Search

he chytrid fungus Batrachochytrium salamandrivorans (Bsal), recently introduced from Asia to Europe, causes mortality in numerous species of salamanders and newts and has led to catastrophic declines and local extinctions of the European fire salamander (Salamandra salamandra) in the Netherlands, Belgium, and Germany. Due to the continuous spread of the pathogen, Germany can be considered as the current ‘hotspot’ of Bsal-driven salamander declines. The pathogen was detected in 2015 in the Eifel Mountains where it probably has been present at least since 2004. Moreover, Bsal was found in 2017 in the Ruhr District where it also might occur since 2004. The Ruhr District is a heavily urbanized and industrialized region in western Germany, which offers an unprecedented opportunity to monitor range expansion and infection dynamics of Bsal in an area affected by intense human activities. We here review the current knowledge on Bsal in the Ruhr District where the pathogen by now has been recorded based on qPCR data from 18 sites distributed over eight cities. Transect counts (adult salamanders) and larval removal-sampling at two sites where Bsal was recorded in 2017 and 2018, confirm that fire salamander populations at the affected sites have declined dramatically. However, single negative-tested individuals were still observed and reproduction could be ascertained. Moreover, we successfully detected Bsal by analysing environmental DNA (eDNA) from samples obtained from a standing water body as well as a stream. Detailed monitoring of a site in Essen (Kruppwald) from January to May 2019 provided data on infection and disease dynamics during an acute Bsal-outbreak in a population of European fire salamanders. After initial observation of single dead infected salamanders in January and February 2019, the maximum Bsal loads in the population ranged from 7.90E+03 ITS copies in early March to 2.29E+09 ITS copies at the end of March. Prevalence of infection ranged from 4% to 50% and significantly increased over time; prevalence of externally visible disease symptoms peaked on May 2 and May 8. Single dead salamanders were encountered throughout the monitoring period. Recaptures of two infected salamanders indicated an increase of Bsal load by about one order of magnitude within one week. Infected salamanders showed small-sized regular round ulcerations usually of 0.25–1 mm but sometimes up to 2.5 mm in diameter, which gave the impression of outward growth from the centre of each ulceration. Among salamander individuals monitored in the Kruppwald, such ulcerations were only found in infected salamanders, but we found no significant correlation between the intensity of the ulcerations and Bsal load. Heat treatment proved effective to cure even deep ulcerations when salamanders were kept for 10 days at 25–27°C or 14 days at 25°C, but infection persisted and ulcerations reappeared six weeks after the end of the treatment; only heat treatment at 25°C for 21 days proved effective to reliably clear the infection in three tested salamanders. Key words. Amphibia, Caudata, Salamandra salamandra, European fire salamander, Bsal, chytridiomycosis, heat treatment, emerging infectious disease, amphibian disease, eDNA. Deutsche Bundesstiftung Umwelt

Lycian salamanders (genus Lyciasalamandra) constitute an exceptional case of micro-endemism of an amphibian species on the Asian Minor mainland. These viviparous salamanders are confined to karstic limestone formations along the southern Anatolian coast and some islands. We here study the genetic differentiation within and among 118 populations of all seven Lyciasalamandra species across the entire genus’ distribution. Based on circa 900 base pairs of fragments of the mitochondrial 16SrDNA and ATPase genes, we analysed the spatial haplotype distribution as well as the genetic structure and demographic history of populations. We used 253 geo-referenced populations and CHELSA climate data to infer species distribution models which we projected on climatic conditions of the Last Glacial Maximum (LGM). Within all but one species, distinct phyloclades were identified, which only in parts matched current taxonomy. Most haplotypes (78%) were private to single populations. Sometimes population genetic parameters showed contradicting results, although in several cases they indicated recent population expansion of phyloclades. Climatic suitability of localities currently inhabited by salamanders was significantly lower during the LGM compared to recent climate. All data indicated a strong degree of isolation among Lyciasalamandra populations, even within phyloclades. Given the sometimes high degree of haplotype differentiation between adjacent populations, they must have survived periods of deteriorated climates during the Quaternary on the spot. However, the alternative explanation of male biased dispersal combined with a pronounced female philopatry can only be excluded if independent nuclear data confirm this result. © 2020 Veith et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Channing, A., Hillers, A., Lötters, S., Rödel, O., Schick, S., Conradie, W., Rödder, D., Wagner, P., Dehling, J. M., Du Preez, L. H., Kielgast, J., Burger, M. (2013): Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species. Zootaxa 3620 (3): 301-350, DOI: http://dx.doi.org/10.11646/zootaxa.3620.3.1

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

Hyperolius poweri Loveridge, 1938 Power's Long Reed Frog (Fig. 9) Genetic material. ZMB 77312 ��� 3 (Port Edward, South Africa); PEM A 9545 ��� 6 (Mkambati Nature Reserve, South Africa) (Fig. 1). Diagnosis: The advertisement call (Fig. 13) consists of an initial brief note with seven pulses, followed by five slower pulses, with a duration of 0.12 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni, H. lupiroensis, and H. nasutus. It differs from species producing a call over 0.2 s; H. benguellensis, H. inyangae and H. viridis. See Table 3 for a summary of call parameters. The snout is bluntly rounded, distinguishing it from those with truncated, shark-like or sharply rounded snouts: H. acuticeps, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae, H. lupiroensis, H. nasutus, and H. rwandae sp. nov. There is a phalanx free of web on the first and third toes, with slightly more than a phalanx free on the fourth toe. The second and fifth toes have about half a phalanx free of web. It can be distinguished from the species that have at least one toe webbed to the disc: H. adspersus, H. benguellensis, H. friedemanni, H. jacobseni, H. lupiroensis, H. nasutus and H. rwandae sp. nov. It differs from the species that have the fifth toe with one or more phalanges free of web: H. acuticeps, H. dartevellei, H. howelli, and H. inyangae. Description of a specimen from Mkambati. This is a male, PEM A 9545, collected at the Mkombati Nature Reserve, Eastern Cape Province, South Africa by J. Venter and W. Conradie, 8 February 2011. Body long and slender, widest at mid-body, slightly tapering to groin; head comparatively small (HL/SUL 0.30, HW/SUL 0.30), not wider than trunk, length subequal to width (HL/HW 0.98); snout long (SL/HL 0.49), sharply rounded in dorsal view, blunt in profile (Fig. 6), projecting beyond lower jaw, wider than long (SL/EE 0.75); canthus rostralis distinct, rounded, slightly concave from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed laterally; situated much closer to tip of snout than to eye (EN/NS 1.46), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.16); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.34); eye diameter shorter than snout (ED/SL 0.70); interorbital distance wider than upper eyelid (IO/EW 1.14), and greater than internarial distance (IO/ NN 1.09); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth; vomer processes and teeth absent; tongue long 5.1, and narrow (2.3 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular, mostly unpigmented and translucent when fully inflated; gular flap consisting of two areas of thickened skin, immediately adjacent to each other; anterior part cream-coloured, larger, more granular, and thicker than posterior whitecoloured part; in resting position only a narrow band of the posterior part visible from below; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like. Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs and gular smooth. Fore limbs slender; hand moderately large (HND/SUL 0.27); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 ��� 2 II 2.5��� 3 III 3 ��� 2.5 IV; thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.5); tibio-tarsal articulation passing level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.52), subsequal to thigh (TFL/THL 1.04); heels overlapping each other when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.81); relative length of toes: II 1 ���1.5 II 0.75��� 2 III 1��� 2 IV 1.5 ��� 0.5 V; inner metatarsal tubercle small, oval, prominent; outer one indistinct. Colouration in life. In life the body is dark green with pale flecks, and fine brown pigment spots. The lateral stripes are shiny white, with a subdermal paradorsal band visible as an irregular pale green band. The toes have reddish tips. Colouration in preservative. In preservative the lateral stripes are shiny white, originating at the nostrils, being pale and subdermal before running over the eyes, and extending back to the groin. The back is densely covered in small chromatophores, with very dark pigment over the snout. The gular region is pale with a few dark spots Eggs and tadpoles. The eggs are white with a grey animal pole, less than 1 mm in diameter, within capsules 2.2 mm in diameter (Wager 1986). Clutch size is about 200, with the eggs being deposited in small groups attached to vegetation under water (Wager 1986). Wager (1986) described the tadpoles. Habitat. The frogs are found on reeds and other emergent vegetation around pools and swamps. Distribution. This species is only confirmed from the east coast of South Africa, from Mkambati in the south, northwards to the Mozambique border. The northern extent of the distribution is unknown. Remarks. The species is only known from the north-eastern coastal strip of South Africa. Due to the disturbed coastal habitat, this species should be regarded as Data Deficient in terms of the IUCN criteria, until further studies are carried out. Holotype. ZMB 77221, adult male, from a pond in farmland on the eastern outskirts of Butare, Huye District, South Province, Rwanda (2 �� 37 ' 10.79 '' S, 29 �� 45 '08.45'' E), collected 13 September 2010 by J.M. Dehling. Genetic material. ZMB 77221 ��� 2 (Butare, Rwanda); ZMB 77223 ��� 4 (Mugesera wetland, Rwanda); ZMB 77225 (Akagera wetland, Rwanda) (Fig. 1). Paratypes. ZMB 77222, adult male, same data as holotype; ZMB 77423 ���24, 77426��� 29, six adult males, ZMB 77425, adult female, all from farmland on the eastern outskirts of Butare, Huye District, South Province, Rwanda, collected in October 2009 by K. L��mkemann, K. Rosar and C. Schwartz; ZMB 77686 ��� 89, four adult males, from farmland on the eastern outskirts of Butare (2 �� 35 ' 44.1 '' S, 29 �� 45 ' 25.6 '' E), collected 27 February 2012 by J.M. Dehling; ZMB 77223, adult female, from the Mugesera wetland south of Lac Mugesera, Bugesera District, East Province, Rwanda (2 �� 12 ' 18.92 '' S, 30 �� 16 ' 18.18 '' E), collected 27 March 2011 by J.M. Dehling; ZMB 77224, adult male, from the Mugesera wetland, Bugesera District, East Province, Rwanda (2 �� 12 ' 15.95 '' S, 30 �� 15 ' 49.25 '' S), collected 27 March 2011 by B. Dumbo and J.M. Dehling; ZMB 77683 juvenile, ZMB 77684 adult female, ZMB 77685 adult male, all from the Mugesera wetland, Bugesera Province, southeastern Rwanda, collected 26 February 2012 by J.M. Dehling; ZMB 77225, adult male, from a wetland of the Akagera River, Kihere District, East Province, Rwanda (2 �� 13 ' 27.63 " S, 30 �� 49 ' 39.06 " E), collected 31 March 2011 by J.M. Dehling; ZMB 77746 ��� 48, three adult males, from a swamp in farmland on the eastern outskirts of Ruhengeri, Musanze District, North Province, Rwanda (1 �� 30 ' 25.73 " S, 29 �� 39 ' 12.11 " E), collected 30 March 2012 by J.M. Dehling. Diagnosis: The advertisement call (Fig. 13) consists of an initial brief note of 13 pulses, followed by three slower pulses, with a duration of 0.14 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni, H. lupiroensis, and H. nasutus. It differs from species producing a call over 0.2 s: H. benguellensis, H. inyangae and H. viridis. It differs from the species that have five or more slower pulses: H. friedemanni, H. igbettensis and H. poweri. The initial note of the call of H. howelli consists of only eight pulses, distinguishing it from H. rwandae with 13. See Table 3 for a summary of call parameters. The snout is sharply rounded in profile, which distinguishes it from those species with truncated, shark-like, or bluntly rounded snouts: H. adspersus, H. benguellensis, H. dartevellei, H. howelli, H. igbettensis, H. inyangae, H. jacobseni, H. poweri, and H. viridis. The third and fifth toes webbed three-fourth the way between disc and distal subarticular tubercle, distinguishing it from the species where the webbing does not reach beyond the distal subarticular tubercles of the third and/or fifth toe: H. acuticeps, H. benguellensis, H. dartevellei, H. howelli, H. igbettensis, H. inyangae, H. nasutus, H. poweri, and H. viridis. It differs from H. friedemanni which has all the toes webbed to the disc, and from H. lupiroensis and H. nasutus which have three phalanges free of web on the inner side of the fourth toe. Standard measurements of the holotype are compared with the other species in Appendix 2. Description of Holotype. Body long and slender, widest at temporal region, slightly tapering to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.30), not wider than trunk, longer than wide (HL/HW 1.10); snout long (SL/HL 0.44), pointed in dorsal view, acute in profile (Fig. 6), considerably projecting beyond lower jaw, wider than long (SL/EE 0.77); canthus rostralis distinct, moderately sharp, almost straight-lined from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 1.42), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.13); eyes directed anterolaterally, moderately protruding, relatively small (ED/HL 0.31); eye diameter shorter than snout (ED/SL 0.70); interorbital distance much wider than upper eyelid (IO/EW 1.71), and greater than internarial distance (IO/NN 1.16); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae small, oval, located far anterolaterally at margins of roof of the mouth, concealed by upper jaw for about the half in ventral view; vomer processes and teeth absent; tongue long 4.9, and narrow (2.4 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular, mostly unpigmented and translucent when fully inflated; gular flap consisting of two medially arranged, subcircular areas of thickened skin, immediately adjacent to each other; anterior part cream-coloured, larger, more granular, and thicker than posterior white-coloured part; in resting position only anterior part visible from ventral; vocal sac aperture on each side of the mouth, situated lateral from and close to base of tongue, slit-like, long, directed posterolaterally. Dorsal surfaces of head, trunk and limbs generally appearing smooth but with many densely and more or less evenly scattered tiny, low, spine-like tubercles, hardly visible with the naked eye; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.29); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 2 +��� 2 II 2���2.75 III 2 ��� 2 - IV (after Myers & Duellman [1982]); thenar tubercle indistinct, low; palmar tubercles absent; metacarpals without supernumerary tubercles; nuptial pads or asperities absent. Hind limbs slender, moderately long (LEG/SUL 1.63); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.54), longer than thigh (TFL/THL 1.11); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.81); relative length of toes: II 1.5���2 + II 1.25���2 + III 1.25��� 2 IV 2 -��� 1.25 V; inner metatarsal tubercle small, oval, prominent; outer one larger, almost circular, low and less distinct. Colouration in life. Generally weakly pigmented and skin more or less translucent. Dorsum and dorsal surface of head and limbs yellowish green; lateral sides of head and scapular region light green; light, yellowishwhite, moderately broad dorsolateral stripe running along each side of the body from lateral edge of upper eyelid to groin, continued as faint, hardly discernible line from eyelid to tip of snout; very small dark brown to black dots and larger brown to reddish brown specks on dorsum, most densely along both sides of canthus rostralis and upper eyelid and to lesser extent on both sides of dorsolateral stripe; dots roundish, specks shaped like stars or neurons with many dendrites; distal portions of fingers and toes, especially the tips, yellow; ventral side and parts of dorsal side of thigh and upper arm largely unpigmented, appearing bluish-green; peritoneum white, shining through the translucent belly skin; most of internal organs covered with silvery-white tissue (only visible when dissected). Iris reddish-brown during the night, yellowish-brown during the day. Colouration in preservative. All colours have faded to yellow; gular flap whitish-yellow. Paratype variation. The paratypes are similar to the holotype in measurements (Appendix 2). Female type specimens (SUL 18.2���20.4, mean 19.2, n= 3) are about as large as males (SVL 18.4 ���22.0, mean 19.5, n= 15). Colouration of male paratypes is similar to that of the holotype. In some specimens, however, the pattern of dots and speckles is more pronounced. In others, the lateral stripe is less distinct. The light canthal stripe is completely absent in ten male paratypes and in seven paratypes as faintly visible as in the holotype. All females observed in the field, including the female paratypes, lack the light dorsolateral and canthal stripes, gular sacs and flaps, and the spiny dorsal tubercles (Fig. 8). In life, the flanks of the body turn reddish in active males, especially those which are calling. Eggs and tadpoles. Several females with enlarged ovarian eggs were observed but only three of them were collected (ZMB 77143, 77425, 77684). Their ovaries contain about 80 enlarged eggs with a diameter of ca. 0.7���0.8. Eggs are darkly pigmented on the animal pole and white on the vegetative pole. Tadpoles are unknown. Habitat. We found the species only in open habitats, in natural wetlands (Mugesera, Akagera) as well as at the edge of ponds and other lenthic water bodies in cultivated areas. Specimens were observed perching on leaves of vegetation between 5 cm and 1.2 m above the ground or the water level. Males called from elevated positions, sometimes in close proximity to each other (ca. 15 cm). Several males were found engaged in combat. They were holding, pushing, and kicking each other, apparently fighting over an apparently favoured calling site. They also emitted aggressive calls which differed markedly from the advertisement call. The male aggression call is shown in Fig. 14. The following species were found sympatricaly or even syntopically with the new species: Afrixalus quadrivittatus, Amietia cf. angolensis, Amietophrynus kisoloensis, A. regularis, Hyperolius cinnamomeoventris, H. kivuensis, H. lateralis, H. viridiflavus, Kassina senegalensis, Leptopelis kivuensis, Phrynobatrachus cf. mababiensis, P. natalensis, Phrynobatrachus sp., Ptychadena anchietae, P. porosissima, P. cf. mascareniensis, Ptychadena sp. and Xenopus victorianus. Distribution. We observed the species at three further locations in Rwanda, near Gitarama (2 ��05' 57.14 '' S, 29 �� 46 ' 41.94 '' E, Muhanga District, Southern Province, central Rwanda) and west of Kigali (1 �� 57 ' 49.11 '' S, 30 ��00'05.87'' E, Kamonyi District, Southern Province, central Rwanda; and 1 �� 56 ' 59.33 '' S, 30 ��00' 48.97 '' E, Nyarugenge District, Kigali Province, central Rwanda). The localities from where the species is known are in the northern, central, southern and eastern parts of Rwanda. Elevations of the sites ranged from 1300 m (Akagera wetland) to 1800 m (Ruhengeri). Population size was high at all sites. Because the locations in Butare, Mugesera, and Akagera are only 17 km and 15 km from the border with Burundi and 1.6 km from the border with Tanzania, respectively, and especially because the wetlands of Mugesera and Akagera continue into Burundi and Tanzania, respectively, we assume that the species occurs in these countries as well. Etymology. The species epithet derives from Rwanda. It is a noun in genitive singular. Remarks. Although the species is so far only known from several localities in Rwanda, it is probably more widespread. The species occurs in both natural and cultivated areas. Therefore, we propose that it should be classified as Least Concern under the current criteria of the IUCN redlist., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., Wagner, P., Dehling, J. M., Du Preez, L. H., Kielgast, J. & Burger, M., 2013, Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species, pp. 301-350 in Zootaxa 3620 (3) on pages 335-339, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

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

Hyperolius igbettensis Schi��tz, 1963 Igbetti Long Reed Frog (Fig. 12) Genetic material. Two samples without vouchers, and ZMB 76542 ��� 43 (Lamto, Ivory Coast); ZMB 77415 (K��rouane, Guinea); ZMB 77416 (Konsankoro, Guinea); ZMB 77410 (Dantilla, Guinea) (Fig. 1). Diagnosis. The advertisement call (Fig. 10) consists of an initial brief note with 12 pulses, followed by five slower pulses, with a duration of 0.12 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. jacobseni sp. nov., H. lupiroensis sp. nov. and H. nasutus. It differs from species producing a call over 0.2 s; H. benguellensis, H. inyangae sp. nov. and H. viridis, and from those where the initial note consists of less than 10 pulses: H. friedemanni, H. howelli, and H. poweri. The snout is bluntly round in profile, which distinguishes it from species with truncated, shark-like, or sharply rounded profiles; H. acuticeps, H. adspersus, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae sp. nov., H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov. The foot is webbed with one or more phalanges free of web on the first four toes, and half free on the fifth toe. This distinguishes it from species where at least one toe is webbed to the disc, at least on one side: H. adspersus, H. benguellensis, H. friedemanni, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus and H. rwandae sp. nov. It differs from species that have at least one phalanx free on the fifth toe; H. acuticeps, H. dartevellei, H. howelli, H. inyangae and H. nasutus. Finally, it differs from the two species that have one or less phalanges of the second toe free of web: H. poweri and H. viridis. Description of a Dantilla specimen. This is based on ZMB 77410, an adult female. The ranges are given from three specimens (ZMB 77410 ��� 412; 1 female, 2 males), with single measurements from the sequenced specimen. Elongate, fragile frogs; Body long and slender (SUL 21.1), widest at temporal region (HW 5.8 ���7.0; 7.0), slightly tapering to groin; head comparatively small (HL/SUL 0.33, HW/SUL 0.26), not wider than trunk, longer than wide (HL/HW 1.27); snout long (SL/HL 0.43), subelliptical in dorsal view and protruding in lateral view (Fig. 6), projecting beyond lower jaw, wider than long (SL/EE 0.73); canthus rostralis indistinct, roundish, straight-lined from eye to just beyond nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril round, directed dorsolaterally; situated much closer to tip of snout than to eye (EN/NS 1.91), separated from each other by distance slightly less than distance between eye and nostril (NN/EN 0.90); eyes directed anterolaterally, protruding, relatively small (ED/HL 0.29); eye diameter shorter than snout (ED/SL 0.67); interorbital distance narrower than upper eyelid (IO/EW 0.8), and greater than internarial distance (IO/NN 1.11); tympanum barely visible, very small with tympanum-eye distance equal to half diameter of eye; upper jaw with dentition; choanae small, oval, located far anterolaterally at margins of roof of the mouth, concealed by upper jaw for about the half in ventral view; vomer processes and teeth absent; tongue long 5.0, and wide (3.6 at widest point), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single of males, median, subgular, mostly unpigmented and translucent when fully inflated; gular gland large covering 2 / 3 to almost entire throat, dilatable skin visible posterior to gland; width of male gular flap 3.6���4.8; gular flap consisting of two medially arranged, heart shaped and triangular areas of thickened skin, immediately adjacent to each other; anterior, heart shaped, light yellow, larger, more granular, and thicker than posterior, triangular white-coloured part; in resting position only anterior part visible from ventral; Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs and gular smooth, lower belly slightly more areolate; a few warts in angle of mouth; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.28); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: 1 ) I 1 +��� 1.25 II 0.25��� 2 III 2 ��� 1 IV 1 ��� 0.3 V; no visible internal or external metatarsal tubercles. Colouration in life. The basic colour of live frogs ranges from a light bluish green, to grass green or almost green-brown; shanks, lower and upper arms are almost transparent blue-green; flanks, back and thighs darker green with many small dark spots, sometimes arranged along vertebral line into a broken line; eyelids usually lighter than rest of head and dorsum, yellowish to reddish brown; sometimes head darker (reddish brown) than rest of dorsal surfaces; in some animals, mostly males, light white to yellow dorsolateral stripe, rarely bordered by two dark lines; dark canthal stripe, reddish iris bordered by narrow blue line; gular gland of males yellowish or like rest of vocal sac skin light green-blue; ventral surfaces light, belly whitish, often almost transparent; toe and finger tips yellow to orange; females are usually more ���transparent��� than males, with eggs visible through the body wall. Colouration in preservative. Very pale beige in preservation with small dark spots scattered over back and extremities, with or without a distinct white dorsolateral band. Eggs and tadpoles. Unknown. Remarks. The biology of this species was discussed by R��del et al. (2006). The species is now known from Guinea to Cameroon, and perhaps occurs further east (Amiet 2006 a). We suggest that the IUCN status of Least Concern be maintained., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., Wagner, P., Dehling, J. M., Du Preez, L. H., Kielgast, J. & Burger, M., 2013, Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species, pp. 301-350 in Zootaxa 3620 (3) on pages 324-326, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

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

Hyperolius dartevellei Laurent, 1943 Dartevelle's Reed Frog (Fig. 9) Synonomy. Hyperolius sagitta Laurent, 1943 Genetic material. ZMB 77303 Ikelenge, Zambia; USNM 576167 ��� 70 (Impongui, Republic of Congo); field numbers A 27, CRT 3577 -9, 3604- 6 (Congo River near Yekela, DRC); CRT 3730, 3798 (Congo River, near Nganda Kona, DRC); CRT 3838 - 9 (Congo River near Ngengele, DRC); CRT 3975 ��� 89 (Congo River near Bomani, DRC); CRT 4024, 4027 (Congo River, near Lulu, DRC); CRT 4205 ��� 10 (Congo River, near Lieki, DRC) (Fig. 1). Diagnosis. A typical advertisement call (Fig. 5) consists of 13 pulses in 0.1 s, with an emphasised frequency of 4.8 kHz. It differs from those species with a brief note consisting of a few initial pulses, followed by a number of pulses at a much slower pulse rate, such as H. benguellensis, H. friedemanni sp. nov., H. howelli sp. nov., H. igbettensis, H. inyangae sp. nov., H. rwandae sp. nov., H. viridis and H. poweri, and those with a longer call consisting of multiple pulses that may change tempo, such as H. acuticeps, H. jacobseni sp. nov., and H. nasutus. See Table 3 for a summary of call parameters. The advertisement call structure is similar to that of H. adspersus and H. lupiroensis sp. nov., while the 16 S sequence of H. lupiroensis sp. nov. differs by more than 11 %. The snout is truncated, distinguishing it from the species with shark-like or rounded snout profiles: H. acuticeps, H. adspersus, H. benguellensis, H. friedemanni sp. nov., H. howelli sp. nov., H. igbettensis, H. inyangae sp. nov., H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus, H. poweri, H. rwandae sp. nov. and H. viridis. The webbing shows a phalanx free on the first, third and fifth toes, with half a phalanx free on the other two. It can be distinguished from the species that have less than a phalanx of the fifth toe free: H. adspersus, H. friedemanni sp. nov., H. igbettensis, H. jacobseni sp. nov., H. lupiroensis sp. nov., H. nasutus, H. poweri, H. rwandae sp. nov. and H. viridis. It differs from the species that have one phalanx or more of the fourth toe free of web: H. acuticeps, H. benguellensis, H. howelli sp. nov., and H. inyangae sp. nov. Description of a Carumbo specimen. An adult male PEM A 10059 (measurements presented in Appendix 2) measuring 18.6 mm SUL; body long and slender, widest just behind orbital region, tapering to groin; head relatively small (HL/SUL 0.32, HW/SUL 0.34), not much wider than long (HL/HW 0.95); snout long (SL/HL 0.46), bluntly pointed in dorsal view (Fig. 6), protruding just beyond lower jaw, wider than long (SL/EE 0.72); canthus rostralis distinct; loreal large and oval in shape; nostril directed dorsolaterally, moderately large vertical slit (0.4 mm in length), situated much closer to tip of snout than to eye (EN/NS 1.60), separated from each other by distance greater than distance between eye and nostril (NN/EN 1.60); eyes large (ED 1.8), directed anterolaterally, protruding outwards and forward, pupil is horizontal to circular, visible from below, eye diameter shorter than snout (ED/SL 0.64); interorbital distance much wider than upper eyelid (IO/EW 1.50), and greater than internarial distance (IO/NN 1.41); tympanum not visible externally; upper jaw with dentition; teeth on premaxilla larger than those on maxilla; choanae large, oval, vomer processes and teeth absent; tongue long 3.9 and broad 2.8, mostly free except for first quater, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular; large granular gular flap covering thin vocal sac (5.9 wide) Dorsal surfaces of head, trunk and limbs generally appearing smooth but with many densely and more or less evenly scattered tiny, asperities; ventral surface of limbs and gular smooth, chin and abdomen slightly more areolate; supratympanic fold absent. Fore limbs slender; hand moderately large (HND/SUL 0.26); tips of fingers enlarged into broad oval disks, no circummarginal groove; relative length of fingers: II 1.5 ��� 0.25 II 0.25 ��� 0.25 III 0.25 ��� 0.25 IV (after Myers & Duellman 1982) thenar tubercle indistinct; palmar tubercles absent. Hind limbs slender, moderately long (LEG/SUL 1.50); tibio-tarsal articulation reaching to level of tip of snout when legs are adpressed to body; tibiofibula moderately long (TFL/SUL 0.52), longer than thigh (TFL/THL 1.07); heels overlapping each other considerably when knees are flexed and thighs are held laterally at right angle to body; foot shorter than tibiofibula (FOT/TFL 0.77); relative length of toes: II 0.25��� 1 II 0.25��� 1 III 0.25��� 1 IV 1 ��� 0.25 V; inner metatarsal absent; outer metatarsal tubercle large, almost circular, low and not distinct. Colouration in life. Below translucent silvery-white, above uniform translucent green to brown, scattered darker spots, clear yellow-white dorsolateral line from snout to vent, forming a light canthus on the snout, darker pigmentation anterior-lateral from snout tip to above eye, upper jaw nearly free of any pigmentation, eye iris is yellow to brown; dorsal surface of arms and legs with scattered dark spots, inner thighs unmarked. Colouration in preservative. All colours have faded to a beige yellow with brown dorsal spots still visible. Eggs and tadpoles. Unknown. Habitat: Specimens were collected in the grassland floodplain wetlands surrounding a large natural lake (350 ha) at daytime. Specimens were found half a meter to a meter above water level on vegetation. The only other amphibians found were Phrynobatrachus mababiensis. Additional material was collected at a small pond (Hyperolius species were present in the same area, Hyperolius angolensis and Hyperolius cf. cinereus, both species were calling further away and higher up the vegetation. Hoplobatrachus occipitalis was present in the pond. Distribution. Southern Cameroon, east and south through Gabon to the lower Congo Basin and the most northern parts of Angola, and the north-western Zambian highlands. Remarks: The synonomy of H. granulatus (the holotype RMCA- 152 was examined) is supported by the absence of dorso-lateral stripes and a short rounded snout. The species is presently only confirmed from northern Angola, the Cabinda enclave, and Gabon. There is little doubt that existing records refer to this species, and we suggest that its conservation status of Least Concern remains unchanged., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., Wagner, P., Dehling, J. M., Du Preez, L. H., Kielgast, J. & Burger, M., 2013, Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species, pp. 301-350 in Zootaxa 3620 (3) on pages 319-320, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Hyperolius viridis Schi��tz, 1975 Robust Long Reed Frog (Fig. 9) Genetic material. ZMB 76096 (Vintukutu Forest Reserve, Malawi); ZMB 76102 (Kaningina Forest Reserve, Malawi) (Fig. 1). Diagnosis. The advertisement call (Fig. 15) consists of a brief initial note consisting of 26 pulses followed by five slower pulses, with a duration of 0.41 s. It can be distinguished from species producing only a single note, and those producing only a buzz: H. acuticeps, H. adspersus, H. dartevellei, H. jacobseni, H. lupiroensis, and H. nasutus. It can be distinguished from the other species producing a two-part call, which have a duration less than 0.4 s: H. friedemanni, H. howelli, H. igbettensis, H. inyangae, H. poweri and H. rwandae. It differs from H. benguellensis which only has five pulses in the initial note. See Table 3 for a summary of call parameters. The snout is bluntly rounded, distinguishing it from those with truncated, shark-like or sharply rounded snouts: H. acuticeps, H. benguellensis, H. dartevellei, H. friedemanni, H. howelli, H. inyangae, H. lupiroensis, H. nasutus, and H. rwandae. The webbing has one phalanx free on the first to third toes, just more than one free on the fourth toe, and half a phalanx free on the fifth toe. It can be distinguished from the species that are webbed to the disc on the fifth toe: H. adspersus, H. friedemanni, H. jacobseni, H. lupiroensis, and H. rwandae. It differs from the species that have more than half a phalanx free of web on the fifth toe: H. acuticeps, H. benguellensis, H. dartevellei, H. howelli, and H. inyangae. It can be distinguished from the remaining species that are webbed to the disc on the third or fourth toes: H. adspersus and H. nasutus. Our specimens show the stocky build noted by Schi��tz (1975). Description of a Vintukutu specimen. An adult male ZMB 76096, from Vintukutu Forest Reserve, Malawi. Body short and compact, widest at mid-body, tapering to head and neck; head very small (HL/SUL 0.22, HW/SUL 0.34), narrower than mid part of trunk, wider than long (HL/HW 0.64); snout short (SL/HL 0.59), suboivoid in dorsal view, almost truncate in profile (Fig. 6), only slightly protruding beyond lower jaw, almost as long as wide (SL/EE 0.96); canthus rostralis distinct, rounded, slightly concave between eye to nostril, slightly convex near tip of snout; loreal region almost vertical, slightly concave; nostril directed dorsolaterally; situated closer to tip of snout than to eye (EN/NS 1.18), separated from each other by distance equal to distance between eye and nostril (NN/EN 0.94); eyes directed anterolaterally, moderately protruding, relatively large (ED/HL 0.59); eye diameter equal to snout length(ED/SL 1.0); interorbital distance much narrower than upper eyelid (IO/EW 0.36), but greater than internarial distance (IO/NN 1.3); tympanum not visible externally; upper jaw with dentition; choanae small, round, located far anterolaterally at margins of roof of the mouth, completely concealed by upper jaw in ventral view; vomer processes and teeth absent; tongue slightly longer than wide (2.1), free for about three-fourths of length, bifurcated distally for about one-fourth of length; median lingual process absent; vocal sac single, median, subgular,; The gular flap is large (almost completely covering the throat), glandular and wider (4.7) than long (4.1), white in preservative with many minute melanophores. The skin of the dorsum and upper limbs appears smooth, finely granular under dissecting microscope; flat granular belly; supratympanic fold absent. Fore limbs slender; hand small (HND/SUL 0.19); tips of fingers enlarged into broad oval disks, each with circummarginal groove; relative length of fingers: II 0.5��� 1 II 0.25��� 1 III 0.25��� 1 IV 1 ��� 0.25 V; inner metatarsal tubercle small, oval, not very prominent; outer one not discernible. Colouration in life. The dorsal and ventral surfaces are white, dorsal surfaces (including thighs) densely covered with minute melanophores. Colouration in preservative. All colours have faded to yellow; gular flap whitish. Eggs and tadpoles. Unknown. Remarks. The species is known from southern Tanzania and northern Malawi. We suggest that the IUCN status of Data Deficient be maintained until further studies are undertaken., Published as part of Channing, A., Hillers, A., L��tters, S., R��del, O., Schick, S., Conradie, W., R��dder, D., Wagner, P., Dehling, J. M., Du Preez, L. H., Kielgast, J. & Burger, M., 2013, Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species, pp. 301-350 in Zootaxa 3620 (3) on pages 339-341, DOI: 10.11646/zootaxa.3620.3.1, http://zenodo.org/record/215983

Within the framework of the present study we test whether climatic niche similarity can be identified in a monophyletic group of species inhabiting remarkably restricted ranges by pooling presence data of all species into a single concatenated data set and subsequently jackknifing single species. We expect that, when the jackknifed species differs markedly in its climatic niche from all other species, this approach will result in increased niche homogeneity, allowing assessments of niche divergence patterns. To test our novel jackknife approach, we developed species distribution models for all members of Lycian salamanders (genus Lyciasalamandra), native to Turkey and the adjacent Aegean islands using Maxent. Degrees of niche similarity among species were assessed using Schoener's index. Significance of results was tested using null-models. The degree of niche similarity was generally high among all seven species, with only L. helverseni differing significantly from the others. Carstic lime stones providing specific microhabitat features may explain the high degree of niche similarity detected, since the variables with the highest explanative power in our models (i.e. mean temperature, and precipitation of the coldest quarter) corresponded well with salamander natural history observations, supporting the biologically plausibility of the results. We conclude that the jackknife approach presented here for the first time allows testing for niche similarity in species inhabiting restricted ranges and with few species records available. Our results strongly support the view that detailed natural history information and knowledge of microhabitats is crucial when assessing possible climate change impacts on species. © Gesellschaft für Biologische Systematik 2011.

We reviewed the pertinent faunistic and taxonomic knowledge available from literature and scientific collections on the amphibians from the central and western highlands of Kenya. Fifty-four anuran species in 19 genera and 12 families were recognized. Higher taxa are those also found in adjacent African regions. Exceptions are some genera otherwise known from extreme East Africa which, along with the order Gymnophiona, are absent from the central and western highlands of Kenya. The species Ptychadena schillukorum, P. taenioscelis and Afrixalus osorioi are reported for the first time from Kenyan territory. The taxonomy of several species is poorly resolved, underlining the need for more alpha-taxonomic effort. This should include the application of modern methods, as molecular markers to uncover cryptic species diversity. We calculated 9.04 ± 6.11 (1-30) localities per species (n = 54) and 3.98 ± 2.88 (1-25) species per locality (n = 127), showing that sampling for both species and localities is of different quality and apparently far from complete. Sampling bias due to accessibility may play a role, and more comprehensive field surveys are suggested. It may be considered that certain amphibians reach their geographic range limits just at the periphery of the Kenyan central and western highlands (e.g. 'Congolian' forest taxa) or are restricted to a small distribution within that region.

A molecular phylogeny based on a fragment of the mitochondrial 16S rRNA gene and morphological comparisons suggest that only one polymorphic species of large Dendrobates, commonly assigned to D. tinctorius, occurs on the eastern Guiana Shield and its vicinities. The oldest available name for this taxon is D. tinctorius (Schneider, 1799). We place D. azureus Hoogmoed, 1969 as its junior synonym. Data for D. machadoi Bokermann, 1958 are not available but we suspect that this name has to be treated in the same way. Dendrobates tinctorius exhibits numerous variants differing in color and pattern.

In their Policy Forum "Confronting amphibian declines and extinctions" (7 July, p. 48), J. R. Mendelson III and colleagues offer a strategy for "stopping" the widespread losses of frogs, toads, and salamanders. Disease research and captive breeding figure prominently in their call for action.

A new species of arboreal Leptopelis is described from Kakamega Forest, western Kenya. It is a small, brown forest species formerly referred to L. modestus, but distinguished by differences in advertisement call and the sequence of the mitochondrial 16S rRNA gene. The specific allocation of certain related populations of Leptopelis in East and West Africa is briefly discussed. Key words: Amphibia: Systematics, Bioacoustics, DNA taxonomy

The amphibian diversity in different vegetation communities was investigated in Meru National Park (Kenya), using transect sampling, drift-fences, pitfalls and opportunistic collection. Acacia wooded grassland had highest amphibian diversity (H' = 2.071, D = 6.74), Acacia-Commiphora bushland ranked second (H' = 1.858, D = 5.88) and Combretum wooded grassland ranked third (H' = 1.581, D = 5.08). Acacia wooded grassland had the highest amphibian abundance (n = 173 individuals) and species richness (10 species). Combretum wooded grassland had eight species with less abundance (n = 113 individuals), while the Acacia - Commiphora bushland had seven species only, with higher abundance (n = 144 individuals). Detrended Correspondence Analysis showed the plots, which occur close together with similar species composition. Correspondence Analysis was performed to investigate the association of amphibian species with plant assemblages. There was a positive linear correlation between amphibian species richness and plant species diversity in all vegetation communities. However, as the Spearman Rank Test could not establish any significant correlation between plant and amphibian diversities in the respective habitats, the amphibians recorded in the present study area cannot be used as surrogate organisms to evaluate environmental changes.

Hyperolius mosaicus Perret (figure 1) This reed frog is known from forest in southern Cameroon and the Monts Cristal in Gabon (Perret, 1966; Frétey and Blanc, 2000; Lötters et al., 2001). From the intervening area, no records are available (e.g. De la Riva, 1994). In the past there has been some taxonomic confusion between this species and the similar H. acutirostris Buchholz and Peters, from Cameroon (cf. Perret, 1966), but the two species are well distinguishable (see below). Our specimens from Gabon closely coincide with the types (cf. Lötters et al., 2001). Diagnosis. (1) SVL of males 24.0¡ 0.76 mm (23.1–24.5 mm, n ~3), SVL of females unknown (most probably slightly larger than males); (2) TIBL/SVL 0.50¡0.01 (0.49–0.50, n ~3), HW/SVL 0.34¡0.01 (0.33–0.34, n ~3); (3) dorsal surface with small scattered warts; (4) snout shape dorsally and laterally rounded, nares visible from above; (5) E-N/EYE 0.97¡0.1 (0.9–1.1, n ~3), canthus rostralis straight from tip of snout to nostril and concave from nostril to eye; (6) tympanum distinct, TYMP/EYE 0.43¡0.08 (0.4–0.5, n ~3); (7) FOOT/TIBL 0.8¡0.04 (0.8–0.9, n ~3); (8) foot webbing formula: 1(1), 2i(1) 2e(K), 3i(1K–2) 3e(1), 4i(1K–2) 4e(1), 5(0–K); (9) PhJ is dorsally brown with up to seven light brown areas (i.e. at snout, at shoulders, on mid-dorsum and on posterior back), usually with tiny white spots, ventrally yellowish with black areas, sole and palm black with yellow toe and finger tips; females were not found but in Cameroon PhF is similar to PhJ (Schiøtz, 1999); the gular flap of males is yellowish; the iris is reddish brown (information provided is based on collected and some non-collected males); (10) for sequence of 560 bp fragment of mitochondrial DNA of the 16S ribosomal gene see GenBank under AY323923 (~ZFMK 73140); (11) LTRF unknown for material from Gabon; Cameroonian tadpoles have 1/3 (1) (cf. Perret, 1959: 720). Similar species are the partly sympatric H. acutirostris and H. endjami Amiet from Cameroon (Perret, 1975; Amiet, 1980; Schiøtz, 1999). The former always has a light band in front of the urostyle and the gular flap in adult males is comparatively smaller. Hyperolius endjami lacks white dorsal tubercles and adult males have, in relation, a larger gular flap. Life history. This species is nocturnal and arboreal. At the Monts Cristal, calling males were observed after sunset on vegetation below 1 m above ground in a semi-cleared area within the forest. This site was far away from open waters but single large trees were around. According to Perret (1959), H. mosaicus, in Cameroon, breeds in tree holes. We expect the same in Gabon and that therefore H. mosaicus may be a prolonged breeder. The advertisement call (figure 2A; table 1) is a short 'click' consisting of a single pulsed note, very similar to what we know from Cameroonian specimens (cf. Amiet in Schiøtz, 1999)., Published as part of Lötters, S., Schick, S., Scheelke, K., Teege, P., Kosuch, J., Rotich, D. & Veith, M., 2004, Bio-sketches and partitioning of sympatric reed frogs, genus Hyperolius (Amphibia; Hyperoliidae), in two humid tropical African forest regions, pp. 1969-1997 in Journal of Natural History 38 (15) on pages 1972-1973, DOI: 10.1080/00222930310001613584, http://zenodo.org/record/4676048, {"references":["PERRET, J. - L., 1966, Les amphibiens du Cameroun, Zoologisches Jahrbuch fur Systematik, 8, 289 - 464.","FRETEY, T. and BLANC, T. C., 2000, Les amphibiens d'Afrique centrale (Libreville: ADIE), 39 pp.","LOTTERS, S., GOSSMAN, V., OBAME, F. and BOHME, W., 2001, Zur Herpetofauna Gabuns. Teil I: Einleitung, Untersuchungsgebiet und Methodik, kommentierte Artenliste der gefundenen Froschlurche, Herpetofauna, 23, 19 - 34.","DE LA RIVA, I., 1994, Anfibios anuros del Parque Nacional de Monte Alen, Rio Muni, Guinea Equatorial, Revista Espanola de Herpetologia, 8, 123 - 139.","SCHIOTZ, A., 1999, Treefrogs of Africa (Frankfurt: Chimaira), 350 pp.","PERRET, J. - L., 1959, Batraciens noeveaux du Cameroun, Revue Suisse de Zoologie, 66, 711 - 721.","PERRET, J. - L., 1975, Revision critique de quelques types de reptiles et batraciens africains, Revue Suisse de Zoologie, 82, 185 - 192.","AMIET, J. - L., 1980, Un Hyperolius nouveau du Cameroun: Hyperolius endjami n. sp, Revue Suisse de Zoologie, 87, 445 - 460."]}

Hyperolius cf. cinnamomeoventris Bocage (figure 7) The reed frog H. cinnamomeoventris was originally described from Angola. Records referred to this species are known in addition from Gabon to western Kenya (Perret, 1966; Inger, 1968; Laurent, 1972; Schiøtz, 1999; Frétey and Blanc, 2000). Frogs are found in both savannah and forest. Different authors (e.g. Perret, 1966; Schiøtz, 1975, 1999; Lötters et al., 2001) have pointed out that probably more than one species may be hidden behind this one name. Our unpublished comparisons of specimens from different localities (see Appendix) as well as sequences of mitochondrial DNA of the 16S ribosomal gene of populations from Gabon and Kenya indicate that more than one species is involved. As a result, we here only tentatively refer Kenyan material to H. cinnamomeoventris sensu stricto. Detailed studies of taxonomic relationships of populations and type specimens including those of current synonyms are necessary. Diagnosis. SVL males 23.7¡ 1.2 mm (22.4–24.6 mm, n ~3) (112 noncollected males had mean SVL 23.3¡ 1.2 mm, range 21.0– 25.7 mm), SVL of preserved females unknown (six non-collected females had mean SVL 28.0¡ 1.16 mm, range 26.3–29.4 mm); (2) TIBL/SVL 0.46¡0.01 (0.46–0.47, n ~3), HW/SVL 0.31¡0.02 (0.3–0.33, n ~3); (3) dorsal surface finely coarse, tuberculate below eye; (4) snout shape dorsally and laterally rounded, nares visible from above; (5) E-N/EYE 0.89¡0.05 (0.84–0.95, n ~3), canthus rostralis convex from tip of snout to nostril and straight or slightly concave from nostril to eye; (6) tympanum distinct or covered by thick skin, TYMP/EYE 0.36–0.42 (n ~2); (7) FOOT/TIBL 0.85¡0.04 (0.84–0.95, n ~3); (8) foot webbing formula: 1(1), 2i(1–2) 2e(K), 3i(1–2) 3e(K), 4i(1K) 4e(1), 5(0–K); (9) PhJ is dorsally tan to brown, with more or less intense, diffuse dark brown spotting, and always a cream to white (rarely yellowish) dorsolateral line from near nostrils (before nostril it is more diffuse, there is no evident U-shape) to the groin which usually is bordered by dark brown (Schiøtz, 1975: 123 mentioned that this line can be absent), a mid-dorsal line of the same colour may be present, ventral sides are cream; PhF is dorsally uniformly green, delimited from the ventral side by an irregular black lateral line, always with a black canthal and usually a black labial line (lines may continue behind eye and/or on arm); the inner femur always has a reddish area; the gular flap of males is yellow; the iris is bronze to dark brown (information provided is based on collected and numerous noncollected specimens of both sexes); (10) for sequence of 560 bp fragment of mitochondrial DNA of the 16S ribosomal gene see GenBank under AY323925 (~NMK A/3858/2); (11) LTRF 1/3. There are some species that can look similar to H. cf. cinnamomeoventris, i.e. H. kivuensis Ahl, H. lateralis Laurent and H. schoutendeni Laurent (Inger, 1968; Schiøtz, 1975, 1999). The former two occur in sympatry with H. cf. cinnamomeoventris at the Kakamega Forest (see below). Hyperolius schoutendeni from the Democratic Republic of Congo is a little-known species which seems almost indistinguishable from sympatric H. cf. cinnamomeoventris PhJ (both phases are equal in H. schoutendeni) but a distinct species, as figured out by Inger (1968) and Schiøtz (1975, 1999). We observed that probably the extension of foot webbing is a useful character to distinguish cinnamomeoventris -like forms. We examined the female holotype of H. schoutendeni (SVL 26.4 mm) which has more webbing on toes 4 and 5 (foot webbing formula: 1(1), 2i(1) 2e(K), 3i(1K) 3e(1), 4i(2) 4e(1K), 5(1)). Life history. Hyperolius cf. cinnamomeoventris is a nocturnal arboreal reed frog found in primary forest and sometimes also in semi-cleared areas (e.g. on bushes). In the Kakamega Forest, it could be observed (and males could be heard calling also away from water) from March to October. Breeding is limited to the period from March to June and takes place in swampy areas or at ponds. This species tends to be intermediate between explosive and prolonged breeding strategies. One single clutch is attached to vegetation above the water surface, containing ca 114 half-pigmented eggs, each about 1.55 mm in diameter (without jelly). Tadpoles can be classified as lentic and omnivorous. As shown in figure 2E and table 1, advertisement calls are short 'cracks', each of one pulsed note. They are thus comparable to vocalizations assigned to H. cinnamomeoventris from a nearby site in Uganda (Schiøtz, 1975). Nevertheless, in the figure by Schiøtz (1975), frequency range and dominant frequency are higher (with overlap). 'Clicks' reported by Largen and Dowsett-Lemaire (1991) from the Republic of Congo also seem comparable. However, rattles of four to six elements (notes?) as found by these authors have never been heard by us in Kenyan specimens, providing additional evidence that more than one species is involved., Published as part of Lötters, S., Schick, S., Scheelke, K., Teege, P., Kosuch, J., Rotich, D. & Veith, M., 2004, Bio-sketches and partitioning of sympatric reed frogs, genus Hyperolius (Amphibia; Hyperoliidae), in two humid tropical African forest regions, pp. 1969-1997 in Journal of Natural History 38 (15) on pages 1985-1986, DOI: 10.1080/00222930310001613584, http://zenodo.org/record/4676048, {"references":["PERRET, J. - L., 1966, Les amphibiens du Cameroun, Zoologisches Jahrbuch fur Systematik, 8, 289 - 464.","INGER, R. F., 1968, Amphibia, Exploration au Parc national de la Garamba, 52, 1 - 190.","LAURENT, R. F., 1972, Amphibiens, Exploration du Parc National des Virunga, deuxieme serie, 22, 1 - 125 zpls I - XI.","SCHIOTZ, A., 1999, Treefrogs of Africa (Frankfurt: Chimaira), 350 pp.","FRETEY, T. and BLANC, T. C., 2000, Les amphibiens d'Afrique centrale (Libreville: ADIE), 39 pp.","SCHIOTZ, A., 1975, The Treefrogs of Eastern Africa (Copenhagen: Steenstrupia), 232 pp.","LOTTERS, S., GOSSMAN, V., OBAME, F. and BOHME, W., 2001, Zur Herpetofauna Gabuns. Teil I: Einleitung, Untersuchungsgebiet und Methodik, kommentierte Artenliste der gefundenen Froschlurche, Herpetofauna, 23, 19 - 34.","LARGEN, M. J. and DOWSETT-LEMAIRE, F., 1991, Amphibians (Anura) of the Kouilou River basin, Republique du Congo, Tauraco Research Report, 4, 145 - 168."]}

Hyperolius lateralis Laurent (figure 9) This reed frog is known to range from north-western Tanzania to the eastern Democratic Republic of Congo and to western Kenya; it occurs in bush land and forest (Schiøtz, 1975, 1999). Several subspecies from Rwanda, Burundi and western Uganda were named by Laurent (e.g. 1951). According to Schiøtz (1975, 1999), they seem to reflect variation among populations of a single taxon only. We examined the female holotypes of the forms capnodogaster, pulcherrimus and subolivaceus and have the impression that they could represent distinct taxa. Comprehensive investigation remains to be carried out including also the similar H. castaneus Ahl and its junior synonyms (from eastern Democratic Republic of Congo, Rwanda and western Uganda). Kenyan material is referable to the nominotypical form and well coincides with the female holotype (SVL 25.0 mm). Diagnosis. SVL males 23.6¡ 0.62 mm (22.5–24.2 mm, n ~7); according to Schiøtz (1975: 129), SVL of one female 25.0 mm (a non-collected female had SVL 24.3 mm); (2) TIBL / SVL 0.47¡0.01 (0.46–0.49, n ~7), HW/ SVL 0.32¡0.02 (0.3–0.35, n ~7); (3) dorsal surface finely coarse, tuberculate below eye and tympanic area; (4) snout shape dorsally and laterally rounded, nares visible from above; (5) E-N/EYE 0.68¡0.08 (0.58–0.76, n ~7), canthus rostralis straight from tip of snout to nostril and concave from nostril to eye; (6) tympanum distinct or covered by thick skin, TYMP /EYE 0.31–0.36 (n ~2); (7) FOOT / TIBL 0.86¡0.03 (0.85–0.92, n ~7); (8) foot webbing formula: 1(K–1), 2i(1) 2e(K), 3i(1) 3e(K), 4i(1) 4e(1), 5(K); (9) PhJ and PhF are dorsally olive, brownish or yellowish green, with or without diffuse dark brown and/or bright yellow spotting, sometimes with a faint, thin light dorsolateral stripe, and in PhF a cream and/or bright yellow irregular lateral line or lateral marbling from below or behind eye to groin which always is bordered by dark brown; a dark brown canthal line may be present, ventral sides are translucent pink to reddish including sole and palm, webbing as well as parts of the upper sides of toes and fingers; the gular flap of males is dark yellowish golden; the iris is light brownish grey (information provided is based on collected and numerous non-collected specimens of both sexes); (10) for sequence of 560 bp fragment of mitochondrial DNA of the 16S ribosomal gene see GenBank under AY323924 (~ NMK A/3925/1); (11) LTRF 1/3. Hyperolius lateralis is similar to H. cinnamomeoventris and H. castaneus, which are both sympatric in part. According to Schiøtz (1975, 1999), the latter is very variable and can be indistinguishable from H. lateralis PhJ (H. castaneus does not fall into PhJ and PhF). It seems that a dark canthal line is always present in H. castaneus (versus occasionally present in H. lateralis). For differentiation from H. cf. cinnamomeoventris see below. Life history. As elsewhere, in the Kakamega Forest, H. lateralis is a nocturnal arboreal species. We observed it between April and June only. Males and females were found in disturbed primary forest, and at its edge in swampy areas or at ponds where males called from vegetation v 1.5 m above ground. This reed frog tends to be intermediate between explosive and prolonged breeding strategies. Two clutches were fixed to vegetation above the water surface, containing 94 and 179 eggs (half black, half cream) each ca 1.6–1.9 mm in diameter (without jelly). Larvae are lentic and omnivorous. The advertisement of this species can be described as a 'squashed peep'. It consists of a single note. Graphic and numerical characteristics (figure 2G; table 1) are comparable to data provided by Schiøtz (1975). However, frequency range and dominant frequency are somewhat higher, with overlap., Published as part of Lötters, S., Schick, S., Scheelke, K., Teege, P., Kosuch, J., Rotich, D. & Veith, M., 2004, Bio-sketches and partitioning of sympatric reed frogs, genus Hyperolius (Amphibia; Hyperoliidae), in two humid tropical African forest regions, pp. 1969-1997 in Journal of Natural History 38 (15) on pages 1989-1990, DOI: 10.1080/00222930310001613584, http://zenodo.org/record/4676048, {"references":["SCHIOTZ, A., 1975, The Treefrogs of Eastern Africa (Copenhagen: Steenstrupia), 232 pp.","SCHIOTZ, A., 1999, Treefrogs of Africa (Frankfurt: Chimaira), 350 pp."]}

Lötters, S., Schick, S., Scheelke, K., Teege, P., Kosuch, J., Rotich, D., Veith, M. (2004): Bio-sketches and partitioning of sympatric reed frogs, genus Hyperolius (Amphibia; Hyperoliidae), in two humid tropical African forest regions. Journal of Natural History 38 (15): 1969-1997, DOI: 10.1080/00222930310001613584, URL: http://dx.doi.org/10.1080/00222930310001613584