Megophrys vegrandis, Mahony, Stephen, Teeling, Emma C. & Biju, S. D., 2013

Mahony, Stephen, Teeling, Emma C. & Biju, S. D., 2013, Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region, Zootaxa 3722 (2), pp. 143-169 : 158-163

Megophrys vegrandis sp. nov.

( Figures 3 , 8 & 9 ; Table 1 )

“ Xenophrys boettgeri ?”: Athreya (2006: 100)

“ Xenophrys cf. boettgeri ”: Athreya (2006: 139, 143) “ Xenophrys boettgeri ”: Mathew and Sen (2010: 54, fig. 89)

Holotype. Adult male (ZSI A 11605 ), from a tributary of the Sessa Nadi (river), Sessa village (27°06.067’N 92°31.642’E, 1,110 m asl), Bhalukpong Forest Division, West Kameng district, Arunachal Pradesh state, northeast India, collected by Systematics Lab members, 8 August 2009.

Paratypes by original designation. Two adult males (ZSI A 11604, BNHS 5597), collected with holotype.

Referred specimen. One adult male (SDB.DU 2009.1274), and one metamorphosed juvenile (SDB.DU 2009.1275), collected with holotype.

Diagnosis. Megophrys vegrandis sp. nov. is diagnosable from geographically relevant congeners by the following combination of characters: small-sized, slender species, adult male size amongst the smallest of all known Megophrys species, SVL 27.5–30.6 mm (N=4); tympanum circular, not concealed by supratympanic ridge; vomerine ridges and vomerine teeth absent; dorsolateral folds extending for approximately 20–40 percent of trunk length; length of finger I 50 –65 percent of length of finger II; webbing between toes rudimentary; lateral fringes on toes present; nuptial pads, subarticular tubercles and protruding projection posterior to cloaca of males absent; groin and ventral thigh colouration not contrasting with surrounding regions on males; ventral surface of thighs mottled and blotched with dark brown.

Comparisons. Megophrys vegrandis sp. nov. differs from the following large species (in parenthesis) by considerably smaller adult-male size, SVL 27.5–30.6 mm, N=4 (vs. adult-male SVL> 53 mm: M. gigantica , N=6; M. glandulosa , N=10; M. jingdongensis , N=2; M. lekaguli , N=8; M. major , N=8; M. medogensis , N=16; M. omeimontis , N=10; M. robusta , N=4; M. spinata , N=2); from the following small to medium-sized species (in parenthesis) by smaller adult-male size, SVL 27.5–30.6 mm, N=4 (vs. Megophrys ancrae sp. nov. SVL 39.1–45.0 mm, N=8; M. binchuanensis SVL 32.0–36.0 mm, N=4; M. binlingensis SVL 45.1–51.0 mm, N=3; M. daweimontis SVL 34–37 mm, N=18; M. megacephala SVL 45.9–53.4 mm, N=7; M. minor SVL 32.2–40.5 mm, N=15; M. pachyproctus SVL 35.3–36.2 mm, N=2; M. palpebralespinosa SVL 36.2–38.0 mm, N=2; M. parva SVL 33.9–41.4 mm, N=4; Megophrys oropedion sp. nov. male SVL 32.8–39.2 mm, N=7; M. takensis SVL 47.3–53.0 mm, N=3; M. wawuensis SVL 34.4–42.8 mm, N=4; M. zhangi SVL 32.5–37.2 mm, N=3), further from Megophrys ancrae sp. nov., M. megacephala , M. minor , Megophrys oropedion sp. nov., M. parva , and M. takensis by lateral fringes on toes present (vs. absent), further from M. binchuanensis by subarticular tubercles absent (vs. present), groin colouration not contrasting to surrounding regions on males (vs. red on males), further from M. binlingensis by vomerine ridges absent (vs. present, slender), further from M. daweimontis and M. wawuensis by FIL 50–65% FIIL (vs. FIIL<FIL), further from M. pachyproctus by protruding projection posterior to cloaca of male absent (vs. present), FIL 50–65% FIIL (vs. FIL subequal to FIIL), tympanum circular (vs. oval), and lateral fringes on toes present (vs. absent), further from M. palpebralespinosa by webbing between toes rudimentary (vs. approx. half digit length), further from M. zhangi by ventral thighs mottled and blotched with dark brown (vs. immaculate); from M. nankiangensis and M. shapingensis by presence of clearly defined tympanum (vs. tympanum concealed by supratympanic fold); from M. wushanensis by subarticular tubercles absent (vs. present), groin colouration not contrasting with surrounding regions on males (vs. red on males); from M. serchhipii and M. zunhebotoensis by lateral fringes on toes present (vs. absent), and vomerine teeth absent (vs. present); from M. wuliangshanensis by lateral fringes on toes present (vs. absent), dorsolateral folds extend approximately 20–40 percent body length (vs. most of body length), ventral thigh colouration not contrasting with surrounding regions on males (vs. red on males).

Holotype description (measurements in mm). Mature male (SVL 29.2) ( Figure 8 , 9 ). Head small (HW 10.2, HL 10.2, IFE 5.2, IBE 9.0), as wide as long; snout rounded in dorsal view, obtusely protruding in lateral view, without rostral appendage ( Figure 8C ); loreal region vertical and concave; canthus rostralis blunt; dorsal region of snout slightly concave ( Figure 9C ); eye (EL 3.6) twice as long as maximum tympanum diameter ( TYD 1.8), and shorter than snout (SL 4.0); eye-tympanum distance (TYE 1.5) shorter than maximum tympanum diameter; tympanum circular, with upper margin concealed by supratympanic ridge ( Figure 8C ); pupil in life oval, horizontally orientated when dilated; nostril oriented laterally, situated mid-way between eye and snout (EN 2.0, SN 2.0); internarial distance (IN 3.3) subequal to eyelid width (UEW 3.3), and narrower than narrowest point between upper eyelids (IUE 3.7); pineal ocellus not visible externally; vomerine ridges and vomerine teeth absent; tongue moderately large, weakly notched posteriorly, with no medial lingual process.

Forelimbs moderately long and thin, forearm (FAL 7.6) slightly enlarged relative to upper forelimb, shorter than hand (HAL 9.4); fingers short and broadly flattened but without lateral fringes ( Figure 8D ), finger length formula I<II=IV<III (FIL 2.6, FIIL 4.6, FIIIL 6.3, FIVL 4.6); interdigital webbing, subarticular and supernumerary tubercles absent; thenar and palmar tubercles barely distinguishable; finger tips slightly expanded and rounded with oval pads; terminal grooves absent. Hindlimbs relatively long and thin ( Figure 8A ), shanks overlap when thighs are held at right angle to body, thigh (TL 15.0) shorter than shank (SHL 15.8), and longer than foot (FOL 13.6); toes short and flattened with broad lateral fringes that extend to tips of all digits ( Figure 8E ); relative toe lengths I<II<V=III<IV; tips not dilated, but with distinct pads; terminal groves absent; base of toes with rudimentary webbing ( Figure 8E ); outer metatarsal tubercle, subarticular and supernumerary tubercles absent; inner metatarsal tubercle present but barely distinguishable; ridge of callous tissue present on ventral surface of all digits.

Skin of dorsal and ventral surfaces of head, body and limbs primarily smooth; flanks with small scattered tubercles ( Figure 9A ); tympanum smooth, borders slightly raised, surrounding area posterior to eye granular ( Figure 8C ); small tubercle present on outer edge of upper eyelid; supratympanic fold narrow without widening posteriorly, extends from orbit and curves down broadly through upper border of tympanum terminating above shoulder ( Figure 8C ); short thin dorsolateral fold extending from behind supratympanic fold to approximately one third distance to groin; weak “V”-shaped parietoscapular ridge present, its two sides extending posteriorly from above tympanum and meeting medially beyond level of axilla; second inverted “V”-shaped ridge present on mid-dorsum ( Figure 8A ); pectoral glands small and raised slightly, positioned on level with axilla ( Figure 8B ); femoral glands small slightly raised, positioned sub-equally distant from knee and cloaca. Skin asperities absent on all surfaces.

Colour in preservative ( Figure 8 ): Dorsal and lateral surfaces of head and body primarily light grey-brown; brown triangular marking with light central blotch between eyes, and large brown, roughly “X”-shaped blotch present on dorsal surface of body; lower borders of dorsolateral and supratympanic folds, and flank tubercles dark brown; front of snout mottled with dark brown; wide oblique dark brown bar below eye and dark brown blotch covers tympanum; lateral surfaces of forearm and dorsal surface of hands and feet with dark brown spots and speckles; forelimbs brown above with feint dark brown transverse stripes on forearm; dorsal surface of upper forelimb pale relative to remaining dorsal trunk colour; dorsal surface of hindlimbs light grey-brown with oblique dark brown crossbars. Gular region, chest and anterior part of abdomen primarily greyish-yellow with barely discernible mottling and lighter patches along margin of lower mandibles; light colouration of chest darkens slightly posteriorly on abdomen which is mottled with light brown; broad dark brown row of blotches present on ventrolateral surfaces of abdomen; ventral surfaces of thighs and shanks with light and dark brown mottling; ventral surfaces of tarsus and feet dark grey-brown; area surrounding cloaca and posterior surfaces of thighs dark brown; ventral surface of hands grey-brown; pectoral and femoral glands lighter than surrounding area. Colour in life ( Figure 9 ): Dorsum of head, body, hindlimbs and forearm light olive brown; above mentioned dorsal markings of head, body and limbs mid-brown; lateral surfaces of body and hindlimbs light mauve-grey; dorsal surfaces of upper forelimbs plain yellow; all dorsal and lateral tubercles, ridges and folds on head, body and limbs, orange; ventral surfaces of tips of fingers I and II and tips of all toes, orange. Iris metallic yellowish-orange.

Variation. See Table 1 for morphometric characters of four adult males. Dorsal markings of paratypes agree with holotype, however overall colouration is darker. The length of the dorsolateral folds on paratypes and the adult referred specimen varies from approximately 20–40 percent trunk length. The supratympanic fold does not obscure the upper border of the tympanum on ZSI A 11604 and SDB.DU 2009.1274. Pineal body externally visible on ZSI A 11604 and BNHS 5597. BNHS 5597 has very sparse, small clear asperities on the dorsal folds of the body and hindlimbs and a few on the outer margin of the lower jaw, whereas SDB.DU 2009.1274 has sparse asperities on the posterior part of the dorsum only. ZSI A 11604, BNHS 5597, and SDB.DU 2009.1274 possess a short ridge on the upper eyelid as opposed to the tubercle present on the holotype. Dark blotches on the ventral surfaces of the body and limbs vary between specimens with respect to density and shade. The dorsal surface of the upper forelimbs vary between yellow and light brown in life. On the metamorph (SDB.DU 2009.1275), the dorsal and lateral surfaces of the head and body are mottled brown with faintly darker brown triangle between the eyes. Limbs greyish-yellow with feint brown crossbars. Lateral head markings like on the adults. Throat and abdomen pale grey with dark brown longitudinal stripe on each side of the lower flanks, on posterior thighs across the cloacal region and on the ventral tarsus. Pectoral and femoral glands visible as white spots.

Secondary sexual characters. Males without nuptial pads on fingers; external vocal sac indistinct; internal vocal slits present near the rear of the lower mandible; protruding fleshy projection posterior to cloaca absent. Females currently unknown.

Etymology. The specific epithet “ vegrandis ” is a Latin word meaning “diminutive” or “tiny” with reference to the small adult male size of this species.

Distribution. The species is known from the type locality near Sessa village, in West Kameng district, in western Arunachal Pradesh state. We provisionally assign another population to this species from Sessni (27°02.844’N 92°25.086’E, 1,250 m asl), Eaglenest Wildlife Sanctuary, approximately 12.4 km west of the type locality (Athreya 2006: see remarks below). It is likely that the species is more widespread in suitable habitat at similar altitudes in western Arunachal Pradesh.

Habitat and natural history. The primary habitat at the type locality is referred to as subtropical forest (Athreya 2006). The type specimens were collected from the banks of a shallow stream (max. depth 10 cm) that flowed through an area of dense bushy vegetation surrounded by disturbed, selectively logged forest ( Figure 4A ). The vegetation formed a low complete canopy (1–2 m height) over the stream. The stream bed consisted of gravel and rocks, with leaf litter gathered in snags and side pools. No individuals of this species were found (or heard) from the banks of the larger Sessa Nadi river. All males were found calling from atop large boulders bordering the stream where individuals were spaced a minimum of three meters apart along the stream bank. Calling began at dusk on a dry cloudless night. The newly metamorphosed specimen was collected on the gravel bank of the stream. The vigor of male vocalisations indicates that the breeding season includes early August.

Remarks. Athreya (2006) provided a selection of photographs, some basic measurements and observations of living individuals from Sessni, Eaglenest Wildlife Sanctuary which they identified as “ Xenophrys cf. boettgeri ” (pg. 143; and pg. 100, plate 2 as “ Xenophrys boettgeri ?”) and which appear to be conspecific with Megophrys vegrandis sp. nov. Mathew and Sen (2010: 54, fig. 89) also show a figure of a Megophrys vegrandis sp. nov. individual from Sessni, and provided a brief description as “ Xenophrys boettgeri ”, which they claim to be found in Arunachal Pradesh and Sikkim. Several of the characters they provide in the description (“tympanum...as large as eye”; “TTA [tibiotarsal articulation] reaches tip of snout”), however, neither match M. boettgeri s.s., or the photographed Megophrys vegrandis sp. nov, thus it is unclear what species they are basing their description. Morphologically, M. boettgeri and Megophrys vegrandis sp. nov. differ considerably (e.g., SVL of adult males 34.5–37.8 mm in M. boettgeri [vs. 27.5–30.6 mm in Megophrys vegrandis ], lateral fringes on toes absent [vs. present], subarticular tubercles present on fingers [vs. absent], dorsolateral folds absent [vs. present anteriorly]) and thus these species can hardly be confused. Many other reports of M. boettgeri exist in the Indian literature primarily based on a misunderstanding that originated with Annandale (1917). Annandale regarded tadpoles of M. boettgeri (= M. brachykolos Inger & Romer—fide Inger & Romer 1961) from Hong Kong, to be superficially similar to tadpoles from Arunachal Pradesh which he had earlier assigned to Megalophrys kempii Annandale (Annandale 1912) . He conjectured that Megalophrys kempii may therefore be synonymous with M. boettgeri , strangely disregarding the clear morphological differences between the adults of both species. Many subsequent authors listed M. boettgeri as present in Arunachal Pradesh based directly or indirectly on this weakly supported synonymy and not based on subsequent specimen collections (i.e., Bordoloi et al. 2000; Chanda 1990, 1992, 1994, 2002; Pillai & Chanda 1976; Sarkar & Ray 2006). Delorme et al. (2006) referred Megalophrys kempii to the rhacophorid genus Philautus Gistel based on a morphological examination of the holotype. The tadpoles that Annandale (1912, 1917) provisionally assigned to “ P.” kempii are therefore referable to an unknown species of Megophrys , thus rendering inaccurate all of the aforementioned publications that list M. boettgeri from Arunachal Pradesh.

Dutta (1997) recognised “ P.” kempii (as Megophrys kempii ) as valid, but unusually gives Sikkim as the range of M. boettgeri in India, and states that “After, Boulenger (1894) the species has not been recorded from India ”. The reference for this citation is not in the bibliography section, and here is regarded questionable as it predates the description of M. boettgeri (Boulenger 1899) , and furthermore Sikkim is not included in the distribution of this species in Boulenger’s subsequent review of the group (Boulenger 1908). Pawar & Birand (2001) reported sighting M. cf. boettgeri from Mouling National Park in Arunachal Pradesh, from an unspecified source from Bhalpakram National Park in Meghalaya, as well as other localities in Meghalaya and Nagaland. Pawar & Birand (2001) recognised that further research was required to confirm the identity of the Indian populations of M. boettgeri . In the absence of referred voucher specimens, images, or any verifiable evidence of this species from their listed localities, we regard their report as anecdotal, and most likely misidentifications of another/other species.

In summary, all literature reports of M. boettgeri from India were either based on Megophrys vegrandis sp. nov. (see above chresonymy), “ Philautus ” kempii (Bordoloi et al. 2000; Chanda 1990, 1992, 1994, 2002; Pillai & Chanda 1976; Sarkar & Ray 2006), or anecdotal records (Dutta 1997; Pawar & Birand 2001; Mathew & Sen 2010). No Indian voucher specimens referable to M. boettgeri have been specifically identified in literature, nor are any currently present in major museums, e.g., BMNH, ZSI, SDB.DU (Mahony pers. obs.). Outside India, the closest reported locality for M. boettgeri is from eastern Guangxi Province, China, ca. 1,400 km east of the eastern-most border of northeast India (Fei et al. 2009), thus in the absence of any solid evidence to support the presence of this species in India, it should no longer be included in the Indian amphibian checklists. The geographical distribution of M. boettgeri is here restricted to eastern China (Fei et al. 2009), where based on current understanding it should be considered endemic.

(ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) (ind) 2.2 (ind) 2.2 (ind) 2.2 (ind) 2.5 (ind) (ind) (ind) (ind)

Figures

FIGURE 3. Map of northeastern India (shaded grey) and adjoining countries (white) showing the collection localities of the three new species. Blue spots— Megophrys ancrae sp. nov., orange spots— Megophrys oropedion sp. nov., green spot— Megophrys vegrandis sp. nov.

FIGURE 8. A – E. Holotype of Megophrys vegrandis sp. nov. (ZSI A 11605, adult male) in preservation. A. dorsal view, B. ventral view, C. profile view of head, D. ventral view of left hand, E. ventral view of left foot.

FIGURE 9. A – D. Holotype of Megophrys vegrandis sp. nov. (ZSI A 11605, adult male) in life. A. dorsolateral view, B. lateral view, C. head viewed from the front, D. ventral view.

FIGURE 4. A – D. Habitat images at the collection localities of the three new species: A. for Megophrys vegrandis sp. nov. showing the point where the side stream meets the Sessa river (dark opening in vegetation, centre frame on the bank of the river), B. for Megophrys oropedion sp. nov. showing the type locality showing Um Risa stream, in Malki Forest, C. for Megophrys oropedion sp. nov. showing a collection stream in Mawphlang Sacred Forest, D. for Megophrys ancrae sp. nov. showing the stream at the type locality in Deban, Namdapha National Park and Tiger Reserve.

Tables

TABLE 1. Morphometric data for the type series and adult referred specimens of Megophrys ancrae sp. nov., Megophrys oropedion sp. nov., and Megophrys vegrandis sp.. Measurements in mm. HT. holotype, PT. paratype, RS. referred specimen, (ind). indistinct, * measurements taken on left side.

sp No. Status	Megophrys ancrae sp. nov. ZSI A ZSI A BNHS BNHS 11606 11607 5598 5600 HT PT PT PT	BNHS 5601 PT	BNHS 5602 PT	SDB2009.SDB 2009. BNHS 727 730 5599 RS RS PT	ZSI A 11608 PT	ZSI A 11601 HT	Megophrys oropedion sp. nov. ZSI A BNHS BNHS SDB 2009. SDB 2009 11602 5595 5596 300.301 PT PT PT RS RS	SDB 2009. 1164 RS	ZSI A 11603 PT	SDB 2009. 299 RS	ZSI A 11605 HT	Megophrys sp. nov. ZSI A BNHS 11604 5597 PT PT	vegrandis SDB 2009. 1274 RS
Sex SVL HW	M M M M 45.3 40.2 40.2 43.7 14.6 13.6 13.3 13.3	M 39.1 13.1	M 43.9 13.7	M M F 42.5 43.0 48.9 15.2 14.6 15.3	F 38.0 12.0	M 32.8 12.0	M M M M M 33.9 33.5 33.2 35.0 39.2 12.2 12.9 12.2 12.2 14.6	M 38 12.9	F 44.1 15.0	F 48.7 17.6	M 29.2 10.2	M M 30.6 27.5 10.2 9.6	M 28.9 10.3
HL IFE IBE	15.6 13.8 14.3 14.0 7.5 7.5 7.6 6.6 12.6 12.5 11.8 11.8	13.4 7.0 11.4	14.4 7.0 12.2	15.9 15.1 15.8 8.1 7.5 8.4 12.1 12.0 12.8	12.4 6.5 10.1	12.0 6.0 10.1	12.0 12.3 12.1 12.0 13.6 5.6 6.6 5.8 6.3 6.8 10.0 10.7 9.9 9.5 11.7	12.6 6.2 10.4	14.5 7.6 12.0	17.0 8.5 13.8	10.2 5.2 9.0	10.2 9.9 5.3 5.1 9.1 8.6	10.3 5.5 8.8
EL TYD TYE	5.4 4.9 5.3 5.3 2.7 2.9 2.7 2.9 2.7 2.6 2.3 2.0	4.6 2.8 2.4	5.1 2.7 2.7	5.7 5.3 6.1 3.6 3.3 3.0 2.7 2.3 3.0	4.4 2.5 2.1	4.4 2.7 1.8	4.1 4.1 4.2 4.1 5.2 2.5 2.5 2.4 2.2 3.3 1.8 2.2 1.9 1.8 2.8	4.6 3.0 2.2	5.0 3.1 2.7	5.8 4.0 2.9	3.6 1.8 1.5	3.7 3.4 1.9 1.5 1.5 1.4	3.6 2.0 1.2
SL EN SN	5.5 5.0 5.9 5.0 2.5 2.1 2.8 1.8 3.4 2.9 3.0 2.9	5.8 2.5 3.0	4.9 2.3 2.9	6.1 5.9 5.4 2.7 2.6 2.6 3.3 3.2 3.2	4.5 2.3 2.5	4.5 2.2 2.3	4.2 4.8 4.5 4.5 5.2 2.0 2.1 1.9 1.9 2.0 2.3 2.9 2.3 2.6 2.9	4.8 2.1 2.8	5.9 2.9 3.0	6.0 2.6 3.5	4.0 2.0 2.0	4.0 4.0 2.1 2.0 2.1 2.0	3.8 1.8 1.8
IUE IN UEW	4.2 4.6 4.5 3.7 4.7 4.8 4.2 4.4 4.6 4.6 4.4 4.3	4.0 4.3 4.0	4.3 4.2 4.0	4.7 4.3 4.4 4.6 5.2 4.6 4.6 4.6 5.1	3.5 3.6 3.6	3.4 3.8 3.8	3.5 3.5 3.4 3.7 3.5 3.9 4.1 3.4 4.1 4.4 3.9 3.9 3.7 3.7 3.8	3.6 4.3 4.0	4.2 4.6 4.6	4.6 5.3 5.2	3.7 3.3 3.3	3.2 3.1 3.4 3.0 3.6 3.0	3.0 3.1 3.5
FAL HAL FIL	10.3 9.6 11.2 10.7 13.1 11.3 11.7 11.4 5.2 4.2 4.5 4.8	9.8 11.2 4.5	10.6 12.6 4.9	10.8 11.3 11.6 12.0 12.9 12.8 5.4 6.0 5.1	9.3 10.2 4.3	7.6 10.1 3.9	7.8 9.0 8.1 8.2 8.7* 9.4 10.3 10.4 10.5 11.9* 4.1 4.1 4.0 4.1 5.6*	9.2 10.4 3.6	9.4 11.9 4.6	10.2 12.8 4.7	7.6 9.4 2.6	7.6 7.3 9.4 9.0 2.8 2.6	7.3 9.1 2.6
FIIL FIIIL FIVL	5.7 5.5 5.0 5.3 9.7 8.2 8.9 8.4 6.8 5.7 5.4 5.7	4.7 8.3 5.2	6.0 9.2 6.8	5.7 6.1 5.4 9.8 9.1 10.0 6.7 6.5 6.9	4.6 8.2 5.6	4.0 6.3 4.0	4.1 5.0 4.1 4.1 4.4* 6.9 7.2 7.1 7.6 7.8* 4.0 4.6 4.7 4.6 4.9*	4.2 7.0 4.5	5.6 7.4 5.0	5.8 9.0 6.0	4.6 6.3 4.6	4.3 4.2 6.1 6.0 4.3 4.1	4.0 5.8 4.1
SHL TL TFOL	22.4 19.3 21.2 20.1 21.8 18.5 19.5 18.0 30.3 27.5 29.3 27.9	18.5 16.6 26.4	20.5 18.0 28.8	23.5 22.7 24.0 21.6 20.7 21.7 32.9 30.4 33.6	18.7 15.6 24.0	14.4 13.9 21.9	14.7 16.2 14.9 15.4 18.1 14.7 16.2 14.0 14.6 16.7 22.5 24.9 23.2 22.9 27.3	16.7 16.4 25.1	18.8 17.9 28.7	20.3 19.6 31.2	15.8 15.0 21.6	15.0 14.3 14.3 13.5 21.1 20.5	15.5 14.6 21.5
FOL	19.7 18.0 18.3 17.9	16.9	18.5	21.0 19.4 21.4	15.8	14.5	15.1 16.0 15.4 14.8 17.9	16.4	18.4	20.1	13.6	13.8 13.2	13.5