Hime surrubea, Gomon, Martin F. & Struthers, Carl D., 2015

Gomon, Martin F. & Struthers, Carl D., 2015, Three new species of the Indo-Pacific fish genus Hime (Aulopidae, Aulopiformes), all resembling the type species H. japonica (Günther 1877), Zootaxa 4044 (3), pp. 371-390 : 373-380

Hime surrubea sp. nov.

Current English vernacular name: Japanese Thread-sail Fish Proposed new English vernacular name: Rosy Flagfin Figs 1–3 ; Tbls 1–3

Hime japonicus (non Günther, 1877): Strasburg, 1966 (in part): 91, fig. 1 (description, central Hawaiian Islands, USNM 198224, 219 mm SL); Tinker, 1978: 90, text figure (description); Borets, 1986: 9–10, Table 1 (Hawaiian Islands, Equator Seamount, 29° 42' N, 179° 22' E, 85–160 m); Parin & Kotlyar, 1989 (in part): 412 (ZIL 39717, 191 mm SL, Maro Reef, 25° 28' N, 170° 34' W, 275 m; ZIL 47257, 238 mm SL, 28° 34' N, 176° 32' W, 85 m); Chave & Mundy, 1994: table 1 (leeward and windward Hawaii, Cross Seamount, 343–510 m, sand bottom); Thompson, 1998 (in part): 44 (comparative data); Mundy, 2005 (in part): 192–193 (checklist).

Undescribed species of Aulopus: Struhsaker, 1973: 49 ; Randall, 1976: 51; Randall, 1981: 211 (new species); Tinker, 1982: appendix XXXIV.

Holotype. BPBM 25117 (156, male) Hawaii, Maui, Pailolo Channel, 21° 02' N, 156° 45' W, 241–254 m, R/V "Townsend Cromwell", TC33-15, 31 October 1967.

Paratypes. 26 specimens, 90.0– 173 mm SL: AMS I.37811-001 (153, male) Hawaii, 21° 01' 36" N, 156° 43' 24" W, 20° 57' 12" N, 156° 42' 24" W, 212–216 m, bottom trawl, R/V "Townsend Cromwell", TC 40-52, 17 November 1968; AMS I.37813-001 (140, female) Hawaii, 21° 03' 12" N, 156° 45' W, 20° 01' 06" N, 156° 51' W, 229 m, bottom trawl, R/V “Townsend Cromwell”, TC 40-95, 26 November 1968; BPBM 24177 (2, 142–142, male, female) Hawaii, Pailolo Channel, 21° 01' N, 156° 45' W, 210 m, R/V "Townsend Cromwell", TC40-47, 16 November 1968; BPBM 41170 (4, 90.0–135, 3 males, 1 female) same collection data as holotype BPBM 25117; BPBM 25124 (2, 119–166, females) Hawaii, Pailolo Channel, 21° 01' N, 156° 43' W, 212 m, R/V "Townsend Cromwell", TC40-52, 17 November 1968; BPBM 25130 (4, 127–170, males) Hawaii, Pailolo Channel, 21° 03' N, 156° 43' W, 212 m, R/V "Townsend Cromwell", TC40-69, 20 November 1968; BPBM 25114 (2, 119–163, males) Hawaii, Maui, Pailolo Channel, 21° 02' N, 156° 45' W, 241–254 m, R/V "Townsend Cromwell", TC 33-15, 31 October 1967; NMNZ P.056101 (153, male) same collection data as paratypes BPBM 25130; NMNZ P.056102 (127, female) same collection data as paratypes USNM 384078; NMV A31122 -001 (132, female) same collection data as paratype AMS I.37813-001; NMV A31123 -001 (148, male) same collection data as paratype AMS I.37811- 001; USNM 384078 (5, 122–173, females) Hawaii, Pailolo Channel, Molokai, 21° 7.0' N, 156° 46.5' W, 229–243 m, R/V "Townsend Cromwell", TC 40-55, 18 November 1968; USNM 358123 (141, male) Hawaii, Pailolo Channel, 21° 00' 24'' N, 156° 46' 54'' W, 234 m, R/V "Townsend Cromwell", TC40-62, 19 November 1968.

Other material. AMS I.37809-001 (112) Hawaii, 20° 59' 30" N, 156° 47' 24" W, 21° 00' 18" N, 156° 45' 30" W, 223–238 m, bottom trawl, R/V "Townsend Cromwell", TC 33-8, 13 November 1967; AMS I.37796-001 (91) Hawaii, 21° 01' 24" N, 156° 46' 30" W, 21° 00' 24" N, 156° 45' 24" W, 210–241 m, bottom trawl, R/V "Townsend Cromwell", TC 33-50, 13 November 1967; AMS I.37811-003 (3, 148–163, males) same collection data as paratype AMS I.37811-001; AMS I.37793-001 (27, 114–161) Hawaii, 21° 03' 06" N, 156° 45' 24" W, 21° 07' " N, 156° 50' 18" W, 229 m, bottom trawl, R/V "Townsend Cromwell", TC 40-63, 19 November 1968; AMS I.37807-001 (36, 89–155) Hawaii, 21° 03' 06" N, 156° 45' W, 21° 01' 06" N, 156° 50' 36" W, 210–240 m, bottom trawl, R/V "Townsend Cromwell", TC 40-72, 21 November 1968; AMS I.37813-002 (30, 112–155, females) same collection data as paratype AMS I.37813-001; BPBM 25119 (2, 117–162, females) Hawaii, Pailolo Channel, 21° 01' N, 156° 44' W, 220 m, R/V "Townsend Cromwell", TC40-42, 16 November 1968; BPBM 41179 (147, male, aberrant dorsal fin) ex BPBM 24177; USNM 431306 (17, 121–154, females) same collection data as paratypes USNM 384078; USNM 358119 (143, female) Hawaii, Pailolo Channel, between Molokai and Maui and off South Coast of Lanai, 21° 3.10' N, 156° 44.04' W, 245 m, R/V "Townsend Cromwell", TC 33-18, 1 November 1967; USNM 198224 (219, female) Hawaii, between Lanai and Kahoolawe, handline, K. Sakamoto, 15 April 1963.

Diagnosis. Dorsal fin with 16 or 17 rays, of moderate height, no rays filamentous in either sex, third and fourth rays longest, only slightly longer than shortest rays at middle of fin, 16.9–23.1% SL in males and females, distal margin nearly straight in both sexes, only slightly concave; caudal peduncle length 13.4–18.7% SL; distance from anus to anal fin origin 1.5–2.2 in distance from pelvic fin origin to anus; head length 30.1–33.3% SL; orbital diameter 10.1–11.8% SL; interorbital width 4.0–5.5% SL; snout length 7.6–9.0% SL; upper jaw length 14.5–16.2% SL; pectoral fin length 19.1–22.1% SL; pelvic fin length 23.7–29.5% SL; scales between anus and anal fin origin 9–11; reddish overall with deep red saddles.

Description. (See Tables 1 –3 for further meristic and morphometric details) Dorsal fin rays 16 (16 or 17, usually 16); anal fin rays 10 (9 or 10); caudal fin rays 10 + 19 + 9 (9–11 + 19 + 8–9); pectoral fin rays 11; pelvic fin rays 9; vertebrae 27+16 = 43 (26–28 + 15–17 = 43); lateral line scales 42 (42 or 43, usually 42) + 1; scales above lateral line 4.5; scales below lateral line 4.5; predorsal scales 13 (11–14); gill rakers 4 + 14 = 18 (4 or 5 + 14–16 = 18–21); no pyloric caeca (based on USNM 384078, 167 mm SL, female).

Hime spp. nov.

H. surrubea sp. nov. H. capitonis sp. nov. H. caudizoma sp. nov.

Holotype Paratypes Holotype Paratypes Holotype Paratypes

n = 26 n = 19 n = 7

n range mean±s.d. n range mean±s.d. n range mean±s.d.

Fin rays

Body elongate, moderately thick, tapering slightly from eyes to posterior end of dorsal fin; dorsal profile of head and nape mostly straight, inclined to dorsal fin origin, sloping more gradually to base of tail with straight dorsal profile; ventral profile of head and body mostly straight; caudal peduncle of moderate length and moderately narrow; body deepest at anterior part of dorsal fin, depth at dorsal fin origin 20.4 (18.2–22.0) % SL, depth at anal fin origin 10.6 (10.0–13.6) % SL; anus closer to anal fin origin than pelvic fin origin, distance from anus to anal fin origin 1.7 (1.5–2.2) in distance from pelvic fin origin to anus. Head of moderate size, length 30.9 (30.1–33.3) % SL, bluntly pointed; snout moderately short, distinctly shorter than eye diameter, broadly rounded from above; eye large, orbital diameter 10.2 (10.1–11.8) % SL and 33.0 (31.2–38.1) % HL, slightly larger in females than males (10.3–11.8, mean 11.1 vs 10.1–11.6, mean 10.7, % SL respectively); top of eye at or extending slightly above dorsal profile of head; bony interorbital distinctly concave, narrow, interorbital width 5.0 (4.0–5.5) % SL and 16.1 (12.3–16.5) % HL; postorbital nearly half head length. Anterior nostril small, about midway between tip of snout and orbit, posterior nostril much larger, about three or four times diameter of anterior nostril, long narrow, lanceolate skin flap with irregular margin based on septum separating nostrils. Edge of preopercle smooth, posterior margin straight, angle blunt; opercular margin nearly straight; without well-developed membranous lobe; branchiostegal membranes overlapping ventrally, free from isthmus. Mouth of moderate size, terminal, lower jaw projecting little in advance of upper, profile of upper jaw with distinct notch at symphysis when viewed from above; posterior end of maxilla broad, posterior margin nearly straight, extending just past vertical through centre of eye, dorsal margin covering lower half of slender suborbital, separated from eye by narrow strip of skin with mouth closed; upper lip narrow, not completely covering tooth band in upper jaw laterally with mouth closed. Teeth in jaws small, caniniform, in about two rows anteriorly and one row posteriorly in upper jaw, those mesially longest, small teeth extending onto side of jaw near front, distinct hiatus of teeth at symphysis; band of about two rows of teeth laterally in lower jaw, additional row of much smaller teeth basally on outer edge at front; narrow hiatus in dentition at symphysis, teeth largest mesially. About two or three rows of small canines on palatines and traversing vomer, those medially slightly smaller. Tongue with prominent lanceolate patch of numerous, uniformly small teeth in about five poorly defined rows with few naked patches mesially near posterior end; dentate patch approaching anterior edge of tongue. Gill rakers moderately long.

Scales of moderate size, finely spiniform along margins on much of head and body apart from cycloid scales on chest, belly and cheek, not extending onto fins; those on chest and belly ctenoid. Predorsal scales extending forward to about vertical through posterior extent of eye or midway between that point and vertical through posterior edge of preopercle, covering opercle and preopercle forward to posterior end of maxilla; axial scale present at origin of pectoral and pelvic fins. Lateral line midlaterally on side, originating just below horizontal through upper margin of eye; lateral line scales corresponding with oblique scale rows above and below lateral line; single pore on each lateral line scale.

Dorsal fin originating just in advance of vertical through pelvic fin origin, distance from snout tip to dorsal fin origin 35.8 (34.1–38.3) % SL; dorsal fin base of moderate length; first dorsal fin ray unbranched, others branched; fin distinctly higher in males than females, membranes deeply incised, none of the rays filamentous, third or fourth rays longest, longest 21.6 (18.3–22.9 in males, 16.9–23.1 in females) % SL, decreasing only slightly in length to middle of fin with posterior rays progressively longer, penultimate ray 21.0 (17.5–22.1 in males, 9.4–15.2 in females) % SL, posterior lobe reaching to or past adipose fin origin in males, but usually reaching just more than half way to adipose fin in females ( Figs 1 I and 2). Adipose fin rather small but prominent, positioned above posterior end of anal fin base. Anal fin originating slightly in advance of vertical through midpoint between dorsal fin insertion and hypural crease, with short base; first one or two rays unbranched, others branched; relatively deep in males, posteriormost rays much longer than anterior rays and posterior lobe approaching hypural crease in large individuals; fin smaller in females, lengths of posterior rays subequal; posterior lobe reaching little more than half way to hypural crease. Caudal fin deeply forked, shortest rays at middle of fin about a third length of longest rays extending to corners, lobes nearly pointed, upper lobe slightly longer than lower; rays near ventral margin of fin more densely segmented than those near dorsal margin of fin; unbranched ray at ventral margin not with opaque fleshy covering. Pectoral fin of moderate size, length 20.8 (19.1–21.6 in males, 19.7–22.1 in females) % SL, tip reaching past middle of dorsal fin base; origin of fin distinctly in advance of vertical through dorsal fin origin; first two rays simple, others branched, third longest, but just slightly longer than second, subsequent rays progressively shorter. Pelvic fin moderately large, posterior tip of retracted fin reaching past anus, more so in males than females; first four rays not especially thickened and cylindrical, densely segmented, first ray unbranched, second to fourth each with pair of simple thickened branches, subsequent rays progressively shorter, flattened with typical multibranching, except for more slender, inner-most unbranched ray. A species of moderately small size, largest verified specimen 219 mm SL ( Strasburg, 1966).

Fresh colour: (Adapted from Strasburg, 1966 and Fig. 2 ). Upper side brownish-red, with dark-brown saddles (having more intensely pigmented broad anterior and posterior margins) running to midside beneath anterior part of dorsal fin, posterior part of dorsal fin, adipose fin, and just before caudal base; interspaces between saddles streaked with yellow pigment above lateral line; lower side silvery-white, blotched irregularly with vermilion, two largest blotches centred below middle two saddles; throat, breast, and belly white, ventral side of caudal peduncle and gill membranes lemon; top of head olive-brown, cheeks silvery with red blotches, snout and opercle olivebrown with red blotches, iris yellow. Dorsal fin grey, females with round vermilion spots arranged in four irregular diagonal rows, largest spots about size of pupil; males with large spots centrally and basally on fin distinctly ocellated, having yellow centres and red margins; adipose fin brown with red distal tip. Anal fin pale lemon in females; broad more distinctly yellow stripe near base of males. Lower lobe of caudal fin yellow, with upper and lower rays streaked with red; upper lobe of grey, blotched irregularly with lemon and red, tip distinctly white, at least in some males. Pectoral fin hyaline with five vermilion bands. Pelvic fin lemon with orange blotches.

TABLE 2. Selected proportional measurements for types of Hime surrubea sp. nov., H. capitonis sp. nov. and H. caudizoma sp. nov. Numbers of specimens reported for measurements indicated below “n” with values for damaged features omitted.

Hime surrubea sp. nov. Hime capitonis sp. nov. Hime caudizoma sp. nov.

Holotype Paratypes Holotype Paratypes Holotype Paratypes

BPBM Males Females MNHN Males Females MZB Males Females

25117 n = 13 n = 14 2003–1495 n = 9 n = 11 22096 n = 3 n = 4

Standard Length (mm) 156 127–170 90.0–173 149 84.7–173 70.8–146 152 139–183 138–159. ± s d 0.8 0.7 0.9 0.9 1.1 0.7 0.8 0.6 0.4 0.5 damaged for Females 4 = n 159 mean ± 8.7 9.7 14.3 ± 15.1 16.1 ± 16.8 16.1 ± 16.8 ± 16.2 17.3 ± 14.9 15.6 ± 13.8 14.5 7.1 ± 7.6 ± 8.9 9.4 8.1 ± 8.6 values nov. – 138 range 7.8 – – 13.7 – 14.8 – 14.8 – 14.7 13.9 – – 12.6 6.4 – – 8.3 7.7 – ” with “ n. sp Paratypes n ± s d. 1.0 4 0.6 4 4 0.8 4 0.6 0.7 4 4 0.6 4 0.3 0.8 4 0.6 4 4 0.2 below caudizoma Males 3 = n mean 9.8 ± ± 15.5 ± 17.3 ± 19.2 ± 19.0 18.8 ± ± 20.0 8.3 ± ± 10.8 14.1 ± indicated Hime 183 139 – range – 8.7 10.5 – 16.0 14.8 16.8 – 18.2 19.7 18.5 – – 18.2 19.5 18.3 19.5 – 19.7 – 20.2 7.5 9.1 – 10.1 – 11.3 – 14.4 14.0 measurements for Holotype MZB 22096 152 n male 3 9.0 3 16.9 3 19.7 3 20.9 3 19.2 3 20.1 3 21.0 3 7.4 3 10.6 3 14.6 reported ± s d. 0.7

±

1.6

± ±

1.3 1.2

±

1.1 ±

±

0.6

±

0.6

±

0.9 0.4

±

0.5

±

mean

specimens of Females 11 n = 70.8 – 146 range 11.2 10.1 12.6 – 19.1 16.6 – 22.5 20.1 – 18.4 22.5 19.6 21.6 17.3 – 18.0 – 19.3 15.7 14.4 – 15.7 13.6 13.0 11.9 – 13.9 6.5 8.4 – 5.4 8.5 8.0 – 9.1 8.0 – 9.0 7.4

.

.

Numbers nov sp.

Paratypes n d. 10 10 9 10 10 11 11 11 11 11 nov. sp capitonis Males 9 = mean ± s 1.6 10.6 ± ± 18.7 1.4 2.4 20.5 ± 1.5 ± 22.7 ± 1.6 21.3 0.7 16.3 ± ± 2.0 18.9 7.7 0.7 ± 0.8 9.6 ± ± 1.8 11.4 caudizoma . H Hime n – 173 84.7 range – 12.5 8.2 – 21.3 16.3 24.5 17.0 – – 20.7 25.4 24.7 – 18.9 17.4 – 15.0 14.7 – 21.6 6.3 – 8.5 – 11.1 8.5 – 7.6 13.6 and n 9 9 9 9 9 9 9 9 9 7. nov. 1495

sp Holotype MNHN 2003 – 149 male 10.4 19.7 23.7 22.3 20.9 17.0 18.9 8.2 9.5 11.2. capitonis d s. 1.4 2.0 1.4 1.6 1.5 1.1 1.7 1.4 1.1 0.5 ± nov H,. 14 mean 9.4 ± 19.0 ± 20.8 ± 20.2 ± 18.4 ± ± 15.2 13.1 ± ± 6.9 8.9 ± 8.4 ± sp. Females = n 173 – 12.1 21.4 23.1 22.8 21.1 16.8 15.2 8.6 10.3 9.0 surrubea . 90.0 range 7.4 – – 14.7 – 18.8 16.9 – – 16.0 – 13.2 9.4 – 3.8 – 6.9 – – 7.0 Hime of nov sp. Paratypes n 12 14 14 13 14 13 14 14 13 14 in types surrubea 13 ± mean d s. 1.4 ± 9.9 ± 1.2 18.2 20.5 ± 1.5 ± 1.1 21.1 19.3 ± 0.9 1.3 ± 17.3 ± 1.4 19.6 6.9 ± 1.5 0.5 9.8 ± ± 12.0 1.1 fin rays Hime Males = n 127 – 170 range 7.1 – 11.4 20.3 16.6 – – 22.9 18.3 18.8 – 22.9 21.0 17.9 – 20.8 – 15.6 22.1 17.5 – – 3.4 9.6 – 8.8 10.4 – 13.5 9.7 selected for n 12 12 12 13 13 13 13 12 12 13 measurements Holotype BPBM 25117 156 male 11.1 20.3 20.6 21.6 20.1 18.0 21.0 7.4 9.7 13.0. 3 Proportional. omitted) (mm Length % SL ray: length st - 1 ray nd 2 ray - rd ray - 3 th ray 4 - th ray 5 - th 10 - ray ray - penulmate ray length: st ray - 1 nd 2 ray - ray penulmate - TABLE features Standard Dorsal-fin Anal-fin

Preserved colours: body pale, slightly duskier on dorsum, darker blotches corresponding to saddles described above, most prominent a rectangular blotch below anterior fifth of dorsal fin base at level of upper third of eye, several others at same level, somewhat paired, below nape, below rear end of dorsal fin base, and below adipose fin; second row of horizontally aligned blotches closer to dorsal profile of side, just above and behind each blotch of first row, blotches appearing somewhat like obliquely slanted ‘H’ shaped marks in some; opercle mostly dark dusky; fins pale.

Etymology. The name surrubea, Latin for “reddish”, refers to the reddish hue that is a characteristic of this species

Distribution. Apparently confined to the Hawaiian Ridge in the north-central Pacific ( Fig. 3 ) from at least the Equator Seamount (29° 42' N, 179° 22' E; Borets, 1986 as H. japonicus ) to off the island of Hawaii (20° 01' N, 156° 44' W) at depths of 170–510 m ( Chave & Mundy, 1994). A demersal species on sandy bottom.

Comments. The first published record of a specimen attributable to this species is that of Strasburg (1966), who identified as an immature female of Hime japonicus (USNM 198224) a 219 mm SL fish taken on handline at a depth of 780 feet (238 m) between Lanai and Kahoolawe, islands near the center of the main Hawaiian group. He provided brief morphological details and a colour description, stating it agreed “with that of the Japanese specimen figured by Kamohara (1955: pl. 9), except that his (Kamohara’s) fish has only weak indications of spotting on the dorsal, caudal, and pectoral fins, and none at all on the cheeks and opercles”. The implied absence of these markings in the illustration is likely attributable to the rapid fading of red pigmentation after death, which Strasburg mentioned in describing the colour of his specimen in preservation, and as evidenced by the presence of red markings in the colour images of fresh female Hime japonica in Masuda et al. (1975: P.20-A), Masuda et al. (1984: pl. 61B) and Matsunuma et al. (2008: fig. 2). Although Strasburg described the top of the head, snout and opercle of his specimen as olive-brown, his reference to the upper sides as brownish-red identifies the reddish accent that is a prominent feature of this species. By contrast, the dorsum of H. japonica is tan to pale brown with considerably darker brown markings.

As implied in the above synonymy, a number of authors waivered over the recognition of this species almost from its discovery. Evidently, Struhsaker after first considering it new seemed to question that opinion as a description was never published. Of those investigating the problem, Carl Hubbs and Doug Hoese appear to have been closest to providing a name as revealed in historical correspondence (circa 1969–1972; Hoese, pers. comm.), but a description never eventuated. Interestingly in his correspondence, Hubbs regarded Hime as a natural assemblage separate from Aulopus and Latropriscis, but considered the three to be subgeneric assemblages within Aulopus rather than distinct genera.

In general form H. surrubea is very similar to H. japonica , especially in the size and shape of the dorsal fin of males, the feature that varies most noticeably between species of the genus ( Fig. 1 ). The new species, however, has a shorter caudal peduncle (length 13.4–18.7, mean 16.0 vs 16.8–19.1, mean 17.9% SL; morphometric values for H. japonica for this and the following characters are mostly those of Gomon et al., 2013: table 3), slightly larger head (length 30.1–33.3, mean 31.8 vs 29.1–31.5, mean 30.3% SL), larger eye, especially relative to head size (orbital diameter 10.1–11.8, mean 10.9 vs 8.4–10.5, mean 9.5% SL, and 31.2– 38.1, mean 34.3 vs 28.8–35.5, mean 31.4% HL), and slightly smaller pectoral fin (19.1–22.1, mean 20.6 vs 20.9–23.7, mean 22.0% SL) and pelvic fin (23.7–29.5, mean 26.1 vs 21.7–30.3, mean 26.1% SL). The pelvic fin of males and females of H. surrubea are of similar size, while in H. japonica the fin of females is consistently smaller than that of males (21.7–26.1, mean 24.6 vs 25.9–30.3, mean 27.7% SL, respectively).

The English vernacular for H. surrubea currently in the literature, “Japanese Thread-sail Fish”, was obviously intended for H. japonica , with which it was thought to be conspecific. To eliminate confusion, we propose “Rosy Flagfin” as a replacement name for the Hawaiian species, which more accurately reflects its appearance. Species of the family are known as flagfins, threadsails, sergeant bakers and aulopus in various parts of the world.

The extremely large numbers of specimens of this species in museum collections, most having been taken with demersal trawls during surveys of the Hawaiian Islands by the R/V Townsend Cromwell in the late 1960s, imply a habitat preference for soft substrate by this species. Most other species are represented by far fewer collection specimens with associated collection data indicating a hard, less easily sampled substrate.

Figures

FIGURE 1. Dorsal fin profiles in males and females of Hime species: A, H. curtirostris, B, H. japonica, C, H. microps, D, H. capitonis sp. nov., E, H. caudizoma sp. nov., F, H. diactithrix, G, H. formosana, H, H. pyrhistion, and I, H. surrubea sp. nov.

FIGURE 2. Hime surrubea sp. nov., BPBM 25114, paratype, 163 mm SL, male, Hawaii, Maui, Pailolo Channel, 21 ° 02 ' N, 156 ° 45 ' W, 241 – 254 m. Photo by J. Randall.

FIGURE 3. Collection localities for specimens examined and verified identifications of species of Hime: H. capitonis sp. nov. (yellow squares), H. caudizoma sp. nov. (blue circle), H. microps (green circles) and H. surrubea sp. nov. (red triangles). Coordinates at 20 degree intervals.