taxonID	type	description	language	source
03C087ACFF8BFF91AB84F8A3FEE5CFF7.taxon	materials_examined	Holotype. BPBM 1187, collected by J. L. Gressitt, Bomberai, 2.766 º S, 132.788 º E, Fakfak Division, West Papua Province, Indonesia, 3 June 1959.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8BFF91AB84F8A3FEE5CFF7.taxon	diagnosis	Diagnosis. A species characterized by its unique combination of small size (SV = 15.7 mm) with finger discs larger than toe discs (3 rdF / 4 thT = 1.30); first finger of normal size, with disc that lacks a circum-marginal groove; moderately large finger discs (3 rdF / SV = 0.046); short legs (TL / SV = 0.46); short and narrow snout (EN / SV = 0.089, SN / EY = 0.88, EN / IN = 1.08); hidden tympanum; large eye (EY / SV = 0.15); face dark brown posteriorly, tan with dark-brown flecks anteriorly; boldly blotched with dark-brown spots laterally and ventrally (Fig. 5). Comparisons with other species. The new species differs from all Papuan congeners except C. daymani Zweifel, 1956 b and C. kethuk in having the snout shorter than the eye. It differs from C. daymani in its narrower snout (IN / SV = 0.90 – 0.108, EN / IN = 0.78 – 0.88 in C. daymani); it differs from C. kethuk in lacking (vs. having in C. kethuk) basal webbing on the toes, having a normal-sized (vs. reduced in C. kethuk) first finger with (vs. lacking in C. kethuk) a disc, and being blotched dorsally (vs. having a dorsal hourglass pattern in C. kethuk). Cophixalus pictus also differs from all Papuan congeners in its boldly blotched lateral and ventral pattern. Cophixalus pulchellus Kraus and Allison, 2000 is boldly blotched dorsally with black on silver-gray, but has a dark venter and disc of third finger smaller than that of fourth toe (3 rdF / 4 thT = 0.94 in C. pulchellus); C. bewaniensis has a single large dark-brown blotch laterally but has the venter flecked with brown and has a reduced first finger lacking a disc. It is uncertain if the hidden tympanum of the holotype of C. pictus is a preservation artifact, but this seems unlikely. If it is a reliable feature, then this character also serves to distinguish C. pictus from all Papuan congeners except C. cryptotympanum Zweifel, 1956 b, which, in addition to the characters mentioned above, differs in having longer legs (TL / SV = 0.50 – 0.52 in C. cryptotympanum), a broader snout (EN / IN = 0.80 – 0.91 in C. cryptotympanum), and a light postocular stripe.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8BFF91AB84F8A3FEE5CFF7.taxon	description	Description of holotype. Sex uncertain because of state of preservation of internal features, but likely a juvenile male; eggs and enlarged oviducts clearly absent. Head moderately wide (HW / SV = 0.43), with steeply oblique loreal region and inflated upper lip; canthus rostralis rounded, straight when viewed from above; nostrils directed laterally, closer to tip of snout than to eyes; internarial distance narrower than distance from naris to eye (EN / IN = 1.08, IN / SV = 0.083, EN / SV = 0.089); snout slightly rounded when viewed from the side, rounded when viewed from above; eyes large (EY / SV = 0.15); eyelid approximately same width as interorbital distance; tympanum hidden. Dorsal, lateral, and ventral surfaces smooth; supratympanic fold absent. Fingers unwebbed, all bearing well-developed discs; relative lengths 3> 4> 2> 1. Discs of F 3 and F 4 with poorly developed circum-marginal grooves, approximately 2 times widths of penultimate phalanges (3 rdF / SV = 0.046); discs of F 1 and F 2 lacking such grooves, that of F 2 approximately 1.5 times width of penultimate phalanx, that of F 1 barely wider than penultimate phalanx. Subarticular tubercles poorly developed, metacarpal tubercles not evident. Toes unwebbed, bearing discs, with terminal grooves on T 2 – T 5; relative lengths 4> 3> 5> 2> 1. Toe discs smaller than those of fingers (3 rdF / 4 thT = 1.30); disc of fourth toe approximately 1.5 times as wide as penultimate phalanx (4 thT / SV = 0.036); disc of first toe approximately same width as penultimate phalanx. Subarticular and metatarsal tubercles not obvious. Hind legs rather short (TL / SV = 0.46). Dorsum and upper sides brown, lighter lower on sides, with few dark-brown flecks on head and shoulders; sides with numerous dark-brown blotches, larger and more rounded ventrally (Fig. 5). Front and rear of thighs brown with large dark-brown blotches. Posterior of face dark brown with few tiny tan flecks; snout tan with few dark-brown flecks (Fig. 5). Chin, throat, and chest dark brown with few tiny tan flecks, mostly near jaw margins; symphysis tan. Abdomen tan with dark-brown blotches and minutely stippled with black. Iris dark brown. Measurements (in mm). — SV = 15.7, TL = 7.3, HW = 6.8, IN = 1.3, EN = 1.4, SN = 2.1, EY = 2.4, 3 rdF = 0.73, 4 thT = 0.56.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8BFF91AB84F8A3FEE5CFF7.taxon	etymology	Etymology. The name is a masculine Latin adjective meaning “ painted ” and is in reference to the boldly blotched color pattern of the species. Range. Known only from the type locality in western New Guinea (Fig. 6). The coordinates are along the coast, and the entire peninsula is of low elevation, so presumably this species inhabits lowland rainforest.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8EFF8AAB84F90EFDF4CFF0.taxon	materials_examined	Holotype. BPBM 19323 (field tag FK 9108), collected by F. Kraus and B. Iova, W slope Mt. Obree, 9.4601 º S, 148.0304 º E, 1800 – 2040 m, Central Province, Papua New Guinea, 7 February 2004. Paratypes (n = 21). BPBM 19320 – 21, same data as holotype except collected 4 February by F. Kraus; BPBM 19322, same data as holotype except collected 5 February by F. Kraus; BPBM 19324 – 37, PNGNM 24108 – 11, same data as holotype.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8EFF8AAB84F90EFDF4CFF0.taxon	diagnosis	Diagnosis. A species characterized by its unique combination of modest size (male SV = 18.9 – 22.7 mm, female SV = 27.2 – 27.5 mm); basal webbing between toes; finger discs larger than toe discs (3 rdF / 4 thT = 1.07 – 1.37); first finger of normal size, bearing a disc; dorsum pustulose, irregularly smudged / mottled with dark brown; and venter straw yellow densely and evenly flecked with dark brown. Comparisons with other species. The new species differs from all other Papuan members of the genus except C. kethuk and C. tagulensis in having webbing between the toes. Both of those species are of smaller size (SV 12.4 – 15.0 mm in C. kethuk and 13.5 – 18.5 mm in C. tagulensis) and have finger discs smaller than toe discs. Further, C. kethuk lacks a disc on the first finger and has a smooth dorsum, and C. tagulensis has the toes halfwebbed and the venter evenly stippled, not flecked, with dark brown.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8EFF8AAB84F90EFDF4CFF0.taxon	description	Description of holotype. An adult male with small right-lateral incision, vocal slits. Head moderately wide (HW / SV = 0.39), with oblique loreal region; canthus rounded, straight when viewed from above (Fig. 7 A); nostrils directed laterally, closer to tip of snout than to eyes; internarial distance larger than distance from naris to eye (EN / IN = 0.90, IN / SV = 0.093, EN / SV = 0.084); snout truncate when viewed from the side (Fig. 7 B), shallowly angulate when viewed from above; eyes of moderate size (EY / SV = 0.12); eyelid approximately 2 / 3 width of interorbital distance; tympanum small (TY / SV = 0.026) and indistinct. Dorsal and lateral surfaces pustulose; ventral surfaces granulose; supratympanic fold small but distinct. Fingers with trace of basal webbing; relative lengths 3> 4> 2> 1; first finger well developed (Fig. 7 C). Third finger disc approximately twice width of penultimate phalanx (3 rdF / SV = 0.077); that of first finger barely wider than penultimate phalanx. Subarticular and metacarpal tubercles low and poorly developed but distinct. Toes with basal webbing and fringes along T 3 and T 4 that extend to the disc, bearing discs with terminal grooves; relative lengths 4> 3> 5> 2> 1 (Fig. 7 D). Toe discs smaller than those of fingers (3 rdF / 4 thT = 1.30); disc of fourth toe less than 1.5 times width of penultimate phalanx (4 thT / SV = 0.059); disc of first toe approximately same width as penultimate phalanx. Subarticular tubercles poorly developed; inner metatarsal tubercle low and elongate, outer lacking. Hind legs moderately long (TL / SV = 0.48). Dorsal ground color dark tan, irregularly mottled, smudged, and freckled with dark brown, this concentrated dorsolaterally, on snout, on posterior of head, and in a suprascapular chevron. A vaguely defined tan postocular stripe runs below the darker supratympanic fold. Rear of thighs tan mottled with dark brown; front of thighs same but with less dark mottling. Ventral surfaces straw yellow densely and evenly flecked with dark brown and with scattered gray-white punctations across abdomen. Iris black flecked with silver. Measurements (in mm). — SV = 22.7, TL = 11.0, HW = 8.8, HL = 7.4, IN = 2.1, EN = 1.9, SN = 3.0, EY = 2.7, TY = 0.6, 3 rdF = 1.75, 4 thT = 1.35. Variation. Mensural variation for the type series is shown in Table 5. Adult males and juveniles appear to have slightly larger snouts than do adult females (Table 5), but the difference is not great. Otherwise, no clear ontogenetic or sexually dimorphic differences are apparent in the type series. All specimens have clearly developed basal webbing between the toes, with that of the largest females appearing only slightly more conspicuous than what is seen in the other specimens. There is little color variation. Most specimens have the gray-brown appearance of the holotype, but three are lighter brown. Most have the dark suprascapular chevron, but this is often indistinct. The dark ventral flecking varies only slightly in intensity, with only four specimens having less contrast with the ground color than does the holotype. Color in life. From field notes for holotype BPBM 19323 (Fig. 8 A): “ Dorsum mud-brown, reddish middorsally, with gray suprascapular chevron and dark brown flecks all over. Iris mud-brown. Venter pale purple, except chin and throat, which are pale straw; entire venter speckled with dark brown flecks. Rear of thighs mudbrown with dark-brown mottling. One dark band across the forearm, another above the wrist. ” BPBM 19321: “ Dorsum green-gray with gray scapular mark and irregular gray or black flecking, with pustules, which are tan. Limbs tan with black flecking; rear of thighs same. Venter flecked light and dark gray with scattered white flecks on abdomen. Posterior belly and under legs light straw. Iris light brown. ” BPBM 19320 was yellow-brown above; BPBM 19330 had a narrow, tan mid-dorsal stripe. Call. This species calls at night from the edges of streams. Most calling animals were perched on leaves 4 – 30 cm above ground, but one specimen was calling from under a rock. The call is a single, relatively long (0.99 – 1.35 s), highly pulsed note lacking harmonic structure and frequency modulation (Fig. 9 C) and delivered at intervals of 2.7 to 50 s apart (Table 6). To the human ear the call sounds similar to a squeaky door hinge or a throaty snore. The number of pulses / call varied from 44 – 63, and pulse rate varied from 41.5 – 57.4 / s. Notes increase in amplitude relatively rapidly and maintain approximately maximum amplitude for most of the note before decreasing again relatively rapidly (Fig. 9 A). Notes have lesser-amplitude pulses at the beginning and end of the note, giving the waveform a rounded rectanglar shape, with most notes showing approximately equal-amplitude pulses for most of their duration (Fig. 10 A), but for the holotype notes frequently increase to peak amplitude more gradually than they decrease, giving the amplitude envelope a slight skew to the left (Fig. 10 B). Pulsing is invariably slower at the beginning, and quicker at the end, of each note. Dominant frequency varies from 2.80 – 3.96 kHz (Table 6, Fig. 9 B).	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF8EFF8AAB84F90EFDF4CFF0.taxon	etymology	Etymology. The trivial epithet is a masculine latinized Greek adjective meaning “ web-footed ”. Range. Known only from a single mid-elevation site at 1800 – 2040 m on the western slope of Mt. Obree, Central Province, Papua New Guinea (Fig. 2). Ecological notes. Animals were moderately common along the edges of a small, first-order stream having a bed of ca. 1 – 3 m width and containing water ca. 2 – 15 cm deep. Streambed consisted mostly of gravel and small rocks and occurred in relatively open forest with a canopy of approximately 30 m height. Only one animal was found higher than 30 cm above the ground or farther than 1 m from the edge of the stream. When disturbed, animals jumped to the rocks or gravel beneath and pressed themselves close to it. In such instances, their color matched the substrate well. This is the only Cophixalus I have seen leap into a stream in an attempt at escape. This and the partially webbed feet suggest the species spends some amount of its time in streams, if only to escape predators. The smallest mature male is 18.9 mm SV; an immature male is 16.5 mm. The smallest mature female is 27.2 mm; four immature females range from 19.9 – 22.4 mm. Three unsexed juveniles range from 13.4 – 14.8 mm SV. Syntopic microhylids include Callulops doriae, Cophixalus ateles, Cophixalus iovaorum, Hylophorbus sp., and Liophryne magnitympanum.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF90FF8CAB84FF09FA67CBC4.taxon	materials_examined	Holotype. BPBM 18433 (field tag FK 8479), collected by F. Kraus at NW slope Mt. Shungol, 5.35 km NW of summit, 6.8188 º S, 146.6933 º E, 780 m, Morobe Province, Papua New Guinea, 21 October 2003. Paratypes (n = 10). BPBM 18432, 18434 – 37, same data as holotype; BPBM 18429 – 30, NW slope Mt. Shungol, 5.6 km NW of summit, 6.8162 º S, 146.6915 º E, 750 m, 16 October; BPBM 18431, same data as 18429 except collected 20 October; BPBM 18438 – 39, same data as 18429 except collected 23 October. Referred specimens. BPBM 13445 – 49. Papua New Guinea: Morobe Province: Oomsis Forestry Camp, 6.6984 º S, 146.8157 º E, 500 – 530 m.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF90FF8CAB84FF09FA67CBC4.taxon	diagnosis	Diagnosis. A species characterized by its unique combination of small size (male SV = 16.8 – 20.5 mm, female SV = 20.1 – 21.0 mm); finger discs smaller than toe discs (3 rdF / 4 thT = 0.65 – 0.77); first finger of reduced size, but still elongate and functional, lacking a flattened disc and terminal groove; leg relatively long (TL / SV = 0.53 – 0.61); snout short and broad (EN / IN = 0.71 – 0.91); skin smooth; face and postocular area black with a white line extending from snout tip, along canthus, and above eye, and another white line extending from behind eye, through tympanum, and down forearm; rear of thighs dark brown; and relatively slow peeping call with a dominant frequency of 1900 – 2000 Hz. Comparisons with other species. Cophixalus albolineatus differs from all other members of the genus in having a conspicuous white line extending from the eye down the forearm; it also differs from all other Papuan Cophixalus except for C. ateles, C. desticans, C. iovaorum, C. kethuk, C. pipilans, C. shellyi, and C. tomaiodactylus in the combination of having the finger discs smaller than the toe discs and having a reduced, though functional, first finger that lacks an expanded disc or terminal groove. Cophixalus bewaniensis, C. humicola, and C. tridactylus also have finger discs smaller than toe discs and a reduced first finger, but in those species the first finger is miniaturized to a non-functional nub easily distinguished from the elongate, functional finger of C. albolineatus. From C. ateles and C. tomaiodactylus, C. albolineatus further differs in having a black face and a smooth dorsum (vs. pustulose or with dorsolateral row of pustules). Cophixalus iovaorum and C. kethuk further differ from the new species in having an hourglass pattern dorsally, marked by black dorsolateral lines. Cophixalus iovaorum also differs in its smaller size (SV = 13.2 – 17.2 mm) and shorter leg (TL / SV = 0.41 – 0.49), and C. kethuk has webbing between the toes, which is lacking in C. albolineatus. From C. desticans, the new species further differs in its larger size (male SV = 13.1 – 16.2 mm in C. desticans) and smooth skin (with dorsal ridges in C. desticans). Cophixalus albolineatus further differs from C. pipilans in lacking a dark postorbital bar, lacking a black dorsolateral line, and having a slower call with a lower dominant frequency (4900 – 5300 Hz in C. pipilans); it further differs from C. shellyi in its longer leg (TL / SV = 0.44 – 0.53 in C. shellyi) and having a much slower call with a lower dominant frequency (ca. 5200 Hz in C. shellyi).	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF90FF8CAB84FF09FA67CBC4.taxon	description	Description of holotype. An adult female with small right-lateral incision. Head rather wide (HW / SV = 0.41), with vertical, slightly concave loreal region; canthus rounded, straight when viewed from above (Fig. 11 A); nostrils directed laterally, closer to tip of snout than to eyes; internarial distance larger than distance from naris to eye (EN / IN = 0.77, IN / SV = 0.127, EN / SV = 0.098); snout projecting when viewed from the side (Fig. 11 B), shallowly angulate when viewed from above; eyes of moderate size (EY / SV = 0.12); eyelid slightly more than half width of interorbital distance; tympanum fairly large (TY / SV = 0.078), with a distinct annulus. Dorsal, lateral, and ventral surfaces smooth; supratympanic fold absent. Fingers without webbing; relative lengths 3> 4> 2> 1. First finger small, with rounded tip but no expanded disc or circum-marginal groove; expanded discs and circummarginal grooves on remaining fingers (Fig. 11 C); third finger disc approximately twice width of penultimate phalanx (3 rdF / SV = 0.049). Subarticular and metacarpal tubercles low and poorly developed but distinct. Toes without webbing or fringes, bearing discs with terminal grooves; relative lengths 4> 3> 5> 2> 1 (Fig. 11 D). Toe discs larger than those of fingers (3 rdF / 4 thT = 0.74); disc of fourth toe more than twice width of penultimate phalanx (4 thT / SV = 0.066); disc of first toe slightly broader than penultimate phalanx. Subarticular tubercles poorly developed; inner metatarsal tubercle low and elongate, outer lacking. Hind legs long (TL / SV = 0.57). In preservative, dorsum medium brown with few dark-brown spots; sides pale straw yellow with reticulum of brown; face, lower jaw, and tympanum dark brown with a postocular pale straw-yellow stripe extending from behind eye, through tympanum, and down forearm, subtended on forearm by field of dark brown. Palmar and plantar surfaces dark brown. Ventral surfaces pale straw yellow flecked with brown, flecking denser anteriorly and sparser on abdomen and under legs. Rear of thighs brown flecked with dark brown. Iris black. Measurements (in mm). — SV = 20.4, TL = 11.7, HW = 8.3, HL = 7.5, IN = 2.6, EN = 2.0, SN = 3.1, EY = 2.5, TY = 1.6, 3 rdF = 1.00, 4 thT = 1.35. Variation. Females average slightly larger than males (Table 7), and there is some indication that females may have larger discs than males, but sample sizes are insufficient to be certain of this point. Otherwise, mensural variation among the type series is rather slight. Color pattern varies slightly: three juveniles and one adult have a narrow light-tan vertebral line; two adults have more extensive dark-brown mottling dorsally than do the remaining specimens, and they have a dark-brown, backward-pointing triangle between the eyes. The width of the white postocular line varies from half to two-thirds the width of the tympanum. And the rear of the thighs varies from light brown distinctly blotched with dark brown to fairly uniformly dark brown. Color in life. From field notes for paratype BPBM 18429 (Fig. 8 B): “ Dorsum brown, face and tympanic region black with a white stripe running from snout along canthus and upper eyelid to forearm insertion and down forearm. Chin to chest black, with few white flecks on chest; belly gray with few black and white flecks. Rear of thighs dark brown; tan line above anus. Iris dark brown rimmed with orange around pupil. ” Call. This species calls primarily from 1 – 2 h before dark to 1 – 2 h after dark, with the frequency of calling decreasing noticeably after dark. It may call briefly during daytime, especially following rain. The call is a rapid series of 2 – 23 raspy chirps emitted at a rate of 0.32 – 0.95 notes / s (mean = 0.71 notes / s for six call series); calls range from 3 – 25 s in duration (Table 8). Each note is brief, with a mean duration of 0.100 s (range 0.076 – 0.113 s). The interval between notes was considerably longer, averaging 1.265 s and ranging from 0.517 – 4.702 s. There was little variation among individuals in note duration, but internote duration varied considerably (Table 8). Each note increases sharply to maximum amplitude and decreases at an accelerating rate, creating an approximately semi-lunar amplitude envelope (Figs. 12 A, 13 A). The note has a well-developed harmonic structure and is frequency modulated, with a downward slur of approximately 200 – 300 Hz (Fig. 12 C). The dominant frequency of the calls varied within a very narrow window (Fig. 12 B), averaging 1972 Hz and ranging from 1911 – 2010 Hz. Specimen Call Temperature Number Call Note duration Internote Repetition Dominant series (˚ C) of notes duration (s) (s) duration (s) rate (notes / s) frequency (kHz) BPBM a 22.5 17 18.6 0.094 ± 0.0013 1.043 ± 0.1138 0.92 1.95 ± 0.0027 18432 (0.084 – 0.101) (0.525 – 1.992) (1.93 – 1.97) b “ 3 9.4 0.083 ± 0.0033 4.446 ± 0.2565 0.32 1.92 ± 0.0052 (0.078 – 0.089) (4.189 – 4.702) (1.91 – 1.93) c “ 2 3.6 0.084 ± 0.0075 3.302 0.55 1.92 ± 0.0005 (0.076 – 0.091) (1.92) BPBM a “ 23 25.1 0.103 ± 0.0008 1.029 ± 0.1821 0.92 1.98 ± 0.0022 18433 (0.097 – 0.110) (0.517 – 3.781) (1.96 – 2.01) b “ 12 20.0 0.099 ± 0.0012 1.730 ± 0.1498 0.60 1.98 ± 0.0017 (0.090 – 0.104) (0.916 – 2.532) (1.97 – 1.99) c “ 19 20.0 0.106 ± 0.0010 0.998 ± 0.1112 0.95 1.98 ± 0.0023 (0.099 – 0.113) (0.567 – 2.476) (1.96 – 2.00) Etymology. The trivial epithet is a Latin combinatorial adjective from “ albus ” for “ white ” and “ linea ” for “ line ”. It refers to the two distinctive white lines on the head and extending down the forearm in this species. Range. Known from the northwestern slope of Mt. Shungol, Madang Province, Papua New Guinea, and from Oomsis, approximately 18 km to the NE (Fig. 2). Ecological notes. This species inhabits leaf litter, from which it begins calling in late afternoon, continuing for the first couple hours of dark. Habitat at the area of collection was medium-crowned lowland hill forest (Paijmans, 1975, 1976) with a canopy ~ 30 m high and emergents to ~ 50 m. Specimens were collected along the narrow (ca. 30 – 50 m-wide) valley bottom bounded by the Dunch River and adjacent steep hills. This valley bottom appeared to be a perched, rocky floodplain, had a series of tributary streams and braided rivulets crossing the area, and had a rich soil with a moderately dense understory, especially of urticaceous herbs. The smallest mature male is 16.0 mm SV; an immature male is 14.0 mm. The smallest mature female is 20.1 mm; one immature female is 14.8 mm. Syntopic microhylids include Albericus exclamitans, Austrochaperina parkeri, Austrochaperina palmipes, Callulops personata, Cophixalus cheesmanae, Copiula fistulans, Hylophorbus sp., Mantophryne lateralis, and Oreophryne geislerorum.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF90FF8CAB84FF09FA67CBC4.taxon	discussion	Remarks. Specimens BPBM 13445 – 49 were erroneously identified many years ago as Cophixalus pipilans by R. Zweifel, author of that species; they have been catalogued under that name since that time. Identity between this older sample and the type series of C. albolineatus led me to apply the name C. pipilans to this latter sample for several years too. I subsequently used these specimens in diagnosing several new species of Cophixalus from C. pipilans (Kraus and Allison, 2009). Because the original identification was in error, those diagnoses vis-à-vis C. pipilans were also sometimes incorrect in particulars. The only newly described species in Kraus and Allison (2009) seriously affected by this error was C. desticans, which is actually more similar to true C. pipilans (subsequently collected by me and verified against the type series, see Appendix) than first presumed. Based on these specimens of C. pipilans and recordings I made for four animals, C. desticans may be correctly diagnosed from C. pipilans in having a dorsum with many parallel dermal ridges (dorsum smooth in C. pipilans), a broad dark lateral band (vs. a narrow lateral stripe or series of dashes in C. pipilans), rear of thighs brown (vs. dark orange in C. pipilans), and a slower advertisement call (0.4 notes / s in C. desticans vs. 1.5 – 2.0 notes / in C. pipilans) with a higher dominant frequency (5900 – 6400 Hz in C. desticans vs. 4900 – 5300 Hz in C. pipilans) and a rounded waveform lacking a spike in amplitude (waveform with a sharp amplitude spike in the first 0.01 s of call in C. pipilans). None of the other species described by Kraus and Allison (2009) is liable to confusion with C. pipilans.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF94FF80AB84FC69FB39C97C.taxon	materials_examined	Holotype. BPBM 37720 (field tag FK 14753), collected by F. Kraus along Upper Muniai Stream, below Bomagai Camp, Woodlark Island, 9.1216 ° S, 152.7486 ° E, 67 m, Milne Bay Province, Papua New Guinea, 12 October 2010. Paratypes (n = 31). BPBM 37718 – 19, 37721 – 25, PNGNM 24116, same data as holotype; BPBM 37726 – 29, same data as holotype, except collected 13 October 2010; BPBM 37731 – 35, 39040, PNGNM 24117 – 18, Thompson’s Creek, 9.15591 ° S, 152.7430 ° E, 104 m, 15 October; PNGNM 24119 – 20, same data as holotype, except collected 16 October 2010; BPBM 37737 – 38, same data as holotype, except collected 18 October 2010; BPBM 37736, below Bomagai Camp E, 9.1222 ° S, 152.7535 ° E, 38 m, 17 October; BPBM 37739, same data as BPBM 37736, except collected 18 October 2010; BPBM 40005 – 06, same data as holotype, except collected 11 October 2011; BPBM 40007, same data as holotype, except collected 12 October 2011; BPBM 40008 – 09, same data as holotype, except collected 16 October 2011.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF94FF80AB84FC69FB39C97C.taxon	diagnosis	Diagnosis. A species characterized by its unique combination of moderate size (male SV = 23.2 – 27.5 mm, female SV = 26.1 – 28.3 mm) with finger discs and toe discs of approximately same size (3 rdF / 4 thT = 0.92 – 1.13); well-developed first finger bearing disc with circum-marginal groove; dorsum with several rows of well-developed warts; top of head lighter than surrounding skin; body with irregular brown markings on a lighter brown, olivebrown, or yellow-brown ground color; and call consisting of a train of fairly slow mechanical-sounding beeps. Comparisons with other species. The combination of larger size (SV> 22 mm), finger discs subequal to or smaller than toe discs (3 rdF / 4 thT <1.14), and first finger of normal size and bearing a disc with a circum-marginal groove distinguishes this species from all congeners except C. cupricarenus Kraus and Allison, 2009, endemic to nearby Rossel Island. Cophixalus clapporum differs from that species in particulars of color pattern and call: C. cupricarenus invariably has a copper-colored or russet triangle on the top of the head, whereas C. clapporum may have that color as a head triangle but usually the top of the head is tan or light brown; C. cupricarenus has black paravertebral, dorsolateral, and lateral stripes or series of dashes, whereas C. clapporum has brown blotches or other irregular markings but lacks stripes other than a short postocular dash that extends no further posteriorly than the tympanum; C. cupricarenus has a black canthal stripe that extends from the eye to the tip of the snout, whereas C. clapporum has no more than a brown canthal blotch that does not extend anterior to the naris; C. cupricarenus often has a white vertebral line, whereas C. clapporum never shows that pattern element; and C. cupricarenus has a call with a repetition rate of 5.0 – 5.2 notes / s and a dominant frequency of 3100 – 3140 Hz, whereas C. clapporum has a slower, lower-pitched call with a repetition rate of 1.82 – 2.15 notes / s and a dominant frequency of 2250 – 2460 Hz.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF94FF80AB84FC69FB39C97C.taxon	description	Description of holotype. An adult male with small right-lateral incision. Head moderately wide (HW / SV = 0.38), with steeply oblique, slightly concave loreal region; canthus rostralis inflated, shallowly inflated when viewed from above (Fig. 14 A); nostrils directed laterally, much closer to tip of snout than to eyes; internarial distance larger than distance from naris to eye (EN / IN = 0.85, IN / SV = 0.11, EN / SV = 0.094); snout projecting when viewed from the side (Fig. 14 B), shallowly angulate when viewed from above; eyes moderately large (EY / SV = 0.13); eyelid approximately 60 % width of interorbital distance; tympanum rather indistinct and of moderate size (TY / SV = 0.053), with a slightly raised annulus anteriorly but visible posteriorly only when skin is slightly dried. Skin minutely rugose, with five rows of enlarged pustules dorsally; ventral surfaces smooth. Short supratympanic fold present. Fingers unwebbed, bearing discs with terminal grooves; relative lengths 3> 4> 2> 1; first finger and disc well-developed (Fig. 14 C). Finger discs approximately 2.5 times widths of penultimate phalanges (3 rdF / SV = 0.056), except for the first finger disc, which is approximately 1.5 times width of penultimate phalanx. Subarticular tubercles low but distinct; metacarpal tubercles oval, well-developed, but low. Toes unwebbed, bearing discs with terminal grooves; relative lengths 4> 3> 5> 2> 1 (Fig. 14 D). Toe discs approximately same size as those of fingers (3 rdF / 4 thT = 0.95); disc of fourth toe approximately twice width of penultimate phalanx (4 thT / SV = 0.058); disc of first toe approximately 1.5 times width of penultimate phalanx. Subarticular tubercles well-developed; inner metatarsal tubercle narrow and elongate, outer lacking. Hind legs moderately long (TL / SV = 0.51). In preservative, dorsum medium brown, heavily and vaguely marked with dark brown, including a vaguely defined suprascapular X; top of head somewhat lighter, followed posteriorly by dark backward-pointing triangle. Sides with greater extent of light ground and many dark-brown flecks and stipples. Tympanum light tan with small brown flecks; supratympanic fold and angle of jaw dark brown; face dark brown with few lighter-brown spots on upper jaw. Rear of thighs tan with large dark-brown blotches. Venter pale straw heavily flecked with brown, sparser on abdomen than on chin, throat, and under legs. Palmar and plantar surfaces dark brown. Iris black stippled with brass. Measurements (in mm). — SV = 24.5, TL = 12.6, HW = 9.2, HL = 8.0, IN = 2.7, EN = 2.3, SN = 3.6, EY = 3.1, TY = 1.3, 3 rdF = 1.36, 4 thT = 1.43. Variation. Mensural variation for the species is not large, and there are no obvious differences between the sexes except that females appear to average slightly larger than males (Table 9). There is, however, some indication that juveniles have relatively longer snouts and heads, wider tympana and heads, and larger eyes than do adults (Table 9). In preserative, dorsa vary from light gray-brown to very dark brown; dark markings are typically small spots or irregular marks, but the contrast with the ground color varies from very little to moderate. The darker specimens are similar to the holotype in having very obscure markings; the lightest specimens appear as light-brown frogs with distinct, scattered brown spots. Ventral patterning varies from dispersed punctations to heavily flecked with brown, such that overall appearance varies from pale straw to dark brown, with most specimens having the overall appearance of venters heavily flecked with brown. Most specimens have the rear of the thighs with a highly contrasting pattern of dark-brown blotches on a pale-straw ground, but three are generally suffused with dark brown instead. The tympanum is always lighter than the surrounding ground color, the iris always black with brass or silver punctations. Color in life. In life, BPBM 37718 was noted to be “ Olive green and brown above, top of head russet brown, black behind eye but tympanum tan. Rear of thighs black with olive-green blotches. Venter charcoal gray with light-gray flecks on posterior belly and smaller gray-white punctations throughout. Iris olive brown with a narrow orange rim around the pupil. Bottoms of hands and feet black. ” BPBM 37726 was noted to be: “ Yellow brown above with darker brown markings, top of snout tan. Rear of thighs dark brown flecked with yellow brown. Venter light gray with dark-gray flecks on throat and silver-white punctations / flecks on abdomen. Iris black reticulated or spotted with gold. ” BPBM 37719 was tan and dark brown above, with black and tan on rear of thighs, and abdomen pale-gray heavily flecked with charcoal gray; BPBM 37721 was olive and orange brown dorsally, with darker orange brown on top of head; BPBM 37722 was olive and yellow brown dorsally and tan on top of head. All animals had distinct rows of large tubercles in life (Fig. 8 C, D). Call. Animals called at night from the forest floor or perched on leaf litter, following either wet or dry days. To the human ear, the call sounds similar to a mechanical beep; when two frogs alternate calling and each calls at a different frequency, the combined effect is similar to the sound of a European police siren. The call is a series of well-tuned beeps or honks, with a series of six or seven harmonics (Fig. 15 C). Seven calls recorded by me include from 10 – 33 notes and range in length from 5.1 – 16.1 s, with notes emitted at a rate of 1.82 – 2.15 / s (Table 10). Each note is relatively brief (Fig. 15 A, C), with a mean duration of 0.220 s (range 0.195 – 0.343 s) among the seven calls. The first note of each call is approximately 30 – 50 % longer than subsequent notes. The interval between notes is only slightly longer, averaging 0.292 s among the seven calls and ranging from 0.211 – 0.811 s (Table 10). Each note begins abruptly and declines smoothly during its last 200 ms or so, producing an amplitude envelope sharply truncated on the left and rounded on the right (Figs. 13 B, 15 A). The dominant frequency of the calls varied within a very narrow window (Fig. 15 B), averaging 2330 Hz and ranging from 2250 – 2460 Hz (Table 10). Below the dominant frequency is another peak with almost as much energy (Fig. 15 C), which averages 1770 Hz and ranges from 1660 – 1937 Hz. Specimen Call Temperature Number Call Note duration (s) Internote duration Repetition Dominant series (˚ C) of notes duration (s) (s) rate (notes / s) frequency (kHz) BPBM a 26.5 21 11.0 0.227 ± 0.0050 0.304 ± 0.0079 1.91 2.42 ± 0.0050 40005 (0.214 – 0.324) (0.271 – 0.412) (2.39 – 2.46) b “ 26 14.3 0.225 ± 0.0037 0.332 ± 0.0234 1.82 2.41 ± 0.0056 (0.210 – 0.309) (0.246 – 0.811) (2.39 – 2.46) BPBM a 25.2 27 13.4 0.216 ± 0.0039 0.289 ± 0.0104 2.01 2.27 ± 0.0043 40006 (0.200 – 0.311) (0.242 – 0.490) (2.25 – 2.32) b “ 25 12.7 0.218 ± 0.0044 0.297 ± 0.0067 1.97 2.29 ± 0.0034 (0.204 – 0.319) (0.259 – 0.377) (2.25 – 2.32) BPBM a 27.0 31 14.4 0.213 ± 0.0048 0.256 ± 0.0069 2.15 2.30 ± 0.0038 40007 (0.195 – 0.343) (0.211 – 0.363) (2.25 – 2.32) b “ 33 16.1 0.212 ± 0.0038 0.277 ± 0.0109 2.05 2.31 ± 0.0033 (0.195 – 0.316) (0.225 – 0.547) (2.25 – 2.35) c “ 10 5.1 0.231 ± 0.0110 0.292 ± 0.0312 1.96 2.32 ± 0.0035 (0.205 – 0.308) (0.223 – 0.534) (2.32 – 2.35) Etymology. The name is a genitive honorific for Eleanor Clapp, who first brought my attention to this frog, and George Clapp, who graciously hosted my visits to Woodlark Island. Range. Known only from Woodlark Island, Milne Bay Province, Papua New Guinea (Fig. 2), where it is likely endemic, although it may occupy some of the adjacent islets. Animals were heard calling at several sites around the center of the island, and the species undoubtedly occurs throughout Woodlark. Ecological notes. This species inhabits mature lowland rainforest, with canopy heights at approximately 25 – 35 m. Animals were always found on or near the ground in lowland primary rainforest, most in a gully adjacent to a small, slow-moving stream. Several frogs were found sheltering during daytime under rocks immediately adjacent to a slow-moving lowland stream. The smallest mature male is 23.2 mm SV; two immature males are 16.4 and 17.6 mm. The smallest mature female is 26.1 mm; four immature females range from 16.9 – 23.9 mm. Syntopic microhylids include Barygenys sp. nov., the following species of Cophixalus, Copiula sp. nov., Genyophryne thomsoni, Hylophorbus myopicus, Mantophryne sp. nov., and Oreophryne sp. nov.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF98FF87AB84FBEAFB39CA5C.taxon	materials_examined	Holotype. BPBM 37730 (field tag FK 14787), collected by F. Kraus along Upper Muniai Stream, below Bomagai Camp, Woodlark Island, 9.1216 ° S, 152.7486 ° E, 67 m, Milne Bay Province, Papua New Guinea, 13 October 2010. Paratype (n = 8). BPBM 37740, same data as holotype, except collected 19 October 2010; BPBM 40010 – 13, same data as holotype, except collected 13 October 2011; BPBM 40014 – 15, PNGNM 24142, same data as holotype, except collected 17 October 2011.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF98FF87AB84FBEAFB39CA5C.taxon	diagnosis	Diagnosis. A species characterized by its unique combination of small size (male SV = 13.6 – 14.3 mm, female SV = 14.8 – 16.1 mm); tympanum hidden; finger discs smaller than toe discs (3 rdF / 4 thT = 0.65 – 0.77); first finger of reduced size, but still elongate and functional, lacking a flattened disc and terminal groove; leg short (TL / SV = 0.42 – 0.48); snout short and broad (EN / IN = 0.79 – 0.87), shorter than or equal to eye diameter (SN / EY = 0.86 – 1.00); skin smooth; face and side black with pale blue flecks and spots; and forearms typically orange. Comparisons with other species. Cophixalus amabilis differs from all other Papuan members of the genus except C. daymani, C. pictus sp. nov., and some C. kethuk in having the snout shorter than the eye, and it differs from all Papuan Cophixalus except C. cryptotympanum and C. pictus in having the tympanum hidden. It differs from each of these species except C. kethuk in having the finger discs smaller than the toe discs (3 rdF / 4 thT = 1.2 – 1.4 in C. cryptotympanum, 1.3 – 2.0 in C. daymani, 1.3 in C. pictus). Cophixalus kethuk differs in having basal webbing between the toes, which is lacking in C. amabilis. The color pattern of C. amabilis — in particular its black face and sides spotted with pale blue and its frequently orange forearms — is unique within the genus.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF98FF87AB84FBEAFB39CA5C.taxon	description	Description of holotype. An adult female with right-lateral incision. Head of modest width (HW / SV = 0.36), with steeply oblique, slightly swollen loreal region; canthus rounded, slightly convex when viewed from above (Fig. 16 A); nostrils directed laterally, closer to tip of snout than to eyes; internarial distance larger than distance from naris to eye (EN / IN = 0.86, IN / SV = 0.095, EN / SV = 0.081); snout rounded when viewed from the side or from above (Fig. 16 A, C), smaller than eye (SN / EY = 0.95); eyes of moderate size (EY / SV = 0.13); eyelid approximately two-thirds width of interorbital distance; tympanum hidden (Fig. 16 C). Dorsal, lateral, and ventral surfaces smooth; supratympanic fold absent. Fingers without webbing; relative lengths 3> 4> 2> 1 (Fig. 16 D). First finger small, with rounded tip but no expanded disc or circum-marginal groove; small expanded discs on remaining fingers but only F 3 and F 4 with circum-marginal grooves; third finger disc barely wider than penultimate phalanx (3 rdF / SV = 0.028). Subarticular and metacarpal tubercles low and very poorly developed. Toes without webbing or fringes, bearing discs with terminal grooves; relative lengths 4> 3> 5> 2> 1 (Fig. 16 E). Toe discs larger than those of fingers (3 rdF / 4 thT = 0.65); disc of fourth toe approximately 1.5 times width of penultimate phalanx (4 thT / SV = 0.043); disc of first toe same width as penultimate phalanx. Subarticular tubercles poorly developed; inner metatarsal tubercle low and circular, outer lacking. Hind legs short (TL / SV = 0.44). In preservative, dorsum brown irregularly marked with dark brown (Fig. 16 A); rear of thighs same. Sides and face dark brown flecked with pale gray; forearms pale straw yellow. Ventral surfaces brown spotted with pale gray (Fig. 16 B). Iris black. Measurements (in mm). — SV = 14.8, TL = 6.5, HW = 5.4, IN = 1.4, EN = 1.2, SN = 1.8, EY = 1.9, 3 rdF = 0.41, 4 thT = 0.63. Variation. As for many frogs, females attain a larger size than males (Table 11). Sample sizes are small, but, based on minimal overlap of mensural ranges between the sexes, there are tentative suggestions that males may have larger internarial distances, tympana, and finger discs than do females (Table 11). In life, one paratype (BPBM 37740) is quite different to the holotype in dorsal pattern (compare Fig. 8 E vs. 8 F), with the dark dorsal markings considerably more dissected and dispersed. The remainder of the paratypes are more like the holotype in generally having larger brown markings dorsally, although they always have a vague X-shaped blotch mid-dorsally in the scapular region. The orange forearms seen in the holotype (Fig. 8 E) were less distinct in BPBM 37740 (Fig. 8 E) and in a few of the other paratypes; in two specimens there is no pale area whatsoever on the upper arms, and these were also less distinct in life than those of the holotype. Pale lateral flecking on the body and face is evident in all specimens, and ventrally all paratypes are similar to the holotype. Color in life. From field notes for holotype BPBM 37730 (Fig. 8 E): “ Dorsum brown with central orange diamond margined with black, forming a large X on the two posterior margins and a chevron on the anterior two. Dorsum with scattered blue-white punctations. Face and sides of body black with scattered blue-white flecks. Rear of thighs brown and dark brown with scattered blue-white flecks. Dark-brown chevron between eyes. Upper arms orange, lower arms brown and black. Iris metallic orange. All ventral surfaces charcoal gray densely flecked with blue white. ” Field notes for paratype BPBM 37740 (Fig. 8 F) state: “ Dorsum metallic orange tan heavily maculated with dark brown, sides black flecked with blue white. Rear of thighs as dorsum, forearms copper orange. Venter dark gray flecked throughout with light blue gray. Iris dark brown with red rim around pupil. ” Call. This species calls at night from among leaf litter in lowland rainforest; the call sounds like a drawn-out, raspy snore. This description is based on 37 calls from a single frog. The call is a single, highly pulsed note lacking harmonic structure and frequency modulation (Fig. 17 A, C), averaging 1.12 ± 0.0131 s in length, and ranging from 0.97 – 1.31 s in length. Calls were delivered at intervals averaging 10.5 ± 2.4733 s apart, and ranging from 1.8 – 57.0 s apart. The number of pulses / call averaged 57.1 ± 0.6666 and varied from 50 – 66, and pulse rate average 50.9 ± 0.4259 / s and varied from 43.5 – 54.9 / s. Each note increases and decreases relatively evenly, giving the waveform a smoothly rounded, bilaterally symmetrical shape (Figs. 10 C, 17 A). Dominant frequency varied from 4460 – 5220 Hz and averaged 5080 ± 0.0219 Hz (Fig. 17 B).	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
03C087ACFF98FF87AB84FBEAFB39CA5C.taxon	etymology	Etymology. The trivial epithet is a masculine Latin adjective meaning “ lovely ”. Range. Known only from the type locality on Woodlark Island, Milne Bay Province, Papua New Guinea (Fig. 2). Almost certainly endemic but widespread across Woodlark Island. Ecological notes. This species inhabits mature lowland rainforest, with canopy heights at approximately 25 – 35 m. All specimens were collected in a gully adjacent to a small, slow-moving stream in lowland primary rainforest or on its bounding ridgeline, usually in microhabitats of steep aspect. Animals were found nocturnally active on the forest floor and were moderately common but difficult to locate because they would typically terminate calling well before they could be closely approached. Syntopic microhylids include Barygenys sp. nov., Cophixalus clapporum sp. nov., Copiula sp. nov., Genyophryne thomsoni, Hylophorbus myopicus, Mantophryne sp. nov., and Oreophryne sp. nov.	en	Kraus, Fred (2012): Papuan frogs of the genus Cophixalus (Anura: Microhylidae): new synonyms, new species, and a dichotomous key. Zootaxa 3559: 1-36, DOI: 10.5281/zenodo.282919
