A New Bent-toed Gecko (Cyrtodactylus: Squamata: Gekkonidae) from the Island of Tanahjampea, South Sulawesi, Indonesia Author Riyanto, Awal Author Hamidy, Amir Author Mcguire, Jimmy A. text Zootaxa 2018 2018-06-29 4442 1 122 136 journal article 29799 10.11646/zootaxa.4442.1.6 42d91fbf-7d4f-4439-b0e2-bdbbeffad5cd 1175-5326 1301328 1F705557-D894-42D8-951A-710C3241DAC5 Cyrtodactylus tanahjampea sp. nov. English Common name: Tanahjampea bent-toed gecko Indonesian name: Cicak Jari-lengkung Tanahjampea Figs. 2–3 Holotype . MZB .Lace.5675 ( Fig.2 ), adult male, Telkom tower road on Pasimasunggu district , Kepulauan Selayar regency, South Sulawesi Province , Tanahjampea Island , Indonesia ( 07°5’09.4” S , 120°39’29.2” E , 316 m above sea level ) collected by J. A. McGuire , Christopher J. Hayden , and Ted Townsend on 18 October 2005 , 9:07 pm. Paratype . MZB . Lace. 5671–72 , MZB . Lace. 5674, adult females, vicinity of Makminasa, Pasimasunggu district, Kepulauan Selayar regency, South Sulawesi Province , Tanahjampea Island , Indonesia ( 07o05’04.9” S , 120o40’58.3” E , 195 m above sea level ), collected by Christopher J. Hayden on 16 October 2005 11:00 pm ; MZB . Lace. 5676, adult female, vicinity of Makminasa, Pasimasunggu district, Kepulauan Selayar regency, Sulawesi Selatan province , Tanahjampea Island , Indonesia ( 07o05’20.3” S , 120o41’05.4” E , 103 m above sea level ), collected by J. A. McGuire on 16 October 2005 , 10:10 pm ; MVZ 267742, MVZ 267744 ( Fig.3 ), adult females, vicinity of Makminasa , Pasimasunggu district , Kepulauan Selayar regency, Sulawesi Selatan province , Tanahjampea Island , Indonesia ( 07o05’04.9” S , 120o40’58.3” E , 195 m above sea level ), collected by Christopher J. Hayden on October 2005, 11 :00 pm ; MVZ 267749–50 , adult males, Telkom tower road, Pasimasunggu district, Kepulauan Selayar regency, Sulawesi Selatan province , Tanahjampea Island, Indonesia ( 07o05’09.4” S , 120o39’29.2” E , 316 m above sea level ), collected by J. A. McGuire , Christopher J. Hayden, and Ted Townsend on 18 October 2005 , 8:30 pm ; MVZ 267752 , adult female, Telkom tower road, Pasimasunggu district, Kepulauan Selayar regency, Sulawesi Selatan province , Indonesia ( 07o05’12.3” S , 120o39’11.1” E , 251 m above sea level ), collected by J. A. McGuire , Christopher J. Hayden, and Ted Townsend on 18 October 2005 , 9:07 pm. Diagnosis. Cyrtodactylus tanahjampea sp. nov. is a medium size species reaching 76.1 mm and can be readily distinguished from the congeners on Sulawesi , the Moluccas , and the Lesser Sunda Islands by the following unique combination of characters: (1) brachium and antebrachium tuberculated, (2) ventrolateral folds with tubercles, (3) 20–23 irregularly aligned rows of keeled tubercles, (4) 31–34 paravertebral tubercles, (5) 29–34 ventral scales between ventrolateral folds, (6) no precloacal depression, (7) enlarged precloacofemoral scales in a continuous series, (8) males with 20–24 precloacofemoral pores in wide Ʌ-shape, (9) enlarged post precloacal scales present, (10) 19–21 fourth toe subdigital lamellae, (11) absence of enlarged transversely median subcaudals, (12) tail not prehensile, (13) tubercles extend over anterior about 71 % of original tail’s length, and (14) the original tails reach 147 % of SVL. Description of holotype . An adult male SVL 69.9 mm ; head triangular in dorsal view ( Fig.4A ), distinct from neck; HeadL 32.6% of SVL, HeadW 63.0% of HeadL, HeadH 38.5% of HeadL and HeadH 11.0% of SVL; prefrontal region concave, canthus rostralis rounded ( Fig.4B ); SnoutL 38.8% of HeadL, OD 26.5% of HeadL, EarL 9.5% of HeadL and OD 93.0% of EyeEar. Scales on snout and forehead small, rounded, granular, homogeneous; scales on snout larger than those on occipital region. Pupil vertical, supraciliaries short and ear opening oval. Rostral rectangular, height 37.9 % of width; incompletely divided dorsally by a Y-shaped shallow groove; bordered posterolaterally by first supralabials and naris, and dorsally by three postrostral scales ( Fig. 4C ); naris oval, bordered anteriorly by rostral, anterodorsally by one postrostral, posteriorly by five scales on right side and by six scales on left side, and ventrally by first supralabials; orbit separated from supralabials by single row of small lorilabial scales; 11 SuL on right and left side; 8 InfL on right and left side. Dorsal surface of head tuberculated, beginning between anterior margins of eyes, subpyramidal and gradually increasing in size toward the posterior margin of head that are subconical in form. Mental triangular, length 89.7% of width; bordered laterally by first infralabials, posteriorly by one pair of enlarged first postmentals, which are in contact medially over 40.2% of their length; second postmentals smaller than first postmentals, hexagonal on right side and pentagonal on left side, with each about half the length of first postmentals, and separated from each other by three small granular scales ( Fig.4D ); gular scales small, granular, grading to slightly larger posteriorly. Forelimbs relatively short, ArmL 20.0% of SVL; dorsal scales on forelimbs slightly larger than those of body, weakly keeled; dorsal surface of antebrachium with keeled tubercles, dorsal surface of brachium with few tubercles, multiple tubercles close to axilla becoming very few close to elbow; palmar scales flat, scales on palmar surfaces granular, juxtaposed; digits well-developed, inflected at basal interphalangeal joints, digits slightly narrower distal to inflection; subdigital scales transversely expanded along the entire length of each digit, but slightly compressed in both length and width immediately distal to interphalangeal inflection; 11 LamF1, 14 LamF2, 16 LamF3, 17 LamF4, and 15 LamF5; claws well developed, sheathed by two dorsal scales and one ventral scale; relative length of fingers IV>III>V>II>I. FIGURE 2. Holotype Cyrtodactylus tanahjampea sp. nov. (MZB.Lace.5675). Photo: A. Riyanto. Hind limbs longer than forelimbs, TibL 16.0% of SVL; dorsal surface of thigh and tibia covered by granular scales interspersed with larger, conical tubercles; scales on ventral surface of thigh and tibia smooth, flat, and subimbricate; enlarged precloacofemoral scales in continuous series; no precloacal depression; 24 precloacofemoral pores in wide Ʌ-shape arrangement; 5 rows of enlarged postprecloacal scales ( Fig.5A ); plantar scales slightly raised, juxtaposed; digits well-developed, inflected at basal metapodial-phalangeal joints, digits slightly narrower distal to inflection; subdigital scales transversely expanded along the entire length of each digit, but slightly compressed in both length and width immediately distal to interphalangeal inflection; 12 LamT1, 15 LamT2, 18 LamT3, 20 LamT4, and 18 LamT5; claws well-developed, sheathed by two dorsal scales and one ventral scale; relative length of toes IV>V> III> II> I. Body elongate, AGL 44.9% of SVL; ventrolateral folds bearing small, scattered conical tubercles; ventral region with relatively homogeneous, smooth scales; dorsal scales small, granular, with scattered irregular, relatively enlarged keeled tubercles; 20 irregular rows of dorsal tubercles at midbody; 32 PVT; 33 VentS, much larger than dorsals, smooth, round, subimbricate, largest posteriorly. Tail regenerated, TailL 75.8 mm with original part 30.8 mm and regenerated part 45 mm , subcylindrical, tapering to the tip; dorsal caudal scales flat, round, hexagonal, or pentagonal, and juxtaposed; large keeled tubercles present on lateral and dorsal surface of original part, lack on regenerated part; tubercles on caudals almost same size as tubercles on the dorsal surfaces of thighs and tibia, these tubercles almost twice larger than tubercle size on the dorsal surface of trunk; no enlarged transverse median subcaudals, on original part are small hexagonal scales almost the same size while those on the regenerated part consist of a mixture between cycloid and rectangular subcaudals of various size; three postcloacal tubercles on each side of tail base. FIGURE 3. Live photo of the paratype Cyrtodactylus tanahjampea sp. nov. (MVZ.267744). Photo: J A. McGuire. Coloration in preservative. Head olive-brown dorsally, labials and ciliaries with a pattern of light and dark mottling; a broad olive-brown stripe from nostril to anterior eye; neck with a triangular, blackish-olive, darkly bordered spot; broad, olive-brown postocular stripe to neck; throat yellow. Variation. The new species is sexually dimorphic with pores present in males that are lacking in females ( Fig.5B ). Four of the paratypes have original tails, and based on these it can be seen that caudal tubercles distally graduate to smaller size, disappearing at 71 % of the original tail length. For other detailed measurements and detailed character states for the entire type series see Table 1. Etymology. The specific name tanahjampea refers to Tanahjampea Island, the only locality at which this species is known to occur. Natural history. Specimens were collected at night in secondary forest ( Fig.6 ) and selectively logged forest on a variety of natural microhabitats including on boulders/rock-faces adjacent to running and dry stream-beds, on large tree stumps, and on the trunks of large trees. Specimens were also collected on a concrete bridge foundation and on a concrete rock wall immediately adjacent to secondary forest. Species comparisons. Cyrtodactylus tanahjampea sp. nov. can be readily distinguished from all recognized congeners occurring on Sulawesi , the Moluccas , and the Lesser Sunda Islands (detailed diagnostic characters for the species from these regions are presented in Table 2 ) except for C. halmahericus ( Mertens, 1929 ) from which it differs by possessing EPFS, PP*, FP*, and in lacking enlarged median subcaudal scales on original tails (characters that are occur on males only are donated by *). The new species is easily differentiated from C. halmahericus by having strongly keeled dorsal tubercles (vs. keeled tubercles), fewer EPFS (40–45 vs. 48–53), precloacofemoral pores* (28–34 vs. 48–53) and in lacking a precloacal depression as opposed to having a deep precloacal groove. Further, it differs from C. deveti in its much smaller body size (76.1 vs. 106 mm SVL) and in lacking enlarged transverse median subcaudal scales (as opposed to presence of enlarged transverse median subcaudal scales). The new species is also easily distinguished from C. nuaulu by having EPFS as opposed to presence of EPS, PFP* as opposed to presence of PP*, and in males lacking a precloacal depression as opposed to having a deep precloacal groove. Among the recognized species occurring in the Sulawesi region, Cyrtodactylus tanahjampea is similar to C. fumosus (data from Mecke et al. 2016b ) in body size (< 80 mm SVL), in having of precloacofemoral pores in a continuous series, and in having a blotched dorsal pattern, but differs from that species in the presence of tubercles in the ventrolateral folds, in lacking a precloacal depression, in having more DorsT (20–23 vs. 4–7), and in having fewer VentS (28–34 vs. 37–50). Further, these species are also readily differentiated in the arrangement of their chin shields, with C. tanahjampea having paired elongated primary postmentals that contact medially for 50% of their posterior sections as opposed to having ~70% contact posteromedially in C. fumosus . The new species differs from C. batik (data from Iskandar et al. 2011 ; reexamine specimens see appendix 1) in being much smaller (76.1 vs. 114.6 mm ), in having a blotched as opposed to a banded dorsal color pattern, in having fewer VentS (28–34 vs. 48–57), by the presence of EPFS (vs. lack of EPFS), PFP* (vs. poreless), and in lack enlarged transversely median subcaudal scales (as opposed to presence of enlarged transversely median subcaudals). The new species differs from C. hitchi in having tubercles on the dorsal surface of the brachium as opposed to the brachium untuberculated, EPFS as opposed to presence of EPS, PFP* as opposed to poreless both on precloacal and femoral, blotched as opposed to banded, and in lacking enlarged transverse median subcaudals as opposed to transversely median subcaudals arranged in a single row. Cyrtodactylus tanahjampea is further differentiated from C. jellesmae by having pores, EPFS, fewer VentS (28–34 vs. 40–50), and a blotched as opposed to striped dorsal pattern. It differs from C. tahuna in having EPFS and PFP* as opposed to a discontinuous series of EPS–EFS and a discontinuous series of PP*–FP*. From C. spinosus , the new species can be distinguished by its smaller body size (76.1 vs. 83.2 mm ), its lack of spine-like tubercles scattered over the dorsum, in having PFP* and in lacking a precloacal depression. The new species differs from C. wallacei by its much smaller size (76.1 vs. 114 mm ), by having PFP* as opposed to lacking pores, and in lacking enlarged transversely median subcaudal scales. The new species differs from all recognized species occuring in the Lesser Sundas in having a continuous series of precloacal and femoral pores in males. Cyrtodactylus tanahjampea differs from all species from this region except C. darmandvillei by having tubercles on the dorsal sufaces of the brachium. The new species can easily be distinguished from C. celatus (data from Kathriner et al. 2014 , Rösler & Kaiser 2016 , and Mecke et al. 2016a ) by its much larger body size (76.1 vs. 38–44 mm ), in having EPFS, in the absence of a precloacal depression as opposed to presence of a precloacal groove in males, and in having more lamellae under the fourth toe (18–23 vs. 15–18). From C. darmandvillei (data from Mecke et al. 2016), the new species differs by lacking enlarged transverse median subcaudals as opposed to having enlarged transverse median subcaudals, and in having pores in males as opposed to being poreless in both sexes. From C. gordongekkoi (data from Das 1993 , Biswas 2007 , and Mecke et al. 2016a ), the new species differs by the presence of tubercles on the dorsal surface of the brachium as opposed to absence and having precloacofemoral pores in males as opposed to being poreless in both sexes. From C. laevigatus (data from Kathriner et al. 2014 and Mecke et al. 2016a ), the new species differs in its much larger body size (76.1 vs. 38.5 mm ) and in having precloacofemoral pores in males as opposed to no pores in either sex. From C. tambora , the new species differs in its larger body size (76.1 vs. 39.4 mm ), having precloacofemoral pores in males as opposed to presence of only precloacal pores in males, and in lacking a precloacal depression as opposed to presence of a precloacal groove in males. From C. wetariensis ( Dunn, 1927 ) (data from Mecke et al. 2016a ), the new species differs by having the brachium tuberculated as opposed to lacking tubercles on the dorsal surface of the brachium and by having precloacal pores and femoral pores arranged in a continuous series in males as opposed to being discontinuously arranged. We also compare the new species to the other congeners occuring west and east of the Wallacea region. Enlarged precloacofemoral scales are present in Cyrtodactylus tanahjampea , whereas C. boreoclivus Oliver, Mumpuni, Krey & Richards, 2011 , C. consobrinus (Peters, 1871) , C. elok Dring, 1979 , C. ingeri Hikida, 1990 , C. majulah Grismer, Wood & Lim, 2012 , C. matsuii Hikida, 1990 , C. pubisulcus Inger, 1958 , C. quadrivirgatus Taylor, 1962 , C. rosichonariefi Riyanto, Grismer & Wood, 2015 , C. semiadii Riyanto, Bauer & Yudha, 2014 , C. sermowaiensis (De Rooij, 1915) , and C. yoshii Hikida, 1990 lack such a series. The subcaudal scales are not transversely enlarged in C. tanahjampea , a condition that is shared with C. klakahensis Hartmann, Mecke, Kieckbusch, Mader & Kaiser, 2016 , C. loriae ( Boulenger, 1896 ) , C. marmoratus Gray, 1831 , C. novaeguineae (Schlegel, 1837) , C. petani Riyanto, Grismer & Wood, 2015 , C. psarops Harvey, O’Connell, Barraza, Riyanto, Kurniawan & Smith, 2015 , C. rosichonariefei Riyanto, Grismer & Wood, 2015 , C. semiadii , and C. semicinctus Harvey, O’Connell, Barraza, Riyanto, Kurniawan & Smith, 2015 . In contrast, transversely enlarged subcaudals are present in C. boreoclivus , C. brevipalmatus (Smith, 1923) , C. consobrinus , C. hikidai Riyanto, 2012 , C. ingeri Hikida, 1990 , C. malayanus (de Rooij, 1915) , and C. rex Oliver, Richards, Mumpuni & Rösler, 2016 .
Museum Catalog Number MZB.Lace. MVZ
5675 Holotype 5671 5672 5674 5676 267742 267744 267749 267750 267752
Sex Male Female Female Female Female Female Female Male Male Female
Tuberculation on heađ yes yes yes yes yes yes yes yes yes yes
Tuberculation on upper arm yes yes yes yes yes yes yes yes yes yes
Tuberculation on lower arm yes yes yes yes yes yes yes yes yes yes
Tuberculation on thigh yes yes yes yes yes yes yes yes yes yes
Tuberculation on tibia yes yes yes yes yes yes yes yes yes yes
SuL (right/left) 11(11) 11(11) 10(10) 10(10) 8(8) 10(10) 8(9) 10(10) 10(10) 10(10)
InL (right/left) 8(8) 10 (10) 8(8) 9(9) 8(8) 8(9) 8(8) 9(9) 8(8) 9(8)
DorsT 20 18 18 18 18 21 22 23 21 20
PVT 32 33 31 32 31 34 33 33 32 34
VentS 33 32 34 34 31 32 29 30 30 28
Postcloacal tubercles at siđe of the base of tail 3right, 3 left 3 right, 4 left 3right, 3 left 3 left, 3 left 3right, 3 left
Ventrolateral folđs with tubercles yes yes yes yes yes yes yes yes yes yes
EPS yes yes yes yes yes yes yes yes yes yes
PFP 24 no no no no no no 23 20 no
Precloacal groove no no no no no no no no no no
Enlargeđ postcloacal scales yes yes yes yes yes yes yes yes yes yes
Enlargeđ transverse međian subcauđal scales no no no no no no no no no no
Manus 4>3>2>1>5 4>3>2>1>5 4>3>2>1>5 4>3>2>1>5 4>3>2>1>5 4>3>5>2>1 4>3>5>2>1 4>3>5>2>1 4>3>5>2>1 4>3>5>2>1
Pes 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1 4>5>3>2>1
Lam F1 11 13 12 12 12 15 14 15 13 14
Lam F2 14 15 15 13 15 17 16 17 16 16
Museum Catalog Number MZB.Lace. MVZ
5675 Holotype 5671 5672 5674 5676 267742 267744 267749 267750 267752
Lam F3 16 17 17 17 17 18 17 20 18 17
Lam F4 17 18 17 18 18 18 17 20 17 17
Lam F5 15 15 15 15 16 16 15 17 16 17
Lam T1 12 12 12 13 13 14 14 14 14 14
Lam T2 15 16 15 16 16 17 16 18 16 16
Lam T3 18 19 17 19 21 19 19 20 19 18
Lam T4 20 20 19 19 21 21 21 22 20 20
Lam T5 18 19 17 18 20 20 20 23 19 20
SVL 69.9 61.4 64.3 60.3 63.8 72.8 68.7 76.1 58.4 72.4
TL 75.8 68 77.5 88.4 67.8 83.2 91.8 88.5 79 91.4
TailL/SVL regenerateđ n/a regenerateđ n/a regenerateđ n/a 1.47 regenerateđ n/a regenerateđ n/a 1.34 regenerateđ n/a 1.35 1.26
HL/SVL 0.29 0.30 0.28 0.35 0.28 0.27 0.25 0.25 0.28 0.27
AGL/SVL 0.45 0.47 0.46 0.53 0.44 0.48 0.45 0.43 0.41 0.43
FL/SVL 0.20 0.19 0.19 0.22 0.19 0.14 0.15 0.16 0.16 0.15
TBL/SVL 0.16 0.16 0.15 0.15 0.15 0.16 0.18 0.19 0.18 0.18
HW/HL 0.63 0.31 0.61 0.58 0.63 0.63 0.72 0.69 0.63 0.65
HH/HL 0.39 0.34 0.36 0.37 0.40 0.40 0.45 0.43 0.42 0.38
ES/HL 0.30 0.26 0.26 0.24 0.26 0.32 0.31 0.33 0.30 0.29
ED/ES 0.89 0.98 1.00 0.81 1.04 0.82 1.07 0.87 0.94 1.00
ED/HL 0.27 0.26 0.26 0.20 0.27 0.26 0.34 0.28 0.28 0.29
Earl/HL 0.10 0.13 0.12 0.10 0.11 0.09 0.12 0.09 0.09 0.09
EE/ED 1.08 1.21 1.15 1.38 1.00 1.02 0.86 1.03 0.98 0.91
TABLE 2. Characters used to distinguish Cyrtodactylus species occurring on Sulawesi, the Moluccas, and the Lesser Sunda Islands. The presence of a diagnostic character is coded as ʻ1ʼ, the absence of a character is coded as ʻ0.ʼ For taxa possessing precloacofemoral scales (= scales in a continuous series; column entitled ‘7’), precloacal- and femoral scales (separated from each other by infrascales) are coded as ‘n/a’ (columns entitled ‘5’ and ‘6’). Numbers at the head of the table correspond to characters as follows: 1 = tubercles on upper arm (brachium), 2 = number of ventral scales, 3 = transversely enlarged median subcaudals in original tails, 4 = number of subdigital scales/lamellae under 4th toe, 5 = enlarged precloacal scales (the number of pores is given in parentheses and includes primordia; if pores are present in one sex only, this is indicated either by ‘ϐ’ or ʻ♀ʼ), 6 = enlarged femoral scales (the number of pores is given in parentheses and includes primordia; if pores are present in one sex only, this is indicated either by ‘ϐ’ or ʻ♀ʼ), 7 = enlarged precloacofemoral scales (the number of pores is given in parentheses and includes primordia; if pores are present in males only, this is indicated by ‘ϐ’), 8 = precloacal and femoral pores in a continuous series, 9 = expression of precloacal depression (N = no depression, G = groove, P = pit; if a depression is present in males only, this is indicated by ‘ϐ’), 10 = pattern of dorsum (bd = banded; bl = blotched; mo = mottled; pl = patternless; st = striped), 11 = data derived from literature sources are abbreviated by letters as follows: A = this publication, B = Riyanto et al . (2018), C = Riyanto et al. (2017), D = Mecke et al. (2016a), E = Mecke et al . (2016b), F = Riyanto et al . (2016), G = Rösler & Kaiser (2016), H = Kathriner et al. (2014), I = Iskandar et al . (2011), J = Oliver et al. (2009), K = Hayden et al. (2008) and L = Linkem et al. (2008). If data for a character are not available, this is indicated by a question mark (?). SVL Taxon of adults 1 2 3 4 5 6 7 8 9 10 11 (mm) tanahjampea sp. nov. 76.1 1 28–34 0 18–23 n/a n/a 1 (28–34,ϐ) 1 N bl A batik 103–113 1 48–57 1 24–27 1 0 0 0 N bd A, L celatus 38‾44 0 34–42 0 15–18 1 (4, ϐ) 0 0 0 G (ϐ) bl D, G, H darmandvillei 80–82 1 34– 36 1 23 ‾24 n/a n/a 1 0 N bl D deveti 101–106 0 40– 49 1 25–28 n/a n/a 1(29–31, ϐ) 0 N bd A fumosus 57–78 0 37–50 0 17–23 n/a (10–11, ϐ) n/a (3, ϐ) 1 0 G (ϐ) bl E gordongekkoi 71–73 0 30 0 22–23 n/a n/a 1 0 N bl D halmahericus 75–78 1 39– 42 0 22 n /a n/a 1 (48–53, ϐ) 1 G (ϐ) bl A hitchi 62–79 0 39–45 1 18–21 1 0 0 0 N bd F jellesmae 58–70 0/1 40–54 0 16–23 1 0 0 0 N bl A, D laevigatus 38–47 0 30–34 0 10–15? 0/1? 0 N mo, pl D, H nuaulu 77–88 1 48–55 0 17–20 1 (6, ϐ) 0 0 0 G (ϐ) bl, st J spinosus 83 1 38–44 0 19–21 1 (12–13, ϐ) 1 0 0 P (ϐ) bd A, I tahuna 79 1 49–50 0 20–24 1 (14, ϐ) 1 (5, ϐ) 0 0 P (ϐ) bl B tambora 39–47 0 40 0 16–17 n/a (5–6, ϐ) n/a 1 0 G (ϐ) pl C wallacei 92–114 1 45–49 1 17–25 1 0 0 0 N bd, bl A, K wetarisensis 58–67 0 36–38 0 20–22 n/a (11, ϐ) n/a (12–16, ϐ) 1 0 N bl D