Chapter 6 Herpetofauna of Montane Areas of Tanzania. 2. Altitudinal Distribution of Amphibians on the Uluguru South Mountains Wilirk Ngalason and Felix J. Mkonyi 1 2 'Department of Zoology and Wildlife Conservation, University of Dar es Salaam, P. O. Box 35064, Dar es Salaam, Tanzania ~ Department of Biological Sciences, Dar es Salaam University College of Education, P.O. Box 2329, Dar es Salaam, Tanzania Abstract We present data on the altitudinal distribution of amphibians on the western slopes of Uluguru South Mountains, Tanzania. These mountains form part of the Eastern Arc Mountains, which are known for their biodiversity richness. We used drift fences with pitfall buckets, plots, and audio-visual survey methods. A total of 21 species was found during the survey. Four species previously reported in the Uluguru South Mountains were not found during the present survey. Generally, the number of species was highest at lower altitudes and decreased with increased altitude. Species similarity index showed that the study sites adjacent to each other had greater similarity in amphibian species composition than did sites distant from each other. The Uluguru Mountains are characterized by altitudinal variation in vegetation types and climatic factors such as temperature and rainfall, which may affect the distribution of amphibians on the mountain. Introduction composition (Poynton et al., 2006). The Uluguru Mountains are divided into Uluguru North and Uluguru South. They rise The distribution of animal species largely relates to their to about 2600 m above sea level on the Lukwangule Plateau. habitat requirements. The primary environmental factors that The lower slopes, from 600 to about 1600 m, have been limit the distribution of species include climate, topography, deforested, whereas the mid-altitude portions are still covered vegetation type, microhabitat, soil, and geology (Davis et al., by degraded forest. The only intact portion is located on the 1990; Heyer et al., 1994). Linked to changes in environmental higher altitudes above 2000 m. Annual distribution of rainfall factors is altitude, and this parameter is often used for examining and dry season length are the main climatic factors the distribution and richness of species across habitats. Changes determining forest limits in the Eastern Arc Mountains, with in the richness of amphibians across altitudinal gradients have the exception of frost, which determines the upper altitudinal been examined across the globe (e.g., Fauth et al., 1989; Owen, limits (Lovett, 1993). Forests in the Uluguru Mountains are 1989; Buckley & Jetz, 2007). Declines in amphibian diversity divided into lowland (below 500 m), submontane (800- with increases in altitude correspond to changes in environmen¬ 1500 m), montane (1500-1850 m), and upper montane forest tal factors, wherein areas with less favorable conditions support (1850-2400 m) (Poes, 1976a, b, c). Above the forest limits are fewer species (Poynton, 1962). heath and grassland vegetation on the Lukwangule Plateau. Generally, the distribution of amphibians along an altitu¬ The Uluguru Mountains are characterized by an oceanic dinal gradient is not uniform (Brown & Alcala, 1961; Scott, rainfall pattern. The mountains receive short rains (Vuli) from 1976; Heatwole, 1982). There appear to be some discrete “cut¬ October to December and the heavy long rains (Masika) from off points” at which the lowland species are separated from March to May. Estimated rainfall on the eastern slopes of those in the upland areas. Studies of southern and East Uluguru South is 2500-4000 mm/year, whereas on the western African lowland and mountain systems have shown species slopes, rainfall is estimated at 2000 mm/year. Mist and clouds turnover, changing between different areas with climatic occur throughout the year and are important sources of dry variations that determine these cut-off points (Poynton, season moisture in higher altitude forests (Poes, 1976a). The 1990, 1992, 2000, 2003; Loader et al., 2004). mean annual temperature in the Uluguru South is about The Uluguru Mountains form part of the Eastern Arc 24.3°C with small seasonal variation. At lower altitudes, mean Mountains, which are a chain of isolated crystalline moun¬ temperature ranges from 21.1°C to 26.5°C, whereas frost tains running from southern Kenya through Tanzania in a occurs above 2100 m in the Uluguru Mountains (Poes, 1976b). crescent-shaped arc (Lovett, 1990). The Eastern Arc Moun¬ The altitudinal distribution of African amphibian fauna is tains form one of the global biodiversity hotspots due to their well studied in southern Africa. Some of the comprehensive high species diversity (Myers et al., 2000), a consequence of broad-scale studies in the altitudinal distribution of amphib¬ factors including fragmentation associated with altitude, ians in Africa include those by Poynton and Broadley (1991), zonation of climate, and altitudinal turnover in species Poynton and Boycott (1996), and Poynton (1992, 2003). In the FIELDIANA: LIFE AND EARTH SCIENCES, NO. 4, August 26, 2011, pp. 81-89 81 Uluguru Mountains, the earliest inventory was carried out by Salaam, Tanzania; Natural History Museum, London; and Barbour and Loveridge (1928). Subsequent studies were Museo Tridentino di Scienze Naturali, Trento, Italy. Species carried out by Doggart et al. (2004) followed by other were classified following the taxonomy of Frost et al. (2006). collectors (mainly Frontier Tanzania). Despite these studies, there has been little effort to determine the altitudinal distribution of amphibians in the Uluguru Mountains. This Data Analysis study attempts to determine the altitudinal distribution of amphibians in the Uluguru South Mountains. The core data used in this study come from an altitudinal transect that was Composition of amphibian species in different altitudinal established on the western slopes of this range. The paper also zones was compared using the Jaccard Similarity Index (53) reviews what is known of the Uluguru South Mountains. (Ludwig & Reynolds, 1988). This index uses presence/absence data and is widely used to assess similarity of communities. Sj is frequently multiplied by 100%, and may be represented in terms of dissimilarity (i.e., D} — 1.0 & minus; Sj). The Materials and Methods similarity index values range from 0 (no species common to both communities) to 1.0 (all species found in both Amphibians were studied along an altitudinal gradient on communities). A higher similarity index suggests greater the western slopes of the Uluguru South Mountains from similarity of species among communities. 1000 m to the Lukwangule Plateau at about 2634 m. The It is represented by the formula surveys were conducted between 2001 and 2003. Five altitudinal zones were chosen and surveyed intensively, (a + b + c+n) ’ including 1000, 1200, 1500, 2000, and 2500 m (Figure 1). The lowest forested site is found at 1200 m in the Shikurufumi where Nj is the Jaccard similarity coefficient, a is the Forest Reserve (Fig. 1). Shikurufumi Forest Reserve is a number of species common to (shared by) communities, b forest fragment at the foothill of the western slopes of the is the number of species unique to the first community, c is Uluguru South Mountains. the number of species unique to the second community, At 1000 m, surveys were conducted in September 2001 (dry and n is the number of species unique to community n. season); and at 1200 m the surveys were conducted in March 2002 (wet season) and in August 2002 (dry season). At 1500 m, the surveys were conducted both in cultivated and forested areas. In the cultivated areas, surveys were done in September Results 2001; in forest, they were conducted in March 2002 and August 2002. At 2000 m, the surveys were conducted in The study recorded a total of 21 amphibian species September 2001 and March 2002, and at 2500 m, the surveys belonging to eight families on the western slopes of Uluguru were conducted between March and April 2003 (wet season) South Mountains. Observations revealed differences in species and between June and July 2003 during the dry season. Details composition at different altitudes. Certain species occurred at and locations of each site are given in Table 1. more than one altitudinal zone, whereas for some species, Standard methods for surveying amphibians were used, there was an altitudinal overlap. including drift fences with pitfall buckets, plots, and audio¬ Seven species were recorded at 1000 m, 12 species at 1200 m, visual encounter surveys (Heyer et al., 1994). Six drift fences 11 species at 1500 m, seven species at 2000 m, and five species with pitfall buckets were set at each altitudinal zone for seven at 2500 m. The pattern shows a general decrease in the number days. Each pitfall line consisted of 11 pitfall buckets set at 5-m of species with increased altitude (Table 2). An exception is at intervals, and a drift fence of polythene bisected all of the 1000 m, which had fewer species than did 1500 m. Only one buckets in the line (see Stanley et al., this volume). Pitfall lines species (Arthroleptis affinis) was found to occur on all were set to include different habitat types (i.e., cultivated altitudinal zones on the western slopes of the Uluguru South areas, forests, forest edges, and grasslands). Buckets were Mountains from 1200 m and above. The submontane forest checked at least once every day. portion (800-1500 m) supported more amphibian species than Pitfall lines cannot adequately sample all amphibians; they montane or upper montane forest portions (Table 2). were supplemented with plot method and audio-visual encoun¬ The amphibian species compositions were clearly separated by ter surveys. Ten plots of 5 by 5 m were established at each altitude. The study sites that were adjacent to each other showed altitudinal zone. Plots were systematically set at each zone, and greater similarity in amphibian species composition than did sites the interval between one plot and another was 50 m. Each plot distant from each other. Pairwise comparison of species compo¬ was searched thoroughly for amphibians once every sampling sition between different altitudinal zones is as shown in Table 3. season by four observers. Sampling involved searching for amphibians in leaf litter, between or within grass tussocks, on plant trunk and leaves, and by turning logs and stones. Digging underground (5-10 cm deep) was also done to ensure a Species Accounts thorough search. Audio-visual encounter surveys were con¬ ducted both during the day and night to detect amphibians not Family Arthroleptidae sampled by pitfall lines and plots. Amphibians found were Arthroleptis spp. identified and released after taking some voucher specimens. Material collected has been deposited in the Department of This is one of the most taxonomically difficult genera in Zoology and Wildlife Conservation, University of Dar es East Africa due to its variable coloration and morphology. Its 82 FIELDIANA: LILE AND EARTH SCIENCES 37°30’ 37°35’ 340000 m E 350000 m E Fig. 1. Map of the western slopes of Uluguru South Mountains showing the study sites and elevations (Source: Tanzania Maps [Mgeta], Toposheet 201/1, Y742 Series, 1982). taxonomy is not yet clearly settled. Breeding in known Arthroleptis of finis Ahl, 1939 Arthroleptis species does not depend on water (Channing & Howell, 2006); members lay their eggs in nests under leaf litter This species is widely distributed along the altitudinal and have no free-living tadpole stage. gradient in the Uluguru South Mountains. It has a very NGALASON AND MKONYI: HERPETOFAUNA OF TANZANIA, ULUGURU SOUTH MOUNTAINS 83 Table 1. Location of the study sites on the western slopes of the Uluguru South Mountains, Tanzania. Sampling site Altitude (masl) Coordinates Sampling time Habitat type 1 1000 7°3.844'S, 37°34.254'E 1-10 September 2001 Cultivated land 2 1200 7°10.925'S, 37°30.744'E 1-10 March 2002 Cultivated land and forest edge 12-22 August 2002 3a 1500 7°5.155'S, 37°35.124'E 12-21 September 2001 Cultivated land 3b 1500 7°9.125'S, 37°30.565'E 1-20 March 2002 Forest 12-22 August 2002 4 2000 7°6.350'S, 37°36.700'E 23 September-2 October 2001 Montane forest 24 March-2 April 2002 5 2500 7°6.782'S, 37°36.850'E 11 March-9 April 2003 Forest and grassland 24 June-17 July 2003 1 masl, meters above sea level. wide altitudinal range and occupies habitats from the Arthroleptis stenodactylus Pfeffer, 1893 forest and cultivated areas and from the lowest sites, at 1200 m, to the upper montane forests at 2500 m. Individuals of this species were found at 1000, 1200, and This species is terrestrial, found mostly under or on leaf 1500 m (Table 4). This species inhabits the forest interior. litter in the forest. In other Eastern Arc Mountains it is forest edge, and cultivated areas. In forest, this species dwells known to occur from 190 to 2104 m (Poynton & Loader, mostly under leaf litter; large numbers of individuals were 2008). caught in pitfall buckets. Table 2. Altitudinal distribution of amphibian species on the western slopes of the Uluguru South Mountains. X means species detected; blank means undetected. Altitude (m) Family/Species 1000 1200 1500 2000 2500 Arthroleptidae Arthroleptis affinis X X X X Arthroleptis stenodactylus X X X Arthroleptis xenodactyloides X X X Leptopelis parkeri X Brevicipitidae Breviceps mossambicus X Callulina kreffti X X Probreviceps loveridgei X X Probreviceps uluguruensis X X Spelaeophryne methneri X Bufonidae Amietophrynus gutturalis X X X Nectophrynoides viviparus X X Nectophrynoides laevis X Nectophrynoides sp. X Hyperoliidae Hyperolius puncticulatus X X X Kassina senegalensis X Phrynobatrachidae Phrynobatrachus natalensis X X X Phrynobatrachus udzungwensis X Ptychadenidae Ptychadena oxyrhynchus X Strongylopus fuelleborni X Pyxicephalidae Amietia angolensis X X X X Caeciliidae Scolecomorphus uluguruensis X X No. of species 7 12 11 7 5 84 FIELDIANA: LIFE AND EARTH SCIENCES Table 3. Comparison of amphibian community similarity in Probreviceps uluguruensis is highly variable in color and different altitudinal zones (m) on the western slopes of Uluguru South pattern. Mountains using Jaccard’s Index (Sj). Spelaeophryne methneri Ahl, 1924 1000 1200 1500 2000 2500 Only a single adult female measuring 31 mm snout-vent 1000 1 0.75 (75%) 0.67 (67%) 0 0 1200 1 0.64 (64%) 0.14 (14%) 0.06 (6%) length was collected over the entire sampling period in 1500 1 0.20 (20%) 0.07 (7%) cultivated land at 1200 m. 2000 1 0.33 (33%) 2500 1 Family Bufonidae Amietophrynus gutturalis (Power, 1927) Arthroleptis xenodactyloides Hewitt, 1933 Individuals of this species were found at lower altitudes at Individuals of this species were found at 1000, 1200, and 1500 m and below (Table 4). This species is highly versatile; it 1500 m (Table 4). It is a terrestrial species that inhabits both can inhabit various habitat types from degraded forests, moist the leaf litter in the forest and cultivated areas. savanna, and agricultural areas to human settlements. In the study area, this species was restricted to the cultivated land. Leptopelis parkeri Barbour & Loveridge, 1928 No A. gutturalis were recorded in the forest habitat throughout the study period. Males were found calling at Leptopelis parkeri is a tree frog that inhabits the forest at an night from wetter areas and ephemeral water pools in the altitude of 2000 m. It was found within the leaf axils of wild cultivated land at 1000, 1200, and 1500 m. The calls intensified bananas filled with water. Eggs, recovered with the adult frogs during the wet season, especially after rain. from the leaf axils of wild bananas, could be the eggs of Leptopelis. The eggs were small and pigmented; however, it is Nectophrynoides laevis Menegon et al., 2004 difficult to state with certainty that they were L. parkeri eggs without further investigation. This species exhibits sexual This species is restricted to the upper elevations at 2000 m. dimorphism. Males are grayish to brownish or olive green Nectophrynoides laevis appears to have a restricted distribu¬ with a conspicuous yellow pattern forming irregular transverse tion in the Uluguru South Mountain forest and is only known bands; females possess a uniform olive-green dorsum. from one specimen. Nothing is known about its breeding biology and breeding behavior (Menegon et al., 2004). It is a Brevicipitidae tree frog that inhabits the forest habitat and the type was Breviceps mossambicus Peters, 1854 found on vegetation about 1 m off the ground. Individuals of this species were found in the cultivated areas Nectophrynoides viviparus Tornier, 1905 at 1200 m, especially during the dry season, and were mainly found in the leaf litter. In the Uluguru South, this species is restricted to the upper montane forest from 1900 m to the Lukwangule Plateau. Callulina kreffti Nieden, 1911 Individuals were found under or on leaf litter and in grass tussocks on the plateau. They were also observed climbing and This species was recorded at 1200 and 1500 m. It was found calling on vegetation to about 1 m off the ground. This species both in forest and farmland habitats at 1500 m, and at 1200 m is highly variable in color pattern and morphology. Adver¬ only in the forest edge. Individuals were found in rotten logs tisement calls produced by frogs at 2000 and 2500 m differed and leaf axils of wild and cultivated banana plants. Many dramatically. The advertisement call of the male recorded at individuals were either opportunistically collected or visually 2000 m was low-pitched and composed of a sequence of 23 encountered; none were collected in pitfalls. discrete notes. Mean call duration was 54 ms (range, 43-87, n = 3). The fundamental and dominant frequencies were Probreviceps loveridgei Parker, 1931 dissimilar for each note at 1.698 and 1.727 kHz, respectively. Nectophrynoides viviparus from the Lukwangule Plateau Individuals of this species were found at 1200 and 1500 m in (2500 m) has a characteristic call of high-pitched clicks, the Shikurufumi Forest Reserve. It inhabits both forested and “cluck” “cluck,” composed by a series of six to eight notes. cultivated areas at 1500 m, but only in the forest edge at Mean call duration was 475 ms (range, 128-833, n = 6) with 1200 m. Like many other leaf-litter frogs, breeding does not fundamental and dominant frequencies of 2.386 and take place in the water. The females lay their eggs under leaf 2.427 kHz, respectively. The differences in calls suggest that litter, and the young develop directly into adults (Muller et al., the two forms may be distinct; this requires further 2007). Eggs were recovered from the leaf litter throughout the investigation. study, indicating that this species breeds through the year. Nectophrynoides sp. Probreviceps uluguruensis (Loveridge, 1925) During the study, three specimens were collected in a forest This species was collected from 2000 m to the Lukwangule patch on the Lukwangule Plateau. These individuals were Plateau. It is endemic to the Uluguru Mountains and is found distinct from other known taxa and may tentatively be in leaf litter, underground chambers in the forest, and below considered new species. These were found in a hole in a grass tussocks on the Lukwangule Plateau. Morphologically, standing dead tree. NGALASON AND MKONYI: HERPETOFAUNA OF TANZANIA, ULUGURU SOUTH MOUNTAINS 85 Table 4. Number of individuals of different amphibian species found during the surveys in different altitudinal zones (m) on the Uluguru South Mountains. Blank cells indicate no individuals of the species were detected. 1000 1200 1500 2000 2500 Family/Species Dry Wet Dry Wet Dry Wet Dry Wet Dry Wet Arthroleptidae Arthroleptis affinis 6 5 2 15 3 Arthroleptis stenodactylus 1 31 36 4 11 A rthroleptis xenodactyloides 12 6 17 6 12 Leptopelis parkeri 25 7 Brevicipitidae Breviceps mossambicus 9 5 Callulina kreffti 1 5 Probreviceps loveridgei 5 40 6 9 Probreviceps uluguruensis 16 63 45 26 Spelaeophryne methneri 1 Bufonidae Amietophrynus gutturalis 9 21 1 5 7 Nectophrynoides viviparus 7 59 39 31 Nectophrynoides laevis 1 Nectophrynoides sp. 3 Hyperoliidae Hyperolius puncticulatus 5 14 22 4 27 Kassina senegalensis 1 Phrynobatrachidae Phrynobatrachus natalensis 7 1 10 1 27 Phrynobatrachus udzungwensis 12 Ptychadenidae Ptychadena oxyrhynchus 1 Strongylopus fuelleborni 27 128 Pyxicephalidae Amietia angolensis 8 2 8 4 1 8 10 Caeciliidae Scolecomorphus uluguruensis 3 1 Family Hyperoliidae period many individuals were collected from the forest floor; Hyperolius puncticulatus (Pfeffer, 1893) none were collected in dry season. This tree frog was recorded in cultivated areas at 1000 and Family Ptychadenidae 1200 m and at the forest edge at 1500 m but was not found in Ptychadena oxyrhynchus (Smith, 1849) the forest interior. It is highly variable in color and pattern (Harper et ah, 2010). Males were calling on vegetation about This species was collected in the degraded areas (cultivated 0.5-1 m high above the ground near streams. areas) at 1000 m. It was found in abandoned farms dominated by grasses during the dry season. Kassina senegalensis (Dumeril and Bibron, 1841) Family Pyxicephalidae This species was collected in pitfall buckets at forest edge at Amietia angolensis (Bocage, 1866) 1200 m during the wet season. This is a common, semi-aquatic species that occupied Family Phrynobatrachidae cultivated areas at 1000, 1200, and 1500 m to the forested Phrynobatrachus natalensis (Smith, 1849) habitat at 2000 m. It is highly associated with water, being found in rivers and along streams. This species was found at 1000, 1200, and 1500 m near stream banks in cultivated lands and at forest edge at 1500 m. Strongylopus fuelleborni Nieden, 1911 Phrynobatrachus uzungwensis Grandison & Howell, 1983 This species is restricted to the Lukwangule Plateau grasslands in the Uluguru South Mountains (—2400-2500 m). This species was only found in forest habitat during the wet Most individuals were observed near water and only a few season at 1500 m, specifically in the forest floor leaf litter. were found away from a water source. Males call from the Breeding probably occurs in the rainy season. During this grassland swamps and have a characteristic call of a high- 86 FIELDIANA: LIFE AND EARTH SCIENCES pitched “pip” uttered once or in a short burst of three or four. populations might be genetically distinct. Further investiga¬ This species was removed from synonymy with Rana tion is needed to determine their status. The altitudinal limit merumontana Lonnberg, 1910, by Poynton (2004). for lowland and montane forest herpetofauna has been reported from other areas, including Kibale National Park Family Caeciliidae (Vonesh, 2001), Mahenge Mountains (Loader et al., 2004), Scolecomorphus uluguruensis Barbour & Loveridge, 1928 Udzungwa Mountains (Menegon & Salvidio, 2005), and across southern and eastern Africa (Poynton & Broadley, This caecilian was only found in forests from 1500 to 1991; Poynton, 1992, 2003; Poynton & Boycott, 1996). 2000 m. Individuals were mostly captured in pitfall buckets According to Poynton et al. (2006), the differences between and also found in plots during the wet season. montane and lowland amphibian assemblages appear to be long standing, broadly characterized respectively by cool temperate relicts and widespread generalists rather than local elimination of species common to both. Discussion The number of amphibian species at 1200 m was twice the number of species found at 1000 m. Sampling at 1000 m A total of 21 species were recorded during this study. Four included cultivated areas in which all natural vegetation had additional species of amphibians are known to occur in the been cleared and replaced by crops, whereas at 1200 m, Uluguru South Mountains but were not recorded during the sampling included both cultivated areas and forest edge. Most study. These include: Nectophrynoides cryptus (Howell, 1993), of the species found at 1200 m were forest-dependent species N. pseudotornieri (at 2040 m; Menegon et ah, 2004; Frontier (Howell, 1993) that cannot tolerate disturbed habitats. Tanzania, 2005), Hoplophryne uluguruensis (at 2040 m; Fron¬ Furthermore, the 1000-m zone was only sampled during the tier Tanzania, 2005), and Petropedetes yakusini (900-1820 m; dry season, whereas the 1200-m zone was sampled during both Frontier Tanzania, 2005). The unrecorded species are perhaps dry and wet seasons. These reasons may explain the smaller rare or highly cryptic and therefore were not detected. Of the number of species at 1000 than at 1200 m. The effect of habitat 25 amphibian species known from Uluguru South, five species, degradation on composition and diversity of species has been including N. cryptus, N. laevis, N. pseudotornieri, Probreviceps stated (Hillers et al., 2008). Cultivation in the Uluguru uluguruensis, and Scolecomorphus uluguruensis, are endemic to Mountains is normally associated with the removal of plant the Uluguru Mountains. The other five species found in the cover, affecting the thickness of leaf litter, humidity, and Uluguru, including Callulina kreffti, N. viviparus, P. loveridgei, canopy cover. Species like Probreviceps loveridgei, Spelaeo- P. yakusini, and Leptopelis parkeri, are endemic to the Eastern phrvne methneri, and Breviceps nwssambicus were found at Arc Mountains in general (see Poynton et al., 2006). The 1200 m but not at 1000 m. These species are commonly Eastern Arc Mountains are known for high endemism in many associated with leaf litter (Channing & Howell, 2006); it seems taxonomic groups (Dinesen et al., 2001), explained as the human-induced clearance of fields at 1000 m may have consequences of stable local conditions (Burgess et ah, 2007). eliminated them. Hillers et al. (2008) showed that opening the These stable conditions are the result of historical processes in canopy changes the microclimate and most likely exceeds the the Eastern Arc Mountains that would result in species physiological capability of several frog species, and this can replacement patterns that are congruent across many evolu¬ lead to a reduction in diversity of amphibian species. This tionary lineages (Burgess et al., 2007). study demonstrates rather restricted habitat and altitudinal The data indicate a general decline in the number of distribution of some montane endemics, which raises conser¬ amphibian species with increasing elevation. The decline in vation concerns, especially as the increasing loss of habitat at amphibian species richness and diversity with an increasing lower altitudes and, continuing toward higher altitudes, will elevation in tropical mountain forests has been reported across eventually eliminate some species. the globe (e.g., Brown & Alcala, 1961; Fauth et al., 1989). The decrease in temperature, precipitation, and productivity are thought to be the most critical factors limiting species diversity at higher elevations (Rahbek, 1995), although how the Acknowledgments environmental variables impinge on the life of amphibians is not fully understood (Loader et al., 2004). This study has been made possible by the financial support Many of the species found below 1500 m are widely of DANIDA-Enreca Biodiversity Project. The Catchment distributed throughout lowland areas in East Africa (e.g., Forest Office, Morogoro Region, granted permission to work Arthroleptis stenodactylus, A. xenodactyloides, Amietophrynus in the Uluguru South Forest Reserve. We thank Prof. Kim gutturalis, and Ptychadena oxyrhynchus', see Channing & Howell and Prof. A. Nikundiwe of the Department of Howell, 2006), whereas those found above 1500 m, including Zoology and Wildlife Conservation. University of Dar es N. laevis N. viviparus, L. parkeri, and P. uluguruensis, have a Salaam, for their input during the study. We are grateful to more restricted distribution. This shows distinctiveness of Dr. F. Urasa and Dr. C. Msuya of the Department of Zoology highland amphibian fauna from those at the lowland. Recent and Wildlife Conservation, University of Dar es Salaam, for taxonomic studies elsewhere in the Eastern Arc Mountains their logistical support during field work. We also extend our have shown distinctiveness in some of the isolated popula¬ thanks to Prof. J. C. Poynton of the Natural History Museum. tions, such as Callulina (Loader et ah, 2010). In the Uluguru London, and Prof. A. Channing of the University of Western South Mountains, at least three species of Nectophrynoides are Cape, South Africa, for assisting in identification of the known, one of which, N. viviparus, was found at more than specimens. The authors thank Dr. S. P. Loader and W. T. one altitudinal zone (Table 2). The calls of males at 2000 m Stanley for providing helpful comments during the prepara¬ were different from those at 2500 m, suggesting that the two tion of this paper. Rebecca Banasiak helped prepare Figure 1. 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