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Channing, A., Hillers, A., Lötters, S., Rödel, O., Schick, S., Conradie, W., Rödder, D., 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, DOI: http://dx.doi.org/10.11646/zootaxa.3620.3.1

Hyperolius friedemanni sp. nov. Mercurio & Rödel Friedemann's Long Reed Frog (Fig. 4) Holotype. SMF 85694 (tissue VM 11), an adult male, collected at Karonga, Malawi, 7 February 2007 by V. Mercurio, 9 ° 55 ' 59.6 '' S, 33 ° 56 ' 44.6 '' N, 472 m a.s.l. Paratypes. ZMB 76095 (tissue VM 12), an adult female, with the same details as the holotype; SAIAB 186000, two juvenile specimens (Monkey Bay, Malawi) (Fig. 1). Genetic material. SMF 85694, ZMB 76095 (holotype and paratype) SAIAB 186000 (two specimens) Monkey Bay, Malawi. Diagnosis. The advertisement call (Fig. 8) consists of a brief initial note of eight pulses, followed by six pulses at a slower rate. The duration of the call is 0.12 s. It can be distinguished from species that produce only a buzz, such as H. acuticeps, H. jacobseni sp. nov. and H. nasutus. It can also be distinguished from H. adspersus, H. dartevellei and H. lupiroensis sp. nov., which produce only a brief single note. It differs from those species with calls longer than 0.2 s, such as H. benguellensis, H. inyangae sp. nov. and H. viridis. It can be distinguished from those species where the slower part of the call consists of less than half the pulses of the initial note, such as H. howelli sp. nov., H. igbettensis and H. rwandae sp. nov. Finally, although the structure of the call of H. poweri is similar, the two differ in pitch, H. poweri having the dominant frequency of 5.9 kHz, while H. friedemanni sp. nov. has a dominant frequency of 4.3 kHz. The snout is sharply rounded in profile, which distinguishes it from species with truncated, bluntly rounded, or shark-like snouts; H. adspersus, H. benguellensis, H. howelli sp. nov., H. igbettensis, H. inyangae sp. nov., H. jacobseni sp. nov., H. poweri, H. dartevellei and H. viridis. It is the only species in the study where the webbing reaches the disc on all toes, at least on one side. This distinguishes it from all other species. Description of Holotype. The width of the gular flap is 5.1, hand 5.5. The top of the snout is flat, with the tip of the snout acutely rounded from above and from the side (Fig. 6) (HW/SUL 0.29). The snout is 1.4 x eye. The tympanum is not visible. The nostrils are positioned near the snout tip (EN/SL 0.5), nostril opening rounded, slightly protruding. Fine teeth are present on the upper jaw. The choanae are small, round. The tongue is long, with the posterior as wide as the length, with the terminal 20 % bifurcated. Vomerine processes absent. The hand is 25.5 % of the SUL. A small inner metacarpal tubercle is present. The relative finger lengths are 1 Paratype variation. The female paratype is similar to the holotype. Tympanum not visible. The paratypes from Monkey Bay collected by EN are subadults, with skin that is transparent in preservative, showing large numbers of subdermal parasite eggs. Advertisement call. Recorded at Karonga, on 7 February 2002 at 23: 40 h, 27 °C air temperature, voucher specimen SMF 85694. The call (Fig. 8) consists of the regular repetition of one single biphasic pulsed note with a duration 110–190 ms. Interval between notes is 180–360 ms. The note repetition rate is 1.4 s - 1. The dominant frequency is 3900–4500 Hz. The specimen was calling at night from dense grassy vegetation within a swamp in an exposed position about 400 mm above the water. See Table 3 for a summary of call parameters. Eggs and tadpoles. Unknown. Habitat. Swamp along the lakeshore with abundant grassy vegetation and sandy soil. Other common species were: Afrixalus fornasini, Hyperolius pusillus, H. viridiflavus nyassae, H. tuberilinguis, Phrynobatrachus acridoides, P. mababiensis, Ptychadena cf. mascareniensis, P. anchietae, Kassina senegalensis, Amietophrynus gutturalis, A. maculatus, Xenopus muelleri, Arthroleptis stenodactylus, and Hemisus marmoratus. Etymology. We dedicate this new species to Friedemann Schrenk in recognition of his enthusiastic and tireless work for the research and protection of the natural history heritage of Malawi. Remarks. The species is only known from the shores of Lake Malawi, and we suggest that it be regarded as Data Deficient, in terms of the IUCN criteria. Holotype. SAIAB 118979, collected at Himo Road, Arusha, Tanzania (3 ° 21 ' 29.6 " S; 36 ° 50 ' 15.3 " E), collected 12 April 2008 by L.H. du Preez. Paratypes. SAIAB 118980 – 1, female, and SAIAB 118980 – 2, male, collected at Himo Road, near Arusha, Tanzania (3 ° 21 ' 29.6 " S; 36 ° 50 ' 15.3 " E), collected 12 April 2008 by L.H. du Preez; NMK 39221 from Kakamega Forest. Genetic material. SAIAB 118979 – 80 (Himo Road, Arusha) and a specimen from Madehani, Tanzania (no voucher), NMK 39221 (16 S sequence accessioned as AY323926, 12S sequence determined as part of this study) Kakamega Forest, Kenya (Lötters et al. 2004) (Fig. 1). Diagnosis: The advertisement call (Fig. 10) consists of an initial brief note, followed by three slower pulses, with a duration of 0.12 s. It can be distinguished from species producing only a single note and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni sp. nov., H. lupiroensis sp. nov., and H. nasutus. It differs from species producing a call over 0.2 s: H. benguellensis, H. inyangae sp. nov. and H. viridis. It differs from those species where the slower, pulsed part of the call has five or more pulses: H. friedemanni, H. igbettensis, and H. poweri. The initial note consists of eight pulses, while the superficially similar call of H. rwandae sp. nov. has an initial note consisting of 13 pulses. The shark-like profile of the snout distinguishes it from species with truncated or rounded snouts; H. acuticeps, H. adspersus, H. friedemanni sp. nov., H. igbettensis, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus, H. poweri, H. rwandae sp. nov. and H. viridis. The foot has at least one phalanx free of webbing on every toe. This distinguishes it from species where at least one toe is webbed to the disc, at least on one side: H. adspersus, H. benguellensis, H. friedemanni, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov.. It also differs from those species that have less than one phalanx free, on at least one toe: H. acuticeps, H. igbettensis, H. inyangae sp. nov., H. poweri, H. dartevellei and H. viridis. Description of Holotype. Body slender, widest at temporal region, slightly tapering to groin; head comparatively small (HL/SUL 0.32, HW/SUL 0.31), not wider than trunk, slightly longer than wide (HL/HW 1.03); snout top flat, tip of snout rounded (SL/HL 0.48), from above the snout is triangular with a rounded tip (Fig. 6), considerably projecting beyond lower jaw with a shark-like profile, wider than long (SL/EE 0.76); canthus rostralis rounded, almost straight-lined from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed laterally; situated much closer to tip of snout than to eye (EN/NS 2.00), separated from each other by distance nearly equal to distance between eye and nostril (NN/EN 1.06); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.36); eye diameter much shorter than snout (ED/SL 0.74); interorbital distance much wider than upper eyelid (IO/EW 0.96), and greater than internarial distance (IO/NN 1.59); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth; vomer processes and teeth absent; tongue long 4.6, and narrow (2.3 at widest point), free for about three-fourths of length, bifurcated distally for about one-third of length; median lingual process absent; vocal sac single, median, subgular, yellow in colour; gular flap consisting of two areas of thickened skin, the anterior thicker, cream coloured, and the posterior thinner, smooth and white; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like, long. Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate. Fore limbs slender; hand moderately large (HND/SUL 0.24); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 +– 2 II 2–2.75 III 2 –2.5 IV; thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.36); tibio-tarsal articulation passing level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.56), longer than thigh (TFL/THL 1.27); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.70); relative length of toes: II 1– 2 II 2 – 1 III 1–1.5 IV 1.5 – 1.25 V; inner metatarsal tubercle small, oval, prominent; outer one indistinct. Colouration in life. Holotype was a brown-green overall, with white lateral stripes running from the snout, through the top of the eye, to the groin. The lateral stripe is lined with irregular large melanophores. The top of each eye has a smudge of golden brown. The back has many small dark melanophores, with a few irregularly spaced larger pigment cells. The limb joints are pale green, with the limbs showing a brown tinge. The fingers and toes are green with yellow tips. The skin is smooth above and on the limbs, while the ventrum is rough with large flat granules. Colouration in preservative. The dorsal pattern shows two pale lateral stripes edged with large dark melanophores, filled with opaque white pigment. The head and dorsum is uniformly speckled with small melanophores, with a few irregularly spaced larger pigment cells. A thin dark line runs from the nostril to the eye Paratype variation. The female has a similar body shape to the holotype, Skin texture the same as the holotype. Colour in preservative: pale yellow background with large irregular melanophores on the dorsum, overlaying a uniform fine speckling. A dark line runs from eye to eye below the snout tip, running through the nostril. In life the body is pale green with yellowish sides, with darker leaf green around the eyes. The top of the eye has a brown smudge. The line running from eye to eye below the snout tip is reddish brown, with a faint brown band around the top of the snout. The irregular large black spots are less dense posteriorly. The tibia has many large melanophores, with very small speckles on the forearm. The snout profile is rounded, with the nostrils behind the tip. Paratype measurements are included in Appendix 2. Advertisement call. The call is a harsh insect-like chirp. Males call from elevated positions on vegetation (Fig 5). See Table 3 for a summary of call parameters. Eggs and tadpoles. Lötters et al. (2004) found egg clutches attached to submerged vegetation. The larvae are omnivorous, found in quiet water. Habitat. The type locality was a pond of roughly 20 m x 40 m with deep clear water. Along the periphery were dense stands of Typha sp. where the frogs were present from water level to about one meter above water level. Other species present included Amietia angolensis. In Kakamega, H. cinnamomeoventris, H. kivuensis, H. lateralis and H. viridiflavus were present (Lötters et al. 2004) Etymology. We have pleasure in honouring Kim M. Howell for his contributions to East African zoology, made during a long career at the University of Dar-es-Salaam. Remarks. The species is known from western Kenya, and southern and northern Tanzania. Due to its wide range and large populations, we suggest that it be regarded as Least Concern in terms of the IUCN criteria.

Hyperolius poweri Loveridge, 1938 Power's Long Reed Frog (Fig. 9) Genetic material. ZMB 77312 ��� 3 (Port Edward, South Africa); PEM A 9545 ��� 6 (Mkambati Nature Reserve, South Africa) (Fig. 1). Diagnosis: The advertisement call (Fig. 13) consists of an initial brief note with seven pulses, followed by five slower pulses, with a duration of 0.12 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni, H. lupiroensis, and H. nasutus. It differs from species producing a call over 0.2 s; H. benguellensis, H. inyangae and H. viridis. See Table 3 for a summary of call parameters. The snout is bluntly rounded, distinguishing it from those with truncated, shark-like or sharply rounded snouts: H. acuticeps, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae, H. lupiroensis, H. nasutus, and H. rwandae sp. nov. There is a phalanx free of web on the first and third toes, with slightly more than a phalanx free on the fourth toe. The second and fifth toes have about half a phalanx free of web. It can be distinguished from the species that have at least one toe webbed to the disc: H. adspersus, H. benguellensis, H. friedemanni, H. jacobseni, H. lupiroensis, H. nasutus and H. rwandae sp. nov. It differs from the species that have the fifth toe with one or more phalanges free of web: H. acuticeps, H. dartevellei, H. howelli, and H. inyangae. Description of a specimen from Mkambati. This is a male, PEM A 9545, collected at the Mkombati Nature Reserve, Eastern Cape Province, South Africa by J. Venter and W. Conradie, 8 February 2011. Body long and slender, widest at mid-body, slightly tapering to groin; head comparatively small (HL/SUL 0.30, HW/SUL 0.30), not wider than trunk, length subequal to width (HL/HW 0.98); snout long (SL/HL 0.49), sharply rounded in dorsal view, blunt in profile (Fig. 6), projecting beyond lower jaw, wider than long (SL/EE 0.75); canthus rostralis distinct, rounded, slightly concave from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed laterally; situated much closer to tip of snout than to eye (EN/NS 1.46), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.16); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.34); eye diameter shorter than snout (ED/SL 0.70); interorbital distance wider than upper eyelid (IO/EW 1.14), and greater than internarial distance (IO/ NN 1.09); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth; vomer processes and teeth absent; tongue long 5.1, and narrow (2.3 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular, mostly unpigmented and translucent when fully inflated; gular flap consisting of two areas of thickened skin, immediately adjacent to each other; anterior part cream-coloured, larger, more granular, and thicker than posterior whitecoloured part; in resting position only a narrow band of the posterior part visible from below; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like. Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs and gular smooth. Fore limbs slender; hand moderately large (HND/SUL 0.27); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 ��� 2 II 2.5��� 3 III 3 ��� 2.5 IV; thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.5); tibio-tarsal articulation passing level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.52), subsequal to thigh (TFL/THL 1.04); heels overlapping each other when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.81); relative length of toes: II 1 ���1.5 II 0.75��� 2 III 1��� 2 IV 1.5 ��� 0.5 V; inner metatarsal tubercle small, oval, prominent; outer one indistinct. Colouration in life. In life the body is dark green with pale flecks, and fine brown pigment spots. The lateral stripes are shiny white, with a subdermal paradorsal band visible as an irregular pale green band. The toes have reddish tips. Colouration in preservative. In preservative the lateral stripes are shiny white, originating at the nostrils, being pale and subdermal before running over the eyes, and extending back to the groin. The back is densely covered in small chromatophores, with very dark pigment over the snout. The gular region is pale with a few dark spots Eggs and tadpoles. The eggs are white with a grey animal pole, less than 1 mm in diameter, within capsules 2.2 mm in diameter (Wager 1986). Clutch size is about 200, with the eggs being deposited in small groups attached to vegetation under water (Wager 1986). Wager (1986) described the tadpoles. Habitat. The frogs are found on reeds and other emergent vegetation around pools and swamps. Distribution. This species is only confirmed from the east coast of South Africa, from Mkambati in the south, northwards to the Mozambique border. The northern extent of the distribution is unknown. Remarks. The species is only known from the north-eastern coastal strip of South Africa. Due to the disturbed coastal habitat, this species should be regarded as Data Deficient in terms of the IUCN criteria, until further studies are carried out. Holotype. ZMB 77221, adult male, from a pond in farmland on the eastern outskirts of Butare, Huye District, South Province, Rwanda (2 �� 37 ' 10.79 '' S, 29 �� 45 '08.45'' E), collected 13 September 2010 by J.M. Dehling. Genetic material. ZMB 77221 ��� 2 (Butare, Rwanda); ZMB 77223 ��� 4 (Mugesera wetland, Rwanda); ZMB 77225 (Akagera wetland, Rwanda) (Fig. 1). Paratypes. ZMB 77222, adult male, same data as holotype; ZMB 77423 ���24, 77426��� 29, six adult males, ZMB 77425, adult female, all from farmland on the eastern outskirts of Butare, Huye District, South Province, Rwanda, collected in October 2009 by K. L��mkemann, K. Rosar and C. Schwartz; ZMB 77686 ��� 89, four adult males, from farmland on the eastern outskirts of Butare (2 �� 35 ' 44.1 '' S, 29 �� 45 ' 25.6 '' E), collected 27 February 2012 by J.M. Dehling; ZMB 77223, adult female, from the Mugesera wetland south of Lac Mugesera, Bugesera District, East Province, Rwanda (2 �� 12 ' 18.92 '' S, 30 �� 16 ' 18.18 '' E), collected 27 March 2011 by J.M. Dehling; ZMB 77224, adult male, from the Mugesera wetland, Bugesera District, East Province, Rwanda (2 �� 12 ' 15.95 '' S, 30 �� 15 ' 49.25 '' S), collected 27 March 2011 by B. Dumbo and J.M. Dehling; ZMB 77683 juvenile, ZMB 77684 adult female, ZMB 77685 adult male, all from the Mugesera wetland, Bugesera Province, southeastern Rwanda, collected 26 February 2012 by J.M. Dehling; ZMB 77225, adult male, from a wetland of the Akagera River, Kihere District, East Province, Rwanda (2 �� 13 ' 27.63 " S, 30 �� 49 ' 39.06 " E), collected 31 March 2011 by J.M. Dehling; ZMB 77746 ��� 48, three adult males, from a swamp in farmland on the eastern outskirts of Ruhengeri, Musanze District, North Province, Rwanda (1 �� 30 ' 25.73 " S, 29 �� 39 ' 12.11 " E), collected 30 March 2012 by J.M. Dehling. Diagnosis: The advertisement call (Fig. 13) consists of an initial brief note of 13 pulses, followed by three slower pulses, with a duration of 0.14 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni, H. lupiroensis, and H. nasutus. It differs from species producing a call over 0.2 s: H. benguellensis, H. inyangae and H. viridis. It differs from the species that have five or more slower pulses: H. friedemanni, H. igbettensis and H. poweri. The initial note of the call of H. howelli consists of only eight pulses, distinguishing it from H. rwandae with 13. See Table 3 for a summary of call parameters. The snout is sharply rounded in profile, which distinguishes it from those species with truncated, shark-like, or bluntly rounded snouts: H. adspersus, H. benguellensis, H. dartevellei, H. howelli, H. igbettensis, H. inyangae, H. jacobseni, H. poweri, and H. viridis. The third and fifth toes webbed three-fourth the way between disc and distal subarticular tubercle, distinguishing it from the species where the webbing does not reach beyond the distal subarticular tubercles of the third and/or fifth toe: H. acuticeps, H. benguellensis, H. dartevellei, H. howelli, H. igbettensis, H. inyangae, H. nasutus, H. poweri, and H. viridis. It differs from H. friedemanni which has all the toes webbed to the disc, and from H. lupiroensis and H. nasutus which have three phalanges free of web on the inner side of the fourth toe. Standard measurements of the holotype are compared with the other species in Appendix 2. Description of Holotype. Body long and slender, widest at temporal region, slightly tapering to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.30), not wider than trunk, longer than wide (HL/HW 1.10); snout long (SL/HL 0.44), pointed in dorsal view, acute in profile (Fig. 6), considerably projecting beyond lower jaw, wider than long (SL/EE 0.77); canthus rostralis distinct, moderately sharp, almost straight-lined from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 1.42), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.13); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.31); eye diameter shorter than snout (ED/SL 0.70); interorbital distance much wider than upper eyelid (IO/EW 1.71), and greater than internarial distance (IO/NN 1.16); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth, concealed by upper jaw for about the half in ventral view; vomer processes and teeth absent; tongue long 4.9, and narrow (2.4 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular, mostly unpigmented and translucent when fully inflated; gular flap consisting of two medially arranged, subcircular areas of thickened skin, immediately adjacent to each other; anterior part cream-coloured, larger, more granular, and thicker than posterior white-coloured part; in resting position only anterior part visible from ventral; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like, long, directed posterolaterally. Dorsal surfaces of head, trunk and limbs generally appearing smooth but with many densely and more or less evenly scattered tiny, low, spine-like tubercles, hardly visible with the naked eye; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.29); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 +��� 2 II 2���2.75 III 2 ��� 2 - IV (after Myers & Duellman [1982]); thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.63); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.54), longer than thigh (TFL/THL 1.11); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.81); relative length of toes: II 1.5���2 + II 1.25���2 + III 1.25��� 2 IV 2 -��� 1.25 V; inner metatarsal tubercle small, oval, prominent; outer one larger, almost circular, low and less distinct. Colouration in life. Generally weakly pigmented and skin more or less translucent. Dorsum and dorsal surface of head and limbs yellowish green; lateral sides of head and scapular region light green; light, yellowishwhite, moderately broad dorsolateral stripe running along each side of the body from lateral edge of upper eyelid to groin, continued as faint, hardly discernible line from eyelid to tip of snout; very small dark brown to black dots and larger brown to reddish brown specks on dorsum, most densely along both sides of canthus rostralis and upper eyelid and to lesser extent on both sides of dorsolateral stripe; dots roundish, specks shaped like stars or neurons with many dendrites; distal portions of fingers and toes, especially the tips, yellow; ventral side and parts of dorsal side of thigh and upper arm largely unpigmented, appearing bluish-green; peritoneum white, shining through the translucent belly skin; most of internal organs covered with silvery-white tissue (only visible when dissected). Iris reddish-brown during the night, yellowish-brown during the day. Colouration in preservative. All colours have faded to yellow; gular flap whitish-yellow. Paratype variation. The paratypes are similar to the holotype in measurements (Appendix 2). Female type specimens (SUL 18.2���20.4, mean 19.2, n= 3) are about as large as males (SVL 18.4 ���22.0, mean 19.5, n= 15). Colouration of male paratypes is similar to that of the holotype. In some specimens, however, the pattern of dots and speckles is more pronounced. In others, the lateral stripe is less distinct. The light canthal stripe is completely absent in ten male paratypes and in seven paratypes as faintly visible as in the holotype. All females observed in the field, including the female paratypes, lack the light dorsolateral and canthal stripes, gular sacs and flaps, and the spiny dorsal tubercles (Fig. 8). In life, the flanks of the body turn reddish in active males, especially those which are calling. Eggs and tadpoles. Several females with enlarged ovarian eggs were observed but only three of them were collected (ZMB 77143, 77425, 77684). Their ovaries contain about 80 enlarged eggs with a diameter of ca. 0.7���0.8. Eggs are darkly pigmented on the animal pole and white on the vegetative pole. Tadpoles are unknown. Habitat. We found the species only in open habitats, in natural wetlands (Mugesera, Akagera) as well as at the edge of ponds and other lenthic water bodies in cultivated areas. Specimens were observed perching on leaves of vegetation between 5 cm and 1.2 m above the ground or the water level. Males called from elevated positions, sometimes in close proximity to each other (ca. 15 cm). Several males were found engaged in combat. They were holding, pushing, and kicking each other, apparently fighting over an apparently favoured calling site. They also emitted aggressive calls which differed markedly from the advertisement call. The male aggression call is shown in Fig. 14. The following species were found sympatricaly or even syntopically with the new species: Afrixalus quadrivittatus, Amietia cf. angolensis, Amietophrynus kisoloensis, A. regularis, Hyperolius cinnamomeoventris, H. kivuensis, H. lateralis, H. viridiflavus, Kassina senegalensis, Leptopelis kivuensis, Phrynobatrachus cf. mababiensis, P. natalensis, Phrynobatrachus sp., Ptychadena anchietae, P. porosissima, P. cf. mascareniensis, Ptychadena sp. and Xenopus victorianus. Distribution. We observed the species at three further locations in Rwanda, near Gitarama (2 ��05' 57.14 '' S, 29 �� 46 ' 41.94 '' E, Muhanga District, Southern Province, central Rwanda) and west of Kigali (1 �� 57 ' 49.11 '' S, 30 ��00'05.87'' E, Kamonyi District, Southern Province, central Rwanda; and 1 �� 56 ' 59.33 '' S, 30 ��00' 48.97 '' E, Nyarugenge District, Kigali Province, central Rwanda). The localities from where the species is known are in the northern, central, southern and eastern parts of Rwanda. Elevations of the sites ranged from 1300 m (Akagera wetland) to 1800 m (Ruhengeri). Population size was high at all sites. Because the locations in Butare, Mugesera, and Akagera are only 17 km and 15 km from the border with Burundi and 1.6 km from the border with Tanzania, respectively, and especially because the wetlands of Mugesera and Akagera continue into Burundi and Tanzania, respectively, we assume that the species occurs in these countries as well. Etymology. The species epithet derives from Rwanda. It is a noun in genitive singular. Remarks. Although the species is so far only known from several localities in Rwanda, it is probably more widespread. The species occurs in both natural and cultivated areas. Therefore, we propose that it should be classified as Least Concern under the current criteria of the IUCN redlist., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., 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, pp. 301-350 in Zootaxa 3620 (3) on pages 335-339, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius jacobseni sp. nov. Channing Jacobsen's Long Reed Frog (Fig. 12) Holotype. ZMB 77280, a male, collected near Gatiko, Central African Republic, 5 ° 4 ' 43 " N, 20 ° 40 ' 2 " E, by N. Jacobsen, 29 August 2006. Paratypes. A female, ZMB 77281, with the same details as the holotype; 16 males and one female, ZMB 77282 – 298, collected at the same locality, and within a few days of the holotype. Genetic material. ZMB 77280 – 1 (holotype and paratype) (Fig. 1). Diagnosis. The advertisement call (Fig. 11) consists of a short buzz with five pulses, with a duration of 0.06 s. This distinguishes it from the species with a single unpulsed note, and those with both an initial note and a series of slow pulses: H. adspersus, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. igbettensis, H. inyangae, H. lupiroensis sp. nov., H. poweri, H. rwandae sp. nov. and H. viridis. It can be distinguished from the other species with a buzz call by the number of pulses: 25 pulses in H. acuticeps, and eight pulses in H. nasutus. See Table 3 for a summary of call parameters. The snout is bluntly round in profile, distinguishing it from those with truncated, shark-like or sharply rounded snouts: H. acuticeps, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae, H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov. The toes are webbed with one phalanx of the third and fourth toes free, and the fifth toe webbed to the disc. This pattern distinguishes it from those species that do not have the fifth toe webbed to the disc: H. acuticeps, H. benguellensis, H. dartevellei, H. howelli, H. igbettensis, H. inyangae, H. nasutus, H. poweri, and H. viridis. It differs from H. friedemanni which has all the toes webbed to the disc, and from H. rwandae sp. nov. which has two phalanges of the third toe free. The webbing is similar to that of H. lupiroensis sp. nov. Standard measurements of the holotype are compared with the other species in Appendix 2. Description of Holotype. Body long and slender, widest at temporal region, slightly tapering to groin; head comparatively small (HL/SUL 0.34, HW/SUL 0.28), not wider than trunk, longer than wide (HL/HW 1.22); snout long (SL/HL 0.43), bluntly rounded in dorsal view, truncated in profile (Fig. 6), not significantly projecting beyond lower jaw, wider than long (SL/EE 0.74); canthus rostralis distinct, rounded, strongly concave from eye to nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed laterally; situated much closer to tip of snout than to eye (EN/NS 1.6), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.31); eyes directed anterolaterally, moderately protruding, relatively small (ED/ HL 0.25); eye diameter shorter than snout (ED/SL 0.58); interorbital distance much wider than upper eyelid (IO/ EW 2.9), and greater than internarial distance (IO/NN 1.38); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth; vomer processes and teeth absent; tongue long 4.7, and narrow (2.8 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular; gular flap consisting of two medially arranged, subcircular areas of thickened skin, immediately adjacent to each other; anterior part cream-coloured, larger, more granular, and thicker than posterior white-coloured part; vocal sac aperture on each side of the mouth, slit-like, long. Dorsal surfaces of head, trunk and limbs smooth but with many densely and more or less evenly scattered tiny, low, spine-like tubercles; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate. Fore limbs slender; hand moderately large (HND/SUL 0.28); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 –2.5 II 2.5– 3 III 2.5 – 2 IV; thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.60); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.52), longer than thigh (TFL/THL 1.05); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.78); relative length of toes: II 1 – II 0.25– 1 III 0.25– 1 IV 1 –0 V; inner metatarsal tubercle small, oval, prominent; outer circular, low and less distinct. Colouration in life. The body is an overall yellow-brown, with a green tinge visible through the skin of the sides of the body. The lateral stripes are bright white, edged with brownish pigment spots. Colouration in preservative. Colour in preservative pale yellow, with pigmented snout, a blotch of pigment on top of the eye, and minute black melanophores on the back, more dense anteriorly, with larger brown spots irregularly scattered. White lateral lines run from the top of the eye to the groin, bordered by dark lines of spots and melanophores. The belly is white. Paratype variation. The paratypes are similar in size and proportions to the holotype, with the large female ZMB 77281 having SUL 19.5 and with HW 6.2, with the largest female having SUL 21.5. The male paratypes have a conspicuous muscle (m. ileolumbaris) running from behind the tympanum to the groin, visible under the skin. The inner metatarsal tubercle is flattened, while the outer metatarsal tubercle is absent. The discs on the toes are slightly wider than the width of the toes. Eggs and tadpoles. A female paratype ZMB 77281 contains enlarged ovarian eggs with a diameter of ca. 1.1. Eggs are darkly pigmented on the animal pole and white on the vegetative pole. Tadpoles are unknown. Habitat. The types were found on emergent grass and other plants around temporary pools. Etymology. This species is named for the collector, the South African herpetologist Niels Jacobsen. Remarks. The species is only known from southern Central African Republic, although it is probably widespread. It should be regarded as Data Deficient in terms of the IUCN criteria.

Hyperolius igbettensis Schi��tz, 1963 Igbetti Long Reed Frog (Fig. 12) Genetic material. Two samples without vouchers, and ZMB 76542 ��� 43 (Lamto, Ivory Coast); ZMB 77415 (K��rouane, Guinea); ZMB 77416 (Konsankoro, Guinea); ZMB 77410 (Dantilla, Guinea) (Fig. 1). Diagnosis. The advertisement call (Fig. 10) consists of an initial brief note with 12 pulses, followed by five slower pulses, with a duration of 0.12 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. jacobseni sp. nov., H. lupiroensis sp. nov. and H. nasutus. It differs from species producing a call over 0.2 s; H. benguellensis, H. inyangae sp. nov. and H. viridis, and from those where the initial note consists of less than 10 pulses: H. friedemanni, H. howelli, and H. poweri. The snout is bluntly round in profile, which distinguishes it from species with truncated, shark-like, or sharply rounded profiles; H. acuticeps, H. adspersus, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae sp. nov., H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov. The foot is webbed with one or more phalanges free of web on the first four toes, and half free on the fifth toe. This distinguishes it from species where at least one toe is webbed to the disc, at least on one side: H. adspersus, H. benguellensis, H. friedemanni, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov. It differs from species that have at least one phalanx free on the fifth toe; H. acuticeps, H. dartevellei, H. howelli, H. inyangae and H. nasutus. Finally, it differs from the two species that have one or less phalanges of the second toe free of web: H. poweri and H. viridis. Description of a Dantilla specimen. This is based on ZMB 77410, an adult female. The ranges are given from three specimens (ZMB 77410 ��� 412; 1 female, 2 males), with single measurements from the sequenced specimen. Elongate, fragile frogs; Body long and slender (SUL 21.1), widest at temporal region (HW 5.8 ���7.0; 7.0), slightly tapering to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.26), not wider than trunk, longer than wide (HL/HW 1.27); snout long (SL/HL 0.43), subelliptical in dorsal view and protruding in lateral view (Fig. 6), projecting beyond lower jaw, wider than long (SL/EE 0.73); canthus rostralis indistinct, roundish, straight-lined from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril round, directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 1.91), separated from each other by distance slightly less than distance between eye and nostril (NN/EN 0.90); eyes directed anterolaterally, protruding, relatively small (ED/HL 0.29); eye diameter shorter than snout (ED/SL 0.67); interorbital distance narrower than upper eyelid (IO/EW 0.8), and greater than internarial distance (IO/NN 1.11); tympanum barely visible, very small with tympanum-eye distance equal to half diameter of eye; upper jaw with dentition; choanae small, oval, located far anterolaterally at margins of roof of the mouth, concealed by upper jaw for about the half in ventral view; vomer processes and teeth absent; tongue long 5.0, and wide (3.6 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single of males, median, subgular, mostly unpigmented and translucent when fully inflated; gular gland large covering 2 / 3 to almost entire throat, dilatable skin visible posterior to gland; width of male gular flap 3.6���4.8; gular flap consisting of two medially arranged, heart shaped and triangular areas of thickened skin, immediately adjacent to each other; anterior, heart shaped, light yellow, larger, more granular, and thicker than posterior, triangular white-coloured part; in resting position only anterior part visible from ventral; Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs and gular smooth, lower belly slightly more areolate; a few warts in angle of mouth; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.28); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: 1 ) I 1 +��� 1.25 II 0.25��� 2 III 2 ��� 1 IV 1 ��� 0.3 V; no visible internal or external metatarsal tubercles. Colouration in life. The basic colour of live frogs ranges from a light bluish green, to grass green or almost green-brown; shanks, lower and upper arms are almost transparent blue-green; flanks, back and thighs darker green with many small dark spots, sometimes arranged along vertebral line into a broken line; eyelids usually lighter than rest of head and dorsum, yellowish to reddish brown; sometimes head darker (reddish brown) than rest of dorsal surfaces; in some animals, mostly males, light white to yellow dorsolateral stripe, rarely bordered by two dark lines; dark canthal stripe, reddish iris bordered by narrow blue line; gular gland of males yellowish or like rest of vocal sac skin light green-blue; ventral surfaces light, belly whitish, often almost transparent; toe and finger tips yellow to orange; females are usually more ���transparent��� than males, with eggs visible through the body wall. Colouration in preservative. Very pale beige in preservation with small dark spots scattered over back and extremities, with or without a distinct white dorsolateral band. Eggs and tadpoles. Unknown. Remarks. The biology of this species was discussed by R��del et al. (2006). The species is now known from Guinea to Cameroon, and perhaps occurs further east (Amiet 2006 a). We suggest that the IUCN status of Least Concern be maintained., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., 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, pp. 301-350 in Zootaxa 3620 (3) on pages 324-326, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius inyangae sp. nov. Channing Nyanga Long Reed Frog (Fig. 12) Holotype. ZMB 77276, a male, collected at Rhodes Dam in the Nyanga National Park, Zimbabwe, 18 ° 17 ' 20.3 " S, 32 ° 43 ' 24.4 " E, 14 November 2009. Paratypes. A female, ZMB 77277, and two males, ZMB 77278 – 9, with the same collecting details as the holotype. Genetic material. ZMB 77277 – 8, ZMB 77276 (Rhodes Dam, Nyanga National Park, Zimbabwe); ZMB 76099 - 101 (Kaningina, Malawi). Diagnosis: The advertisement call (Fig. 10) consists of a brief initial note of four pulses, followed by nine slower pulses, with a duration of 0.35 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni sp. nov., H. lupiroensis sp. nov., and H. nasutus. It can be distinguished from those species with short calls under 0.2 s: H. friedemanni, H. howelli, H. igbettensis, H. poweri and H. rwandae sp. nov. It differs from H. viridis, which has an initial note consisting of 26 pulses. See Table 3 for a summary of call parameters. It has a shark-like snout profile, which distinguishes it from those species that have truncated or rounded snouts; H. acuticeps, H. adspersus, H. dartevellei, H. friedemanni, H. igbettensis, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus, H. poweri, H. rwandae sp. nov., and H. viridis. The webbing is characterized by three phalanges free of the fourth toe, and two phalanges free of the fifth toe. This distinguishes it from all other species (which have more webbing). Description of Holotype. Body long and slender, widest at mid-body, slightly tapering to groin; head comparatively small (HL/SUL 0.37, HW/SUL 0.29), not wider than trunk, longer than wide (HL/HW 1.27); snout long (SL/HL 0.46), sharply rounded in dorsal view, acute in profile with a distinct protruding tip (Fig. 6), considerably projecting beyond lower jaw, wider than long (SL/EE 0.82); canthus rostralis distinct, sharp, almost straight-lined from eye to nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed dorsolaterally; situated midway between tip of snout and eye (EN/NS 1.0), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.15); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.23); eye diameter shorter than snout (ED/SL 0.51); interorbital distance much wider than upper eyelid (IO/EW 1.14), and greater than internarial distance (IO/NN 1.04); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, slightly oval, located far anterolaterally at margins of roof of the mouth; vomer processes and teeth absent; tongue long 4.6, and narrow (2.9 at widest point), free for about one-quarter of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular, mostly unpigmented and translucent when fully inflated; gular flap consisting of thickened granular skin, vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like, long, directed posterolaterally. Dorsal surfaces of head, trunk and limbs finely granular with minute tubercles visible under magnification; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate. Fore limbs slender; hand moderately large (HND/SUL 0.24); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 +– 2 II 2– 3 III 2.5 – 2.5 IV (after Myers & Duellman [1982]); thenar tubercle absent; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.42); tibio-tarsal articulation not reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.48), subequal to thigh (TFL/THL 0.97); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.85); relative length of toes: II 1.5 – 1.5 II 0.75– 2 III 1.5– 3 IV 3 – 2 V; inner metatarsal tubercle absent; outer one almost circular, flattened. Colouration in life. head green, overlain with brown pigment which extends over the back and exposed surfaces of limbs. Tibia reddish-brown. Iris and eyelid pale brown. The vocal sac is pale green. Colouration in preservative. a yellow-brown background, covered dorsally with a dense speckling of small black and brown melanophores and chromatophores. No pale lateral stripes, pigmentation over snout and head more dense than dorsum. Upper exposed surfaces of limbs and digits pigmented. Paratype variation. The paratypes are similar to the holotype in measurements (Appendix 2). The two males are similar in proportions, including the sharp protruding snout tip, but both have pale lateral stripes. The female, 21.6 SUL, is gravid, with a mid-body width of 10.2. The female also has a sharp shark-like snout, although it is not as acute as those of the males. Eggs and tadpoles. A female (ZMB 77277) has enlarged ovarian eggs with a diameter of ca. 1.3. Eggs are darkly pigmented on the animal pole and white on the vegetative pole. Tadpoles are unknown. Etymology. The species is named for the Nyanga National Park, Zimbabwe. Remarks. The species is known from the Eastern Highlands of Zimbabwe and northern Malawi. The distribution of this species appears to cover at least 900 km of highlands between the collecting localities. Due to the extensive range we suggest that this species be regarded as Least Concern in terms of the IUCN criteria.

Hyperolius dartevellei Laurent, 1943 Dartevelle's Reed Frog (Fig. 9) Synonomy. Hyperolius sagitta Laurent, 1943 Genetic material. ZMB 77303 Ikelenge, Zambia; USNM 576167 ��� 70 (Impongui, Republic of Congo); field numbers A 27, CRT 3577 -9, 3604- 6 (Congo River near Yekela, DRC); CRT 3730, 3798 (Congo River, near Nganda Kona, DRC); CRT 3838 - 9 (Congo River near Ngengele, DRC); CRT 3975 ��� 89 (Congo River near Bomani, DRC); CRT 4024, 4027 (Congo River, near Lulu, DRC); CRT 4205 ��� 10 (Congo River, near Lieki, DRC) (Fig. 1). Diagnosis. A typical advertisement call (Fig. 5) consists of 13 pulses in 0.1 s, with an emphasised frequency of 4.8 kHz. It differs from those species with a brief note consisting of a few initial pulses, followed by a number of pulses at a much slower pulse rate, such as H. benguellensis, H. friedemanni sp. nov., H. howelli sp. nov., H. igbettensis, H. inyangae sp. nov., H. rwandae sp. nov., H. viridis and H. poweri, and those with a longer call consisting of multiple pulses that may change tempo, such as H. acuticeps, H. jacobseni sp. nov., and H. nasutus. See Table 3 for a summary of call parameters. The advertisement call structure is similar to that of H. adspersus and H. lupiroensis sp. nov., while the 16 S sequence of H. lupiroensis sp. nov. differs by more than 11 %. The snout is truncated, distinguishing it from the species with shark-like or rounded snout profiles: H. acuticeps, H. adspersus, H. benguellensis, H. friedemanni sp. nov., H. howelli sp. nov., H. igbettensis, H. inyangae sp. nov., H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus, H. poweri, H. rwandae sp. nov. and H. viridis. The webbing shows a phalanx free on the first, third and fifth toes, with half a phalanx free on the other two. It can be distinguished from the species that have less than a phalanx of the fifth toe free: H. adspersus, H. friedemanni sp. nov., H. igbettensis, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus, H. poweri, H. rwandae sp. nov. and H. viridis. It differs from the species that have one phalanx or more of the fourth toe free of web: H. acuticeps, H. benguellensis, H. howelli sp. nov., and H. inyangae sp. nov. Description of a Carumbo specimen. An adult male PEM A 10059 (measurements presented in Appendix 2) measuring 18.6 mm SUL; body long and slender, widest just behind orbital region, tapering to groin; head relatively small (HL/SUL 0.32, HW/SUL 0.34), not much wider than long (HL/HW 0.95); snout long (SL/HL 0.46), bluntly pointed in dorsal view (Fig. 6), protruding just beyond lower jaw, wider than long (SL/EE 0.72); canthus rostralis distinct; loreal large and oval in shape; nostril directed dorsolaterally, moderately large vertical slit (0.4 mm in length), situated much closer to tip of snout than to eye (EN/NS 1.60), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.60); eyes large (ED 1.8), directed anterolaterally, protruding outwards and forward, pupil is horizontal to circular, visible from below, eye diameter shorter than snout (ED/SL 0.64); interorbital distance much wider than upper eyelid (IO/EW 1.50), and greater than internarial distance (IO/NN 1.41); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae large, oval, vomer processes and teeth absent; tongue long 3.9 and broad 2.8, mostly free except for first quater, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular; large granular gular flap covering thin vocal sac (5.9 wide) Dorsal surfaces of head, trunk and limbs generally appearing smooth but with many densely and more or less evenly scattered tiny, asperities; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.26); tips of fingers enlarged into broad oval disks, no circummarginal groove; relative length of fingers: II 1.5 ��� 0.25 II 0.25 ��� 0.25 III 0.25 ��� 0.25 IV (after Myers & Duellman 1982) thenar tubercle indistinct; palmar tubercles absent. Hind limbs slender, moderately long (LEG/SUL 1.50); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.52), longer than thigh (TFL/THL 1.07); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.77); relative length of toes: II 0.25��� 1 II 0.25��� 1 III 0.25��� 1 IV 1 ��� 0.25 V; inner metatarsal absent; outer metatarsal tubercle large, almost circular, low and not distinct. Colouration in life. Below translucent silvery-white, above uniform translucent green to brown, scattered darker spots, clear yellow-white dorsolateral line from snout to vent, forming a light canthus on the snout, darker pigmentation anterior-lateral from snout tip to above eye, upper jaw nearly free of any pigmentation, eye iris is yellow to brown; dorsal surface of arms and legs with scattered dark spots, inner thighs unmarked. Colouration in preservative. All colours have faded to a beige yellow with brown dorsal spots still visible. Eggs and tadpoles. Unknown. Habitat: Specimens were collected in the grassland floodplain wetlands surrounding a large natural lake (350 ha) at daytime. Specimens were found half a meter to a meter above water level on vegetation. The only other amphibians found were Phrynobatrachus mababiensis. Additional material was collected at a small pond (Hyperolius species were present in the same area, Hyperolius angolensis and Hyperolius cf. cinereus, both species were calling further away and higher up the vegetation. Hoplobatrachus occipitalis was present in the pond. Distribution. Southern Cameroon, east and south through Gabon to the lower Congo Basin and the most northern parts of Angola, and the north-western Zambian highlands. Remarks: The synonomy of H. granulatus (the holotype RMCA- 152 was examined) is supported by the absence of dorso-lateral stripes and a short rounded snout. The species is presently only confirmed from northern Angola, the Cabinda enclave, and Gabon. There is little doubt that existing records refer to this species, and we suggest that its conservation status of Least Concern remains unchanged., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., 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, pp. 301-350 in Zootaxa 3620 (3) on pages 319-320, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius viridis Schi��tz, 1975 Robust Long Reed Frog (Fig. 9) Genetic material. ZMB 76096 (Vintukutu Forest Reserve, Malawi); ZMB 76102 (Kaningina Forest Reserve, Malawi) (Fig. 1). Diagnosis. The advertisement call (Fig. 15) consists of a brief initial note consisting of 26 pulses followed by five slower pulses, with a duration of 0.41 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni, H. lupiroensis, and H. nasutus. It can be distinguished from the other species producing a two-part call, which have a duration less than 0.4 s: H. friedemanni, H. howelli, H. igbettensis, H. inyangae, H. poweri and H. rwandae. It differs from H. benguellensis which only has five pulses in the initial note. See Table 3 for a summary of call parameters. The snout is bluntly rounded, distinguishing it from those with truncated, shark-like or sharply rounded snouts: H. acuticeps, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae, H. lupiroensis, H. nasutus, and H. rwandae. The webbing has one phalanx free on the first to third toes, just more than one free on the fourth toe, and half a phalanx free on the fifth toe. It can be distinguished from the species that are webbed to the disc on the fifth toe: H. adspersus, H. friedemanni, H. jacobseni, H. lupiroensis, and H. rwandae. It differs from the species that have more than half a phalanx free of web on the fifth toe: H. acuticeps, H. benguellensis, H. dartevellei, H. howelli, and H. inyangae. It can be distinguished from the remaining species that are webbed to the disc on the third or fourth toes: H. adspersus and H. nasutus. Our specimens show the stocky build noted by Schi��tz (1975). Description of a Vintukutu specimen. An adult male ZMB 76096, from Vintukutu Forest Reserve, Malawi. Body short and compact, widest at mid-body, tapering to head and neck; head very small (HL/SUL 0.22, HW/SUL 0.34), narrower than mid part of trunk, wider than long (HL/HW 0.64); snout short (SL/HL 0.59), suboivoid in dorsal view, almost truncate in profile (Fig. 6), only slightly protruding beyond lower jaw, almost as long as wide (SL/EE 0.96); canthus rostralis distinct, rounded, slightly concave between eye to nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed dorsolaterally; situated closer to tip of snout than to eye (EN/NS 1.18), separated from each other by distance equal to distance between eye and nostril (NN/EN 0.94); eyes directed anterolaterally, moderately protruding, relatively large (ED/HL 0.59); eye diameter equal to snout length(ED/SL 1.0); interorbital distance much narrower than upper eyelid (IO/EW 0.36), but greater than internarial distance (IO/NN 1.3); tympanum not visible externally; upper jaw with dentition; choanae small, round, located far anterolaterally at margins of roof of the mouth, completely concealed by upper jaw in ventral view; vomer processes and teeth absent; tongue slightly longer than wide (2.1), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular,; The gular flap is large (almost completely covering the throat), glandular and wider (4.7) than long (4.1), white in preservative with many minute melanophores. The skin of the dorsum and upper limbs appears smooth, finely granular under dissecting microscope; flat granular belly; supratympanic fold absent. Fore limbs slender; hand small (HND/SUL 0.19); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 0.5��� 1 II 0.25��� 1 III 0.25��� 1 IV 1 ��� 0.25 V; inner metatarsal tubercle small, oval, not very prominent; outer one not discernible. Colouration in life. The dorsal and ventral surfaces are white, dorsal surfaces (including thighs) densely covered with minute melanophores. Colouration in preservative. All colours have faded to yellow; gular flap whitish. Eggs and tadpoles. Unknown. Remarks. The species is known from southern Tanzania and northern Malawi. We suggest that the IUCN status of Data Deficient be maintained until further studies are undertaken., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., 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, pp. 301-350 in Zootaxa 3620 (3) on pages 339-341, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983