Southern Highlands glandular tree toad.

Chura manundu wa nyanda za juu kusini.

In the following, we provide a re-description of Nectophrynoides viviparus sensu stricto as revealed by our museomics analysis to comprise the clade from the Southern Highlands; it supersedes previous redescriptions by e.g., Loader et al. (2009), which included non-conspecific members of the complex and cannot therefore be used to distinguish among members of that clade. However, we do not provide a redescription of the lectotype, which was adequately described by Loader et al. (2009).

An adult female specimen in the Museum für Naturkunde, Berlin, Germany, ZMB 21775 collected 2^nd of June 1900 in Ngosi (Ngozi) Crater, Poroto Mountains, Mbeya Region, Tanzania (approximate coordinates: –9.00, 33.56), also known as “Kratersee des Nyisilvulkans” on the original label, by Friedrich Fülleborn (Fig. 7A).

Large series of subadult and adult specimens in the Museum für Naturkunde, Berlin, Germany, ZMB 71524 and 71525 with the same collection data as the lectotype. ZMB 21784, 21788, 25296, 71527 and 71528 collected in “Daressalaam” by Dr. Emil Werth. ZMB 25297 collected in “Amani” by Prof. Dr. Julius Vosseler. ZMB 25261, 25268, 25312, 71529, 71530, 71535 and 71536 collected in “Südliches Deutsch-Ostafrika” (southern Tanzania), ZMB 71187–95, 78704–9 and 78798–803 collected between 26^th and 27^th of October 1899 in “Rugwe (D.O.A.)” (Rungwe, Deutsch-Ostafrika), and ZMB 84908, 84909 and 84910 collected between 26^th and 27^th of October 1899 in “Rugwe-Gebirge”, Tanzania by Friedrich Fülleborn.

One specimen in the Natural History Museum, London, UK, BMNH 1947.2.1945 collected by Friedrich Fülleborn without collection data, is part of the original type series. One specimen in the American Museum of Natural History (AMNH), New York City, New York, USA, AMNH A23562 collected in “Daressalaam”, Tanzania by Dr. Emil Werth, is part of the original type series of Nectophryne werthi (Nieden 1911; Loader et al. 2009).

The paralectotypes ZMB 21784, 21788, 25296, 25297, 71527 and 71528 are also co-types of Nectophryne werthi (Nieden 1911; Loader et al. 2009) and are not conspecific with the rest of the type series of N. viviparus sensu stricto (Fig. 1).

A member of the Nectophrynoides viviparus species complex based on overall body proportions, glandular limbs and large parotoid glands (Fig. 4), as well as genetic affinities based on mitochondrial markers (Fig. 1). This species is characterised by the unique combination of the following set of characters: (1) distinct glandular masses on limbs; (2) medium body size (adult SUL 18.8–37.2 mm, mean 24.76 ± 4.75 mm); (3) fingers slender with rounded discs; and (4) parotoid gland fusiform and widest above arm insertion.

Nectophrynoides viviparus sensu stricto is easily distinguished from N. asperginis, N. cryptus, N. frontierei, N. laevis, N. laticeps, N. minutus, N. paulae, N. poyntoni, N. pseudotornieri, N. tornieri, N. vestergaardi and N. wendyae by having distinct glandular masses on limbs (versus indistinct or absent).

The lectotype is in fair condition, although showing signs of discolouration, softness and other ‘old’ specimen attributes. The paralectotypes range from bad to good condition, the specimens in bad condition are dehydrated, and some have had incisions made on the thigh or inguinal region.

MTSN 9365 and 9383 have been successfully sampled and sequenced (Liedtke et al. 2016). Museomics were done on the following name-bearing types ZMB 21775, 21784, 21788, 25261, 25296, 25297, 25312, 71187 and 71193. Nectophrynoides viviparus sensu stricto is genetically distinct according to Liedtke et al. (2016), who used species delimitation approaches (specifically bGYMC) to examine current bufonid diversity against undescribed diversity. In their analysis, N. viviparus sensu stricto was genetically distinct and identified as “Nectophrynoides viviparus”. MTSN 9365 and 9383 are at least 3.11% genetically different in partial (ca. 562 bp) 16SrRNA from all other Nectophrynoides, with the closest relative being N. saliensissp. nov. (see Table 3). This is rather at the inter-specific level than the infra-specific (population) level; the intra-specific distance between sequenced specimens is 0–0.561%.

The call analysis was carried out on a single audio file consisting of 7 calls with a mean of 13.43 pulses per call in each audio file. The call was recorded 26^th of January 2011 in Mdandu, Livingstone Mountains, the Southern Highlands, Tanzania (–9.7719, 34.7867) at around 2100 m above sea level (a.s.l.) by Michele Menegon near a stream in a closed canopy montane rainforest (Fig. 5). The calls are not associated with any known specimen. However, to our knowledge, this is the only audio file from the Southern Highlands populations, and we therefore cautiously assume that this audio file is a suitable representative of Nectophrynoides viviparus sensu stricto. Audio file containing calls of this species deposited online (https://doi.org/10.5281/zenodo.17277236).

The calls consist of a sequence of 12–15 pulses per call (Fig. 6A). The mean call duration is 0.23 ± 0.03 s (range: 0.20–0.27 s), with a mean call interval of 16.00 ± 5.29 s (7.36–22.27 s). Each call contains a mean of 13.43 ± 1.27 pulses (12–15), with a mean pulse duration of 0.017 ± 0.001 s (0.005–0.019 s). The mean dominant frequency is 1877.71 ± 54.62 Hz (1809–1979 Hz). The call structure is illustrated in a spectrogram and waveform in Fig. 6A. The male advertisement call is monophasic consisting of pulse trains of similar proportions. The first pulse has the highest intensity followed by a series of pulses that slowly decrease in intensity (Fig. 6A). The audio file used for this analysis was sound polluted by a nearby stream. For statistical comparisons between Nectophrynoides viviparus sensu stricto and N. uhehesp. nov. see Table 2. For visual comparisons between N. viviparus sensu stricto and N. uhehesp. nov. see Fig. 6. More behaviour studies and recordings need to be made in the field to rule out certain factors that could shape the call, such as areas of close vicinity with a high competition between males, stress calls, and simplified communicational calls.

The Latin adjective >viviparus, meaning ‘bearing live offspring’. The suggested common name is a reference to the distribution of this species across the Southern Highlands, its glandular skin, and semi-arboreal lifestyle. One of the previous common names of this species was “Morogoro tree toad”, but this is no longer a valid representation of this species since it is not considered to occur in the Morogoro Region or District of eastern Tanzania.

As mentioned in Loader et al. (2009), and with additional collection data from ZMUC specimens, collectors have found this species from approximately 1800 to 2800 m a.s.l. The species (here referring to N. viviparus sensu stricto) has been associated with a range of different habitats such as wet, open, closed, primary, secondary and disturbed forests, ericaceous heathland, montane grassland and bamboo forests. The original description, and observation of toadlets, suggest that this species is ovoviviparous, as in its congeners.

Luhomero glandular tree toad.

Chura manundu wa mlima Luhomero.

This species has previously been referred to as “Nectophrynoides sp06” by Liedtke et al. (2016).

An adult female specimen in the Natural History Museum, London, United Kingdom, BM 1983.6 (KMH 2438) collected on the 25^th of October 1987 in Luhomero Mountains, Udzungwa Mountains National Park, Udzungwa Mountains, Iringa Region, Tanzania (approximate coordinates: –7.78, 36.60) at 2500 m a.s.l. by Jan Kielland (Fig. 7B).

Series of seven subadult and one juvenile specimens in the Museo Tridentino di Scienze Naturali, Trento, Italy, MTSN 8311, 8312, 8397, 8401, 8404, 8405, 8408 and 8409, collected on the 15^th of September 2004 in Luhomero Mountains, Udzungwa Mountains National Park, Udzungwa Mountains, Iringa Region, Tanzania (–7.6965, 36.5722) at 2200 m a.s.l. by Michele Menegon.

A member of the Nectophrynoides viviparus species complex based on overall body proportions, glandular limbs and large parotoid glands (Fig. 8), as well as genetic affinities based on mitochondrial markers (Fig. 1). This species is characterised by the unique combination of the following set of characters: (1) distinct glandular masses on limbs; (2) medium body size (adult SUL 18.4–30.0 mm, mean 21.79 ± 3.77 mm); (3) expanded, rounded finger and toe tips with small discs; (4) parotoid gland rhomboid and slightly pointed posteriorly; (5) relative head width (HW/SUL) 0.35–0.39; and (6) relative head length (HL/SUL) 0.33–0.37.

Nectophrynoides luhomeroensissp. nov. is easily distinguished from N. asperginis, N. cryptus, N. frontierei, N. laevis, N. laticeps, N. minutus, N. paulae, N. poyntoni, N. pseudotornieri, N. tornieri, N. vestergaardi and N. wendyae by having distinct glandular masses on limbs (versus indistinct or absent).

Nectophrynoides luhomeroensissp. nov. is distinguishable from N. viviparus sensu stricto by its slightly smaller body size (SUL 18.4–30.0 mm vs 18.8–37.2 mm), finger- and toe-tips more expanded and less slender and rounded, and parotoid glands rhomboid shaped, slightly pointed posteriorly (Fig. 9B) (vs fusiform shaped and rounded posteriorly [Fig. 9A]). For comparison to the other two new species described herein, refer to the diagnoses of the respective taxa, below.

A medium-sized (SUL: 30 mm) and robust Nectophrynoides with relatively short, muscular, and glandular limbs. The snout is triangular with a rounded tip and extends slightly beyond the upper lip. The canthus rostralis is slightly concave and flattened. The tympanum is distinct. The parotoid glands are distinct and continuous with the dorsal orbits. The parotoid glands extend from the posterior end of the eyes to above the arm insertion in the scapular region forming a rhomboid shape (Fig. 9B). The body has irregular glandular patches scattered across the dorsal and lateral surfaces. The limbs with distinct glandular masses. The length of the foot is greater than the length of the tibia. The hands and feet with rudimentary webbing. The finger and toe-tips are expanded and rounded.

In preserved and alive specimens, the colouration and patterning are highly variable (Fig. 8). Preserved specimens have a cream to dark tawny brown ground colour with darker lateral flanks. The glandular masses are lighter tawny brown with caramel brown patterning or cream brown with little to no patterning.

BM 1983.6 (KMH 2438), an adult female. There are large yolky eggs visible through the abdomen. All measurements are given in mm. A medium-sized and robust specimen (SUL: 30.0, SVL: 30.7). Width of head (HW: 10.7) almost equal to length of head (HL: 10.9). Lower jaw rounded in dorsal and ventral profile with a very slightly blunted snout. Triangular snout slightly rounded anteriorly. In lateral profile, anterior end of snout level with bottom of eye, and inclines to upper jaw. Nostrils situated on either side of snout, at level of eye centre (ND: 2.7), and clearly visible dorsally. Eyes relatively large and bulging in dorsal profile (ED: 3.4). Distance between eye and naris (END: 2.1) greater than distance between naris and tip of snout (NSD: 1.8). In lateral profile, eye and dorsal orbit are continuous with anterior end of snout to scapular region. Canthus rostralis flattened and loreal region slightly concave from top of canthus rostralis to edge of upper jaw. Canthus rostralis visible in dorsal profile. Tympanum and tympanic annulus distinct and rounded. Horizontal diameter of tympanum (TYMP: 1.1) roughly 1/3 of horizontal diameter of eye. Forelimbs muscular and relatively short. Forearm longer than humerus (FOL: 8.0, HUL: 5.9), hand longest (HAL: 9.4). Outer metacarpal tubercle length greater than width (OMCL: 1.9, OMCW: 1.5), inner metacarpal tubercle shortest (IMCL: 1.0). First and third fingertip almost equally expanded (F1W: 0.9, F3W: 0.9). Hindlimbs muscular and relatively long. Tibia and thigh almost equal in length (TIL: 12.3, THL: 12.7), roughly 1/3 longer than metatarsus (ML: 8.0), foot longest (FL: 14.0). Outer metatarsal tubercle length (OMTL: 1.4) shorter than inner metatarsal tubercle (IMTL: 2.1). First toe tip less expanded (T1W: 0.8) than fourth toe tip (T4W: 0.9). Hindlimbs more than twice as long as forelimbs (HIL: 47.0, FORL: 23.3).

Skin texture smooth on glandular and non-glandular surfaces. Dorsal head and dorsum to cloacal region glandular with small pores. Dorsal orbit glandular with medium pores. Dorsum with irregular, large circular glandular masses. Dorsal surface of limbs with glandular masses. Humerus and femur with irregular glandular masses. Forearm, hands, tibia, metatarsus and feet have slightly swollen glandular masses with large pores. Parotoid glands paired and continuous with dorsal orbit. Parotoid glands with large pores and spongy texture. Parotoid glands situated from posterior to eye to scapular region above arm insertion. Parotoid glands rhomboid shaped, widest posterior to eye above angle of jaw and narrows to a slightly pointed shape in scapular region above arm insertion. Parotoid glands extend to lateral surface of tympanic region posterior to tympanum and narrows before arm insertion. Lateral surface of head consists of irregular patches of glandular and non-glandular skin. Posterior and inferior surface of tympanum to posterior end of eye with 10–15 small to medium glandular masses each with a small translucent spine. Flank with glandular patches. Ventral surfaces non-glandular except for femoral area with small, raised bumps. Fingers and toes slender with slightly expanded and rounded digit tips. Hands and feet with distinct tubercles that are raised from the skin. Hands and feet with rudimentary webbing. Feet slightly more webbed extending slightly beyond the first subarticular tubercles.

Dorsal ground colour tawny brown. Head and dorsum with caramel brown patches and spots. Dorsum and femur with tawny brown circular raised glandular bumps without patterning. Parotoid glands, limbs, glandular masses on limbs, hands, and feet tawny brown with caramel brown patches and spots. Femur dark tawny brown close to body and caramel towards knee. Flank ground colour caramel brown with few tawny brown spots toward dorsal margin and cream brown patterning toward ventral margin. Lateral head tawny brown with cream and caramel brown patches. Nostrils caramel brown. Abdomen, pectoral region and chin cream with tawny brown patches and spots. Ventral surface of hands and feet tawny brown with cream tubercles, fingers and toes. Ventral surface of limbs dark tawny brown. Femoral area caramel brown with tawny brown bumps.

No photographs or field notes describing colouration of holotype in life are currently known.

Paratypes are smaller in body size and currently considered subadults. MTSN 8311 with dark tawny brown ground colour covering dorsal and lateral surfaces with tawny brown glands. MTSN 8405 and 8409 with cream ground colour and less distinct tympana. MTSN 8312 and 8405 with several white glandular patches and spots on lateral and dorsal surfaces. Sexual dimorphism was not observed in preserved material; females are expected to be larger than males as seen in congeners, but adult male specimens are needed to confirm this. Photographs and field notes of paratypes, and other individuals, highlight a strong variation in colouration and patterning (Fig. 8).

The holotype is in good condition. The paratypes are generally in fair condition but are soft-fixed, making them fragile and difficult to work with. The condition of paratype MTSN 8312 has deteriorated since measurements were taken; it was then in fair condition but is now poorer.

Paratypes MTSN 8404 and 8405 have been successfully sampled and sequenced (Liedtke et al. 2016). Nectophrynoides luhomeroensissp. nov. is genetically distinct according to Liedtke et al. (2016), who used species delimiting approaches (specifically bGYMC) to examine current bufonid diversity against undescribed diversity. In their analysis, N. luhomeroensissp. nov. was genetically distinct and identified as “Nectophrynoides sp06”. MTSN 8405 is at least 3.13% genetically different in partial (ca. 550 bp) 16SrRNA from all other Nectophrynoides, with the closest relative being N. uhehesp. nov. (see Table 3). This is rather at the inter-specific level than the infra-specific (population) level; the intra-specific distance between sequenced specimens is 0–0.478%.

Not recorded.

The species Nectophrynoides luhomeroensissp. nov. is named after the type locality, which is Luhomero Mountains, within the Udzungwa Mountains National Park, Udzungwa Mountains, Iringa Region, Tanzania. The suggested common name is a reference to the distribution of this species across Luhomero Mountains in Udzungwa Mountains National Park, its glandular skin, and semi-arboreal lifestyle.

The specimens were hand-caught at an elevation of 2200–2500 m a.s.l., and all specimens were found in a forest and grassland mosaic (Fig. 10). Paratypes were found on the moist montane forest floor along the forest edge. The presence of large yolky eggs in the holotype, BM 1983.6, suggest that this species is ovoviviparous, as in its congeners. Two more individuals (JVL 1291–1292) were collected recently by John Lyakurwa, Glory Summay, Christina Kibwe, Anifa John and Pius Mollel in November 2023 (Fig. 8A, B). The two individuals were caught on the ground during the day in a mosaic of shrubs and grasses at 1857 m a.s.l. on the western side of the Luhomero Mountains ~2 km from the Ruipa River (–7.7563, 36.5855). The two individuals show strong resemblance both morphologically and genetically to the type specimens of Nectophrynoides luhomeroensissp. nov. described above.

Udzungwa glandular tree toad.

Chura manundu wa milima ya Udzungwa.

This species has previously been referred to as “Nectophrynoides sp04” by Liedtke et al. (2016).

An adult female specimen in the Natural History Museum of Denmark, Copenhagen, Denmark, ZMUC R131391 (M000044), collected 16^th of December 1997 at Kihanga Stream, Uzungwa Scarp N.F.R., Udzungwa Mountains, Iringa Region, Tanzania (approximate coordinates: –8.37, 35.98) by Mette M. Westergaard (Fig. 7C).

Series of adults and subadults in the Natural History Museum of Denmark, Copenhagen, Denmark: Adult gravid female ZMUC R131389 (M000031), and adults R131390 (M000036) and R131392, all with the same collection data as the holotype. Adult ZMUC H001955 (R970529), collected 26^th of May 1997 in Kihanga Stream, Uzungwa Scarp N.F.R., Udzungwa Mountains, Iringa Region, Tanzania (–8.3683, 35.9783) at 1750 m a.s.l. by David C. Moyer in primary montane forest. Series of adults in the Museo Tridentino di Scienze Naturali, Trento, Italy: MUSE 5247 (MTSN 5247), 5248 (MTSN 5248) and 5249 (MTSN 5249) collected between 4^th and 19^th of January 1999 in Kihanga, Uzungwa Scarp N.F.R., Udzungwa Mountains, Iringa Region, Tanzania (–8.3733, 35.9786) at 1800 m a.s.l. by Michele Menegon in a closed canopy montane rain forest.

A member of the Nectophrynoides viviparus species complex based on overall body proportions, glandular limbs and large parotoid glands (Fig. 11), as well as genetic affinities based on mitochondrial markers (Fig. 1). This species is characterised by the unique combination of the following set of characters: (1) distinct glandular masses on limbs; (2) large body size (adult SUL 17.2–52.5 mm, mean 29.83 ± 8.61 mm); (3) expanded, rounded finger and toe tips with small discs; (4) parotoid gland not continuous with dorsal orbit, kidney shaped; (5) relative head width (HW/SUL) 0.33–0.42; and (6) relative head length (HL/SUL) 0.32–0.41.

Nectophrynoides uhehesp. nov. can be distinguished from N. asperginis, N. cryptus, N. frontierei, N. laevis, N. laticeps, N. minutus, N. paulae, N. poyntoni, N. pseudotornieri, N. tornieri, N. vestergaardi and N. wendyae by its very large body size and having large, distinct glandular masses on limbs (versus indistinct or absent).

Nectophrynoides uhehesp. nov. is distinguishable from N. viviparus sensu stricto by its larger body size (SUL 17.2–52.5 mm vs 18.8–37.2 mm) and having more expanded, distinct limb and parotoid glands. The shape of finger- and toe tips of N. uhehesp. nov. are more expanded than N. viviparus sensu stricto, which has more slender and rounded fingers. The parotoid glands are not continuous with the dorsal orbits, they are large and protruding, forming a rough kidney shape (Fig. 9C), whereas the parotoid glands of N. viviparus sensu stricto are continuous with the dorsal orbits, they are smaller and less protruding, forming a rough fusiform shape (Fig. 9A).

Nectophrynoides uhehesp. nov. is distinguishable from N. luhomeroensis by a larger maximum body size (SUL 17.2–52.5 mm vs 18.4–30.0 mm) and less distinct glandular limbs, and kidney-shaped and more pronounced parotoid glands (Fig. 9C) (vs rhomboid and less pronounced; Fig. 9B).

For distinction from N. saliensissp. nov., refer to the diagnosis of that species, below.

A large and robust Nectophrynoides with relatively short, muscular and very glandular limbs. The snout shape is triangular with a rounded tip, extending slightly beyond the upper lip. The canthus rostralis is slightly concave and flattened. The tympanum is distinct. The parotoid glands are distinct and continuous with the dorsal orbits. The parotoid glands extend from the posterior end of the eyes to above the arm insertion in the scapular region forming a rough kidney shape (Fig. 9C). The body has medium sized irregular glandular bumps and patches scattered across the dorsal and lateral surfaces. The limbs with distinct and expanded glandular masses. The hindlimb is more than twice as long as the forelimb. The length of the foot is greater than the length of the tibia. The hands and feet with rudimentary webbing. The finger and toe-tips are expanded and rounded.

In preserved specimens, the colouration and patterning are highly variable. The ground colour is caramel to very dark tawny brown with cream to tawny brown glandular bumps and patterning. Several specimens with large amounts of patterning in different shapes and sizes of variable colours from white to dark tawny brown. The glandular masses on limbs and the parotoid glands are cream to tawny brown with no patterning, or with caramel to dark tawny brown patterning.

ZMUC R131391 (M000044), an adult female. All measurements are given in mm. Large and robust specimen (SUL: 45.2, SVL: 47.7). Width of head (HW: 17.6) greater than length of head (HL: 16.7). Lower jaw rounded in dorsal and ventral profile with flattened and blunted snout. Very wide triangular snout and very slightly rounded anteriorly. In lateral profile, anterior end of snout is level with bottom of eye. Nostrils situated on either side of snout, at level of eye centre (ND: 4.3), and clearly visible dorsally. Eyes relatively large and bulging in dorsal profile (ED: 5.0). Distance between eye and naris (END: 3.3) greater than distance between naris and tip of snout (NSD: 2.4). In lateral profile, eye and dorsal orbit continuous with anterior end of snout to posterior end of eye. Canthus rostralis flattened and loreal region slightly concave from top of canthus rostralis to edge of upper jaw. Canthus rostralis visible in dorsal profile. Tympanum and tympanic annulus distinct and rounded. Horizontal diameter of tympanum (TYMP: 1.7) almost 1/3 of horizontal diameter of eye. Forelimbs muscular and relatively short. Forearm longer than humerus (FOL: 11.7, HUL: 8.7), hand longest (HAL: 15.0). Outer metacarpal tubercle length almost equal to width (OMCL: 2.3, OMCW: 2.4), inner metacarpal tubercle shortest (IMCL: 1.8). First fingertip less expanded (F1W: 1.1) than third fingertip (F3W: 1.3). Hindlimbs muscular and relatively long. Tibia and thigh almost equal in length (TIL: 19.6, THL: 19.9), almost twice as long as metatarsus (ML: 11.6), foot longest (FL: 24.3). Outer metatarsal tubercle length (OMTL: 2.2) shorter than inner metatarsal tubercle (IMTL: 2.9). First and fourth toe tip equally expanded (T1W:1.2, T4W: 1.2). Hindlimbs more than twice as long as forelimbs (HIL: 75.3, FORL: 35.4).

Skin texture smooth on glandular and non-glandular surfaces. Dorsal head glandular with small pores. Dorsal orbits glandular with large pores. Dorsum with large, irregular, circular glandular bumps. Dorsal surface of limbs with distinct glandular masses. Humerus and femur with irregular glandular masses. Forearm, hands, tibia, metatarsus and feet have distinct, swollen glandular masses with large pores. Parotoid glands paired and continuous with dorsal orbits. Parotoid glands with large pores and spongy texture. Parotoid glands situated from posterior to eye to scapular region above arm insertion. Parotoid glands rough and asymmetrical kidney shape, widest posterior to tympanum above angle of jaw and narrows to a point above arm insertion. Parotoid glands extend to lateral surface of tympanic region posterior to tympanum and narrows before arm insertion. Lateral head consists of irregular patches of glandular and non-glandular skin. Canthus rostralis has glandular skin with small pores. Posterior and inferior surface of tympanum to posterior end of eye has 25 medium to large glandular masses each with a small translucent spine. Flank without glandular patches. Ventral surfaces non-glandular except femoral area with small, raised bumps. Fingers and toes stout with expanded and rounded digits. Hands and feet with distinct, raised tubercles and rudimentary webbing. Feet slightly more webbed extending slightly beyond the first subarticular tubercles.

Dorsal ground colour tawny brown. Head caramel brown. Dorsum tawny brown with large caramel brown raised circular glandular masses. A dark tawny brown indistinct and broken dorsal stripe runs from snout to cloaca. Tawny brown glandular masses on humerus and femur. Parotoid glands and glandular masses on limbs, hands and feet cream to caramel brown. Fingers and toes cream. Flank dark tawny brown. Lateral head tawny brown with cream and caramel brown patches. Nostrils caramel brown. Canthus rostralis, upper and lower lip cream. Dorsal orbits bluish ash grey. Abdomen dark tawny brown. Pectoral region and chin tawny brown with indistinct cream and caramel brown spots. Ventral surface of hands and feet tawny brown with cream tubercles, fingers and toes. Ventral surface of forelimbs dark tawny brown and hindlimbs tawny brown. Femoral area tawny brown with caramel brown bumps.

No photographs or field notes describing colouration of holotype in life are currently known.

MUSE 5247, 5248 and 5249 with irregular patterns and spots covering dorsal and ventral surfaces with white blotches on head, chin, back, abdomen, flank, limbs, hands and feet. ZMUC R131389, R131390 and R131392 are smaller (SUL: 22.79–26.94 vs 45.22) with more slender fingers and toes, and less distinct glandular masses on limbs, hands and feet. There is a large variety in colouration and patterning between individuals (Fig. 11). Sexual dimorphism was not observed in preserved material, but females are expected to be larger than males, like congeners.

The holotype and paratypes are in good condition.

Paratypes MUSE 5248 and 5249 have been successfully sampled and sequenced (Liedtke et al. 2016). Nectophrynoides uhehesp. nov. is genetically distinct according to Liedtke et al. (2016), who used species delimiting approaches (specifically bGYMC) to examine current bufonid diversity against undescribed diversity. In their analysis, N. uhehesp. nov. was genetically distinct and identified as “Nectophrynoides sp04”. The specimen used in the analysis done in Liedtke et al. (2016) was MHNG 2609.071 (field number TZ-088), which is effectively 100% genetically identical to paratype MUSE 5249. MUSE 5249 is at least 3.13% genetically different in partial (ca. 550 bp) 16SrRNA from all other Nectophrynoides, with the closest relative being N. luhomeroensissp. nov. (see Table 3). This is rather at the inter-specific level than the infra-specific (population) level; the infra-specific distance between sequenced specimens is 0–0.481%.

The call analysis was carried out on two audio files both consisting of 11 calls with a mean of 25.27 pulses per call in each file. The two calls were recorded in January 1999 in Site 5, Mkaja, Uzungwa Scarp N.F.R., Udzungwa Mountains, Tanzania (–8.3420, 35.9671) at 1800 m a.s.l. by Michele Menegon in an open montane wetland consisting of an ecotone between bamboo forest and open wetland (Menegon and Salvidio 2005). The two calls are not associated with any known specimens. However, the two calls were recorded nearby to the type locality. Therefore, we assume that these calls are suitable representatives of Nectophrynoides uhehesp. nov. A singular audio file containing calls of this species deposited online (https://doi.org/10.5281/zenodo.17277236).

The two calls were practically identical in all aspects of their call parameters. We cannot confirm whether these are of one or two individuals. The calls consist of a sequence of 16–30 pulses per call (Fig. 6B). The mean call duration is 0.28 ± 0.03 s (range: 0.22–0.32 s), with a mean call interval of 3.24 ± 1.27 s (2.10–5.95 s). Each call contains a mean of 25.27 ± 3.66 pulses (16–30), with a mean pulse duration of 0.011 s +/– 0.001 s (0.011–0.014 s). The mean dominant frequency is 1891.64 ± 7.17 Hz (1883–1904 Hz). The call structure is illustrated in a spectrogram and waveform in Fig. 6B.

The male advertisement call is monophasic consisting of pulse trains of similar proportions. The first pulse has the highest intensity followed by a series of pulses that slowly decreases in intensity. The two recordings show remarkable resemblance, with almost identical results. It is worth noting that both call series begin by a pulse train that has a significantly lower number of pulses. The background noises from the habitat, people talking, and other interferences indicate that these two individual call series were taken separately and illustrate a remarkable consistency within the call structure of this species.

The few recordings highlight the need for further field research and analysis of the bioacoustics in the Nectophrynoides viviparus species complex. We can therefore only hint at the possibility that the call of N. uhehesp. nov. is distinctive, and easily distinguishable from other Nectophrynoides species in the field. The call of the new species is distinct from N. viviparus sensu stricto which has fewer pulses in each pulse train (mean is ~13 pulses), and a longer duration between calls (Fig. 6A). For the statistical analysis between the two species see Table 2. However, the low number of recordings result in uncertainty regarding the actual difference between calls. More behaviour studies and recordings need to be made in the field to rule out certain factors that could shape the calls such as close vicinity with high competition between males, stress calls, and simplified communicational calls.

The species Nectophrynoides uhehesp. nov. is named in honour of the indigenous Hehe tribe, who live in villages surrounding the forests where the species occurs, for their support/involvement in herpetological surveys in the area. The Swahili word “>uhehe” indicates something with affinity to the Hehe tribe. The suggested common name is a reference to the distribution of this species across the Udzungwa Mountains, its glandular skin, and semi-arboreal lifestyle.

Specimens have been collected in several forest fragments across the Udzungwa Mountains, including Kigogo F.R., Kiolela F.R., Kitungulu F.R., Mufindi Scarp F.R., Uzungwa Scarp N.F.R., and Kilombero N.F.R. The type specimens were collected in closed canopy montane rainforest between 1700 and 1800 m a.s.l. near Kihanga stream inside Uzungwa Scarp N.F.R. (Fig. 12). The presence of developed toadlets inside several adult female specimens suggests that this species is ovoviviparous, as in its congeners. More recent observations are known by John Lyakurwa, Michele Menegon and Elena Tonelli in 2014–2016 at multiple locations within the Uzungwa Scarp N.F.R. from 1600 m a.s.l. Individuals were observed on the ground, around small wetlands both inside and bordering the forest, and some specimens were observed up to 2 m above the ground on understory vegetation in the moist forest inside closed canopy montane rainforest.

Mahenge glandular tree toad.

Chura manundu wa Mahenge.

This species has previously been referred to as “Nectophrynoides sp13” by Liedtke et al. (2016).

An adult presumably female specimen in the Museo Tridentino di Scienze Naturali, Trento, Italy, MUSE 13758 (KMH 26644) collected December 2005 in Sali F.R., Mahenge Mountains, Morogoro Region, Tanzania (approximate coordinates: –8.93, 36.65) at 1050–1500 m a.s.l. by Frontier-Tanzania (Menegon et al. 2011a) (Fig. 7D).

Series of adults and subadults: MUSE 13754 (KMH 26637), 13755 (KMH 26638), 13756 (KMH 26639), 13757 (KMH 26641) and 13759 (KMH 26998), all with the same collection data as the holotype.

A member of the Nectophrynoides viviparus species complex based on overall body proportions, glandular limbs and large parotoid glands (Fig. 7D), as well as genetic affinities based on mitochondrial markers (Fig. 1). This species is characterised by the unique combination of the following set of characters: (1) indistinct glandular masses on limbs; (2) medium body size (adult SUL 20.8–34.3 mm, mean 26.30 ± 5.02 mm); (3) weakly expanded rounded finger and toe tips with minute discs; (4) parotoid gland spearhead shaped; (5) relative head width (HW/SUL) 0.40–0.43; and (6) relative head length (HL/SUL) 0.37–0.40.

Nectophrynoides saliensissp. nov. is easily distinguished from N. asperginis, N. cryptus, N. frontierei, N. laevis, N. pseudotornieri, N. wendyae by having a distinct tympanum.

The smooth dorsal surface of Nectophrynoides saliensissp. nov. is covered with glandular patches and warts, indistinct glandular masses on limbs, and the parotoid glands form a large distinct spearhead shape, which distinguishes it from N. laticeps, N. minutus, N. tornieri, N. paulae, N. poyntoni, N. vestergaardi.

Nectophrynoides saliensissp. nov. is distinguishable from N. viviparus sensu stricto by having indistinct limb glands. The finger- and toe-tip expansion of N. saliensissp. nov. are also more expanded than N. viviparus sensu stricto, which has more slender and rounded fingers. The parotoid glands are elongated, and spearhead shaped, narrowing to a thin acuminate shape posteriorly (Fig. 9D), whereas the parotoid of N. viviparus sensu stricto are fusiform shaped and rounded posteriorly (Fig. 9A).

Nectophrynoides saliensissp. nov. is distinguished from Nectophrynoides luhomeroensis by lacking distinct limb glands (vs distinct), differently shaped parotoid glands (spearhead vs rhomboid [Fig. 9D and Fig. 9B, respectively]), larger relative head size (HL/SUL 0.37–0.40 vs 0.33–0.37, and HW/SUL 0.40–0.43 vs 0.35–0.39), and from N. uhehe by a much smaller maximum body size (SUL 34.3 mm vs 52.5 mm), by lacking distinct limb glands (vs distinct), and differently shaped parotoid glands (spearhead vs kidney [Fig. 9D and Fig. 9C, respectively]).

A medium-sized (SUL: 34.3 mm) and robust Nectophrynoides with relatively short, muscular and slightly glandular limbs. The snout shape is triangular with a rounded tip and extending slightly beyond the upper lip. The canthus rostralis is angular. The tympanum is distinct. The parotoid glands are distinct and continuous with the dorsal orbits. The parotoid glands extend from the posterior end of the eyes to above the arm insertion in the scapular region forming a rough spearhead shape (Fig. 9D). The body has small irregular glandular bumps and patches scattered across the dorsal and lateral surfaces. The limbs with indistinct glandular masses. The length of the foot is greater than the length of the tibia. The hands and feet with rudimentary webbing. The finger and toe-tips are expanded and rounded.

In preserved specimens, the colouration and patterning are variable. The ground colour is very dark tawny brown with ash grey glandular bumps. The glandular masses on limbs and the parotoid glands are ash grey with dark tawny brown patterning.

MUSE 13758 (KMH 26644), presumably female adult. The specimen has a mid-ventral incision along the body. All measurements are given in mm. A medium-sized and robust specimen (SUL: 34.3, SVL: 35.0). Width of head (HW: 13.6) almost equal to length of head (HL: 13.1). Lower jaw rounded in dorsal and ventral profile with slightly flattened and blunted snout. Wide triangular snout slightly rounded anteriorly. In lateral profile, anterior end of snout is level with bottom of eye. Nostrils situated on either side of snout, at level of eye centre (ND: 2.3), and clearly visible dorsally. Eyes relatively large and bulging in dorsal profile (ED: 3.6). Distance between eye and naris (END: 3.0) greater than distance between naris and tip of snout (NSD: 1.5). In lateral profile, eye and dorsal orbit continuous with anterior end of snout to scapular region. Canthus rostralis angular and loreal region concave from top of canthus rostralis to edge of upper jaw. Canthus rostralis visible in dorsal profile. Tympanum and tympanic annulus distinct and rounded. Horizontal diameter of tympanum (TYMP: 1.4) less than half of horizontal diameter of eye. Forelimbs muscular and relatively short. Forearm longer than humerus (FOL: 9.3, HUL: 6.8), hand longest (HAL: 11.4). Outer metacarpal tubercle length equal to width (OMCL: 2.0, OMCW: 2.0), length of inner metacarpal tubercle shortest (IMCL: 1.5). First fingertip less expanded (F1W: 0.9) than third fingertip (F3W: 1.0). Hindlimbs muscular and relatively long. Tibia longer than thigh and metatarsus (TIL: 17.8, THL: 17.0, ML: 10.3), foot longest (FL: 18.9). Outer metatarsal tubercle length (OMTL: 1.7) shorter than inner metatarsal tubercle (IMTL: 2.6). First and fourth toe tip equally expanded (T1W:1.0, T4W: 1.0). Hindlimbs more than twice as long as forelimbs (HIL: 64.1, FORL: 27.5).

Skin texture smooth on glandular and non-glandular surfaces. Dorsal head glandular with small pores. Dorsal orbits glandular with medium pores. Dorsum with small, irregular, raised glandular bumps. Front and hind limbs with indistinct glandular masses. Humerus and femur have small irregular glandular bumps. Forearm, hands, tibia, metatarsus and feet have indistinct glandular masses with large pores. Parotoid glands paired and continuous with dorsal orbits. Parotoid glands with medium pores. Parotoid glands situated from posterior to eye to scapular region above arm insertion. Parotoid glands rough spearhead shaped, widest posterior to eye above angle of jaw and tympanum, narrowing to an acuminate shape in scapular region above arm insertion. Parotoid glands extend to lateral surfaces of tympanic region posterior to tympanum and become irregular patches of glandular and non-glandular skin. Lateral head consists of irregular patches of glandular and non-glandular skin. Posterior and inferior surface of tympanum to posterior end of eye has 10–15 small to medium glandular masses each with a small translucent spine. Flank with small, irregular, raised glandular bumps. Ventral surfaces non-glandular except for femoral area with small, raised bumps. Fingers and toes slender with slightly expanded and rounded digit tips. Hands and feet with distinct, raised tubercles and rudimentary webbing. Feet slightly more webbed extending slightly beyond the first subarticular tubercles.

Dorsal ground colour very dark tawny brown. Dorsum and flank with many ash grey, raised, glandular bumps. Dorsal head with a large ash grey glandular patch with very dark tawny brown spots. Limbs, hands and feet with ash grey glandular patches, spots and stripes. Tympanic region tawny brown with ash grey glandular patches. Lateral head ash grey. Canthus rostralis and nostrils tawny brown. Dorsal orbits bluish ash grey. Ventral surface of limbs, abdomen and chin cream. Pectoral regions cream with caramel brown patches. Ventral surfaces of hands and feet caramel brown with cream tubercles, fingers and toes. Femoral area caramel brown with cream bumps.

No photographs or field notes describing colouration of holotype in life are currently known.

MUSE 13755 tawny brown dorsal ground colour with very dark tawny brown patterning on dorsal and ventral surfaces. MUSE 13755 has a less distinct tympanum. MUSE 13755 and 13759 have very slightly pointed snout tips. Sexual dimorphism was not observed in preserved material, but females are expected to be larger than males, like congeners.

The holotype and paratypes are in fair condition but show evidence of previous exposure to unsuitable preservation. The specimens are shrunken, stiff, and partially desiccated, with dried finger- and toe tips.

Holotype MUSE 13758 and paratypes MUSE 13754, 13755, 13757 and 13759 have been successfully sampled and sequenced (Liedtke et al. 2016). Nectophrynoides saliensissp. nov. is genetically distinct according to Liedtke et al. (2016), who used species delimiting approaches (specifically bGYMC) to examine current bufonid diversity against undescribed diversity. In their analysis, N. saliensissp. nov. was genetically distinct and identified as “Nectophrynoides sp13”. MUSE 13758 is 3.11% genetically different in partial (ca. 550 bp) 16SrRNA from all other Nectophrynoides, with the closest relative being Nectophrynoides viviparus sensu stricto (see Table 3). This is rather at the inter-specific level than the infra-specific (population) level; the intra-specific distance between sequenced specimens is 0–0.718%.

Not recorded.

The species Nectophrynoides saliensissp. nov. is named after the location where the species was discovered, which is Sali F.R. in Mahenge Mountains, Tanzania. The suggested common name is a reference to the distribution of this species in the Mahenge Mountains, its glandular skin, and semi-arboreal lifestyle.

There is very limited knowledge of habitat, ecology, and behaviour of this species. The specimens were discovered by Frontier-Tanzania in a dense, low canopy submontane forest near open canopy wetlands in the northern inaccessible forests of Sali F.R. (Fig. 13). A paper published based on the work by Frontier-Tanzania in Mahenge Mountains describes Sali F.R. as a landscape consisting of miombo woodlands at lower elevation, submontane and montane forests, dry grassland, wetlands and large rocky outcrops (Menegon et al. 2011a). Little is known about the reproductive biology of this species but the presence of large, developed embryos visible through the abdomen of paratype MUSE 13757 suggests that the species is ovoviviparous, as in its congeners.