Corresponding author: Alexander Haas (
Academic editor: Andreas Schmidt-Rhaesa
The current account presents the results of a 14-day amphibian survey at Maliau Basin Conservation Area (
To the general public, the amphibian species of East Malaysia (Sabah and Sarawak, Borneo) may seem to be relatively well-known. Particularly, the books by
The State of Sabah has created a number of national parks and conservation areas of various protection levels. Maliau Basin Conservation Area (
The Maliau Basin is an almost circular, saucer-like geological structure, approximately 25 km in diameter, and comprising Neogene mudstone and sandstone deposits (
Considering the relative short period of scientific exploration, it is not surprising that there is still no comprehensive amphibian inventory of the Maliau Basin. A first list of amphibian species was published in
In early 2017, we were granted the opportunity to conduct a rapid amphibian survey of Maliau Basin. Time and manpower were limited, so that a complete list of all
Field work was mainly conducted from 16 February 2017 to 2 March 2017. With respect to a comprehensive species list, additional photographic evidence by one of us (MBA) was added but was not restricted to the survey period and localities. In the following, we refer to these four main collecting sites (camps), including areas that could be reached from those sites during transect walks:
1) Study Center (
2) Agathis Camp (
3) Ginseng Camp (
4) Nepenthes Camp (
Surveys at each of the sites where conducted mostly between 19:00–24:00 h. Specimens were collected during nightly transect walks and included visual and acoustic encounter techniques (
Living specimens were photographed with a Canon 5D IV camera and Cannon 100 mm macro lens (for adult frogs) or a Fujifilm X-T2 camera and Fujinon 60 mm macro lens (tadpoles), some under field but mostly under laboratory conditions with double flash lighting. Photos were taken in RAW format and subsequently developed in Capture One 10 or AfterShot Pro3 software. Voucher specimens were euthanized with an aqueous chlorobutanol solution (adults) or Eugenol (
Taxonomy follows
The following primers were used: For the
Between 16 February–2 March 2017, a total of approximately 170 man-hrs was expended and 44 species of frogs were detected at four study sites within
List of anuran species encountered at
Taxon | Agathis Camp | Ginseng Camp | Nepenthes Camp/Trails | Study Center/Trails/Maliau |
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Snout obliquely protruding in lateral view; snout tip pointed in lateral view; body generally slender; dorsum brown to olive, with blurred dark markings; finely warted dorsally, warts without keratinized spines; dorsolateral warts slightly larger, some fused to form a ridge from above tympanum to shoulder; belly gray with dense brown spots; throat dark brown; light marking under the eye and between angle of jaw and shoulder; toes and fingers possess rounded, slightly swollen tips; iris black with dense golden iridophores; more reddish in anterior and posterior sector; tympanum distinct, small.—This species was common at Fawzy Fall near Nepenthes Camp. Specimens were observed at or in the stream, sitting on low vegetation or rocks.
Genetic barcoding and sequence analysis (see Appendix
A slender toad, variable brownish in color, sometimes with tinges of orange; only faint color markings, indistinct crossbars on thigh and shank; small rounded warts on dorsum, larger flat warts at lateral dorsum and flanks; throat dark with lighter mottling; abdominal skin pebbled, mostly amber in color; snout projecting obliquely in lateral view; finger and toe tips rounded and slightly swollen; toes three-fourth webbed in males slightly less in female; no tarsal ridge; tympanum visible, small; iris golden with black network.—One female was recorded from a rocky tributary of Sungai Maliau. This species occurred in syntopy with
Snout obliquely protruding in lateral view; obtuse, rounded snout tip in lateral view; small conical warts on dorsum and dorsal side of limbs, often with small black spines giving a rough appearance; limbs and fingers long; almost uniformly brown to dark brown above, sometimes with reddish hue; diffuse scattered dark spots on the body; crossbars on thigh and shank may be indistinct in a dark marbled pattern; cream to amber with dark markings ventrally; darker on the throat and chest; toes and fingers bear rounded, slightly swollen tips; iris mostly golden to reddish-golden, with black reticulated pattern and thin reddish ring around pupil; tympanum distinct but small.—Two specimens were collected at the streams of Ginseng Camp, perching on low vegetation.
Snout obliquely protruding in lateral view; snout tip obtuse and rounded in lateral view; body generally slender; dorsum mostly black, with lighter yellowish or reddish warts laterally and in bands of such lighter warts on fore- and hind limbs; with a light cream, gray or reddish patch between shoulders; coarsely warted dorsally, warts with keratinized black spines; belly pebbled in texture, dark brown to black in color, with white markings; ventral side of thigh pebbled with scattered white spots; toes and fingers possess rounded, slightly swollen tips; iris black with some reddish stippling and a reddish ring around the horizontal pupil; tympanum distinct, vertically oval.—This species was common along the river at Ginseng Camp. Specimens were observed sitting on low vegetation close to the river. Encounters included numerous juvenile specimens along the Ginseng Camp stream below Ginseng Falls. One tadpole was captured at the river above Ginseng Fall.
Body stocky; two hard ridges between eyes; parotoid gland narrow, longer than wide, posteriorly followed by row of warts; in general, coarsely warted above; warts pointed, with keratinized spinules; ventral skin coarsely pebbled in texture, cream with black spots in color; brown above with darker markings between eyes and on back; narrow vertebral stripe; limbs with cross bars ; sometimes with orange or reddish warts; ear drum visible but not different in color from surrounding skin; iris golden or reddish-gold with a black network.—
A slender toad with long limbs, long fingers and toes; generally mid to dark brown above with various dark markings, particularly a diagnostic thick dark ‘X’-shaped mark, with cream contour line, on middle of dorsum and a less pronounced one on shoulder to head transition (in our specimen separated into distinct spots); lips with dark bars; belly cream with some dark markings; underside of thigh orange; parotoid glands narrow, elongated and indistinct, extending posteriorly into dorsolateral ridge of warts; iris golden (sometimes reddish in anterior and posterior sector of iris) with black reticulate pattern; ear drum indistinct in color and posteriorly covered by skin; slightly swollen finger and toe tips;—A juvenile was collected at Ginseng Camp, along the rocky stream above Ginseng Falls, perching on low vegetation.
A large and stocky toad; with large, prominent parotoid gland behind eye; large warts all over the dorsal side; warts rounded to conical, bearing spinule; dorsal coloration uniformly grayish to mud brown; dark markings present or absent; ear drum small and indistinct; iris, golden with fine, dark reticulation—
To our knowledge, photos of
Frogs of the
Body slender to stocky; narrowing snout in dorsal view; snout projecting well beyond lower jaw in lateral view; finger and toe tips not expanded; prominent broken longitudinal ridges on back; coloration from shades of gray to brown and greenish hues, with scattered dark blotches; often with vertebral stripe; alternating light and dark marks across lips; belly white or cream underside of hind limb flesh colored; males with dark throat markings; ear drum moderately distinct; iris brass in the upper sector, silvery in the lower; outside the iris the eye bulb is white-bluish.—
A large, heavy frog with muscular, long legs; area between nostril and eye smooth, not forming an edge (i.e., no canthus rostralis); head broad and flat; finger and toe tips slightly bulbous but not expanded; toes fully webbed; dorsal skin finely textured, shagreened, with scattered tubercles and small ridges; multiple conical tubercles on top of the eye; supratympanic fold long, with black line, bending in angled manner around tympanum to shoulder; tympanum moderately distinct (colored as surrounding skin); color homogeneously brown above; some black blotches posterior to shoulder; sometimes red on thigh; dark markings on upper and lower lip; a thin midline cream colored streak can be present or absent; thin banding on thigh; venter white, dotted brown, throat dark mottled; thighs pale with some cream spots; iris densely stippled silver-gold on black background, venation pattern.—This large species was encountered in the riparian habitat along Sungai Maliau, from one of its tributaries.
Although the Creek Frog species complex (formerly known as
Robust large frog with strong hind limbs and broad triangular head; snout long and tapering in dorsal view, projecting well beyond lower jaw in lateral view; mostly smooth, dorsal skin texture shagreened, occasionally with scattered elongated tubercles and short ridges on back, esp. in juveniles; tubercles on top of eye; toes fully webbed; grayish, sandy brown to mid-brown above with scattered dark spots; dark band between eyes; dark streak from eye to snout along an edge between eye and nostril (canthus rostralis); dark markings on lips; dark below supratympanic fold and upper part of ear drum; supratympanic fold being in a moderate angular manner around tympanum; tympanum moderately distinct; hind limb barred above; belly uniform cream; throat faintly mottled; odontoid processes on anterior of lower jaw; iris golden with three more or less pronounced dark radial areas (anterior, posterior and ventral sectors) and fine black venation.—This large river frog is common along lowland rivers of Sabah and Sarawak and was detected at the Sungai Maliau tributaries, Ginseng and Agathis Camps. Often these frogs sit on the ground of the river bank a few meters from the water line. One male was observed building a nesting site, i.e., a circular depression in the fine gravel of one of Sungai Maliau’s tributaries. The male was calling with a deep single note and a female was noticed nearby.
Snout long and tapering in dorsal view; bulging bluntly beyond lower jaw in lateral view; toe tips slightly widened, finger tips rounded and not widened; toes webbed to almost the tip, except for Toe IV; webbing; dorsal side sandy brown, mid-brown or reddish brown, uniform or with irregular markings; supratympanic fold bending around tympanum; tympanum almost entirely covered with dark fleck below supratympanic fold; dark streak from eye to nostril along canthus rostralis; contrasty light and dark markings on upper and lower lips; faint banding on thigh; venter yellowish; throat mottled; dorsal skin texture shagreened with some low tubercles and short ridges on back; odontoid processes on anterior of lower jaw; iris golden with four conspicuous dark radials and fine black venation;—
Limbs slender; head wider than body; eyes large; toes nearly free of webbing; hind limbs relatively short; forelimbs long and thin; supratympanic fold arched around ear drum; upper part of tympanum with a dark mask; skin of dorsum smooth to granulated; dark brown above, with a darker pattern on forehead; ventral side smooth; abdominal skin dark grayish with white spots partially arranged in rows; throat white with grayish markings; iris dark brown to black; beyond the iris the eye is blueish.—The species was present on the forest floor along Sungai Maliau (Wildlife Trail) and at Nepenthes Camp. A calling chorus of this species was noticed at Agathis Camp.
Snout rounded; vocal sacs of males subgular and bipartite; supratympanic fold angled; tympanum distinct, dark; venter white with darker blotches; generally grayish or brownish to anthrazite above with some light brown flecks and large dark blotches; blotches and flecks not bordered in light color; dorsal surface of upper arm and part of hind limb lightened; thigh with distinct banding; iris grayish with reddish sector dorsally and red iris edge around pupil.—Calling males were detected along a tributary right of Sungai Maliau. This side stream was rocky and had large boulders. Males perched on low vegetation (< 50 cm) up to 10 m away from the waterline, elevated on the steep river banks.
Body robust, particularly in females; relatively short hind limbs; head large, wider than long; prominent triangular, dermal appendages on tip of snout and upper eyelids; ear drum hidden; two pairs of parallel dermal folds on back; dorsal skin mostly smooth with scattered tubercles, some of the conical; tips of fingers and toes rounded, not expanded; toes webbed at base; dorsum brown with darker markings; a dark bar from eye to upper lip; limbs with narrow crossbars; throat reddish-brown to dark brown in males; posterior half of ventral side whitish with dark flecks; iris dark brown to reddish brown, somewhat lighter in upper part.—The species was not active at the time of our survey. One specimen was encountered near Study Center, Belian Trail.
Frogs of the
Broad flat body with long hind limbs; snout contour in dorsal view tapering but narrowly rounded; finger tips moderately widened, toe tips clearly widened into discs, somewhat truncated digital discs; fully webbed toes, except for fourth toe; dorsal skin texture relatively smooth, with scattered fine tubercles; color brown above, with dark and light-edged polygonal marking from shoulder to mid-back; banding pattern on limbs; venter yellow, ventral side of limbs dull yellow; throat dark and finely mottled brown; iris brass with dark reticulate pattern; ear drum invisible;—This microhylid ground-dwelling species was encountered at and around swampy areas and roadside ditches along Belian Trail.
Stocky, broad, flat body, short limbs; narrow head relative to body, short pointed snout; all fingers short except third; outer finger and toe very short, nub-like; webbing reduced; dorsal skin texture granulated to moderately tuberculate; color red-brown above, with faint dark markings; toes and fingers bright orange to red; inguinal spot light colored and distinctly edged dark; row of well-defined cream tubercles from above ear drum towards groin; dark-edged cream inguinal spot; similar smaller spots on rear of thigh; iris bronze with fine reticulate pattern; pupil horizontal; ear drum indistinct.—The species was tentatively assigned to
Body stocky, head small in relation to body, snout pointed but short; limbs moderately long; finger and toe tips not expanded, third finger conspicuously long; toe webbing variable, approx. half of toes; dorsal skin structure finely pebbled in females, finely spinose in males; dorsal skin color sand to brown often with darker markings and black spots; a narrow light line from snout tip to groin separates the dorsum from the darker or orange flanks; dark spot in groin region; abdomen light gray, throat orange to rusty; pupil horizontal, iris gold-brown.—The species was recorded by photograph for Nepenthes Camp; no specimen was collected.
Body squat and clumsy; hind limbs relatively short; head significantly narrower than body, snout short, eyes relatively small; tympanum invisible; fingers and toes long, distally expanded, toes more so; toe webbed at base; dorsal skin granular with scattered tubercles; even lower jaw with rough skin; dorsal skin coloration quite variable, usually marbled in shades of brown and gray, often darker in central part of dorsum; pupil elliptical, horizontal in orientation; iris dark amber.—The species is mostly seen during breeding activities when the loud deep calls of the males can be heard. The specimen in Fig.
Short, narrow, somewhat obtuse snout; toes and fingers relatively large, fleshy, and bear expanded digital discs; toes webbed half; dorsal skin texture warty; color mostly brown and gray, variable pattern but commonly with a large dark constricted marking on the dorsum; fingers and toes often with yellowish or orange; iris black with some bronze stippling; pupil horizontal; ear drum rather indistinct.—
Frogs of the
Slender body; head long and narrow; long hind limbs with great jumping performance; fingers and toes elongate and with slightly swollen tips; toes half webbed; long snout, pointed in dorsal view; snout projecting lower jaw, narrowly rounded in lateral view; dorsal skin texture finely pebbled or granulated; dorsal side uniform mid-brown to marbled brown; dorsal side of hind limbs with darker bars; black streak at side of head starts from snout, broadens to band under canthus rostralis and continues behind eye covering the ear drum and beyond; dorsolateral fold present, thin but continuous from eye to pelvic region; tympanum large and distinct; upper lip als uniform white streak that continues onto rictal ridge behind jaw joint; venter cream, throat cream with darker mottling; thighs in pale flesh color below and weak banding above; iris dark reddish-brown below, golden above.—This species commonly follows human alteration of landscapes such as rural areas, road ditches and clearings; it does not enter the forest. At Maliau Basin it is present in large numbers at the pond of the Study Center and in ditches along the roads.
Moderately stocky with moderately long limbs; toes webbed to less than half of their length; finger and toe tips slightly expanded, narrowed to the tip; snout protruding beyond lower jaw and rounded in lateral view; area between eye and nostril smooth; skin texture smooth; a mahogany-brown area is delimited by a circumferential thin white to orange line on the dorsum and head; cream streak on upper lip from under the eye to shoulder; remaining parts of body mostly black; arms and legs variable, with black banding on brown background or black with banding in light gray; flanks gray with or without black dots; tympanum large and distinct; under side dark gray with white spots on venter and vermiculated pattern on thighs, zebra pattern on underside of shank; webbing; ear drum black, distinct, and large; iris dark brown to black, except for golden upper fifth;—The species was present with adults and larvae in a pond near Nepenthes Camp. For size and altitudinal range see
A relatively large species in the genus; body slender; limbs, fingers, and toes long; head long and narrow; snout pointed; snout projects with sharp tip beyond lower jaw in lateral view; snout longer than eye diameter; canthus rostralis present; fingers and toes long, bearing significantly expanded tips in form of adhesive disks with circummarginal grooves; tympanum very distinct and large (in males); dorsal skin texture finely granular; brown above with scattered dark spots; flanks often green otherwise brown; upper lip all cream in color, cream streak extends beyond jaw joint to shoulder; ventral side cream on venter and throat; under side of thighs flesh colored; iris golden with fine black venation and more reddish hues in anterior and posterior sectors.—
Body moderately stocky but with strong and long hind limbs; toes and fingers with terminal expanded and slightly pointed discs; toes fully webbed; dorsal skin texture mostly smooth, flanks with low tuberculation; color dark brown on dorsum, separated from lighter flank by ochre to orange dorsolateral stripe; dorsolateral fold broken; dark blotches expand from eye posteriorly, ventrally and anteriorly; hind limbs distinctly banded; ochre to orange line along upper side of shank; ventral side uniform yellow; tympanum distinct but small, sunken in, transparent; iris bicolored, dark reddish-brown below and beige to golden in the upper third.—The species has large tadpoles that are highly adapted to fast currents. The tadpoles can hold on to rocks in very fast currents by applying their conspicuous abdominal sucker. Because of this larval adaptation, this species is linked to clear cascading streams, however, individuals can occasionally be found at some distance from the stream. We recorded the species from the Fawzy Waterfall area.
Body moderately robust with long and muscular hind limbs; head and snout relatively long and moderately narrow; finger and toe tips only moderately expanded to rounded discs; toes webbed to half; dorsal skin texture smooth to granular, areolate at the flanks; color predominantly green above; dorsolateral fold thick, continuous and cream in color, bordered in dark brown; upper lip cream without marking; upper lip streak continues posteriorly in a lower longitudinal stripe at lower flank; upper side of limbs sandy brown; throat, chest and abdomen ventrally all white; thighs flesh to orange in color tympanum conspicuous, large, dark in color; iris golden in upper part, darker below, with dark mid-horizontal band.—At Maliau, the species is abundant at the Study Center Laguna and roadside ditches.
Slender (males) to moderately stocky (females) body with long slender legs; snout moderately long; tips of toes and fingers expanded, slightly rhomboidal; toes fully webbed; skin texture finely shagreened; dorsolateral fold present; color above brown, with scattered dark dots and light markings; upper lip without dark markings; upper side of thigh broadly banded; throat, chest and belly white or cream; males with lateral dark vocal sack areas on throat; rear of thigh mottled dark with light spots; iris golden in upper and lower sectors, reddish in anterior and posterior sectors. —
Body moderately robust with long and muscular hind limbs; toes and fingers long; tips of fingers and toes expanded to large rhomboidal discs; toes fully webbed; webbing dark in color; dorsal skin texture shagreened to granulated; ventral skin texture smooth anteriorly but areolate on posterior belly and thigh; color mostly green above, flanks can be brown; dorsolateral fold present but weakly developed, set off by dark line; upper side of limbs green or brown, with banding; dark stripe from nostril to eye along canthus rostralis; upper lip cream without markings, cream stripe continues to shoulder; ventral side white; tympanum distinct, dark in color, moderate in size; iris silver to gold in upper and lower part with abundant black reticulating pattern, mostly black in anterior and posterior part, with narrow golden ring around pupil,—
Body slender to moderately robust; limbs moderately long; head relatively narrow, moderately long snout; tips of digits slightly swollen; webbing to approximately half the toes; granular dorsal skin texture; color black with yellow to red spots and angular blotches; colored line from snout to eye and along upper eye lid; no continuous stripe from eye to groin; banded pattern on upper side of limbs; venter dark gray with scattered cream spots; tympanum distinct, black; iris coppery, with reddish narrow ring around pupil and abundant black reticulate pattern.—This species prefers rocky streams. It occurs at the Sungai Maliau tributaries, Agathis Camp and Ginseng Camp; however, the species was not abundant during our survey.
Body slender to moderately robust; limbs moderately long; head relatively narrow, moderately long snout; tips of digits slightly swollen; webbing to approximately half the toes; granular dorsal skin texture; color black with yellow to red angular spots or blotches; colored line from snout to eye, along upper eye lid and posterior to groin; banded pattern on upper side of limbs; venter gray, thigh darker; tympanum distinct, blackish; iris coppery, with reddish narrow ring around pupil and abundant black reticulate pattern.—
Body slender; snout long, pointed and protruding in lateral view; long fingers and toes, tips of digits broadly expanded; webbing of toes shiny turquoise blue; upper head and back yellowish brown to olive brown with dark spots on back, light green or yellowish green at the flanks with white dots; limbs brown to green above with more or less pronounced dark banding; lips without bars; outer third of digital discs with light cross band; dorsal skin texture pebbled or slightly warty, especially dorsolaterally; ear drum distinct but small; ventral side of throat, breast and belly white, more yellowish anteriorly; underside of limbs not white but greenish gray; iris gold to copper in upper sector, black in anterior and posterior sector and pale silvery in lower sector, narrow color ring around pupil (copper except for lower pale silver sector).—
Body slender; snout long, narrow, edged, pointed and protruding in lateral view; long fingers and toes, tips of digits broadly expanded; webbing of toes shiny turquoise blue; upper head and back yellowish brown to olive brown with dark spots on back forming marbling, flanks not sharply different from back; limbs brown to olive brown above with distinct dark banding; lips without bars; outer third of digital discs with light cross band; dorsal skin texture pebbled, tuberculate on flank; scattered short longitudinal ridges on dorsum and flank; ear drum distinct but small; ventral side of throat, breast and anterior belly white with diffuse dark spots; underside of limbs not white but brown to gray; iris golden in upper sector, black in anterior and posterior sector and pale golden in lower sector, narrow gold ring around pupil.—The species was abundant at Fawzy Waterfall. The species was distinguished from
Frogs of the
Slender body; snout very short and blunt, eyes slightly forward directed; finger and toe tips broadly rounded; webbing at base of outer fingers; toes half-webbed; dorsal skin texture finely shagreened; wavy edge along outer border of fourth finger and forearm and outer foot; color green above abruptly ending at flanks, brown posteriorly (pelvic region), scattered white dots above; venter white anteriorly and transparent posteriorly; iris silvery below and golden above, with some dark reticulation and golden ring around pupil; pupil horizontally oval; ear drum present but small and inconspicuous due to coloration of skin.—A number of individuals of
Body slender with relatively long fore limbs; head wide and flat; snout short; eyes slightly forward directed; outer fingers webbed at base; toe and finger tips expanded to adhesive discs; dorsal skin texture smooth, venter smooth; color yellow to brown above with dark spots; venter semi-transparent pink or red, with two longitudinal white streaks; brown band runs from tip of snout to flank at half of body, adjoined below by a cream stripe; iris amber, fading into beige at the dorsal and ventral periphery; eye moderately large; pupil without distinctly colored edge; pupil oval horizontally; ear drum present, small, inconspicuous.—This small rhacophorid species was described by
Intensive dip-netting of the pool revealed one
Body slender; head relatively broad, triangular; snout pointed, females with pronounced snout projection; finger and toe tips broadly expanded into adhesive discs; toes webbed to three-fourth; outer fingers webbed at base; forearm and foot bear a wavy fringe of skin along outer side; series of skin projections ventrally along mandible; dorsal skin texture irregular, tuberculate; venter skin texture distinctly areolate; color above can combine brown, gray, and green with various markings, highly variable, resembling bark; throat and venter cream, ventral side of legs gray to brown; eyes large; pupil oval horizontally; iris color gold, often paler around pupil and in lower part; iris moderately reticulated; ear drum small, distinct but not set off in color from surrounding skin.—The species is probably widespread in the Maliau Basin. We encountered it in the low elevation swampy habitats at the Belian Trail areas as well as the more elevated forests around Nepenthes Camp. Males prefer to stay close to muddy forest floor depressions and often call even if such sites are not water-filled. An adult female was found at night close to the trail from Nepenthes Camp to Fawzy Waterfall sitting low on a bush.
Medium sized, slender frog; head relatively wide, triangular and flat, snout long; canthus rostralis edged; hind limbs long; toes and finger bear round expanded discs, each disc bears two white dots; toes approximately half-webbed; dorsal skin structure spinose, spines low; dorsal coloration reddish-brown to orange, skin spines may be pale; white dots scattered over dorsal surface, flanks, and limbs; a row of white spots from snout tip to eye and along edge of upper palpebra; tympanum visible, large; eyes large; pupil rhomboid; iris white in upper sector, red in anterior, posterior, and lower sectors.—The specimen reported (photographic evidence) here was encountered at Nepenthes Camp. The species is a treehole breeder and, thus, independent of streams or ponds and might be widely distributed at MCBA. No specimen was collected.
Body moderately stout; head flat and broad, large eyes; snout long; canthus rostralis edged; toe and finger tips broadly expanded into adhesive discs; toes almost fully webbed; dorsal skin texture granulated, with some tubercules; tubercles can form V-shape on shoulder; skin of venter areolate; color above sandy to mid brown often with faint dark X-marking on back; scattered black spots, particularly from arm pit posteriorly; some banding on limbs; ventral side cream with pale gray-brown areas peripherally; ventral side of thighs darker, not cream; dorsolateral fold absent, supratympanic fold present, bent towards armpit and edged with dark pigmentation; iris color usually with a distinct green hue and with fine black venation; pupil horizontal, thinly edged with golden line; ear drum distinct but not set off from surrounding skin in color or skin texture.—
Body relatively stout, but with long hind limbs; head relatively broad; snout and lower jaw arch broadly rounded; toe and finger tips expanded into adhesive discs; toes half-webbed; dorsal skin texture moderately tuberculate and granulated; dorsal color variable from olive to shades of brown with dark markings; some with X-shape pattern on mid-dorsum; limbs with banding; venter coloration brown; iris coloration golden in the dorsal sector, paler, more silvery in the lower sector and dark brown in the anterior and posterior sectors; ear drum concealed by skin.—This small species of
Body slender with long hind limbs; head relatively broad and triangular; sharp canthus rostralis; toes and fingers expanded into near circular, slightly blunt discs; toes fully webbed, except fourth; dorsal skin texture smooth (lack of tubercles) but finely granulate; ventral skin areolate; supratympanic fold present; dorsal color sandy brown to dark tan, often with four dark longitudinal stripes from head to pelvic region, plus partial dorsolateral stripe; throat, venter and parts of ventral thigh cream; throat may bear scattered darker markings; banding pattern on limbs; iris color sandy brown to amber, golden narrow edge around pupil; pupil horizontal; ear drum distinct.—This species avoids the forest and prefers open agricultural country or forest edges. It benefits from human activities and uses roadside ditches, flooded lawns, large puddles, or shallow agricultural ponds for reproduction. The species is commonly encountered and its calls heard around the Study Center area and its roads.
Body slender with long hind limbs; head broad, rounded triangular in dorsal view; eyes big and noticeably directed forward; sharp canthus rostralis; toes and fingers expanded to broad blunt adhesive discs; not fully webbed toes; dorsal skin texture finely granular, without prominent tubercles; ventral skin of venter and thigh areolate; coloration light gray or ochre to tan above with dark spots or (sometimes) two dark stripes from snout to pelvis; area below supratympanic fold and most or all of tympanum masked in a dark brown marking that extends to flank; banding pattern on limbs; color of throat, venter and parts of ventral thigh is cream; throat and chest with some dark mottling; iris color brass to beige, with darker anterior and posterior sectors; pupil horizontal; ear drum distinct, large. —
Body slender to moderately robust (females); head edgy, broad, triangular; snout pointed; eyes big; saw-edged ridge above ear drum; hind limbs long, bearing distinct heel spur; fingers and toes expanded to large blunt adhesive discs; toes half webbed; color above grayish olive to yellowish light brown, fine longitudinal stripes on dorsum; thighs anteriorly and posteriorly banded with distinct zebra pattern; ventral side dirty white; ear drum visible, not embedded in dark marking; dorsal skin texture generally smooth (no prominent tubercles) but granulated above, distinctly areolate at flanks; venter skin areolate; iris beige, with fine black reticulations; pupil horizontal.—
No specimens were collected. Calls from the canopy, short in duration (approx. 1s) and composed of multiple notes of moderately high pitch in fast sequence (resembling a woodpecker) were heard at the type locality and were interpreted as calls of this species as we have know these calls from other breeding populations of this species (the original species description does not provide a call description). No recordings were made. —
Body slender with long hind limbs; large eyes; pointed snout, angular in lateral view; canthus rostralis sharply edged; toes and fingers expanded into adhesive discs; toes fully webbed except fourth toe; outer fingers webbed at base, inner fingers less; dorsal skin texture granulated; supratympanic fold present; dorsal color tan to mid brown with dark brown or green blotches; white spots or a white blotch below the eye; light blue spots at the otherwise black lower flank esp. towards groin and on anterior and posterior thigh as well as ventral shank; throat and venter white to cream; horizontal pupil; iris dark amber; ear drum visible, distinct.—
Body slender (males) to robust (females); head broad and flat, large eyes; hand and foot large with extensive red webbing; toes and fingers terminally expanded; heal with small skin flap; dorsal skin texture finely granulated; color above brown with dark spots or faint dark marbling and occasional white spots; lower flanks and venter with distinctive marbling in yellow/black or yellow/red, respectively; iris yellowish beige peripherally and more brown towards center; pupil oval and horizontal; ear drum visible, indistinct in color.—Among the species of gliding frogs,
Rhacophorid frogs at
The present study contributes to the amphibian inventory of
Considering the time invested and the experience of the searching participants in our survey the field campaign yielded results in the range that could be expected. For comparison, in 2009, a rapid assessment at Kubah National Park, Sarawak, with a slightly higher man-hrs investment (240 man-hrs) yielded 50 species (Haas, unpubl. data). Although different geographic and geological sites can only be compared with more elaborate study designs, this crude comparison suggests that the diversity encountered at
During the period of our survey, we were struck by the low abundance of frogs and low reproductive activities. This may have been caused by a longer period of dry weather in the Maliau Basin before and during our survey. Low precipitation is not unusual for
Four previous amphibian surveys conducted at
Interestingly,
Our work confirms many species that have already been known from previous studies (Table
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The species list is still preliminary, yet our account fills several gaps of its predecessors (
Accumulative species curve from species encounters at three sites within
In groups of aquatic organisms, it has been argued that the typically black, humic acid rich, nutrient-poor water of the Maliau Basin streams (
We wish to thank Sabah Biodiversity Council (SaBC) for granting permission to access biological resources in Sabah (JKM/MBS.1000-2/2 JLD.5(67)) and for export of parts of the voucher specimens (JKM/MBS.1000-2/3 JLD.3(1)). C.Y. Chung (SaBC) helped us with advice along the permitting process. Permission to work at
Specimens examined from Maliau Basin Conservation Area. GenSeq nomenclature follows Chakrabarty (2013).