliidae) is a speciose genus containing over 140 species of mostly small to medium-sized frogs distributed in sub-Saharan Africa. Its high level of colour polymorphism, together with in anurans relatively rare sexual dichromatism, make systematic studies more difficult. As a result, the knowledge of the diversity and taxonomy of this genus is still limited. Hyperolius robustus known only from a handful of localities in rain forests of the central Congo Basin is one of the least known species. Here, we have used molecular methods for the first time to study the phylogenetic position of this taxon, accompanied by an analysis of phenotype based on external (morphometric) and internal (osteological) morphological characters. Our phylogenetic results undoubtedly placed H. robustus out of Hyperolius into a common clade with sympatric Cryptothylax and West African Morerella. To prevent the uncovered paraphyly, we place H. robustus into a new genus, Congolius. The review of all available data suggests that the new genus is endemic to the central Congolian lowland rain forests. The analysis of phenotype underlined morphological similarity of the new genus to some Hyperolius species. This uniformity of body shape (including cranial shape) indicates that the two genera have either retained ancestral morphology or evolved through convergent evolution under similar ecological pressures in the African rain forests.

Kassina cassinoides (Boulenger, 1903) (Fig. 4F) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 &female; (SVL 53.4 mm), 1 &male; (SVL 44.3 mm); Samba; 12.70145��N, ��� 2.40387��W; 290 m a.s.l.; 13.VII.1962; M. Lamotte leg.;MNHN-RA-1998.2307, 1998.2308 ��� 1 juvenile (SVL 15.6 mm); Kourweogo province, Laye; 12.53084��N, ��� 1.77155��W; 290 m a.s.l.; VIII.1966; M. Lamotte leg.; MNHNRA-1998.2309. FIELD MATERIAL. ��� Burkina Faso ��� 1&male; (SVL 44.9 mm); Pabr�� dam, south-west of this dam; 12.51159��N, ��� 1.58833��W; 290 m a.s.l.; 10.VII.2017; H. J. Ayoro & A. Ohler leg.; under a stone on the edge of the dam; hand capture ��� 1 &male; (SVL 42.4 mm); Arly National Park, close to the Bouliabou pond; 11.57892��N, 1.4638��E; 170 m a.s.l.; 16. VI.2018; H. J. Ayoro leg.; near a small water body; hand capture ��� 1 &male; (SVL 40.1 mm); W National Park, Koubougou River (500 m north-west of the Koubougou forest post office); 11.96860��N, 2.01259��E; 280 ma.s.l.; 18. VI.2018; same collector as for preceding; in a paddy field; hand capture ��� 1 &male; (SVL 44.5 mm); Folonzo village swampy valley; 9.88803��N, ��� 4.61969��W; 230 m a.s.l.; 6.VIII.2018; H. J. Ayoro & A. Ohler leg.; on a shrub, hand capture ��� 2 &male;; Loumbila dam; 12.49837��N, ��� 1.40313��W; 285 m a.s.l.; 15.VII.2017; same collectors as for preceding; in a bush near the dam; acoustic report ��� 1 &male;; Koubri, ���Ancien barrage���; 12.22843��N, ��� 1.34766��W; 280 m a.s.l.; 16.VII.2017; same collectors as for preceding; on a tree near the dam; acoustic report ��� 1&male;; Koubri, downstream of Nabazana dam; 12.20319��N, ��� 1.35105��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; in vegetable garden near the dam; acoustic report ��� 2 &male;; D��dougou; 12.48972��N, ��� 3.43611��W; 300 m a.s.l.; 22.VIII.2017; H. J. Ayoro leg.; under shrubs near the SONATUR city; acoustic report ��� 1 &male;; W National Park, around PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; acoustic report ��� 1 &male;; Como��-L��raba Forest, Sirigou�� swampy valley; 9.91358��N, ��� 4.62686��W; 240 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; on a tree; acoustic report ��� 3 &male;; Kou Forest, on the bank of Kou River in the savannah part of this Forest; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; same collectors as for preceding; on the trees; acoustic report ��� Dano dam; 11.16464��N, ��� 3.08633��W; 340 m a.s.l.; 2011; J. Penner leg.; field direct observation. DESCRIPTION. ��� 7 specimens examined. Larger than the two other Kassina species from Burkina Faso. SVL of the five observed males 40.1-44.9 mm. The female MNHN-RA-1998.2307 with 53.4 mm SVL exceeding the known body size (Schi��tz 1999; R��del 2000; Amiet 2012). This species distinguished known from all other savannah frogs by a double, black vertebral line (Fig. 4F). HABITAT. ��� Kassina cassinoides and K. fusca Schi��tz, 1967 were calling at night from holes at the base of shrubs, at the beginning of rainy season. Three of our vouchers were collected on the ground near water (in about 10-50 m distance) in June and July. One was found on a tree where it was calling (in about 100 m from the water) in August. R��del (2000) also reported that single individuals may call from trees. It is most abundant in denser vegetated parts of dry savannah (R��del 2000; Nago et al. 2006). This species was not found in the Sahelian zone (Dori and Yakouta). Its distribution range covers a large part of the Sudano-Sahelian region in West Africa from Senegal to northeast Cameroon (Schi��tz 1999; B��hme 2005; Amiet 2012)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 561, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","BOHME W. 2005. - Presence of Kassina cassinoides (Boulenger, 1903) in Senegal. Herpetozoa 18: 177 - 178."]}

Kassina senegalensis (Dum��ril & Bibron, 1841) (Fig. 5B, C) COLLECTION MATERIAL. ��� Burkina Faso ��� 2 &male; (SVL 31.5 - 34.0 mm); Kourweogo province, Laye; 12.53079��N, ��� 1.77155��W; 290 m a.s.l.; VIII.1966; M. Lamotte leg.; MNHN-RA-1998.2009, 1998.2010. FIELD MATERIAL. ��� Burkina Faso ��� 1 &male; (SVL 31.3 mm); W National Park, on the edge of the PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; H. J. Ayoro leg.; under shrubs; hand capture ��� 1 &male; (SVL 32.2 mm); Como��-L��raba Forest; 9.93239��N, ��� 4.60655��W; 220 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; under grasses; hand capture ��� 4&male; (SVL 34.9-39.4 mm), 3 subadults (SVL 20.6-22.4mm); Dori, Toroodi (near the road from Dori to Ouagadougou); 14.00385��N, ��� 0.02411��W; 270 m a.s.l.; 24.VIII.2018; H. J. Ayoro leg.; in small holes and in dried footprints of oxen near a water body; hand capture ��� 1 &female; (SVL 32.0 mm), 2 subadults (SVL 24.3-27.6 mm); N���Djomga (on the road from Dori to Gorom-Gorom); 14.0722��N, ��� 0.0486��W; 265 m a.s.l.; 25.VIII.2018; same collector as for preceding; under a heap of dead grasses in a vegetable garden; hand capture ��� 1 subadult (SVL 22.5 mm); Yakouta; 14.06111��N, ��� 0.13185��W; 267 m a.s.l.; 26.VIII.2018; same collector as for preceding; under a shrub; hand capture ��� 1 &male;; Koubri, around ���Ancien barrage���; 12.22828��N, ��� 1.34587��W; 320 m a.s.l; 16.VII.2017; same collector as for preceding; in a bush; acoustic report ��� 2 &male;; Arly National Park, floodplain (500 m north of Bouliabou pond); 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; same collector as for preceding; acoustic report ��� 1 &male;; Kou Forest; in the savannah part of this Forest; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; H. J. Ayoro & A. Ohler leg.; in a bush; acoustic report ��� Dano, Gbagba; 11.18133��N, ��� 3.05163��W; 300 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Yako; 12.96744��N, ��� 2.26155��W; 330 m a.s.l.; 18.VIII.1961 (Schi��tz 1967). DESCRIPTION. ��� 15 specimens examined. Males from Laye, Como��-L��raba forest and W National Park measuring 31.3- 34.0 mm SVL. Males of Dori and Yakouta atteining larger size, 34.9-39.4 mm SVL. These sizes within the known size range of the species (Lamotte & Xavier 1981; Schi��tz 1999; R��del 2000; Channing 2001; Channing & Howell 2006; Amiet 2012). Dorsal colouration of K. senegalensis a light-green, yellow or green olive, usually with three dark longitudinal strips, either thin continuous or broken into dark broad blotches (Fig. 5B, C). HABITAT. ��� Kassina senegalensis are fast moving among grasses and shrubs. Two vouchers were collected under bushy grasses from where they called and were difficult to localize. In Dori and Yakouta area, K. senegalensis was found on the ground under piles of pebbles near water, as well as in small holes or cattle footprints between grasses close to a pond. We heard other males calling from the base of thorny shrubs and piles of dry grass near a millet field. Kassina senegalensis was also found in Sahelian savannahs in Mauritania (Padial et al. 2013). It shelters under rocks or termite mounds during dry periods (Poynton & Broadley 1987; R��del 2000)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 562, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","CHANNING A. 2001. - Amphibians of Central and Southern Africa. Cornell University Press, New York, USA, 470 p.","CHANNING A. & HOWELL K. M. 2006. - Amphibians of East Africa. Cornell University Press, New York, USA, 432 p.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002","POYNTON J. C. & BROADLEY D. G. 1987. - Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae. Annals of Natal Museum 28: 161 - 229."]}

Hyperolius concolor concolor (Hallowell, 1844) (Fig. 4D) COLLECTION MATERIAL. ��� Burkina Faso ��� 2&male;; (SVL 27.2-28.4 mm); Houet province, Sokourani (near Hippopotamus pond); 11.51183��N, ��� 4.11611��W; 320 m a.s.l.; 8.XI.1970; M. Lamotte leg.; MNHNRA-1999.7211, 1999.7212. FIELD MATERIAL. ��� Burkina Faso ��� 2&male; (SVL 27.8-29.5 mm); Como��-L��raba Forest, Sirigou�� swampy valley; 9.88809��N, ��� 4.61954��W; 230 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; on flooded grasses; hand capture ��� 5 &male; (SVL 30.3-31.7 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92522��N, ��� 4.57575��W; 240 m a.s.l.; 4.VIII.2018; same collectors as for preceding; on high, flooded grasses; hand capture ��� 1&male; (SVL 29.7 mm); Kou Forest, near the Guinguette water source; 11.18720��N, ��� 4.44185��W; 340 m a.s.l.; 7.VIII.2018; same collectors as for preceding; on leaves of palm trees; hand capture. DESCRIPTION. ��� 10 specimens examined. Only males collected, with SVL 27.8-31.5 mm. Life colouration yellowish to greenish (Fig. 4D). Usually during the day, males more brownish to yellow, often with some dark dorsal pattern, whereas females showing a green colouration. At night individuals of both sexes with a yellow colour (R��del 2000). In alcohol, specimens whitish with small black spots, visible only with a dissecting microscope. HABITAT. ��� Hyperolius concolor concolor was found on flooded grasses in Como��-L��raba Forest and on palm leaves at the source of the Guinguette River in Kou Forest. Specimens were usually sitting comparatively higher than H. nitidulus, but see R��del et al. (2006) for calling site position in various West African hyperoliids. The species is known to occur in areas of degraded forests and humid savannah; it extends into drier savannah zones along gallery forests (Schi��tz 1999; R��del 2000). It is distributed in West Africa from Sierra Leone to northern Benin (Schi��tz 1999; R��del 2000; Nago et al. 2006)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 560, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O., LAMPERT K. P. & LINSENMAIR K. E. 2006. - Reproductive biology of the West African savannah frog Hyperolius nasutus Gunther, 1864 (Amphibia: Anura: Hyperoliidae). Herpetozoa 19: 3 - 12.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108."]}

Afrixalus weidholzi (Mertens, 1938) (Fig. 4C) FIELD MATERIAL. ��� Burkina Faso ��� 23 &male; (SVL 17.4-21.2 mm); Como��-L��raba Forest; 9.82802��N, ��� 4.62727��W; 230 m a.s.l; 1-7.VIII.2018; H. J. Ayoro & A. Ohler leg.; on grasses in a swampy valley; hand capture ��� 1 &male;; W National Park, Kabougou River (in its protected part); 11.96865��N, 2.0125��E; 280 m a.s.l.; 19. VI.2018; H. J. Ayoro leg.; in a bush near water body; acoustic report ��� 1&male;; Koubri, downstream from the Nabazana dam; 12.20322��N, ��� 1.35101��W; 280 m a.s.l.; 17.VII.2017; H. J. Ayoro & A. Ohler leg.; close to a paddy field; acoustic report. DESCRIPTION. ��� 23 specimens examined. Frogs with SVL from 17.4-21.2 mm in 23 males, smaller than A. vittiger. Specimens with a yellow or a very intense silvery white back (Fig. 4C), often with a fine brown vertebral line, at least in the posterior part of the back. Laterally, a less distinct dark band from the snout to the groin. On the shank, presence of dark brown longitudinal bands. HABITAT. ��� This species was usually collected on grass at a height not exceeding 0.5-1.5 m (Fig. 2D). During the day, resting animals can be found above water on plants (Lamotte & Xavier 1981; Schi��tz 1967, 1999). Males call at night, usually on high grasses, a few meters from the pond edges (R��del et al. 2006). Afrixalus weidholzi is a common species which has a wide range in West African drier savannah zones (R��del 2000; Padial et al. 2013) and occurs occasionally in the same habitats as A. vittiger., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 558-560, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O., LAMPERT K. P. & LINSENMAIR K. E. 2006. - Reproductive biology of the West African savannah frog Hyperolius nasutus Gunther, 1864 (Amphibia: Anura: Hyperoliidae). Herpetozoa 19: 3 - 12.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002"]}

Kassina fusca Schi��tz, 1967 (Fig. 5A) COLLECTION MATERIAL. ��� Burkina Faso ��� 1&male; (SVL 29.5 mm), 1&female; (SVL 37.3 mm); Kourweogo province, Laye; 12.53083��N, ��� 1.77150��W; 290 m a.s.l.; 6.VIII.1966; M. Lamotte leg.;MNHNRA-1998.2011, 1998.2302. FIELD MATERIAL. ��� Burkina Faso ��� 3 &male; (SVL 32.2-32.4 mm), 1&female; (SVL 35.8 mm); Arly National Park, Bongola swampy valley; 11.53252��N, 1.48241��E; 170 m a.s.l.; 13. VI.2018; H. J. Ayoro leg.; at the base of shrubs; hand capture ��� 1&male; (SVL 32.0 mm), 1 &female; (SVL 31.4 mm); Arly National Park, in the floodplain; 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; same collector as for preceding; in a bare space; hand capture ��� 1 &male;; W National Park, around the PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; in a bush near the water body; acoustic report ��� 6 &male; (SVL 32.2-35.5 mm), 4 &female; (SVL 32.8-36.5 mm); W National Park, the edge of a temporary pond (1 km south-west of the Kabougou forest post office); 11.93669��N, 1.99464��E; 310 m a.s.l.; 21. VI.2018; same collector as for preceding; near water body; hand capture ��� 1 &male; (SVL 29.8 mm); Como��-L��raba Forest; 9.88803��N, 4.61969��W; 230 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; near a maize field (50 m of the forest); hand capture ��� 1 &male; (SVL 32.4 mm); Folonzo village swampy valley; 9.95777��N, ��� 4.67455��W; 240 m a.s.l.; 6.VIII.2018; same collectors as for preceding; in a cavity of tree; hand capture ��� 4&male; (SVL 32.8 -35.0 mm), 3 &female; (SVL 41.7-42.7 mm); Koti; 11.38779��N, ��� 3.06401��W; 310 m a.s.l.; 22.VIII.2018; H. J. Ayoro leg.; near a bean field; hand capture ��� 3 &male;; Kou Forest, on the bank of Kou River in the savannah part; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; H. J. Ayoro & A. Ohler leg.; in a bush on the river bank; acoustic report ��� 2 &male;; Koubri, around ���Ancien barrage���; 12.22828��N, ��� 1.34587��W; 320 m a.s.l.; 16.VII.2017; same collectors as for preceding; acoustic report ��� Ioba province, near Dissin; 10.94250��N, ��� 3.15778��W; 260 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l.; 1983, I-III.2008 (B��hme et al. 1996; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011); ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 29 specimens examined. According to Schi��tz (1999) and R��del (2000), SVL of this species ranging from 29- 40.5 mm in both sexes. In the observed specimens, males measure 29.8-35.5 mm and females 31.4-42.7 mm SVL. The back of living specimens of a light green-olive colouration with scattered larger and smaller darker spots (Fig. 5A). HABITAT. ��� This species calls at night from holes at the base of shrubs. After rain these frogs may be found moving on the open ground, between grasses and bushes during the night. This species was not found in Dori and Yakouta in the Sahel region of Burkina Faso. It is a common West African savannah species, with a distribution ranging from southern Senegal to western Nigeria (B��hme et al. 1996; Schi��tz 1999; R��del 2000; Nago et al. 2006; Segniagbeto et al. 2007)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 561-562, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90."]}

Hyperolius lamottei Laurent, 1958 (Fig. 7B) COLLECTION MATERIAL. ��� Burkina Faso ��� 1&female; (SVL 21.6 mm); L��raba province, Katiar�� (north-east of Ferk��) close to Ivorian border; 9.91200��N, ��� 4.94776��W; 250 m a.s.l.; 10.XI.1970; M. Lamotte leg.; MNHN-RA-1997.9889. DESCRIPTION. ��� Female with a SVL of 21.6 mm. A small savannah frog with a SVL of 20-27 mm of both sexes (Lamotte & Xavier 1981; Schiotz 1999). Data based on additional specimens from various West African localities: males smaller than females; Ivory Coast 38&male; (19.5-23.1 mm), 32 &female; (20.3-26.1 mm); Guinea 32 &male; (18.0- 23.4 mm), 27 &female; (20.0- 24.4 mm); Liberia 27 &male; (19.5-23.8 mm), 37&female; (20.3-28.1 mm); Sierra Leone 54 &male; (19.3-23.7 mm), 28 &female; (21.8-27.3 mm). Slender frog, as Afrixalus vittiger. Back with 9 longitudinal bands: four white and five brown (Fig. 7B); see also figures inSchi��tz (1999), R��del & Ernst (2003), Adeba et al. (2010) and Channing et al. (2013). HABITAT. ��� Hyperolius lamottei was collected in Katiar��, north-east of Ferk�� region. This locality is situated in Cascades region in Burkina Faso, characterized by savannah and gallery forest areas with waterfalls, hills and cliffs. Based on the collection data of MNHN specimens from others localities of West Africa, H. lamottei inhabits wet savannah zones and seems to have a preference for montane regions such as the Nimba, Loma and P��ko mountains (Lamotte 1971; R��del & Ernst 2003; R��del et al. 2004), where it inhabits short grassland savannahs, often on granite ground (Schi��tz 1967; R��del & Ernst 2003). The Burkina Faso record is thus very unexpected and future field work should try to confirm its presence at the locality mentioned above and other similar sites., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 560, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. & ERNST R. 2003. - The amphibians of Marahoue and Mont Peko National Parks, Ivory Coast. Herpetozoa 16: 23 - 39.","ADEBA P. J., KOUASSI P. & RODEL M. - O. 2010. - Anuran amphibians in a rapidly changing environment - revisiting Lamto, Cote d'Ivoire, 40 years after the first herpetofaunal investigations. African Journal of Herpetology 59: 1 - 16. https: // doi. org / 10.1080 / 04416651003595337","CHANNING A., HILLERS A., LOTTERS S., RODEL M. - O., SCHICK S., CONRADIE W., RODDER D., MERCURIO V., WAGNER P., DEHLING J. M., DU PREEZ L. H., KIELGAST J. & BURGER M. 2013. - Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species. Zootaxa 3620: 301 - 350. https: // doi. org / 10.11646 / zootaxa. 3620.3.1","LAMOTTE M. 1971. - Le Massif des Monts Loma (Sierra Leone), Fasciule I; XIX. Amphibiens. Memoire de l'Institut fondamental d'Afrique noire (A) 86: 397 - 407.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346."]}

We related the first commented list of the amphibian species of Burkina Faso. To obtain contemporary data, we investigated six sites from July 2017 to September 2018. The survey is unique for West Africa in combining a variety of different habitat types, e.g. Sahelian areas, grassland and woodland savannahs, floodplains, gallery forests and agricultural sites. Animals were found at varying spatial scales by opportunistic visual encounter surveys after sunset supplemented with acoustic searching. Further data were based on museum vouchers collected between the years 1959 and 2011 and deposited in European and American museums, and the literature review focusing on the country species. We listed 36 anuran species from 11 families. Surprisingly seven taxa, namely Arthroleptis poecilonotus Peters, 1863, Afrixalus vittiger (Peters, 1876), Afrixalus weidholzi (Mertens, 1938), Kassina cassinoides (Boulenger, 1903), Ptychadena schillukorum (Werner, 1908), Hyperolius lamottei Laurent, 1958 and Amnirana albolabris (west) (Jongsma et al. 2018), were new country records. A few rare, hard to record or little known species, including Arthroleptis poecilonotus, Hyperolius lamottei, Xenopus tropicalis (Gray, 1864) and Amnirana albolabris (west) can be confirmed. The origin of a specimen of Sclerophrys mauritanica (Schlegel, 1841) mentioned as coming from Burkina Faso deposited in the collections of MNHN must be considered as an error. We briefly discussed the taxonomy and indicated the habitats of all species mentioned in this country so far. The discovery of several new country records highlights that the amphibian fauna of Burkina Faso is still incompletely known and that many poorly explored sites may still harbour a high diversity.

Hyperolius nitidulus Peters, 1875 (Fig. 4E) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 &female; (SVL 24.1 mm), 2&male; (SVL 19-27.5 mm); Passor�� province, Samba; 12.70144��N, ��� 2.40390��W; 290 m a.s.l.; 18.VII.1961; NA leg.; MNHNRA-2000.3113, 2000.3114, 1979.3724 ��� 4 &male; (SVL 18.9-24 mm); Pabr��; 12.51000��N, ��� 1.57953��W; 290 m a.s.l.; 1963; M. Lamotte leg.; MNHN-RA-2000.3115, 2000.3116, 2000.5609, 2000.5606. FIELD MATERIAL. ��� Burkina Faso ��� 4 &male; (SVL 27.3 -29.0 mm); Pabr�� dam, downstream of the dam; 12.51685��N, ��� 1.59687��W; 290 m a.s.l.; 11.VII.2017; H. J. Ayoro & A. Ohler leg.; on grasses; hand capture ��� 1 &male; (SVL 29.4 mm); Loumbila dam; 12.49844��N, ��� 1.40311��W; 280 m a.s.l.; 14.VII.2017; same collectors as for preceding; on aquatic plants near a plantain field; hand capture ��� 8 &male; (SVL 29.3 -32.0 mm); Koubri, ���Ancien barrage���; 12.22878��N, ��� 1.34522��W; 290 m a.s.l.; 16.VII.2017; same collectors as for preceding; in a maize field near water body; hand capture ��� 1 &male;; Deux Bal��s National Park, downstream from Soumbou dam; 11.69833��N, ��� 2.79713��W; 260 m a.s.l.; 8.XI.2017; H.J. Ayoro leg.; on aquatic plants; acoustic report ��� 2&male; (SVL 23.2-24.9 mm); W National Park, near the Kabougou forest post office; 11.95738��N, 2.01381��E; 290 m a.s.l.; 20. VI.2018; same collector as for preceding; on shrubs; hand capture ��� 14 &male; (SVL 25.7-28.4 mm), 1 &female; (SVL 30.6 mm); Como��-L��raba Forest, Sirigou�� swampy valley; 9.88809��N, ��� 4.61954��W; 230 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; around a small pond; hand capture ��� 5 &male; (SVL 26.8-28.4 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92522��N, ��� 4.57575��W; 240 m a.s.l.; 5.VIII.2018; same collectors as for preceding; in an untilled field; hand capture ��� 4 &male; (SVL 25.5 -28.0 mm); near the Como�� River bridge; 9.93239��N, ��� 4.60655��W; 220 m. a.s.l; 5.VIII.2018; same collectors as for preceding; on aquatic plants in small stagnant waters on the trail; hand capture ��� 1 &male; (SVL 26.2 mm); Folonzo village swampy valley; 9.88803��N, ��� 4.61969��W; 230 m a.s.l.; 6.VIII.2018; same collectors as for preceding; near a vegetable garden; hand capture ��� 5 &male;; Kou Forest; 11.18888��N, ��� 4.43944��W; 260 m a.s.l.; 7-9.VIII.2018; same collectors as for preceding; on shrubs and grasses around a water body; acoustic report ��� Ioba province, Bontioli reserve; 10.93682��N, ��� 3.13804��W; 260 m a.s.l.; J. Penner leg. ��� Dano; 11.16681��N, ��� 3.08870��W; 330 m a.s.l. and 11.16464��N, ��� 3.08633��W; 340 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011). DESCRIPTION. ��� 49 specimens examined.Size of vouchers (SVL 23.2- 31.3 for males and 28.8-30.9 mm for females) corresponding the values of Lampert (2001). In life, specimens with a yellowish back framed by dark lateral bands with narrow yellow edges, extending from the eyes back to above the legs (Fig. 4E). Sometimes, back showing a brown and greyish pattern with very small dark spots. Dorsal coloration lighter and with less contrasting pattern at night. HABITAT. ��� Hyperolius nitidulus was found on grasses and bushes in floodplains, in swampy valleys and farmland such as maize and paddy fields, plantain plantations and vegetable gardens. It was also found on plants above water at nearby ponds and rivers. This species inhabits shrublands, grasslands and wetlands in the Guinean and Sudanian savannahs, from central Guinea to Cameroon (Lamotte 1967b; R��del 2000; R��del et al. 2010a)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 560-561, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["LAMPERT K. P. 2001. - Alternative life history strategies in the West African reed frog, Hyperolius nitidulus. Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universitat Wurzburg. Germany, 168 p.","LAMOTTE M. 1967 b. - Les batraciens de la region de Gbakobo (Cote d'Ivoire). Memoire de l'Institut fondamental de l'Afrique noire (A) 29: 218 - 294.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O., SANDBERGER L., PENNER J., MANE Y. & HILLERS A. 2010 a. - The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae). Bonn zoological Bulletin 57: 177 - 188."]}

Afrixalus vittiger (Peters, 1876) (Fig. 4B) COLLECTION MATERIAL. ��� Burkina Faso ��� 2&female; (SVL 21.6-27.4 mm), 1 subadult (SVL 18 mm); Passor�� province, Samba; 12.70144��N, ��� 2.40390��W; 290 m a.s.l.; VII.1962; M. Lamotte leg.; MNHNRA-1979.4741, 1979.4742, 1979.4743. FIELD MATERIAL. ��� Burkina Faso ��� 8 &male; (SVL 24.5 -27.0 mm); Pabr�� dam, downstream of the dam; 12.51679��N, ��� 1.59693��W; 290 m a.s.l.; 11.VII.2017; H. J. Ayoro & A. Ohler leg.; on flooded grasses; hand capture ��� 2 &male; (SVL 23.7-24.5 mm); same locality; 12.VII.2017; same collectors as for preceding; on grasses in irrigation ditch; hand capture ��� 3 &male; (SVL 24.3-25.2 mm), 1 &female; (SVL 26.7 mm); Loumbila dam; 12.49844��N, ��� 1.40311��W; 280 m a.s.l.; 14.VII.2017; same collectors as for preceding; in a plantain plantation downstream from the dam; hand capture ��� 8 &male; (SVL 24.3-26.7 mm); same locality; 15.VII.2017; same collectors as for preceding; on small plants around vegetable gardens; hand capture ��� 4 &male; (SVL 23.4- 25.2 mm); Koubri, ���Ancien barrage���; 12.20322��N, ��� 1.35100��W; 280 m a.s.l.; 16.VII.2017; same collectors as for preceding; in a maize field; hand capture ��� 2 &male; (SVL 22.8-24.3 mm); Koubri, downstream from the Nabazana dam; 12.20322��N, ��� 1.35101��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; on the edge of a paddy field; hand capture ��� 2 &female; (SVL 23.4-27.3 mm); Koubri, downstream from Arzoumbaongo dam; 12.22064��N, ��� 1.29590��W; 270 m a.s.l.; 18.VII.2017; same collectors as for preceding; on shrubs near water body; hand capture ��� 1 subadult (SVL 17.1 mm); Deux Bal��s National Park, downstream from the Soumbou dam; 11.69830��N, ��� 2.79711��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; on dry grasses near the water body; hand capture ��� 3 subadults; Deux Bal��s National Park, edge of Koh��na pond; 11.67661��N, ��� 2.83614��W; 270 m a.s.l.; 9.XI.2017; same collector as for preceding; on the mud near water body; direct observation in field ��� 2 &female; (SVL 21.0- 21.7 mm); Arly National Park, a floodplain; 11.57855��N, 1.46001��E; 260 m a.s.l.; 16-17. VI.2018; same collector as for preceding; on stones near the water body; hand capture ��� 4 &male; (SVL 21.9-24.8 mm), 2 &female; (SVL 21.3-23.1 mm); W National Park, Kabougou River (500 m north-west of the Kabougou forest post office); 11.97060��N, 2.01622��E; 280 m a.s.l.; 18. VI.2018; same collector as for preceding; on grasses; hand capture ��� 11 &male; (SVL 20.9-23.6 mm), 4 &female; (SVL 23.8-24.8 mm); the protected part of Kabougou River (500 m north-east of the Kabougou forest post office); 11.96865��N, 2.0125��E; 280 m a.s.l.; 19. VI.2018; same collector as for preceding; on grasses ��� 2 &male; (SVL 21.3 -22.0 mm); the edge of PK-27 bouli; 11.88744��N, 2.11913��E; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; on grasses; hand capture ��� 4&male; (SVL 22.7-24.1 mm), 1&female; (SVL 25.2 mm); Como��-L��raba Forest, Sirigou�� swampy valley; 9.88811��N, ��� 4.61950��W; 230 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; on shrubs and grasses; hand capture ��� 2 &male; (SVL 22.7-23.5 mm), 1 &female; (SVL 25.9 mm); Kotokoi swampy valley; 9.82802��N, ��� 4.62727��W; 230 m a.s.l.; 3.VIII.2018; same collectors as for preceding; on grasses; hand capture ��� Kou Forest, bank of Kou River; 11.18720��N, ��� 4.44185��W; 340 m a.s.l.; 8.VIII.2018; same collectors as for preceding; on shrubs around the river bank; acoustic report ��� Ioba province, Gbagba (near Dano); 11.18133��N, ��� 3.05165��W; 300 m a.s.l.; 2011; J. Penner leg.; direct observation in field. LITERATURE REPORTS. ��� Burkina Faso ��� Ouagadougou; 12.38009��N, ��� 1.51100��W; 300 m a.s.l.; 30.VII.1992 (B��hme et al. 1996) ��� Yako; 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; 18.VIII.1961 (Schi��tz 1967). DESCRIPTION. ��� 71 specimens examined. Some of these specimens slightly larger than others. Male���s body size between 20.9-27.3 mm while female���s size 21.3-26.7 mm. Agreeing with Pickersgill (2007), indicating males body size from 21-26 mm and 22-28 mm for females. Skin asperities, typical for the genus, present on dorsum but also on belly, prominent on tarsus and feet in males. On back of all specimens, seven vertical distinct bands, four brown separated by three whitish (Fig. 4B). Ventral skin colour whitish. HABITAT. ��� We observed A. vittiger in floodplains and swampy valleys. The species is present in protected areas such as Arly National Park, W National Park, Como��-L��raba Forest and Deux Bal��s National Park. In Koubri, Pabr�� and Loumbila, the species was found in paddy fields, around ponds on grasses and plants surfacing water (Fig. 2C, D, F). It is a common savannah species which has a large distribution range in West Africa (R��del 2000; Leach�� et al. 2006; Nago et al. 2006; Segniagbeto et al. 2007). This species has often been confused with its congener A. fulvovittatus. The two species partly overlap in the forest-savannah ecotone of West Africa. However, Afrixalus fulvovittatus occurs in heavily degraded forest areas whereas A. vittiger mostly lives in savannah areas (Pickersgill 2007; R��del & Glos 2019)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 558, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","PICKERSGILL M. 2007. - A redefinition of Afrixalus fulvovittatus (Cope, 1860) and Afrixalus vittiger (Peters, 1876) (Amphibia, Anura, Hyeroliidae). African Journal of Herpetology 56: 23 - 37. https: // doi. org / 10.1080 / 21564574.2007.9635551","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LEACHE A., RODEL M. - O., LINKEM C. W., DIAZ R. E., HILLERS A. & FUJITA M. K. 2006. - Biodiversity in a forest island: reptiles and amphibians of the West African Togo hills. Amphibian and Reptile Conservation 4: 22 - 45. https: // doi. org / 10.1514 / journal. arc. 0040018","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","RODEL M. - O. & GLOS J. 2019. - Herpetological surveys in two proposed protected areas in Liberia, West Africa. Zoosystematics and Evolution 95: 15 - 35. https: // doi. org / 10.3897 / zse. 95.31726"]}

Heterixalus carbonei The description of the external morphology is based on a specimen in stage 31, field number 61HiwaB7.3.01, from Kirindy Forest (TL 22.56 mm, BL 7.63 mm; Table 1-2, Fig. 2). External morphology. In the dorsal view, the body has an ovoid form. Its widest point lies in the distal half to three quarters of the body. The snout is broadly rounded. In profile, the body is slightly depressed (BW 113% of BH). Its maximal height is in its distal half shortly behind mid-body. The snout is pointed. The eyes are moderately sized (ED 14% of BL) in a lateral position with dorsolateral orientation, and visible in the ventral view. Nares are located dorsolaterally, closer to the snout than to the eyes. They have a dark spot on their back and open both dorso- and anterolaterally in a round opening form. The distance between nostrils is about one quarter of the inter-orbital distance (NN 26% of IOD). The spiracle is sinistral and visible in the dorsal view. Its opening is located at maximum body width, shortly below the level of the proximal end of the tail myotomes axis. The spiracle opens posterodorsally, and its inner wall is free from the body. The vent tube is dextral, very short, and detached from the ventral fin. The coiled gut is visible. The tail is twice as long as the body (TAL 196% of BL). The tail muscle is narrow (TMW 32% of BW) and low (TMH 49% of BH). It rapidly converges (TMHM 22% of THM) until it reaches the pointed tail tip. Dorsal and ventral fins originate pointed at the body tail junction. The dorsal fin diverges to maximum height in the proximal quarter to half of the tail and starts converging towards its pointed tail tip in the distal quarter to half. The ventral fin reaches the comparably lower maximum height in the proximal quarter. Fin forms result in a tail that is at its maximum, which is in the proximal quarter to half of the tail (THM 67% of MTH), as high as the body (MTH 97% of BH). The oral disc type is generalized without emargination and of moderate size (ODW 41% of BW). Its position and orientation are anteroventrally. On the lower labium, there is a single row of very few marginal papillae. Ventrally and laterally, there are some submarginal papillae. The longish and rounded papillae form a moderately sized dorsal gap (DG 55% of ODW). The LTRF is 1/3(1), P3 is much shorter than P1 and P2. Labial teeth have three to four cusps. The upper tooth row has at least half the width of the oral disc (A1 51% of ODW). The same is true for the jaw sheaths (JL 59% of ODW). The jaws sheaths are half keratinized with pointed serrations on their edges. The upper jaw sheath has a broad, arched form, while the lower one is V-shaped. Coloration in preservative. Laterally, the tadpole is faintly pigmented in the anterior half and strongly pigmented in the posterior half. The coiled gut is visible. Dorsally, there is a broad strip of pigmentation that forks anteriorly. The fins and the lateral tail muscle show sparse pigmentation with a web of fine brown lines on the muscle and the dorsal fin. The limb buds are without pigmentation as well as the spiracle, which is transparent. The eyes are black. Ventrally, there is hardly any pigmentation, including the oral disc, which has dark brown pigments solely on the keratinized part of the jaws sheaths and the labial teeth. Variation. The external morphology and general appearance was very similar between specimens. The ratios taken on five tadpoles in stages 27 to 31 vary in the following proportions: BH 104-133% of BW; ED 12-14% of BL; NN 24-46% of IOD; TAL 168-196% of BL; TMW 22-31% of BW; TMH 31-49% of BH; TMHM 21-28% of THM; THM 71-89% of MTH; MTH 68-97% of BH; DF 121-196% of TMHM; VF 126-177% of TMHM; DF 95 116% of VF; ODW 31-41% of BW; DG 39-55% of ODW; A1 51-61% of ODW; JL 51-59% of ODW., Published as part of Griep, Stine & Glos, Julian, 2020, Description of tadpoles of the frogs Heterixalus tricolor, H. carbonei and H luteostriatus (Anura: Hyperoliidae) from western Madagascar, pp. 332-344 in Zootaxa 4767 (2) on pages 338-340, DOI: 10.11646/zootaxa.4767.2.8, http://zenodo.org/record/3771530