Five new microhylid frog species from Enga Province , Papua New Guinea , and remarks on Albericus alpestris ( Anura , Microhylidae )

Five new species of microhylid frogs in the genera Albericus, Cophixalus and Oreophryne are described from high altitudes in the Kaijende Highlands of Enga Province, central Papua New Guinea. A population of short-legged, terrestrial Albericus discovered in grasslands on the Kaijende Highlands seems to be very close to the recently described A. alpestris. Because the systematic status of our new frogs is not yet clear, they are referred to as Albericus cf. alpestris in our paper. Information about the advertisement call of this population, its colour in life, and its habitat is presented for the fi rst time and enables possible inferences about Albericus alpestris sensu stricto.


Introduction
The high-altitude frog fauna of New Guinea is dominated by members of the family Microhylidae, with the highest known record of a frog from the island being for an Oxydactyla stenodactyla that was collected at 4,000 m above sea level on Mount Wilhelm (Zweifel, 2000).Although relatively few species inhabit New Guinea's alpine environments, the faunas there are phylogenetically diverse, with representatives from various genera, including Aphantophryne, Oreophryne, and Oxydactyla, successfully inhabiting habitats above 3,000 m (e.g.Zweifel, 2000;Zweifel, Cogger & Richards, 2005).Many of these species exhibit adaptations to a terrestrial life in montane meadows, having short limbs and reduced fi nger and toe discs in comparison to their lower-elevation forest-dwelling congeners.
In August-September 2005 the Conservation Inter national Rapid Assessment Program (RAP) con-ducted a biological inventory of alpine environments in the Kaijende Highlands near Porgera in Enga Province, Papua New Guinea (Richards, 2007) (Fig. 1).During that survey a number of new species of amphibians were discovered, including at least seven frogs in the family Microhylidae.Two of those species, Oreophryne anamiatoi Kraus & Allison, 2009 and Albericus alpestris Kraus, 2010 have subsequently been described from nearby mountain ranges and here we present descriptions of another fi ve species.

Material and methods
Most frogs were detected at night by tracking their advertisement calls.Some females were also encountered with the aid of head-torches, and a small number of individuals were collected from beneath stones and logs on the forest fl oor.Frogs were anaesthetized with chlorobutanol and most were fi xed in 10 % formalin in the fi eld for 1 -2 days and transferred to 75 % ethanol for permanent storage.Samples of liver tissue were extracted from up to fi ve specimens of each species and stored in 95 % ethanol to permit future DNA analysis.Clearing and double staining of one specimen of each new species as an osteological preparation according to a modifi ed method of Dingerkus and Uhler (1977) was carried out in order to identify their generic allocation.
Measurements of SUL, TaL, and TL to the nearest 0.1 mm were made with a digital calliper, all others were made with an ocular micrometer in a dissecting microscope: SUL snout-urostyle length, from tip of snout to distal tip of urostyle bone (SUL and snout-vent length differ insignifi cantly, but SUL is more accurately measured); TL tibia length, external distance between knee and heel (calliper gently pressed); TaL length of tarsus, external distance, tarsal and heel joints held at right angles; L4T length of fourth toe, from tip of toe to proximal end of inner metatarsal tubercle; L3F length of third fi nger; F3D transverse diameter of third fi nger disc; F1D transverse diameter of fi rst fi nger disc; T4D transverse diameter of fourth toe disc; their relative length 3 > 4 > 2 > 1; no webs; all discs with circummarginal grooves; no subarticular tubercles, weakly expressed inner metacarpal tubercle, no outer metacarpal tubercle.Hind limbs short (TL/SUL 0.38); terminal discs roughly as wide as those on fi ngers; relative length of toes 4 > 5 > 3 > 2 > 1; no webbing between toes; no clearly expressed subarticular, plantar, or metatarsal tubercles.Moderately large tubercles on all dorsal surfaces, rare in middle of dorsum, head, and on fore limbs, and most frequent on body sides, hind limbs, and around tympanum; a ring of characteristic tubercles around tympanum; all ventral surfaces smooth.
Ground colour of all dorsal and ventral surfaces greyish-yellow, dark brown blotches in the scapular region, some diffuse blotches on head, behind tympanum, and on extremities.Besides these blotches, dark brown pigmentation than the holotype.Their dorsal and lateral head surfaces are completely dark brown, and, compared to the holotype, there are more blotches on most of the other dorsal surfaces, and the ventral surfaces are also more densely dotted.Conspicuous in all types are a fi ne whitish band between the eyes, an irregular whitish spot in front of the tympanum, a diagonal white stripe and some white tubercles in the scapular region, a dark brown patch or streak from eye corner to shoulder, and a pigmentless patch below the elbow.The following remarks on colouration in life are based on a photograph of the paratype SAMA R66101 (Fig. 2): ground colour of head and dorsum darker than that of lateral surface of torso and limbs; head and nape predominantly dark brown; upper eyelids, lower loreal region and area behind eyes light grey; dorsum grey brown; fl anks uniform grey; dorsal surfaces of limbs grey with some diffuse red-brown fl ecks especially around tubercles; further anterior markings consist of small white spots in the tympanic region and a short diagonal white stripe above the arm insertion.We did not note the colour characters of ZMB 76959, now an osteological preparation.
Distribution and ecological notes.This species is known only from the type locality (Fig. 7) in mossy montane forest on the Paiela Road near Porgera, Enga Province, where the males were calling from heights of ~ 2 -3 m in Pandanus trees at night.This species was not heard calling in areas of the forest without Pandanus trees.
Vocalisation.Following the terminology of Menzies (1999), the calls are buzzes (Figs. 3a and 3b).Twentyfi ve calls from two males (ZMB 76958 and SAMA numerous dark brown dots present on all dorsal and ventral surfaces except central dorsum and areas around elbow; conspicuous are a transverse whitish band between eyes, a whitish spot in front of the tympanum, and a patch of dark pigmentation along the supratympanic fold.

Morphological characters of the paratypes.
There are three male paratypes, one slightly smaller and two slightly larger than the holotype.All of their body dimensions are similar to those of the holotype (Table 1).Skin structure, distribution of tubercles, and basic colour are also similar in all types.ZMB 76958 and SAMA R66101 exhibit a much more pronounced   . siegfriedi Menzies, 1999;and A. tu ber culus (Richards, Johnston & Burton, 1992) Albericus darlingtoni has a similar call length and pulse repetition rate as A. pandanicolus but differs in dominant frequency (2.5 kHz in A. darlingtoni according to illustrations in Menzies, 1999 and2006, versus 3.3 kHz in A. pandanicolus).The two R66101) were analysed.Air temperature during recording was 17 °C.Calls were uttered at irregular intervals, and the shortest interval between two calls was 4.5 s.Call duration is from 229 ms to 275 ms, mean 247 ± 9.4 ms.Number of pulses per call varies from 46 -63, mean 53, and pulse repetition rate varies from 188 to 243 pulses per s, mean is 215 pulses per s.Frequencies scatter in a broad range between 2.5 kHz and 7 kHz, with dominant frequencies at 3.3 kHz (Fig. 4).Wave forms of the calls show a spindle-like shape, with the weakest pulses at the beginning and towards the end of the calls.
Albericus fafnieri is larger (21 -23 mm SVL), has longer tibiae (0.38 -0.43), and has different adverti se ment calls: the mean call duration is 650 ms and the pulse repetition rate is 70 -170 pulses per s.Albericus laurini has longer hind legs (TL/SUL 0.39 -0.43), different head ratios (among others END/ IND in A. pandanicolus is 0.88 -1.06 and in A. laurini is 1.24 -1.50), and a different call (call duration is 157 -204 ms in A. laurini and 229 -275 ms in A. pan da ni colus).Albericus rhenaurum is smaller (14.9 -15.8 mm SVL), has longer hind legs (TL/SVL 0.42 -0.45), and a different call: the call duration is 470 -510 ms and the note (pulse) repetition rate is 40 -50 pulses per s.Albericus siegfriedi is slightly larger , has longer hind legs (TL/SVL 0.41 -0.45), and a pulse repetition rate of 500 -600 pulses per s.Albericus tuberculus has more pronounced tubercles on body and limbs, longer hind legs (TL/SVL 0.40 -0.46), a different END/IND ratio (1.0 -1.67), and a different call (less than 200 pulses per s).Data for comparisons were taken from the original descriptions of the species (see above).
Etymology.Specimens were found on plants of the genus Pandanus.The specifi c epithet pandanicolus is in accordance with the gender of the genus, a composite masculine substantive in apposition and means "inhabitant of Pandanus plants".

Albericus brevicrus sp. nov.
Plate II, Figs. 5 -9 and Table 2 Holotype.SAMA R66100 (FN SJR 9710), adult male, collected at Paiela Road (05°30.183'S,143°04.866'E;2,900 m asl) near Porgera Town, Enga Province, Papua New Guinea on 5 September 2005, by S. Richards.Paratypes.UPNG 10061 (FN SJR 9711) same data as holotype; SAMA R66097 (FN SJR 9648), R66098 (FN SJR 9651), R 66099 (FN SJR 9656) and ZMB 76742 (FN SJR 9655) collected adjacent to Lake Tawa, Enga Province, Papua New ciated with tubercles and narrow glandular folds.A broad patch of paler pigment extends from posterior corner of eye, below the darker supratympanic band, to a point immediately anterior of the arm insertion.Scattered pale fl ecks are concentrated laterally and ventro-laterally with several fl ecks behind the eye and along the jaw, and on the dorsal surfaces of the thighs.Iris copper with brown fl ecks.

Morphological characters of the paratypes.
For measurements of all types see Table 2. Including the holotype, there are eight adult males with SUL from 14.3 -18.2 mm, one adult female of 17.7 mm SUL, and one probably subadult male of 13.4 mm in which no vocal slits could be seen.Vocal slits were also not seen in the male ZMB 76743 of 14.4 mm SUL.
Ovarian eggs of the one female were unpigmented or thighs, and on distal shanks.Blackish and brown spots on dorsal extremities, encircling light blotches on dorsum, and around anal opening.All ventral surfaces, except those of toe discs, fi nger discs, and proximal fore arm, strongly pigmented with brown, only venter shows some light ground colour between dark brown areas.
Based on photographs in life of the holotype (SAMA R66100), body and limbs generally brown with extensive areas of darker brown pigmentation, focused as follows: in a broad band extending from a line mid-way between the orbits to the posterior edge of the arm insertion; in a broad canthal stripe from the tip of the snout to the eye and extending posterior of eye along the curved glandular supratympanic fold to the axilla; along the edge of the upper and lower jaws; and in a series of small patches and bars asso- transverse stripe above the vent.Their ventral surfaces exhibit a dense pattern of brown spots on a yellowish ground colour.Conspicuous is a fi ne longitudinal line of whitish colour in the middle of the abdomen which crosses the vent opening and extends on the dorsum up to the snout tip.A similar line extends across the chest.Abdomen and ventral surfaces of legs of most paratypes are dark brown with yellowish spots; throat and chest are uniformly brown and without such yellowish spots.Abdomen of paratype ZMB 76743 exhibits more lighter than darker (brown) areas, and its throat and chest are spotted.ZMB 76745 is brown with pale brown snout and pale on dorsum and dorsolaterally; darker brown band between eyes, behind eye to axilla, in patch below eye and on limbs, pale cream stripe on distal edge of hind limbs.UPNG 10060 mottled dark and lighter brown with extensive and measured about one mm in diameter.Four specimens (UPNG 10060, SAMA R66097, R66098, and R66099) exhibit a similar basic colour and a similar mottling as the holotype, colouration of pigmented areas varied from brown to black.One paratype (ZMB 76742) has completely blackish dorsal and lateral surfaces, except a whitish coloured snout dorsum, a light interocular bar, two curved stripes in the scapular region, some whitish spots around elbow and tibio-tarsal articulation, one white spot right of the urostyle, and a transverse bar on the posterior thighs.All dorsal surfaces of paratype ZMB 76743 are darker or lighter brown with irregular blackish spots and stripes.The one female (ZMB 76744) and UPNG 10061 show a reticulate pattern of brown, blackish and greyish spots on all dorsal surfaces except for light yellowish areas between eyes, on fore and hind legs, and a ter in a broad range mainly be tween 2 kHz and 5 kHz, dominant frequency is at 3.5 kHz (Fig. 9).Most calls start with up to 10 very weak pulses and end with one double pulse.
Comparison with other species.Albericus brevicrus belongs to the group of species with a buzz-call (Menzies 1999).Its extremely short hind legs (TL/ SUL 0.30 -0.35, mean 0.33) delineate it from all other species of the genus except A. alpestris.This latter species has even shorter legs (TL/SUL 0.26 -0.33) but a larger body: SUL more than 19 mm versus a maximum SUL of 18 mm in A. brevicrus (for further details see last section below).
Etymology.The specifi c epithet is a composite adjective, derived from the Latin word brévis (-is, -e) for short or small and the Latin crus (crúris) for shank.It refers to the extremely short hind legs of the new species.
grey stippling concentrated predominantly over the paler brown blotches.All three previous specimens with pale cream patch that covers region of the axilla and extends ventrally to angle of the jaws.
In life colour pattern extremely variable.SAMA R66099 (Fig. 5) golden brown on snout anterior of eyes, with additional large golden-brown patches on mid-dorsum, dorso-laterally, postero-dorsally, and in a broad band along distal edge of thigh and around tip of heels.A broad, dark brown band extends from orbits posteriorly above supratympanic fold laterally to a point above arm insertion.Remaining lateral areas transluscent pale grey with stippled dark brown pigment and pale cream, russet and golden-brown pigments, and patches of darker brown mottling.Iris golden-cream with narrow reticulations and a vertical, dark brown triangular notch.
Distribution and ecological notes.Albericus brevicrus is known from wet, mossy mid-montane rainforest adjacent to Lake Tawa (2,200 m asl) and along the Paiela Road (~ 2,900 m asl) in Enga Province, Papua New Guinea (Figs. 6 and 7) where males called from low perches on leaves ~ 20 -30 cm above the ground at night.
Vocalisation.Forty-six calls from three males (SAMA R66097, R66098, and ZMB 76742) were analysed.Air temperature during all recordings was 14 °C.Calls are buzzes (Figs.8a and 8b) and are uttered in series with well-defi ned short intervals between calls.Forty-one inter-call intervals have a mean duration of 3.1 ± 0.72 s, range 2.30 -4.99 s; in ter vals longer than fi ve seconds within one series rarely occur.Mean call duration is 561 ± 49 ms, range 489 -641 ms; mean number of pulses per call is 92 ± 9.5, range 76 -105; mean pulse repetition rate is 165 ± 15.5 pulses per s, range 146 -197 pulses per second.Frequencies scat-   Description of the holotype.For morphological characters see Plate III and Fig. 10 and for body measure ments and body ratios see Table 3. Head wider than long (HL/HW 0.80), no constriction at neck region, snout rounded in dorsal view and truncate in profi le, canthus rostralis clearly expressed, curved outwards above nostrils and continued on tip of snout; loreal region oblique, eyes relatively small (ED/SUL 0.118) with pupils small and almost round; nostrils nearer to tip of snout than to eyes, internarial distance considerably greater than distance between eye and naris (END/IND 0.75); tongue long and narrow without posterior indentation, prepharyngeal ridge weakly expressed and not denticulate, vocal slits present on fl oor of mouth; tympanum barely visible, its diameter less than half eye diameter (TyD/ED 0.38), supratympanic fold pronounced.Forelegs well developed, fi ngers narrow without broadened tips, terminal grooves present on fi ngers 2 -4 but diffi cult to identify on fi nger 1; relative lengths of fi ngers 3 > 4 > 2 > 1, fi nger 1 less than half length of fi nger 2; no clearly expressed subarticular, carpal, or metacarpal tubercles.Hind limbs well developed but with very short thighs and shanks (TL/SUL 0.33); toe tips not broadened, those of toes 2, 3 and 4 with clearly recognizable grooves, grooves on fi ngers 1 and 5 poorly developed; no webbing between toes, relative lengths

Diagnosis.
A small-to medium-sized (SUL of adult males 18.4 -20.3 mm, of adult females 20.9 -23.5 dots; throat medium-brown with fi ne white and larger brown spots.Upper lips brown, loreal region blackish, a blackish stripe inferior to lateral glandular ridges, and with a conspicuous white fl eck which continues onto parts of the tympanum and is bordered by blackish areas behind the eyes.
Based on photographs in life of the holotype, body and limbs generally brown with extensive areas of darker brown pigmentation, focused as follows: in a broad band that extends from a line mid-way between the orbits to the posterior edge of the arm insertion; in a broad canthal stripe from the tip of the snout to the eye and back from eye along the curved glandular supratympanic fold to the axilla; along the edge of the upper and lower jaws; and in a series of small patches and bars associated with tubercles and narrow of toes 4 > 3 > 5 > 2 > 1, toe 1 rudimentary.Posterior dor sum, dorsal surfaces of tibiae, of tarsi, and of lateral body covered with tubercles; head, anterior back, thighs, and forelegs smooth; undulating dorsolateral folds from eyes to lumbar region conspicuous; ventral surfaces smooth.There is an abdominal incision on the right side of body.
Ground colour of dorsum, fl anks, and extremities reddish brown with diffuse blackish spots and reticulations, dorsal surface of snout (back to level between eyes) distinctly lighter than other dorsal surfaces; two light lumbar spots and one triangular light spot above posterior urostyle, some small white spots on lower fl anks; abdomen and ventral surfaces of extremities with a mixture of irregular brownish, blackish, and off-white spots, with conspicuous numerous white .5-17.8 mm.Ovarian eggs of the two adult females are entirely whitish and are 1.5 to 2.0 mm in diameter.Five specimens have a similar ground colour as the holotype and two specimens have a greyish-brown ground colour.Three of the eight frogs show fairly uniform dorsal colouration, the others exhibit more or less numerous irregularly-shaped blackish spots on the dorsum.Characteristic of all specimens are a light snout dorsum, light spots in lumbar region, blackish longitudinal stripes that border the dorsolateral glandular ridges inferiorly, blackish upper half of loreal region, and an elongate whitish spot behind the eye which is superiorly and inferiorly bordered by blackish areas.Occasional tubercles are on the fl anks, shanks, and in the lumbar region; ventral surfaces are commonly light yellow with medium-or glandular folds.A broad patch of paler pigment extends from posterior corner of eye, below the darker supratympanic band, to a point immediately anterior of the arm insertion.Scattered pale fl ecks are concentrated laterally and ventro-laterally with several fl ecks behind the eye and along the jaw, and on the dorsal surfaces of the thighs.Iris copper with brown fl ecks.
Morphological characters of the paratypes.For measurements and body ratios of all types see   Distribution and ecological notes.Cophixalus tenuidactylus was found in alpine grassland around Omy aka Camp at elevations between 3,100 -3,300 m (Fig. 11) and at the edge of montane mossy forest along the Paiela Road at about 2,900 m asl, both locations on the Kaijende Highlands.Males called from within soil and humus below grass and fern mats in the alpine grasslands, and from within raised mossy clumps surrounding small patches of stunted forest within these grasslands.At Paiela Road, specimens were collected from beneath logs and mats of grass at the edge of the forest during the day.No specimens were seen or heard within the forest, and no specimens were found in elevated or in exposed positions.Two calling males traced by their calls deep inside mossy clumps in the grassland were guarding fi ve and eight large, unpigmented eggs in small hollows within the moss clump (Fig. 10).However, calling males were extremely diffi cult to fi nd because indi-   Etymology.The specifi c name is derived from the Latin adjective ténuis (-is, -e) for narrow or slender, and the Latin substantive dáctylus (-i) (Greek ho dáctylos) for digit.It refers to the unusually narrow (for the genus Cophixalus) fi nger and toe tips of the new species.
Plate IV, Figs. 14 -16, and  The wave form of the fi rst part of these double notes looks like an asymmetric spindle and has a lower volume than all remaining notes.The second part is spindle-like and has the same volume as the following notes.Fundamental frequency of the calls is around 1.3 kHz and dominant frequency is concentrated in a small range around 2.6 kHz (the dominant frequency band is synonymic with the fi rst harmonic band).Further harmonics are at 3.9 kHz, 5.2 kHz, 6.5 kHz (scarcely expressed) and 7.8 kHz (Fig. 13).Mean note length is 260 -320 ms with a dominant fre quency above 3.2 kHz and a note repetition rate of about 1.5 notes per s.Cophixalus verecundus utters a "rapid train of click-like pulses giving the aural effect of a soft buzz" (Zweifel & Parker, 1989).Notes (= pulses) are not fi nely tuned but show a wide spectrum of frequencies with energy peaks at about 2.5 and 3.3 kHz.Inter-note intervals are very short, and from this results a note (pulse) repetition rate of 50 -60 per s.Notes (pulses) and inter-note intervals are considera-plantar or subarticular tubercles present.Shanks moderately short (TL/SUL 0.40); some small tubercles around tympana, on body sides, and on dorsal surfaces of extremities; all ventral surfaces smooth.Ground colour of dorsal surfaces pale straw yellow; except on snout dorsum, all dorsal surfaces covered by many dark brown spots of irregular size.Obvious are lumbar "eye spots", a W-shaped mark in scapular region, some elongate whitish dashes bordered by blackish spots, a dark spot posterior of eye, a blackish spot surrounding vent, and an inverted whitish U in loreal region; all ventral surfaces of pale straw colour with many blackish dots.

Morphological characters of the paratypes.
For mensural variation of the type series see Table 4. Paratypes include seven adult males with SUL of 20.2 -22.4 mm, three adult females with SUL of SUL 0.123), eye-naris distance smaller than internarial distance; tongue very broad with a posterior transverse furrow, no posterior notch; two prepharyngeal ridges, both with some indentations; tympanum nearly half the size of eye (TyD/ED 0.45), its superior margin covered by skin, supratympanic fold weakly expressed.Forelegs long and slender, fi ngers long, their relative length 3 > 4 ≈ 2 > 1, tips of fi ngers slightly rounded and very wide (more than twice the width of penultimate phalanges), all with terminal grooves; elongate inner and rounded outer metacarpal tubercles feebly developed, no clearly demarcated subarticular tubercles.Hind limbs also slender and with long toes, their relative length 4 > 5 > 3 > 2 > 1, no webs between toes, toe tips somewhat narrower than fi nger tips (T4D/F3D 0.86), all with terminal grooves; and tip of fi rst toe of the right hind leg missing.With the exception of an inner metatarsal tubercle, no other In life dorsally mottled with darker and lighter brown.ZMB 76749 (Fig. 14) infused with yellowish-orange particularly on posterior and lateral surfaces; ventro-lateral surfaces with translucent, creamy patches with brown stippling and pale, creamy orange mottling; a pale cream inverted 'U' on upper lip, dorsal surfaces of snout also cream but with yellowish tinge and pale yellow and small brown spots; a pale bar from posterior corner of eye to posterior edge of jaw and a narrow brown stripe from tip of snout along canthus to eye and extending, slightly narrower, pos-22.5 -25.1 mm, and one subadult specimen with SUL of 14.9 mm.All paratypes with an off-white dorsal ground colour, this ground colour interspersed with numerous fi ne dark brown dots.These dots can be isolated but can occur in more or less dense groups and thus form more or less densely pigmented spots.Form and arrangements of dots varies between individuals.According to number and dimensions of spots, dorsal surfaces of paratypes appear lighter or darker.One specimen (UPNG 10066) is completely dark brown except on the snout dorsum, a W-shaped mark in the scapular region, and on lumbar spots.Characteristic features of all paratypes are a light snout dorsum, a light W-shaped mark in the scapular region, whitish lumbar spots, an elongate brown spot beneath arm, a blackish spot around vent, an elongate blackish spot above tympanum, and an inverted U-shaped mark of white colour in loreal region.Five and dominant frequency at 2.6 kHz.There are fi ve upper harmonics above the dominant band, and most pronounced of these are the fi rst at 3.9 kHz and the fourth at 7.8 kHz (Fig. 16).
One call with 21 notes (call number 047 of SAMA R66118) differs clearly from the other two calls of this male.Mean note length is 132 ms, mean internote length is 208 ms, note repetition rate is 3.0, and dominant frequency 2.9 kHz.It is impossible to determine whether this call is an aberration, a "normal" variation, or whether it stems from a male of another species.A similar situation concerns a call probably by SAMA R66119 (call 145): this call lasts 4.8 s and has 18 notes.Mean note duration is 103 ms and mean inter-note interval duration is 174; consequently, this results in a note repetition rate of 3.75 notes per s.
Comparison with other species.The procoracoid of O. streiffeleri is connected to the scapula by a cartilaginous bridge.All species without such a cartilaginous bridge will not be considered further here, but we will consider those species where it is not known whether the procoracoid is connected to the scapula by a ligament or by a cartilaginous bridge.Oreophryne brevirostris Zweifel, Cogger & Richards, 2005 is an alpine species (found at 3400 -3500 m asl) with very short legs (TL/SUL less than 0.35) and no whitish mark on lores.A syntype (ZMA 5819) of O. crucifer (Van Kampen, 1913) has webs terior of eye across top of tympanum and half-way to axilla; posterior of thighs reddish orange; a thin, pale mid-dorsal stripe may be present.Eye golden with copper fl ecks.SAMA R66117 similar but less translucent, and with more extensive yellow laterally, on fi ngers and toes, and on venter.SAMA R66118 has a similar general pattern but a much darker brown dorsum colour.
Distribution and ecological notes.This species is known only from mossy montane forest along the Paiela Road near Porgera Town, Enga Province (Fig. 7).Males called from the ground, or from low foliage (< 2m above the ground) during and after heavy rain.The female holotype was collected from a leaf in dense foliage during rain.
Vocalisation.Advertisement calls are a series of peeps (Figs.15a and 15b).Four calls of two males (SAMA R66118, and R66119) were analysed.Air temperature during recording was 17 o C.Call length is 6.4 -7.7 s, mean 6.84 s, and note repetition rate is 2.47 -2.58 notes per s (mean 2.53 notes per s).Notes start explosively, show a slight sound modulation in the course of the note, and cease slowly.There is a very weak frequency modulation from lower to higher in the course of each note.Mean note length varies from 148 to 161 ms (mean of means 155 ms) and mean inter-note interval length varies from 241 to 278 ms (mean of means 258 ms).Total range of note length 142 -183 ms (except fi rst note which is often much shorter, less than 100 ms), total range of internote interval length 223 -495 ms.Total number of analysed notes is 270 and total number of inter-note intervals is 266.Fundamental frequency is at 1.3 kHz high elevations between 3000 and 3500 m asl and has (among other differences) shorter legs than O. streiffeleri (TL/SUL 0.33 -0.38 versus 0.40 -0.46).With a SUL of up to 45 mm, O. idenburgensis is larger than O. streiffeleri.Oreophryne kampeni Parker, 1934 differs from the new species by the presence of distinct digital webs, no white loreal mark, shorter legs (TL/SUL of the holotype of O. kampeni 0.42), distinct T4D/SUL (0.043 versus 0.62 -0.73), distinct T4D/F3D (0.66 versus 0.86 -1.08) and distinct ED/ SUL (0.152 versus 0.112 -0.128).The structure of the shoulder girdle of O. loriae (Boulenger, 1898) is not clear.According to Menzies (1975Menzies ( , 2006) ) the latter species utters "6 to 12 short, rather nasal, buzzes" with a buzz length of between 200 and 300 ms.Oreophryne streiffeleri utters peeping notes, with note length varying between 100 and 200 ms.No inverted U-mark or O-mark on lores.Connection between procoracoid and scapula is cartilaginous.Advertisement call consists of a train of melodious peeps.Duration of the calls 7.2 -10.4 seconds, note repetition rate 2.4 -2.7 notes per s, and dominant frequency at 3.6 kHz.
Description of the holotype.For morphological characters see Plate V and for body measurements and body ratios see Table 5.There is a longitudinal incision in the upper fl ank of the right body side.Head shorter than wide (HL/HW 0.77); neck region slightly constricted; snout tip weakly rounded in dorsal view and truncate in profi le; canthus rostralis nearly straight, loreal region oblique; nostrils lateral and nearer to tip of snout than to eye; eyes medium sized (ED/SUL 0.120), pupil horizontally oval, iris blackish with numerous silvery spots; superior half of tympanum covered by skin, its diameter less than half that of eye (TyD/ED 0.37); eye-naris distance less than internarial distance; tongue moderately broad and posteriorly more than half free, no notch in its posterior margin; prepharyngeal ridge scarcely developed.Forelegs long and slender, fi ngers medium sized, their relative lengths 3 > 4 > 2 > 1, all fi ngers with broadened tips not wider than one and a half width of penultimate phalanges, tip of fi rst fi nger clearly narrower than that of other fi ngers, all tips with terminal grooves; no clearly developed tubercles on fi ngers or hands.Hind legs more robust and with long toes, toe tips rounded and discs slightly wider than width of penultimate phalanges, toe discs approximately as wide as fi nger discs, all discs with terminal grooves, no clearly expressed subarticular or plantar tubercles and no webs; relative length of toes 4 > 5 > 3 > 2 > 1.A fairly large and angular tubercle present between insertion of foreleg and tympanum, tympanic fold inconspicuous, no other conspicuous tubercles or skin ridges.
Ground colour of dorsal surfaces light yellowishgrey and covered by a diffuse network of middlebrown to dark brown spots, these spots tending to be lighter on head posterior to eyes, dorsum, and dorsal surfaces of extremities, and darker on fl anks, on anterior and posterior surfaces of extremities, and on dorsal surface of hand.Dorsal surface of snout especially light, whitish also are "eye-spots" in lumbar region; loreal and subocular region dark brown with some light dashes and spots; conspicuous is the contrast between the dark brown loreal region and the light dorsum of snout.Ground colour of throat, chest, and inferior thighs pale yellow, that of abdomen whitish; all ventral surfaces covered by numerous blackish dots.ger discs and toe discs (T4D/SUL 0.36 -0.48 versus 0.62 -0.73 in O. streiffeleri), a signifi cantly lower ratio T4D/F3D (0.71 -0.91 versus 0.86 -1.08 in the new species, p=0.024), a signifi cant higher ratio END/IND (0.90 -1.00 versus 0.85 -0.95 in the new species, p=0.005), and larger eyes (ED/SUL 0.135 -0.152 versus 0.115 -0.128).Moreover, the note repetition rate in O. pseudasplenicola is 1.3 to 1.4 notes per s, while this rate in the new species is about 2.5 notes per s; dominant frequency in O. pseudasplenicola is 3.4 kHz versus 2.5 kHz in O. streiffeleri.The character of the shoulder girdle of O. wolterstorffi (Werner, 1901) is unclear, but this species differs from the new one, among other characters, by conspicuous webs on the toes.
The presence of a white inverted U-mark or an O-mark on the loreal region and further common morphological and anatomical features as well as similarities in the advertisement calls lead to our conclusion that O. notata, O. asplenicola, O. pseudasplenicola, and O. streiffeleri belong to the same phylogenetic branch.
Comparative morphological and behavioural details in this paragraph were taken from studies of the authors and from the original descriptions of the species whose authors are mentioned when a species name was fi rst mentioned.
Etymology.The specifi c name is in memory of one of the oldest friends of the senior author, Heinz Streif feler (Berlin), who unfortunately died on 21 May, 2011 as this paper was nearing completion.

Oreophryne graminis sp. nov.
Plate V, Figs. 17 -20, and Table 5 of snout is lighter than remaining dorsal surfaces and clearly demarcated from a darker region between eyes in all specimens.Only four specimens have light lumbar spots.Some frogs exhibit shorter or longer white stripes in the loreal region, but none has an inverted U-shaped mark there.Ground colour of ventral surfaces off-white.Throat in all specimens covered with fi ne dark brown dots and is lighter than other ventral surfaces.Venter and extremities may be fi nely dotted (similar to on throat) or covered by a more or less dense network of brownish spots.
Based on photographs of two specimens (SAMA R66124; and ZMB 76754, Fig. 17) in life, grey dorsally and laterally with uneven coppery tinge, or more rusty brown; indistinct brown bar between the eyes with well-defi ned anterior edge but poorly defi ned posteriorly and grading into mottled darker brown patches.Other brown pigmentation variable but may be concentrated in a poorly defi ned canthal stripe Morphological characters of the paratypes.For mensural variation of the paratypes see Table 5. Snout-urostyle length of 11 males, including the holotype, vary from 15.5 to 18.5 mm (mean 17.0 mm, SD 0.91); the only female measures 22.1 mm SUL.The latter has obviously ripe or nearly ripe ova in its ovaries.It differs from all males by its much larger tympana.Mean ratio TyD/ED of the males is 0.38, that of the female is 0.64.It is impossible to decide here whether the large tympanum of the female is an aberration or expression of a real sexual dimorphism.Its other body ratios are within those of the males.Dorsal surfaces of the female are uniformly grey-brown with only a few darker spots on extremities, above tympanum, and on eye lids; the dorsum of her snout is paler than the remaining dorsum.Dorsal surfaces of males vary from uniformly greyish to heavily spotted with irregular dark spots on certain regions, predominantly on fl anks, head sides, and forelimbs.Dorsal surface that is narrowly interrupted behind the eye but then continues to a point above the axilla; in a broad, irregular band of mottled blotches along the upper jaw; in a broken line of patches along the mid-lateral surfaces extending to the inguinal region; in a narrower line ventrolaterally from the axilla to the inguinal re-   Distribution and ecological notes.This species was found only in the montane grasslands that dominate much of the highest altitudes of the Kaijende Highlands (Fig. 18).Males called at night, particularly after rain, from perches up to 30 cm high in Des champsia grass clumps.The species' local distribution appears to be patchy because this species was not seen or heard around the nearby Omyaka Camp.
Vocalisation.Advertisement calls are series of peeps (Figs. 19a and 19b).Four calls by SAMA R66121   The junior author collected 12 specimens of an Albericus species at Omyaka Camp in alpine grassland at 3,200 m asl on the Kaijende Highlands (Fig. 11) of Enga Province, Papua New Guinea (05°31.533'S,143°03.266'E;3,200 m asl) in August 2005.Their mor phology and ecology suggest a close relationship with A. alpestris, and in the absence of information about advertisement calls and colour in life of A. alpestris, we have taken a conservative approach and tentatively assign the Kaijende material to that species as Albericus cf.alpestris.However, we recognise that future collections of A. alpestris from the type locality with associated call data and information about colours in life may reveal species-level differences.Because knowledge of A. alpestris is very scanty, we here provide some information on colour, morphology (Table 6), and the advertisement call of the Kaijende population of this species.Based on photographs of several specimens in life, A. cf.alpestris from the Kai jende Highlands is a striking orange and yellowmottled frog with extensive patches of dark brown pigmentation that, although highly variable in extent, is often focused in a broad band posterior to the orbits, and in large but broken patches elsewhere on the dorsum, lateral surfaces and limbs (Figs. 21,22,23).Some specimens have a narrow, orange-yellow middorsal stripe, and creamy-grey patches laterally and on the limbs.The iris is dark golden brown.Ventral surfaces are predominantly orange with scattered pale fl ecks, except for the under-side of the limbs and a bar across the throat which are translucent with dark stippling (on throat) and dark brown on the posterior of the thighs.There are small patches of red-brown on exposed surfaces of the hind limbs, and pale creamy-168 ms, mean 156 ± 6.24 ms and mean of means of inter-note interval length varies from 231 ms to 271 ms, mean 251 ± 16.2 ms.Absolutely shortest notes are about 50 ms in the case of starting notes and 135 ms in the course of a series, and the absolutely longest note is 185 ms.Absolutely shortest inter-note interval is 197 ms and absolutely longest interval, which was mostly the last interval in a call, is 542 ms.Fundamental frequency is at 1.9 kHz and the dominant frequency is at 3.8 kHz (Fig. 20).There are fi ve upper harmonic bands (including dominant band), the highest and still well pronounced is at 11.4 kHz.All notes exhibit a weak increase of frequency.

Comparison with other species. There is no other
Oreophryne species that matches a combination of the features given under the paragraph "Diagnosis".
Etymology.Gramen is a Latin substantive of neuter gender and means grass.Graminis is the genitive of gramen and means "of the grass", and refers to the preferred habitation of the new species.

Remarks on Albericus alpestris
Albericus alpestris was recently described on the basis of four specimens that were collected in January 1973 and stored since that time in the collection of the Bishop Museum in Honolulu (Kraus, 2010).This species exhibits most anatomical characters of the genus Albericus but, unlike all other known members of the genus, it does not live as an arboreal or scansorial forest dweller, but occupies alpine grasslands.    A. cf.alpestris in the Kaijende Highlands was found only in mossy 'fern-land' above 3,000 m asl (Fig. 11) where males called from holes in the moss at the base of grass tussocks, or from slightly elevat-pestris 0.75 -0.83, in A. cf.alpestris 0.86 -1.00; and ED/SUL in A. al pes tris 0.082 -1.02, in A. cf.alpestris 0.095 -0.120.However, there are clear differences in pigmentation: A. cf.alpestris from the Kaijende Highlands exhibit many more blackish spots and even blackish areas (in preservative) on all dorsal, lateral,   (Günther, 2000).
Although obvious differences in some mensural characters and in colouration between A. alpestris from the type locality and A. cf.alpestris from the Kajiende Highlands were observed, it seems premature on the data currently available to designate the specimens from the Kajiende Highlands as a separate species.ed positions on mossy tree fern (Cyathea) stumps.Calling activity occurred predominantly during a short period beginning at about 5.30 pm and then ceased, or reduced to a very low level, soon after nightfall; sporadic calls were heard throughout the night.However the calling period of this species was longer than that of the sympatric Cophixalus tenuidactylus.The calling period of that species had ceased by around 7.30 pm.Temperatures during this time were generally 10 -13 °C.There are two types

Fig. 1 .
Fig. 1.Maps of Papua New Guinea and of Enga Province, PNG, showing the locations of collection sites mentioned in the text.
Plate II.Holotype of Albericus brevicrus sp.nov.(a) dorsal view, (b) ventral view, (c) lateral view of head, (d) ventral view of left hand, (e) ventral view of right foot.
Plate III.Holotype of Cophixalus tenuidactylus sp.nov.(a) dorsal view, (b) ventral view, (c) lateral view of head, (d) ventral view of left hand, (e) ventral view of left foot.
dark-brown spots, and in a few individuals the ventral surfaces are completely brown.Numerous whitish dots on the ventral surfaces are characteristic of all specimens.The holotype is the darkest individual and one paratype is more golden brown with a greenish tinge (ZMB 76746) and another is almost olive coloured (un-numbered individual) in life.

Fig. 13 .
Fig. 13.Power spectrum of one advertisement call note of Cophixalus tenuidactylus sp.nov.
Plate IV.Holotype of Oreophryne streiffeleri sp.nov.(a) dorsal view, (b) ventral view, (c) lateral view of head, (d) ventral view of left hand, (e) ventral view of right foot.uniform light ventral surface, and in all other frogs the ventral surfaces are more or less densely covered by dark brown dots; consequently, those frogs exhibit ventral surfaces from light brown to dark brown.Inferior tarsi are dark brown or blackish in most specimens.Distribution of tubercles in paratypes is much the same as in the holotype.

Fig. 16 .
Fig. 16.Power spectrum of one advertisement call note of Oreo phryne streiffeleri sp.nov.
Plate V. Holotype of Oreophryne graminis sp.nov.(a) dorsal view, (b) ventral view, (c) lateral view of head, (d) ventral view of left hand, (e) ventral view of left foot. in other scattered patches on the dorsum.Ventrally transluscent creamy pink on throat with pale brown stippling and splotches of opaque white, abdomen darker than throat, with scattered pale blotch es.Hidden surfaces of thighs translucent with brown stippling and rusty blotches.

Fig. 19 .b
Fig. 19.Wave form (a) and audiospectrogram (b) of seventeen notes of an advertisement call of Oreophryne graminis sp.nov.

Table 1 .
Body measurements and body ratios of the type series of Albericus pandanicolus sp.nov.SAMA R66102 is the holotype, ZMB 76959 is now an osteological preparation.All types are adult males.All measurements are in mm; abbreviations are explained in "Material and methods".
Description of the holotype.For morphological characters see Plate II and for body measurements and body ratios see Table2.Head in the region of tympana clearly broader than long (HL/HW 0.75); neck region constricted; snout from above mucronate, round in profi le; canthus rostralis clearly expressed and curved; eyes large and protruding, pupil horizontally oval; loreal region oblique, nostrils anterolaterally directed, neither visible from above nor from below, nearer to tip of snout than to eyes; internarial distance wider than distance from eye to naris (END/ IND 0.85); only lower part of tympanum visible, its horizontal diameter less than half of eye diameter (TyD/ED 0.41), supratympanic fold consists of large tubercles; tongue longitudinally oval, posteriad somewhat broadened and without posterior notch, its lateral and posterior regions free, prepharyngeal ridge of strong and irregularly shaped protuberances, vocal slits fairly long.Fingers of medium size, all with terminal discs and circum-marginal grooves, disc of fi rst fi nger conspicuously smaller than

Table 2 .
Body measurements and body ratios of the type series of Albericus brevicrus sp.nov.SAMA R66100 is the holotype; ZMB 76745 is now an osteological preparation; ZMB 76744 is a female, all others are males.UPNG 10060 is a subadult with small testes and without vocal slits.

Table 3 .
Body measurements and body ratios of the type series of Cophixalus tenuidactylus sp.nov.SAMA R66105 is the holotype, ZMB 76746 is now an osteological preparation.SAMA R66103, SAMA R66105, and ZMB 76746 are adult males, SAMA R66104 and ZMB 76747 are adult females, and ZMB 76748, SAMA R66106, and UPNG 10062 are probably subadult specimens.
Günther, 2006b)only infrequently, and they hid in thick mossy humus under clumps of grass.As a result, few individuals were found despite intensive searching during the survey period.Egg guarding seems to be common among New Guinean microhylids (e.g.Günther, 2006b).

Table 4
and body ratios see Table4.There is a longitudinal cut in the left side of the posterior abdomen.Head clearly broader than long (HL/HW 0.74), weak constriction at neck region, snout blunt in dorsal view and truncate in profi le, canthus rostralis slightly bent, loreal region oblique and slightly concave, nostrils lateral and near tip of snout; eyes medium-sized (ED/ per second, mean rate 18.0 notes per second.Except for the introductory note, all wave forms of the notes exhibit the shape of a cross section of a discus.Most introductory notes are double notes.Both parts of the introductory notes are longer than the remaining notes.

Comparison with other species.
Nearly alAllison, 1982 andgenus Cophixalus possessParker, 1989nger and toe tips.Exceptions in New Guinea are C. sphagnico laZweifel &Allison, 1982 and C. verecundus Zweifel &Parker, 1989.The third case is the new species C. tenuidactylus.These three species are morpho lo gi cally similar but they differ conspicuously in their advertisement calls.Cophixalus sphagnicola utters long, high-pitched, and fi nely tuned peeps.

Table 5 .
Body measurements and body ratios of the type series of Oreophryne graminis sp.nov.SAMA R66123 is the holotype, ZMB 76754 is now an osteological preparation, ZMB 76755 is a female, and all others are males.