The Nguru mountains of Tanzania, an outstanding hotspot of herpetofaunal diversity Michele Menegon1,2, Nike Doggart3, Nisha Owen4,5 1 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali, Via Calepina 14, I–38100 Trento, Italy 2 Udzungwa Mountains Monitoring Centre, c/o Udzungwa Mountains National Park, P.O. Box 99 Mang’ula, Tanzania 3 Tanzania Forest Conservation Group, P.O. Box 23410, Dar es Salaam, Tanzania 4 Frontier Tanzania Forest Research Program, P.O. Box 9473, Dar es Salaam, Tanzania 5 Society for Environmental Exploration/Frontier, 50-52 Rivington Street, London EC2A 3QP Submitted on 2008, 7th March; revised on 2008, 22nd August; accepted on 2008, 30th September. Abstract. Despite the vicinity of a major road, the rainforests of the South Nguru Mountains in eastern Tanzania were virtually unexplored until 2004, particularly from a herpetological point of view. Several surveys were conducted between 2004 and 2006 with the aim of providing a comprehensive list of the amphibian and reptile species of this overlooked hotspot of biological diversity. The surveys resulted in this assess- ment of the herpetofaunal diversity, with 92 species recorded, of which 15 represent new records for this area, and the discovery of 16 species new to science, all of which are likely to be strictly endemic to the Nguru Mountains. Pressure on the forests, par- ticularly the lowland forests, remains high. A conservation planning process is now underway that is attempting to address the loss of these critically important forests. These results, documenting the high species richness and the outstanding number of putative endemics of the forests, strongly highlight the biological importance of the South Nguru Mountains and place them among the most important sites for the con- servation of herpetofauna in Africa. Keywords. Herpetofaunal diversity, new species, endemism, zoogeography, Eastern Arc Mountains. INTRODUCTION In the last few years increased interest in the forests of the Eastern Arc Mountains has resulted in the biological exploration of several poorly known mountain blocks. Previously unsurveyed montane ranges and forest reserves have recently been explored by different Acta Herpetologica 3(2): 107-127, 2008 ISSN 1827-9643 (online) © 2008 Firenze University Press 108 M. Menegon, N. Doggart and N. Owen groups of researchers and faunal lists are being completed. Among the ranges explored in the last few years, the South Nguru Mountains stand out due to their exceptional amphib- ian and reptile species richness, number of undescribed species and endemism ratio. The present paper outlines the results of several surveys conducted by two different groups of researchers and highlights the richness and uniqueness of the South Nguru Mountains herpetofauna, furthermore it stresses the need for a rapid taxonomic and conservation assessment of the newly discovered and as yet undescribed species in order to properly assess the biological value of the South Nguru Mountains forests. The South Nguru Mountains are part of the Eastern Arc Mountains of Tanzania and Kenya. This chain of isolated massifs is part of one of the world’s most important biodiver- sity hotspots (Mittermeier et al., 2004). The Eastern Arc Mountain forests are biologically exceptional due to the high concentration of endemic species that have been recorded in the area (Howell, 1993; Lovett, 1993; Burgess et al., 1998; Burgess et al., 2007). The biodi- versity of the South Nguru Mountains is less well known than some of the other Eastern Arc massifs, such as the East Usambara and Uluguru Mountains (Burgess et al., 2007). As stated by Lovett and Thomas (1988) the interest in the South Nguru Mountains is three- fold: they are in the central part of the Eastern Arc Mountain chain and should possess many rare species; the mountains have continuous forest cover from less than 400 m to 2400 m altitude; and are still poorly known biologically. At a national level, the South Nguru forests are important as the main catchment area for the Wami River which provides water for the town of Chalinze, the Mtibwa sugar plan- tation and many villages. Locally, the forests are important as a source of wood products, medicinal plants and for their cultural values. Most biological research in the South Nguru Mountains has concentrated on plants. Lovett and Thomas (1988) surveyed both Kanga and Nguru South Forest Reserves provid- ing a brief account of the vegetation. Botanical surveys conducted by Pócs et al. (1990) in the South Nguru Mountains provided additional data supplemented by subsequent surveys carried out by Lovett and Pócs (1993). These surveys indicated a high number of rare and endemic plants in Nguru South and Kanga Forest Reserves. Pócs (1998) recorded three endemic bryophyte species in the South Nguru Mountains. Results of several bird surveys were published in the past (Moreau, 1940; Sclater and Moreau, 1932-1933; Stu- art and van der Willingen, 1978), but it was not until 1996 that Romdal (2001) carried out a systematic ornithological survey covering various altitudes in the Ngurus (Baker and Baker, 2002). Until the publication of results of herpetological surveys carried out in the South Ngu- ru Mountains by Emmrich (1994), very little information was available on the herpetofau- na. Burgess et al. (1998) reported the collection of an undescribed Rhampholeon species, subsequently described by Mariaux and Tilbury (2006). Nine species of small mammals (shrews and small rodents), of which three are endemic to the Eastern Arc Mountains, were recorded in the South Nguru Mountains by Stanley et al. (1998). A few further records are available for other mammals. This paper demonstrates that the herpetofau- nal value of the South Ngurus is exceptional, particularly given that the survey intensity remains low relative to other areas of the Eastern Arc Mountains such as the Uluguru and East Usambara Mountains. 109Herpetofauna of Nguru Mountains STUDy AREA The South Nguru Mountains lie between S 05°53’S-S 06°17’ and E 037°27–E 037°45’ in Mvomero District, Morogoro Region, Tanzania (Fig. 1). Nguru South Forest Reserve (see Fig. 2) was gazetted during the German colonial period and covers an area of 18 797 ha with an altitu- dinal range between 760 and 2400 m. It is contiguous with Mkindo Forest Reserve which covers 5244 ha and extends down to 380 m. The estimated mean rainfall on the eastern slopes of these mountains is around 2000 mm/year increasing up to 3000-4000 mm at 2000 m (Lovett and Pócs, 1993). Kanga Forest Reserve was gazetted in 1954 and covers an area of 6664 ha with an altitu- dinal range between 500 and 2019 m. The estimated rainfall is 1500-2000 mm/year (Lovett and Pócs, 1993). Within the South Nguru landscape, there are 56 villages. The majority of people in the South Nguru landscape depend on agriculture for their livelihood. At the base of the mountains, the Mtibwa sugar plantation covers an extensive area. Farmers are also involved in smallholder sugar and rice cultivation. At higher altitudes, farmers grow bananas, yams, sweet potatoes, maize and coffee. Many farmers also have plots within the Nguru South Forest Reserve where they are growing cardamom and yams. Cardamom is one of the main sources of cash income for the com- munities immediately adjacent to the reserves. Cardamom cultivation is now widespread within Nguru South Forest Reserve. Fig. 1. Location of the sampling sites within the Nguru South and Kanga Forest Reserves. 110 M. Menegon, N. Doggart and N. Owen MATERIALS AND METHODS Between October 2004 and August 2006 a total of 13 sites were surveyed in the South Nguru Mountain forests, seven sites in Nguru South Forest Reserve and six in Kanga Forest Reserve. Two addi- tional sites were surveyed opportunistically in Nguru South Forest Reserve during January 2006 (see Table 1). Sampling sites were located in submontane and montane forests between 750 and 2200 m asl. Further records were obtained during random sampling in the village belt and from local people. Sites were sampled by opportunistic search, visual and acoustical monitoring. Searches were conducted both during the day and night to sample the highest number of species. Sixteen Visual Encounter Survey (VES) quadrats were conducted within a 10x10 m area, requiring one man-hour each per unit. Four VES transects of 100 m were also conducted, each requiring four man-hours. Another sampling technique used was pitfall trapping with drift fences (Heyer et al., 1993). They consisted of three 50 m linear transects each containing eleven 10 litre plastic buckets positioned 5 m apart, sunk into the ground with a plastic sheet ‘drift fence’ stretching between each bucket. Each line was placed 50 m apart, encompassing a range of micro-habitats. A total of 264 bucket pitfall trap-nights were conducted in each work unit. Specimens have also been sampled opportunistically during forest walks. Further records (mainly concerning snakes) were obtained from local people living in the villages at the forest edge. A collection of specimens opportunistically killed by villagers in crop fields surrounding the forest, were identified by the authors and deposited in the collection. The abundance of individuals of different species was also recorded. Sampling intensity was com- parable in sites 1-4. Voucher specimens were collected and, when possible, frog calls were recorded by means of a Sony TCM directional microphone and a Sharp Minidisc or an analogue Marantz PMD-222 audiocassette recorder and a Sennheiser K6-ME66 directional microphone. Specimens, Fig. 2. The montane rainforest of the Nguru South Mountains, one of the sampled habitats. 111Herpetofauna of Nguru Mountains photographs and sound recordings were deposited in both the Museo Tridentino di Scienze Natu- rali, Trento, Italy, and in the collections of the University of Dar es Salaam, Tanzania. The major- ity of species collected have been identified by the lead author, on the grounds of morphological, molecular and bioacoustic analysis and through comparison with material held in the herpetological collections of the University of Dar es Salaam, Tanzania, the Natural Sciences Museum of Trento, Italy and the Natural History Museum, London, UK. Molecular analysis were carried out at the Cen- tre for Alpine Ecology, Trento, Italy; the Natural History Museum, London, UK and the Institute of Biogeography, Basel, Switzerland. Identifications or confirmation of identification were also provided by Simon Loader, Institute of Biogeography, Basel; John Poynton, Natural History Museum, London and Van Wallach, Museum of Comparative Zoology, Harvard University, Cambridge, USA. Table 1. Location and main characteristics of the sampling sites. Sampling site UTM co-ordinates Elevation range m Main vegetation type Main habitats investigated Nguru Site 1 ‘Maskati’ 37M0333779 9329236 1900-2200 Montane and upper montane rain forest Open canopy forest, small bogs Nguru Site 2 ‘Pemba’ 37M0336825 9333210 950-1050 Submontane forest Closed canopy forest, small wetland, ecotone Kanga Site 3 ‘Kanga’ 37M0358812 9336174 750-900 Transitional and Submontane forest Closed canopy forest, Kanga Site 4 “Difinga” 37M0356024 9340888 1000-1300 Submontane rain forest Closed canopy forest, FT Kanga Site 5 37M0355845 9340733 1000-1100 Submontane rain forest Closed canopy forest, FT Kanga Site 6 37M0358539 9341552 800-950 Lowland and submontane forest Half-closed, canopy forest FT Kanga Site 7 37M0356729 9345887 1100-1250 Submontane rain forest High dense closed- canopy forest FT Kanga Site 8 37M0358571 9336368 800-900 Lowland and submontane forest Half-closed canopy forest FT Nguru Site 9 37M0333711 9329248 1950-2050 Montane and upper montane forest Open canopy forest, small bogs FT Nguru Site 10 37M0333540 9325622 1950-2020 Dry upper montane forest Open canopy forest, heath FT Nguru Site 11 37M0334554 9323454 1850-1950 Upper montane forest Open canopy forest, FT Nguru Site 12 37M0337300 9322700 1700-1850 Submontane and montane forest Dense closed-canopy forest FT Nguru Site 13 37M0361810 9324701 1800-1950 Montane forest Dense closed-canopy forest FT Nguru Site 14 37M0335146 9329820 1700 Montane forest Closed canopy forest FT Nguru Site 15 37M0336314 9332299 1180 Submontane forest Open canopy forest Village belt (two sites) 1000 – 2000 Farmland Synanthropic habitats 112 M. Menegon, N. Doggart and N. Owen The nonparametric species richness estimator based on abundance data Chao 1, computed along log-linear 95% confidence interval, and the abundance-based coverage estimator ACE, were used incorporating abundance data, represented by the number of individuals per species (Colwell and Coddington, 1994; Chao et al., 2000; Magurran, 2004). The shared species estimator V (Chao et al., 2000) has been used to estimate the number of shared species between sites. Analyses were performed using the sofware EstimateS (Magurran, 2004). The species complementarity between sites has been described as an expression of pattern diversity (Whittaker, 1972) by means of a Bray- Curtis similarity analysis expressed as a percentage (Legendre and Legendre, 1998; Magurran, 2004). The analysis of similarity has been performed using the software Biodiversity Pro. Only the records from the lead author’s direct collections in sites 1-4 were used in the statistical analysis. Amphibian taxonomy follows Frost et al., (2006), Reptile taxonomy follows Spawls et al. (2004), except for Colu- broidea taxonomy that follows Vidal et al. (2007) and Lawson et al. (2005) and Scincidae taxonomy that follows Brandley et al. (2005). RESULTS Species composition and richness. A total of 41 species of amphibian from 12 families and 51 species of reptile from 11 families were recorded in Nguru South Forest Reserve, Kanga Forest Reserve and farm- land adjacent to the two forest reserves during the survey activity. Three species previously documented for the area were not found during this study. By combining previous species records with those from the present study, the total number of amphibian and reptile species recorded from the South Nguru Mountains is 92 species, from 23 families (appendix 1). The majority, 76 species out of a total of 89 species collected during this survey, were recorded from forest, and of these, based on the known information on their ecology, 67 species are considered forest associated based upon available literature (Howell, 1993, Menegon and Sal- vidio, 2005; Burgess et al., 2007). The remaining species were collected from the village belt. Biological notes During the surveys we observed the activities of some species whose behaviour is poorly known. These observations are summarised as follows: Hoplophryne sp. nov. was active during and immediately after heavy rain in undisturbed forest during the day. At these times, individuals of Hoplophryne sp. nov. move around on soft soil and under the leaf litter, they can be detected by searching gently with a stick among the leaf litter. Female Hyperolius spinigularis were observed returning to their egg clutches and squirting water over the eggs. This was previously observed by Stevens (1971), and may be done to avoid desiccation and possibly parasitism of the eggs (J. Vonesh, pers. comm.). The geckos Urocotyledon wolterstorffii and Cnemaspis africana were observed sharing an egg-laying site on dry, sandy soil under a large overhanging rock. Two eggs of Urocotyledon wolter- storffii and several eggs of Cnemaspis africana were found under wood debris. Several females Cnemaspis africana were probably using the site. One of the eggs of Urocotyledon was 12 mm in length. After 16 days in controlled conditions a hatchling 21.25 + 23.37 mm long (Head body + Tail length) emerged. 113Herpetofauna of Nguru Mountains Range extensions of species endemic to the Eastern Arc Mountains The known distributions of 15 species endemic or near endemic to the Eastern Arc were extended by our findings (see Table 2). New species The present study recorded 15 amphibian taxa and one reptile taxon that are suffi- ciently distinct from the other known taxa that they may be considered new species based upon the available data (see Table 3). The discovery of numerous species new to science, Table 2. Reptile and amphibian species endemic to the Eastern Arc whose documented ranges have been extended by the present study. Species Known distribution prior to the present study References for previous records Amietophrynus brauni Usambara, Uluguru and Udzungwa Mountains Channing and Howell, 2006 Hyperolius spinigularis Usambara and Uluguru and Udzungwa Mountains and Mulanje Mts. Malawi Channing and Howell, 2006 Leptopelis vermiculatus Usambara, Udzungwa and Rungwe Mountains Channing and Howell, 2006 Leptopelis cf. parkeri Usambaras, Uluguru and Udzungwa Mts. Channing and Howell, 2006 Phrynobatrachus uzungwensis Nguu, Uluguru and Udzungwa Mountains Menegon et al., 2003; Channing and Howell, 2006 Probreviceps macrodactylus macrodactylus* Usambara Mountains Channing and Howell, 2006 Urocotyledon wolterstorffii East Usambara and Uluguru Mountains Spawls et al., 2004 Kinyongia oxyrhina Uluguru and Udzungwa Mountains Klaver and Böhme, 1988; Spawls et al., 2002 Chamaeleo werneri Uluguru and Udzungwa Mountains Spawls et al., 2004 Rhinotyphlops gierrai East and West Usambara Mountains Broadley and Wallach, 2000 Buhoma vauerocegae Usambara and Uluguru Mountains Spawls et al., 2004 Dipsadoboa werneri Usambara Mountains and the Uzungwa Scarp Forest Reserve in the southern Udzungwa Mountains Rasmussen, 1997; Menegon and Salvidio, 2005 Xyeledontophis uluguruensis Uluguru Mountains Broadley and Wallach (2002) Elapsoidea nigra North Pare, Usambara, Magrotto and Uluguru Mountains Broadley and Howell, 1991; Spawls et al. 2004 Atheris ceratophorus East Usambara, Uluguru and Udzungwa Mountains Spawls et al., 2004 * The record of this species from North Pare Mountains (Channing and Howell, 2006) is actually due to a misidentification of an undescribed species of Callulina. 114 M. Menegon, N. Doggart and N. Owen Table 3. Reptile and amphibian taxa considered new species based upon the available data. The occur- rence and occupancy areas were made without considering elevational constraints, in some cases the area may be an over-estimate. Taxa Range based on current information Estimated maximum area of occupancy Estimated maximum extent of occurrence Amphibia Anura Arthroleptidae Arthroleptis sp. nov. Nguru South and Mkindo Forest Reserves (278 km2) 240.3 km2 240.3 km2 Leptopelis sp. nov. Kanga Forest Reserve 66 km2 66 km2 Bufonidae Nectophrynoides sp. nov. 1 Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Nectophrynoides sp. nov. 2 Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Nectophrynoides sp. nov. 3 Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Nectophrynoides sp. nov. 4 Kanga Forest Reserve 66 km2 66 km2 Brevicipitidae Callulina sp. nov. 1 Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Callulina sp. nov. 2 Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Callulina sp. nov. 3 Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Callulina sp. nov. 4 Kanga Forest Reserve 66 km2 66 km2 Probreviceps sp. nov. Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Microhylidae Hoplophryne sp. nov. Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 Petropedetidae Petropedetes sp. nov. Nguru South, Mkindo and Kanga Forest Reserves 306.3 km2 57 km2 GyMNOPHIONA Caecilidae Boulengerula sp. nov. Nguru South, Mkindo and Kanga Forest Reserves 306.3 km2 57 km2 Scolecomorphus sp. nov. Nguru South and Mkindo Forest Reserves 278 km2 278 km2 Reptilia Sauria Chamaleonidae Rhampholeon sp. nov. Nguru South and Mkindo Forest Reserves 240.3 km2 240.3 km2 115Herpetofauna of Nguru Mountains which are likely to be narrowly endemic to the Nguru Mountains only, is among the most interesting results of this study and make the South Nguru Mountains one of the most important ranges for herpetofaunal diversity and conservation in Africa. The taxonomic status of all the species collected during the surveys has been assessed in the context of a more extensive work on the genera, involving molecular, osteological, bioacoustic and morphological analysis conducted between 2005 and 2008 (Loader et al. unpublished; Poynton et al. unpublished; Menegon et al. unpublished). Formal description of some of the unnamed taxa has started. Zoogeography As with other Eastern Arc Mountains, the herpetofauna of the South Nguru Moun- tains contains a combination of Afromontane (Eastern Arc forests endemic and near endemic species), lowland/coastal and widespread species associated with savannah and woodland (see Table 4). While this is a strong indication that the forests of these areas were once connected, more detailed taxonomic work is still needed to fully understand the historical biogeography of the Eastern Arc Mountains. Endemism The proportion of amphibian and reptile species from the South Nguru Moun- tains that are endemic to these mountains is extremely high. Based on the results of this study, 23% (n = 18) of the herpetofauna species are strictly endemic to the South Nguru Mountains, of which only two were described. A further 33.3 % (n = 26) of the species have ranges restricted to the Eastern Arc Mountains. The percentage of endemic or near endemic species is directly related to elevation, a pattern that has also been documented for other Eastern Arc forest localities (Menegon and Salvidio, 2005). Based on three col- lecting localities, endemic species values at different elevation are as follows: Kanga, 750 m = 55.1%; Pemba, 1000 m = 59.4%; Maskati, 2000 m = 72.7%. This highlights the unique- ness of the montane herpetofauna community. Species richness and pattern diversity Nonparametric estimators were used to estimate the species richness of sites and the number of shared species between sites in order to assess the total richness of the area and the differential diversity across sites and along an altitudinal gradient. In order to avoid biases due to different collecting methods and timing, only records compiled from the lead author’s direct sampling were used to calculate relative abundance and diversity estimates. Kanga is estimated to be the richest site (59.26 ± 2.49 ACE mean; 54.5 ± 2.79 Chao 1 95% CI lower bound), followed by Pemba (48.46 ± 1.61 ACE mean, 43.05 ± 2.68 Chao 1 95% CI lower bound) and Maskati (30.73 ± 2.49 ACE mean, 26.11 ± 2.79 Chao 1 95% CI lower bound), this result is probably due to the low elevation component of the Kanga species assemblage, following the assumption that species richness decreases with increas- 116 M. Menegon, N. Doggart and N. Owen Table 4. Biogeographic affinities of the recorded species. Bibliographic records and species collected by authors were considered. Five species not yet identified to species level have not been included in the table, species identified as undescribed are included. Nguru south and Kanga forest endemics no. of species 18 % on total recorded 23 Arthroleptis sp. nov., Nectophrynoides sp. nov. 1, Nectophrynoides sp. nov. 2, Nectophrynoides sp. nov. 3, Nectophrynoides sp. nov. 4, Leptopelis sp. nov., Callulina sp. nov. 1, Callulina sp. nov. 2, Callulina sp. nov. 3, Callulina sp. nov. 4, Probreviceps sp. nov., Hoplophryne sp. nov., Petropedetes sp. nov., Scolecomorphus sp. nov., Boulengerula sp. nov, Rhampholeon acuminatus. Rhampholeon sp. nov., Kinyongia fisheri Eastern Arc forest endemics or near-endemics no. of species 26 % on total recorded 33.3 Arthroleptis affinis, Amietophrynus brauni, Afrixalus uluguruensis, Hyperolius mitchelli, Hyperolius spinigularis, Hyperolius sp. (puncticulatus-like), Leptopelis cf. parkeri, Leptopelis cf. uluguruensis, Leptopelis vermiculatus, Probreviceps macrodactylus macrodactylus, Phrynobatrachus uzungwensis, Chamaeleo werneri, Chamaeleo deremensis, Kinyongia oxyrhina, Agama montana, Urocotyledon wolterstorffii, Scelotes uluguruensis, Proscelotes cf. eggeli, Leptosiaphos kilimensis, Typhlops gierrai, Buhoma vauerocegae, Rhinotyphlops gierrai, Thelotornis usambaricus, Crotaphopeltis tornieri, Dipsadoboa werneri, Elapsoidea nigra, Atheris ceratophorus East African coastal mosaic species no. of species 18 % on total recorded 23 Arthroleptis xenodactylus, Arthroleptis xenodactyloides, Afrixalus stuhlmanni complex. Afrixalus stuhlmanni stuhlmanni, Afrixalus fornasinii*, Leptopelis flavomaculatus, Gastropholis prasina*, Chamaeleo melleri, Rieppeleon brevicaudatus, Rieppeleon brachyurus, Cnemaspis africana, Holaspis laevis, Melanoseps loveridgei*, Aparallactus guentheri, Lycophidion capense loveridgei, Natriciteres sylvatica, Philothamnus macrops, Dendroaspis angusticeps. West African forest and savannah species no. of species 2 % on total recorded 2.5 Lycophidion meleagre, Bitis gabonica East/south African savannah species no. of species 8 % on total recorded 10.2 Schismaderma carens, Ptychadena anchietae, Chiromantis xerampelina Xenopus cf. petersii, Lygosoma afrum*, Rhinotyplops mucruso, Aparallactus jacksoni, Atractaspis bibroni, Philothamnus punctatus, Widely distributed savannah species no. of species 15 % on total recorded 19.2 Ami e t o p hr y nu s g ut t ura li s , Ami e t i a ang o l e n s i s , Phrynobatrachus parvulus, Chamaeleo dilepis, Agama agama, Trachylepis varia, Trachylepis maculilabris, Trachylepis striata, Varanus niloticus, Typhlops lineolatus, Crotaphopeltis hotamboeia, Dispholidus typus, Lamprophis fuliginosus, Prosymna stuhlmanni, Philothamnus hoplogaster, Bitis arietans. * Bibliographic records 117Herpetofauna of Nguru Mountains ing elevation (MacArthur, 1972; Stevens, 1992). The same pattern of species richness is obtained by considering the Simpson index of diversity: Simpson Mean (D): Kanga = 33.43; Pemba = 24.93; Maskati = 14.29. According to the Bray-Curtis indices of similarity the site at highest elevation (Maska- ti) shows the greatest difference from other sites (25% of similarity), with medium altitude sites Pemba and Kanga showing closer faunal similarities (38% of similarity), in spite of the latter two being positioned on two separate massifs (see Fig. 4). This pattern of diver- sity highlights the unique nature of the montane and upper montane forest herpetofauna and demonstrates how altitude influences the composition of assemblages in Eastern Arc forests. Based upon predictions of species richness, estimates suggest an extremely diverse forest associated herpetofauna (possibly > 110 species for the entire landscape). Further comparisons with other Eastern Arc Mountains will be necessary to see how significant these estimates are, but it is likely to be higher than most areas based on our current understanding of other Eastern Arc Mountains. Species richness comparison The South Nguru Mountain forests support a rich forest associated herpetofauna of about 67 species. This value is higher than all the considered sites in Africa (see Table 5) Fig. 4. Species overlap among sampled sites. The graph shows the high complementarity of the species com- position in the sampled sites and how sites at same altitude on different mountain blocks (Kanga and Pem- ba) have a more similar species composition than sites at different altitudes on the same mountain block. 118 M. Menegon, N. Doggart and N. Owen and comparable with the one recorded for the Virunga National Park (De Witte, 1941), but in an area about more than 20 times smaller. Detailed comparison between sites is complicated by the differences in: total area and elevational ranges of the available sites, and biases in data collecting effort. Despite this, a preliminary comparison between sites with similar geographic features places Nguru South as being among the richest sites in Africa, comparable with the Mesoamerican site of Monteverde in Costa Rica. DISCUSSION Before the surveys described in this paper, the South Nguru Mountains were believed to have no narrow endemic species (Burgess et al., 2007); the results presented in this paper stress the need for a rapid taxonomic assessment of the newly discovered species in order Table 5. Comparison of the “forest-associated” herpetofauna species richness in some tropical mid-eleva- tion-montane forests of Africa, Central America, and the Philippines (modified from Vonesh 1998). Site Elevation m Area (ha) Amphibians Reptiles Total Spp /area Reference Usambara Mts., Tanzania 600-2286 22100 23 29 52 0.23 Howell, 1993; Poynton et al., 2007 Uzungwa Scarp FR, Tanzania 600-2000 21000 28 29 57 0.27 Menegon and Salvidio, 2005 Kibale N.P., Ugandaa 1400-1550 56000 15 29 44 0.07 Vonesh, 1998 Bwindi N.P., Uganda 1200-2600 31000 20 21 41 0.13 Drewes, 1991 Virungas N.P., Congoc 1300-3000 300000 27 40 67 0.022 DeWitte, 1941 Korup N.P., Cameroond 1080-1768 124000 33 11 44 0.035 Lawson, 1993 Monteverde, Costa Ricae 1300-1470 12000 25 36 61 0.5 Timmerman, 1981 Cuernos de Negros, Philippine 1050-1350 10000 7 13 20 0.2 Brown and Alcala, 1961 Nguru South, Tanzania 750 - 2200 12000 33 34 67 0.55 This paper a) only Kibale species restricted to the forest interior are included; b) species listed as extralimital by Drewes and Vindum (1991) are excluded; c) only forest species found above 1300 m are included; d) only species found at Mt. yuhan (to 1079 m); Rumpi Hills (1000-1768 m) and Nta Ali (to 1200 m) are includ- ed. Lawson (1993) comments that these elevations were under sampled; e only species restricted to Tim- merman’s (1981) pre-montane and lower montane zones (2-5) are included; f only specimens in submon- tane and montane forest zones are included from Brown and Alcala (1961), the source for closed forest area in East Usambara and South Nguru Mountains is Newmark (2002) for the Uzungwa Scarp the closed forest area is assumed to be the same of the gazetted area. 119Herpetofauna of Nguru Mountains to further define the biological value of South Nguru Mountain forests where 16 out of 18 strictly endemic species have no name or IUCN Red List assessment. Nguru South, Kan- ga and Mkindo are categorised as National Protection Forest Reserves. As such they are reserved ‘for the purposes of protection of water sheds, soil conservation and the protection of wild plants’ (Government of Tanzania, 2002). Kanga and Nguru South Forest Reserves are managed directly by the Morogoro Regional Catchment Forest Office while Mkindo is man- aged jointly by the four surrounding communities and the Morogoro Regional Catchment Forest Office. The miombo woodland in the lowlands between Kanga and Nguru South is on village land and does not lie within a protected area. Although it is intended that all for- est reserves will have a management plan, only Mkindo has one at present. However, despite the legal protection afforded by the Forest Act (2002) enforcement is weak and sporadic. Some of the key threats to the herpetofauna of the South Ngurus are forest loss and deg- radation as a result of fire, selective logging, encroachment from agricultural land and the removal of the forest shrub and herb layer for the cultivation of cardamom and yams. Many of the species of reptile and amphibian recorded during the survey are associated with for- ests and several species are associated with the shrub layer of the forest. This includes Cal- lulina spp., Nectophrynoides spp. and Rhampholeon sp. that were only found in the shrub layer of undisturbed forest. While the clearance of the forest is likely to have the most signif- icant negative impact on all forest species in the Ngurus, cardamom cultivation is particular- ly problematic in the forest close to Ubiri, Mafuta, Kwelikwiji, Mhonda, Digoma, Digarama, Maskati, Pemba and Kigugu villages. Cardamom cultivation could therefore potentially also have a significant impact, as this agricultural practice will affect many endemic species close- ly associated with the forest shrub layer. The long term impacts of cardamom cultivation in forest areas are not fully understood, but are likely to impede regeneration of forest plants. The distinct nature of the herpetofauna species assemblages at high altitude and the remarkable number of putative endemics in the South Nguru Mountains, highlights the conservation importance of the South Nguru Mountain forests within the Eastern Arc Mountains. The high elevational turnover of species indicates the importance of conserv- ing forest at all altitudes. Areas of forest or marginal habitats, even at low elevation, might also be vital in generating high species diversity, and need to be considered in the devel- opment of a conservation strategy for the area. Differences also exist between assemblag- es at sites at similar altitudes in adjoining mountain fragments within the South Nguru Mountain forests indicating the fine scale geographic turnover in the herpetofauna of the area (Fig. 3). This suggests that not only is it important to conserve the forest along the elevational gradient but it is also important to conserve fragments in geographically com- plex terrains, where isolated populations and therefore potentially new species may exist. Since 2004, the Tanzania Forest Conservation Group (TFCG), a Tanzanian non-gov- ernmental organisation, has been working with stakeholders to improve conservation plan- ning in the South Ngurus. TFCG, through a partnership programme known as PEMA, has facilitated a dialogue between communities and the government to develop a vision for natural resource management in the South Nguru landscape. Villagers and government have identified a series of actions required to address the issue of forest loss. This includes a combination of direct forest management activities such as developing and implementing forest management plans and boundary demarcation; and activities aimed at reducing local people’s dependence on unsustainable activities such as the current methods of cultivating 120 M. Menegon, N. Doggart and N. Owen cardamom. The programme represents an opportunity to reverse the current trend of for- est loss and degradation. To succeed the programme will need sustained commitment from the Government of Tanzania, civil society organisations, the local communities and devel- opment partners to conserve the unique biodiversity of this area. ACKNOWLEDGEMENTS Thanks to Abrahaman Mndeme, David Loserian and Ernest Benson. Moses Mwangoka kindly provided information on vegetation of the sampled sites, Simon Loader provided results of prelimi- Fig. 3. Some species recorded during the present study. Clockwise: Rhampholeon acuminatus; Kinyongia fisheri fisheri; Dipsadoboa werneri; an undescribed species of Callulina; an undescribed species of Leptope- lis and an undescribed giant species of Nectophrynoides. 121Herpetofauna of Nguru Mountains nary molecular investigation on some of the species collected and useful suggestions on an early draft of this manuscript, John Poynton confirmed the identification of some of the amphibian speci- mens, Van Wallach kindly provided identifications for the Typhlopidae, Joe Beraducci kindly pro- vided data on some snakes species, David Moyer, Aaron Bauer, Neil Burgess and Kim Howell helped us in many ways. Thanks to an anonymous referee for his valuable comments on our work. 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Leptopelis uluguruensis Barbour and Loveridge, 1928 Leptopelis flavomaculatus (Günther, 1864) Leptopelis vermiculatus (Boulenger, 1909) Leptopelis cf. parkeri Bufonidae Amietophrynus brauni (Nieden, 1910) Amietophrynus gutturalis (Power, 1927) Nectophrynoides sp. nov. 1 Nectophrynoides sp. nov. 2 Nectophrynoides sp. nov. 3 Nectophrynoides sp. nov. 4 Schismaderma carens (Smith, 1848) Hyperolidae Afrixalus uluguruensis (Barbour and Loveridge, 1928) Afrixalus stuhlmanni stuhlmanni (Pfeffer, 1893) Afrixalus stuhlmanni complex Afrixalus fornasinii (Bianconi, 1940) Hyperolius mitchelli (Loveridge, 1953) Hyperolius spinigularis (Stevens, 1971) Hyperolius sp. (puncticulatus – like) Hyperolius sp. Brevicipitidae Callulina sp. nov. 1 Callulina sp. nov. 2 Callulina sp. nov. 3 Callulina sp. nov. 4 Probreviceps macrodactylus macrodactylus Nieden, 1926 Probreviceps sp. nov. Microhylidae Hoplophryninae Hoplophryne sp. nov. Rhacophoridae Rhacophorinae Chiromantis xerampelina Peters, 1854 Pyxicephalidae Cacosterninae Amietia angolensis (Bocage, 1866) Ptychadenidae Ptychadena anchietae (Bocage, 1868) 125Herpetofauna of Nguru Mountains Petropedetidae Petropedetes sp. nov. Phrynobatrachidae Phrynobatrachus uzungwensis Grandison and Howell, 1983 Phrynobatrachus parvulus (Boulenger, 1905) Phrynobatrachus sp. Pipidae Xenopus cf. petersii GyMNOPHIONA Caecilidae Boulengerula sp. nov. Scolecomorphus sp. nov. REPTILIA SAURIA Chamaleonidae Chamaeleoninae Chamaeleo werneri (Tornier, 1899) Chamaeleo deremensis Matschie, 1892 Chameleo melleri Gray, 1865 Chamaeleo dilepis Leach, 1819 Kinyongia fischeri (Reichenow, 1887) Kinyongia oxyrhina (Klaver and Böhme, 1988) Brookesiinae Rieppeleon brachyurus (Günther, 1893) Rieppeleon brevicaudatus (Matschie, 1892) Rhampholeon acuminatus Mariaux and Tilbury, 2006 Rhampholeon sp. nov. Agamidae Agama montana Barbour and Loveridge, 1928 Agama agama (Linnaeus, 1758) Gekkonidae Cnemaspis africana Werner, 1895 Urocotyledon wolterstorffi (Tornier, 1900) Lacertidae Holaspis laevis Werner, 1895 Gastropholis prasina Werner 1904 Scincidae Lygosominae Leptosiaphos kilimensis Stejneger 1891 Lygosoma afrum Peters 1854 Trachylepis varia (Peters, 1867) Trachylepis maculilabris (Gray, 1845) Trachylepis striata (Peters, 1844) Scincinae Scelotes uluguruensis (Barbour and Loveridge, 1928) Proscelotes cf. eggeli Tornier 1902 Melanoseps loveridgei Brygoo and Roux-Estève, 1982 Varanidae Varanus niloticus Linnaeus, 1758 SERPENTES Typhlopidae Rhinotyphlops mucruso (Peters, 1854) 126 M. Menegon, N. Doggart and N. Owen Typhlops lineolatus Jan, 1864 Typhlops gierrai Mocquard, 1897 Colubridae Colubrinae Crotaphopeltis tornieri (Werner, 1897) Crotaphopeltis hotamboeia Laurent, 1968 Dipsadoboa werneri Boulenger, 1897 Thelotornis usambaricus Broadley, 2001 Xyeledontophis uluguruensis Broadley and Wallach, 2002 Dispholidus typus Smith, 1829 Philothamnus hoplogaster Günther, 1863 Philothamnus punctatus Peters,, 1867 Philothamnus macrops (Boulenger, 1895) Lamprophiidae Lamprophinae Lamprophis fuliginosus Boie, 1827 Lycophidion capense loveridgei Laurent, 1968 Lycophidion meleagre Boulenger, 1893 Atractaspidinae Aparallactus jacksoni Günther, 1888 Aparallactus guentheri Boulenger, 1895 Atractaspis bibronii Smith,1849 Incertae sedis Buhoma vauerocegae Tornier, 1902 Prosymna stuhlmanni (Pfeffer, 1893) Natricidae Natriciteres sylvatica Broadley, 1966 Elapidae Elapinae Elapsoidea nigra Günther, 1888 Dendroaspis angusticeps Smith,1849 Viperidae Viperinae Atheris ceratophora Werner, 1895 Bitis arietans (Merrem, 1820) Bitis gabonica Duméril, Bibron and Duméril, 1854 Appendix 2. Voucher specimens collection number Amphibia Arthroleptis sp.1 – MTSN8135; Arthroleptis xenodactyloides – MTSN8529; Arthroleptis sp. 2 – MTSN8168; Arthroleptis sp. nov – MTSN8142; Leptopelis cf. barbouri – MTSN8494; Leptopelis ulugu- ruensis – MTSN8227; Leptopelis flavomaculatus – MTSN8239, Leptopelis vermiculatus – MTSN8230; Amietophrynus brauni – MTSN8251; Amietophrynus gutturalis – MTSN8355; Nectophrynoides tornieri – MTSN8344; Nectophrynoides sp. nov. 1 – MTSN8199; Nectophrynoides sp. nov. 2 – MTSN8547; Nec- tophrynoides sp. nov. 3 - KMH35971; Nectophrynoides sp. nov. 4 - Schismaderma carens – MTSN8231; Afrixalus sp. – MTSN8375; Afrixalus uluguruensis – MTSN8163; Afrixalus stuhlmanni stuhlmanni – MTSN8275; Hyperolius mitchelli – MTSN8277; Hyperolius spinigularis – MTSN8265; Hyperolius puncticulatus MTSN8438; Chiromantis xerampelina KMH26922; Hyperolius sp. – MTSN8361; Cal- 127Herpetofauna of Nguru Mountains lulina sp. nov. 1 – MTSN8131; Callulina sp. nov. 2 – MTSN8138; Callulina sp. nov. 3 – MTSN8205; Callulina sp. nov 4 – Probreviceps sp. nov. KMH26923; MTSN8242; Probreviceps macrodactylus mac- rodactylus – MTSN8507; Hoplophryne uluguruensis – MTSN8144, Amietia angolensis – MTSN8174; Ptychadena anchietae – MTSN8339; Petropedetes sp. nov. – MTSN8364; Phrynobatrachus uzungwen- sis – MTSN8371, Phrynobatrachus cf. parvulus – MTSN8372; Phrynobatrachus sp. – MTSNpending, Xenopus cf. petersii – MTSN8249; Boulengerula sp. – MTSN8302; Scolecomorphus sp. – MTSN8421. Reptilia Chamaeleo werneri – MTSN8451; Chamaeleo deremensis – MTSN8195; Chameleo melleri – pictured and released; Chamaeleo dilepis - pictured and released; Bradypodion fischeri - MTSN8318; Bradypodion oxyrhinum – MTSN8416; Rieppeleon brachyurus – MTSN8431; Rieppeleon brevicaudatus – MTSN8220; Rhampholeon sp. nov. – MTSN pending; Rhampholeon sp. – MTSN8539; Agama montana – MTSN8186; Agama agama – MTSN8346, Cnemaspis africana – MTSN8214; Urocotyledon wolterstorffi – MTSN8222; Holaspis laevis – obs. and pictured; Trachylepis varia – obs. and pictured; Trachylepis maculilabris – obs. and pictured; Trachylepis striata – MTSNpending; Proscelotes eggeli - KMH26709; Scelotes ulugu- ruensis – MTSN8423; Varanus niloticus – obs.; Rhinotyphlops mucruso – MTSN8335; Typhlops gierrai – MTSN8433; Thelotornis usambaricus – MTSN8400, Buhoma vauerocegae - KMH26702; Dipsadoboa werneri – MTSN8505; Crotaphopeltis tornieri – MTSN8235; Crotaphopeltis hotamboeia - 8334; Dispholidus typus – MTSN8410, Lamprophis fuliginosus – MTSN8387; Lycophidion capense loveridgei – MTSN8345; Lycophidion meleagre – MTSN8414; Natriciteres sylvatica – MTSN8439; Philothamnus cf. hoplogaster – MTSN8178; Philothamnus punctatus – MTSN8190; Philothamnus macrops – MTSN8200; Prosymna stuhl- manni – MTSN8343; Aparallactus jacksoni – MTSN8352; Aparallactus guentheri – MTSN8341; Atractaspis bibronii – MTSN8354; Elapsoidea nigra – MTSN8349; Dendroaspis angusticeps – Atheris ceratophorus - KMH23977; MTSN8206; Bitis arietans – MTSN8209.