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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., 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 327-329, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius lamottei Laurent, 1958 Lamotte's Reed Frog (Fig. 12) Synonymy: Hyperolius nasutus Channing et al., 2002 (part) Genetic material. ZMB 76536 ��� 7 (Loma Mountains, Sierra Leone); ZMB 76535 (Nimini Forest Reserve, Sierra Leone); ZMB 76525 (Korombadou/ Tourou, Guinea); ZMB 76532 (Mont B��ro Forest Reserve, Guinea); ZMB 76526 ��� 27 (Nimba Mountains, Guinea); three samples (no vouchers), (Mare d'hivenage, Nimba, Guinea); ZMB 76516 (Savanne de But, Nimba, Guinea); ZMB 76523 ��� 24 (Nimba Mountain, Guinea) (Fig. 1). Diagnosis. It is distinguished on the overall yellow background colour pattern, rounded body shape and advertisement call from the species in the H. nasutus clade. The call (Fig. 12), recorded at Lamto by Arne Schi��tz, is a brief unpulsed whistle that has a duration of 0.08 s, and a dominant frequency of 3.5 kHz. See Table 3 for a summary of call parameters. It is known mostly from high altitude grassland, in Senegal, Sierra Leone, Guinea, Liberia and Ivory Coast (e.g. R��del & Ernst 2003, R��del et al. 2004, Adeba et al. 2010). The night-time colour pattern is shown in Figure 12, and the day-time pattern is illustrated in Schi��tz (1999) and R��del & Ernst (2003). In our dataset, H. lamottei is outside the H. nasutus group, with a genetic distance for the 16 S fragment of 13.2���16.4 % to the species inside the group. Tissues were available from near the type locality. Description of Nimba material. A male ZBM 76526, from the type locality, Mount Nimba, Guinea: Body long and slender, widest at temporal region, almost parallel to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.28), not wider than trunk, longer than wide (HL/HW 1.14); snout long (SL/HL 0.44), subelliptical in dorsal view, protruding in profile (Fig. 6), considerably projecting beyond lower jaw, almost as wide as long (SL/ EE 0.97); canthus rostralis indistinct, round, very slightly concave from eye to 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 1.6), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.19); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.38); eye diameter shorter than snout (ED/SL 0.87); interorbital distance almost equalling upper eyelid (IO/EW 0,94), and greater than internarial distance (IO/NN 1.53); tympanum not visible externally; upper jaw with dentition; choanae small, oval, located far anterolaterally at margins of roof of the mouth, completely concealed by upper jaw in ventral view; vomer processes and teeth absent; tongue broad and heart shaped, 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; The gular flap is glandular, white in preservative, with folded skin posteriorly. width of gular flap 4.6, gular flap consisting of one round areas of thickened skin, cream-coloured; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like, long, directed posterolaterally. The skin of the dorsum and upper limbs is smooth, with a flat granular belly; supratympanic fold absent. Fore limbs slender; hand short (HND/SUL 0.18); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 1.5���2.5 II 1���2 + III 1 +��� 2 IV 1���1.25 V; inner and outer metatarsal tubercle indiscernible. Colouration in life. The back and flanks are dark beige to yellow and densely covered with minute melanophores, bands of more densely arranged melanophores border broad white dorsolateral bands; the back with a narrow black vertebral line, a white longitudinal band on upper surfaces of tibia, slightly bordered darker, not very distinct. The colour pattern may be either fainter or with much more contrasting darker stripes and lines in different individuals. Colouration in preservative. All colours fade, but the pattern remains distinct. Eggs and tadpoles. The eggs were described by Schi��tz (1967). The tadpole was described by Arnoult & Lamotte (1958). See R��del (2000). Habitat. Humid savanna habitats close to forest belt, mostly in mountainous areas. Reaching altitudes of above 100 m asl. Often in areas with low grasses and rocky ground. Distribution. Recorded from Senegal, though Sierra Leone, Liberia, southern Guinea into western Ivory Cost (R��del 2000, R��del & Ernst (2003). A population from Central Ivory Coast, Lamto Reserve, may be extinct (Adeba et al. 2010). Remarks. Channing et al. (2002) suggested that this species should be regarded as a junior synonym of H. nasutus, based on similarity of colour pattern and advertisement call structure, which view was disputed by R��del & Agyei (2003) and Schi��tz (2006 b). Molecular evidence presented in this paper shows that it is not in the H. nasutus species group. The sequences of 13 specimens of Hyperolius lamottei all group together forming a clade outside the nasutus group, with little variation. The intraspecific differences in the sample range from 0.0��� 0.63 % for 16 S. The conservation status of this species is Least Concern (IUCN 2011)., 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 329-330, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius acuticeps Ahl, 1931 Sharp-headed Long Reed Frog (Fig. 4) Genetic material. MCZ A- 137085 ��� 86 (Chelinda Camp, Nyika Plateau, Malawi); ZMB 76103, 76107, 76109 (Chongoni Forest Reserve, Malawi); ZMB 76097 ��� 98 (Kaningina Forest Reserve, Malawi) (Fig. 1). Diagnosis. The illustrated advertisement call (Fig. 5) has a duration of 0.22 s, consisting of 25 pulses, with a slower pulse rate at the end. This differs from the brief single notes of H. adspersus, H. lupiroensis sp. nov., and the 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. The structure of the call of H. jacobseni sp. nov. and H. nasutus is similar to that of H. acuticeps, but the former consists of only five pulses with a duration of 0.07 s, while the latter consists of eight pulses in 0.1 s. The call of H. dartevellei consists of 13 pulses in 0.1 s. See Table 3 for a summary of call parameters. The snout is sharply rounded in profile and from above, differing from the truncated snout of H. dartevellei and the sharp, shark-like profile of H. benguellensis, H. inyangae sp. nov. or the bluntly rounded snout profile of H. adspersus, H. igbettensis, H. jacobseni sp. nov. and H. poweri. Although the webbing is variable, a typical specimen has one phalanx of the fifth toe free, while the fourth toe is webbed with half to just more than the first phalanx free, and the third toe likewise. It can be distinguished from species with less than one phalanx of the fifth toe free, such as H. adspersus, H. friedemanni sp. nov., H. igbettensis, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. poweri, H. rwandae sp. nov. and H. viridis. It differs from those species with more than one phalanx free; H. howelli sp. nov. and H. inyangae sp. nov. It differs from H. benguellensis which has webbing on the third toe extending to the disc, from H. nasutus which has webbing on the fourth toe reaching the disc, and from H. dartevellei which has half a phalanx of the fourth toe free. Description of a Chelinda specimen. This description is based on a female MCZ A- 137085 from Chelinda on Nyika Plateau. 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)although slightly wider than long in specimen MCZ A- 137086; snout long (SL/HL 0.43), pointed in dorsal view, acute in profile (Fig. 6), considerably projecting beyond lower jaw, wider than long (SL/EE 0.72); canthus rostralis distinct, rounded, slightly concave from eye to nostril, concave 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 1.43), separated from each other by distance less than distance between eye and nostril (NN/EN 0.9); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.26); eye diameter shorter than snout (ED/SL 0.61); interorbital distance as wide as upper eyelid (IO/EW 1.0), and greater than internarial distance (IO/NN 1.67); 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 4.8, and wide (3.7 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent. Dorsal surfaces of head, trunk and limbs smooth; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolatet. 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 1��� 2 II 1���2.5 III 2 ��� 1 IV (after Myers & Duellman [1982]); thenar tubercle indistinct, low; palmar tubercles absent; inner metacarpal tubercle small, rounded, outer metacarpal tubercle absent. Hind limbs slender, moderately long (LEG/SUL 1.45); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.49), 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 0.82); relative length of toes: II 1.5��� 2 II 1��� 2 III 1��� 2 IV 2 ��� 1.5 V; inner metatarsal tubercle small, oval, prominent; outer one larger, almost circular, low and almost indistinct. Measurements of a second specimen from Chelinda MCZ A- 137086 are included in Appendix 2. Colouration in preservative. The pattern elements that remain after the green has faded are a dark vertebral stripe made up of a single row of chromatophores, with pale lateral bands bordered by irregular dark spots, and a row of dark chromatophores forming a dorsolateral stripe. A male is illustrated in Fig. 4. Eggs and tadpoles. Unknown. Distribution. This species is presently only confirmed from northern Malawi on the Nyika Plateau and Chongoni and Kaningina Forest Reserves, and the type locality in southern Tanzania. Remarks. This study restricts the distribution of the species from the wide range presently attributed to it (Schi��tz et al. 2004), suggesting that its conservation status should be changed from Least Concern to Data Deficient, pending the collection of more data., 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 310-314, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

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 nasicus Laurent, 1943 Pointed long reed frog (Fig. 12) Genetic material. SAIAB A- 136 - 1 (2 specimens, Elephant's Camp, Mozambique); SAIAB A- 188 (Satellite Camp, Mozambique); SAIAB KU 95952, KU 96401, KU 98212 (Quantum Mine & Kalumbila River, Zambia); ZMB 77308 (Kisanfu River, DRC); ZMB 77309 Fungurume, DRC; ZMB 77310 (Nyanga Flats, Zimbabwe); ZMB 77301 ���2, 77304���7, 77314 Ikelenge, Zambia. Diagnosis. The advertisement call (Fig. 11) consists of a brief chirp-like note of about 10���20 constant-rate pulses with a duration of 0.09��� 0.2 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. lamottei, and H. lupiroensis 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 a number of slower pulses: H. friedemanni, H. igbettensis H. howelli and H. poweri. The call is similar in structure to that of H. nasutus, but the latter call has a shorter duration and a higher pulse rate. See Table 3 for a summary of call parameters. The snout is sharply rounded from above, with a shark-like tip when viewed from the side, which distinguishes it from those species without truncated, shark-like, or bluntly rounded snouts: H. adspersus, H. dartevellei, H. friedemanni, H. howelli, H. igbettensis, H. inyangae, H. jacobseni, H. lamottei, H. lupiroensis, H. nasutus, H. poweri, H. rwandae and H. viridis. The first, third and fifth toes have one phalanx free of webbing, distinguishing it from all the other species. Measurements of some individuals are compared with the other species in Appendix 2. Description of a Nyanga Flats specimen. This is a male (ZMB 77310). Body long and slender, widest at midbody, slightly tapering to groin; head comparatively small (HL/SUL 0.32, HW/SUL 0.27), not wider than trunk, longer than wide (HL/HW 1.22); snout long (SL/HL 0.45), pointed in dorsal view, acute in profile (Fig. 6), with a slightly upturned tip, considerably projecting beyond lower jaw, wider than long (SL/EE 0.78); canthus rostralis distinct, sharply rounded, 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 1.7), separated from each other by distance equal to distance between eye and nostril (NN/EN 1.0); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.27); eye diameter less than snout length (ED/SL 0.61); interorbital distance subequal to upper eyelid (IO/EW 1.04), 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 small, oval, located far anterolaterally at margins of roof of the mouth; vomer processes and teeth absent; tongue long 5.4, and narrow (2.2 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; gular flap consisting of thickened skin; anterior part cream-coloured, larger, more granular, and thicker than posterior whitecoloured part; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slitlike, 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 0.28); 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; thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.41); tibio-tarsal articulation not reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.48), longer than thigh (TFL/THL 1.14); 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 0.84); relative length of toes: II 1 ��� 1 II 0.5��� 1 III 1 ��� 1 IV 1 ��� 1 V; inner metatarsal tubercle small, oval, prominent; outer one indistinct. Colouration in life. Generally weakly pigmented. The frog was translucent green, with the pale dorsolateral bands originating at the snout tip. A thin dark middorsal line runs from the snout tip to between the eyes. The iris is dark brown. Colouration in preservative. The back is pale yellow with dense small dark chromatophores. The pale dorsolateral bands originate at the nostril, run over the eye and continue to the groin. They are not bordered. A darkly pigmented band runs from the nostril to the eye. Ventrally unpigmented. Eggs and tadpoles. Unknown. Habitat. This species has been found in open savanna, through densely vegetated areas along the Congo River, to flooded swamp forest. They call from emergent vegetation bordering pools or flowing water up to a meter above the water surface. Distribution. This species has been confirmed on molecular data from Mozambique, north-western Zambia, eastern Zimbabwe and Democratic Republic of Congo (Fig. 1)., 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 333-334, 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., 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 326-327, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius lupiroensis sp. nov. Channing Lupiro Long Reed Frog (Fig. 12) Holotype. ZMB 77299, a gravid female collected near Lupiro, 8 �� 25 ' 29.3 " S, 36 �� 41 ' 33.1 " E, Ifakara district, Tanzania, by A. Danby, 9 July 2007. Paratype. ZMB 77300, a subadult male, with the same collecting details as the holotype. Genetic material. The holotype and paratype (Fig. 1). Diagnosis. The advertisement call (Fig. 11) consists of a single unpulsed note, with a duration of 0.06 s. It can be distinguished from the species with calls consisting of a few initial pulses, followed by a number of pulses at a much slower pulse rate, such as H. benguellensis, H. friedemanni, H. howelli, H. igbettensis, H. inyangae, H. rwandae sp. nov., H. viridis, and H. poweri. It is different from the longer calls consisting of a number of pulses at a more or less constant rate, such as H. acuticeps, H. dartevellei, H. jacobseni, and H. nasutus. 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. benguellensis, H. howelli, H. igbettensis, H. inyangae, H. jacobseni, H. poweri, H. dartevellei and H. viridis. 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, from H. rwandae sp. nov. which has two phalanges of the third toe free, and from H. adspersus which has the fourth toe webbed nearly to the disc. Description of Holotype. Body long and slender, widest at mid-belly, slightly tapering to groin; head relatively small (HL/SUL 0.32, HW/SUL 0.23), not wider than trunk, appreciably longer than wide (HL/HW 1.38); snout long (SL/HL 0.48), bluntly rounded in dorsal view, rounded in profile (Fig. 6), slightly projecting beyond lower jaw, longer than wide (SL/EE 1.1); canthus rostralis distinct, rounded, almost straight-lined from eye to nostril; loreal region almost vertical, slightly concave; nostril a thin slit directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 2.25), separated from each other by distance almost equal to the distance between eye and nostril (NN/EN 0.94); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.35); eye diameter shorter than snout (ED/SL 0.72); interorbital distance about equal to upper eyelid (IO/EW 0.96), and greater than internarial distance (IO/NN 1.44); 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 4.0, and narrow (2.5 at widest point), free for about three-fourths of length, bifurcated distally for about one-third of its length; median lingual process absent. 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 0.26); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2.5��� 3 II 2��� 3 III 2.5 ��� 2.5 IV; thenar tubercle small, distinct; palmar tubercles absent; metacarpals without supernumerary tubercles. Hind limbs slender, moderately long (LEG/SUL 1.64); tibio-tarsal articulation passing level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.52), subequal to thigh (TFL/THL 1.03); 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 0.75; relative length of toes: II 1 ��� 1 II 0.5���2 + III 0.25��� 2 IV 1 ���0 V; inner metatarsal tubercle small, oval, distinct; outer metatarsal tubercle absent. 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 subadult male is considerably smaller than the type, SUL 14.2, and has bright white lateral stripes originating at the nostrils and running back through the top of the eye to the groin. A dark line runs below the pale band from the nostril to the eye. The pale bands have dark borders between the eye and the groin. The back is uniformly speckled, giving the back a brown colour in preservative. Eggs and tadpoles. The type contains enlarged ovarian 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. Remarks. Although it is common for males to have lateral stripes and the females to be spotted in this group, the gravid holotype shows distinct pale lateral stripes. The species is only known from eastern Tanzania, and should be regarded as Data Deficient in terms of the IUCN criteria., 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 330-331, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius benguellensis (Bocage, 1893) Benguella Long Reed Frog (Fig. 4) Synonym. Hyperolius oxyrhynchus (Boulenger, 1901) Genetic material. ZMB 77271 ��� 2, ZMB 77318 (Humpata, Angola); ZMB 77273 ��� 4 (Bicuar National Park, Angola); ZMB 77275 (Zootecnica Plateau, Humpata, Angola); AACRG 1030 (Kaparotta, Botswana); GenBank AF 215224, AF 215442 (Rundu, Namibia) (Fig. 1). Diagnosis. The advertisement call (Fig. 8) is a brief note consisting of five pulses, followed by 14 pulses at a slower rate, with a duration of 0.41 s. It can be distinguished from the brief calls consisting only of a single note, H. adspersus and H. lupiroensis sp. nov. and those consisting only of a series of pulses, H. acuticeps, H. jacobseni sp. nov., H. nasutus and H. dartevellei. The other species with advertisement calls consisting of an initial note followed by some discrete pulses can be distinguished either by their short duration, less than 0.2 s, as in H. friedemanni sp. nov., H. howelli sp. nov., H. igbettensis, H. poweri and H. rwandae sp. nov., or by the lower number of slow pulses, less than 10, as in H. inyangae and H. viridis. See Table 3 for a summary of call parameters. The snout is shark-like in profile, protruding forward of the mouth in a straight line, before forming a sharp tip. It can be distinguished from the truncated, sharply- or bluntly rounded snout profiles as in 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. Description of a Humpata specimen. Body long and slender, widest at mid-body, slightly tapering to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.30), not wider than trunk, longer than wide (HL/HW 1.11); snout long (SL/HL 0.42), bluntly rounded in dorsal view, acute, sharklike in profile (Fig. 6), considerably projecting beyond lower jaw, wider than long (SL/EE 0.68); canthus rostralis distinct, moderately sharp, slightly concave 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 1.64), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.11); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.33); eye diameter shorter than snout (ED/SL 0.78); interorbital distance much wider than upper eyelid (IO/EW 1.71), and greater than internarial distance (IO/ NN 2.1); 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 5.8, and narrow (2.7 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, mostly unpigmented and translucent when fully inflated; gular flap cream-coloured, granular; 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 smooth; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate. Fore limbs slender; hand moderately large (HND/SUL 0.30); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 ���2.5 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 1.55); tibio-tarsal articulation reaching to level of snout tip when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.53), shorter than thigh (TFL/THL 0.91); 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 0.83); relative length of toes: II 1 ��� 1 II 0.75��� 1 III 0��� 1 IV 1 ��� 1 V; inner metatarsal tubercle small, oval, prominent; outer one low, almost indistinct. Colouration in life. Skin more or less translucent. Dorsum and dorsal surface of head and limbs dark green (Fig 4); lateral sides of head and scapular region dark green; light, yellowish-white, moderately broad dorsolateral stripe running along each side of the body from snout tip, over the eye to vent; Pale paravertebral stripes originating on posterior of snout, diverging to level of eyes, and then running parallel to vent; small dark brown to black dots and on dorsum, most densely bordering both sides of dorsolateral stripes; distal portions of fingers and toes, especially the tips, yellow; ventral side and parts of dorsal side of thigh and upper arm largely unpigmented but with irregular dark spots, appearing bluish-green; peritoneum white, shining through the translucent belly skin. Iris reddish-brown. Colouration in preservative. All colours have faded to yellow; gular flap whitish-yellow. Variation. The female ZMB 77271 is similar to the male in measurements (Appendix 2). The female is larger than the male (SUL 23.5). Colouration is variable; ZMB 77271 has a pale green dorsum with irregular darker marks, and yellow eyelids, with fingers and toes also yellow, and a white underside. AC 3073 has pale dorsolateral stripes on a dark green background. Eggs and tadpoles. Unknown. Habitat. We found the species only in open grassy habitats, along stream banks and man-made impoundments where sedges and other tall emergent vegetation were present (Humpata, Bicuar NP). Specimens were observed on leaves and stems of vegetation between 5 cm and 1.0 m above water level. Males called from elevated positions. The following species were found sympatrically or even syntopically with the new species: Hyperolius marmoratus and Xenopus laevis. Remarks. The species is confirmed using molecular data from Rundu in the Caprivi Strip of Namibia, northern Botswana, and southern Angola. The type locality of H. benguellensis is Caota, Angola. The type series (MBL 17.220-223; now Museu Bocage, National Museum of Natural History, University of Lisbon) has been destroyed (Frost 2011). Our specimens agree with the original description of a sharp snout and small dorsal speckles. The specimens from Humpata show a range of colour patterns, from a uniform finely spotted dorsum, to pale dorsolateral stripes, to dorsolateral and paravertebral stripes. This variation was absent from the type description. The genetic material is from the same drainage basin as the type. Hyperolius oxyrhynchus is regarded as a synonym as the type description matches this species. The species is presently only confirmed from southern Angola, northern Namibia, and northern Botswana. We suggest that the conservation status 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 317-318, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

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 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 adspersus Peters, 1877 Sprinkled Long Reed Frog (Fig. 4) Synonymy. Hyperolius granulatus (Boulenger, 1901). Genetic material. USNM 578140, 578144, 578166 (Plain of Vera, 15 km south east of Gamba, Gabon); USNM 578157 (Uemba Road, 2 km south of Gamba, Gabon); USNM 578142 (Setecama Road, 3 km west of Gamba, Gabon); USNM 578165 (National Forestry School, Gabon) (Fig. 1). Specimens examined as above, including the type (ZMB 917). Diagnosis. The advertisement call consists of a brief note, duration 0.04 s, and indistinguishable pulses (Fig. 5). It can be distinguished from the call consisting of a brief note comprising 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. It differs from the longer calls consisting of a number of pulses at a more or less constant rate, such as H. acuticeps, H. jacobseni sp. nov., H. nasutus, H. nasicus and H. dartevellei. See Table 3 for a summary of call parameters. The snout is truncated to bluntly rounded, which differs from the sharp, shark-like profile of H. benguellensis and H. inyangae sp. nov., and the sharply rounded snout profile of H. acuticeps, H. friedemanni sp. nov., H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov. The fifth toe has about half a phalanx free of web. This distinguishes it from those species that have one or more phalanges of the fifth toe free, H. benguellensis, H. howelli, H. inyangae, H. lamottei and H. nasicus; and those with the fifth toe fully webbed, H. friedemanni, H. jacobseni, H. lupiroensis and H. rwandae.It has one phalanx free of webbing on the fourth toe, which distinguishes it from H. dartevellei, which has less than a phalanx free; nasutus which is webbed to the disc at least on one side; and H. poweri and H. viridis which have more than one phalanx free. It has no more than one phalanx free on the second toe, at least on one side, which distinguishes it from H. igbettensis which has more than one phalanx free of the second toe at least on one side Description of a specimen from Gamba, Gabon. An adult male USNM 578142 (measurements presented in Appendix 2) measuring 19.5 mm SUL; body long and slender, widest just behind orbital region, tapering to groin; head relatively small (HL/SUL 0.27, HW/SUL 0.33), wider than long (HL/HW 0.82); snout long (SL/HL 0.51, truncated in dorsal view (Fig. 6), just protruding just beyond lower jaw, wider than long (SL/EE 0.64); canthus rostralis rounded; loreal large and oval in shape; nostril directed laterally, eliptical slit, situated just behind the tip of the snout (EN/NS 1.60), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.19); eyes large (ED 2.0), directed anterolaterally, protruding outwards and forward, pupil is horizontal to circular, visible from below, eye diameter shorter than snout (ED/SL 0.74); interorbital distance much wider than upper eyelid (IO/EW 1.46), and equal to internarial distance (IO/NN 1.0); 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 and broad, mostly free except for first quarter, 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 (4.3 wide). Dorsal surfaces of head, trunk and limbs generally appearing smooth but with many densely and more or less evenly scattered tiny, melanophores; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.28); tips of fingers enlarged into broad oval disks, thin circummarginal groove; relative length of fingers: II 2.5 ��� 2.5 II 2 ���2.25 III 2.25 ��� 2 IV; thenar tubercle indistinct; palmar tubercles absent. Hind limbs slender, moderately long; tibiofibula moderately long (TFL/SUL 0.54), longer than thigh (TFL/ THL 1.08); 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��� 2 II 1��� 2 III 1��� 2 IV 2 ��� 1 V; inner metatarsal flat; outer metatarsal tubercle absent. Colouration in life. No information, but see the photo (Fig. 4) of an unvouchered individual from the Bateka Nature Reserve, Gabon. Colouration in preservative. All colours have faded to a beige yellow with evenly spaced black dorsal melanophores. Eggs and tadpoles. Unknown. Habitat. Specimens were collected in grassland. Distribution. Southern Cameroon, east and south through Gabon to the lower Congo Basin. 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 314-317, 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

Hyperolius nasutus G��nther, 1865 Large-nosed Long Reed Frog (Fig. 9) Synonymy. Hyperolius punctulatus (Bocage, 1895) Genetic material. ZMB 77311 (Calandula, Angola); AC 2990 (Kangandala, Angola); LdP field specimen (Xigera, Botswana); SAIAB 186001 (Vumbura, Botswana) (Fig. 1). Diagnosis. The advertisement call (Fig. 13) consists of a buzz with eight pulses, with a duration of 0.1 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. friedemanni, H. howelli, H. igbettensis, H. inyangae, H. lupiroensis, H. poweri, H. rwandae sp. nov. and H. viridis. It can be distinguished from the other species that have a buzz call by the number of pulses: 13 pulses in H. dartevellei, 25 pulses in H. acuticeps, and five pulses in H. jacobseni. 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 second to fifth toes are webbed to the disc, or just below the disc. This distinguishes it from those species with half or more of a phalanx of the fourth toe free: H. acuticeps, H. benguellensis, H. howelli, H. igbettensis, H. inyangae, H. jacobseni, H. lupiroensis, H. poweri, H. rwandae sp. nov., and H. viridis. Description of a Vumbura specimen. A male, SAIAB 186001 has the body long and slender, widest at temporal region, slightly tapering to groin; head comparatively small (HL/SUL 0.28, HW/SUL 0.31), not wider than trunk, longer than wide (HL/HW 0.88); snout short (SL/HL 0.30), rounded to trapezium shaped in dorsal view (Fig. 6), (SL/EE 0.53); canthus rostralis fairly distinct, straight-lined from eye to nostril; nostril oval (0.13 X 0.10), directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 1.20), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.58); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.46); eye diameter exceeding than snout length (ED/SL 1.50); interorbital distance much wider than upper eyelid (IO/EW 1.91), and greater than internarial distance (IO/NN 2.32); tympanum not visible externally; 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, free for about three-fourths of length, bifurcated distally for about one-fourth of length; vocal sac single, median, glandular, pale yellow, roughly triangular; Dorsal surfaces of head, trunk and limbs finely granulated Fore limbs slender; hand moderately large (HND/SUL 0.25); tips of fingers enlarged into broad rounded disks; relative finger lengths are II 2 +��� 2 II 2.5 ��� 2 III 1.5��� 2 IV; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.48); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.48), longer than thigh (TFL/THL 1.14); foot shorter than tibiofibula (FOT/TFL 0.70); relative length of toes: II 1 ���1.5 II 2 ��� 1.5 III 2.5 ��� 2 IV 2.5 ��� 2.5 V; inner metatarsal tubercle small, oval, prominent; outer one larger, almost circular, low and less distinct. Eggs and tadpoles. Unknown. Colouration in life. Dorsal surface light green, skin more or less translucent. Dorsal surface of body and limbs with very small dark brown to black dots and slightly larger brown to reddish brown specks on dorsum; dots roundish, specks shaped like stars or neurons with many dendrites, limbs green; lateral sides of snout and area above eyes reddish brown, moderately broad dorsolateral stripe running along each side of the body from groin to eye extending to the snout as a thin white line; distal portions of fingers and toes, especially the tips, yellow; ventral side and parts of dorsal side of thigh and upper arm largely unpigmented. Iris reddish-brown. Colouration in preservative. The back is yellow brown with darker uniform speckles that extend on to the upper surfaces of the limbs. Remarks. Specimens are similar to the type description, and one was collected from the type locality Calandula (Duque de Bragan��a). The back is covered with small and large spots. The synonymy of H. punctulatus is supported by the spotted back. The species is known from northern Botswana and northern Angola. Due to the extensive range and large population numbers, 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 334-335, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

PhD (Zoology), North-West University, Potchefstroom Campus Blood parasites have been recorded in a variety of vertebrate and invertebrate hosts, inhabiting both aquatic and terrestrial environments. Until this study, only a few blood parasite surveys had been carried out on frogs in sub-Saharan Africa. Thus information on the diversity of these parasites remained limited. To increase our knowledge of frog blood parasites, a large multi-approach study on the diversity, evolutionary biology, and ecology of frog blood parasites was undertaken. The majority of the fieldwork took place in northern KwaZulu-Natal (KZN), South Africa, focussing specifically on the area adjacent to the Phongolo River and its associated floodplain. However, samples also included those collected from frogs in the southern regions of the Kruger National Park, South Africa, and from frogs in Belgium. The latter was fortuitous, as Europe is the type locality for many frog blood parasite species and genera. These samples provided essential data for phylogenetic comparisons between the African and European species. Presently this is the largest multi-species, generic and family amphibian blood parasite survey to be completed, including a total of 643 anurans of 38 species, 20 genera and 13 families. The study was divided into three main components for the collection, analysis and reporting of data. The first component was to determine the frog blood parasite diversity, the second to determine phylogenetic relationships in conjunction with the former component, and lastly the ecological and host-vector-parasite relationships. Blood samples were drawn from the femoral artery of each frog and thin blood smears prepared for screening and morphometrics; the remaining blood fixed in 70% molecular grade EtOH for later molecular analysis. Giemsa stained blood smears were screened microscopically for the presence of any blood inhabiting organisms. Positive infections were then further analysed according to the aims of the respective chapters. Analyses included both morphological and molecular aspects. Morphology was used for the description and identification of species, and molecular analyses were used to assist with the morphologybased descriptions, as well as to allow for phylogenetic relationship comparisons of the blood parasites with one another. In the present study, three new species of Hepatozoon were described from hyperoliid frogs (Afrixalus fornasini, Hyperolius argus, and Hyperolius marmoratus), namely Hepatozoon involucrum Netherlands, Cook & Smit, 2018; Hepatozoon tenuis Netherlands, Cook & Smit, 2018; and Hepatozoon thori Netherlands, Cook & Smit, 2018. Phylogenetic relationships show that species of Hepatozoon isolated from African frogs form as a monophyletic group, separate from the species of Hepatozoon isolated from European and North American frogs. Two species of Dactylosoma Labbé, 1894, were found parasitising three species of frogs namely, Ptychadena anchietae and Sclerophrys gutturalis from South Africa, as well as Pelophylax lessonae from Belgium. Based on morphometrics and molecular findings a new dactylosomatid, Dactylosoma sp. 1, is described form Pty. anchietae and Scl. gutturalis. The species of Dactylosoma isolated from Pel. lessonae conforms morphologically with Dactylosoma splendens Labbé 1894, thus placing in question the validity of D. splendens synonymy with D. ranarum (Kruse, 1890). Phylogenetic analysis shows species of anuran Dactylosoma as a monophyletic group, separate from the other haemogregarine groups. Five species of frogs from South Africa and two from Belgium were found parasitised with haemococcidia. Based on morphological, morphometric and molecular findings Lankesterella minima (Chaussat 1850) is redescribed from Pelophylax kl. esculentus (Linnaeus, 1758) and Pel. lessonae (Camerano, 1882) from Belgium. Additionally, two new species of Lankesterella were described, namely Lankesterella sp. 1 in Pel. lessonae from Belgium, and Lankesterella sp. 2 in Afr. delicates and Afr. fornasini from South Africa. Furthermore, a new genus of haemococcidia, with a new species combination, is described from Phr. mababiensis, Pty. anchietae, and Pyx. edulis from South Africa; as well as a new species, haemococcidia sp. 2, described from Afr. fornasini from South Africa. This is the first study to provide molecular data for species of haemococcidia from African and European anurans. A new species of amphibian filarial nematode (Onchocercidae: Waltonellinae) was described from the toads Scl. gutturalis and Scl. garmani. The life history of this nematode was elucidated from its natural mosquito vectors Uranotaenia (Pseudoficalbia) mashonaensis and Uranotaenia (Pfc.) montana. All stages of development were characterised using morphological and molecular methods. This study is the first to elucidate the life history of an amphibian filarial nematode from southern Africa, and provide data on its phylogenetic placement within the Onchocercidae. In addition to the taxonomic and phylogenetic perspective of this study, this study also aimed at exploring the potential of frog blood parasites as indicators of environmental health. For this, blood parasites infecting grass frogs (Ptychadena Boulenger, 1917) from the Phongolo River system in South Africa were used as a case study. In general, findings indicate that frogs from more impacted sites harboured more blood parasites than from lessimpacted sites. In summary, this study explored the efficacy of a large multi-species, multi-approach survey on the diversity of frog blood parasites from northern KwaZulu-Natal, South Africa. Based on the results several new species of frog blood parasites from different taxa were discovered and described, greatly contributing to knowledge and species records on the overall diversity of frog blood parasites from South Africa. Furthermore, this study provides the first molecular data for species of Dactylosoma and Lankesterella for frogs from Africa, as well as the first molecular data for a filarial nematode for frogs from South Africa. The phylogenetic relationships of species of Hepatozoon, Dactylosoma, Lankesterella, and the filarial nematode were also characterised based on comparisons to other available molecular data. From an ecological perspective, blood parasites from this study adhere to several criteria of what is considered a good indicator and thus demonstrate potential as indicators for healthy ecosystems and intact food webs. The results of this study establish a foundation for future research into the blood parasite biodiversity in northern KZN, an area that this study has highlighted as not only rich in anuran diversity, but also rich in anuran blood parasite diversity. Furthermore, this study provides a baseline for future taxonomic and ecological studies on these parasite groups, not only in South Africa but globally as well. DAAD-NRF, VLIR-UOS university scholarship Doctoral