Nesometra integra, Pratama & Virgili & Reimer & Fujita, 2024

Subfamily Antedoninae Norman, 1865 View in CoL Genus Nesometra Virgili, Poliseno, Fujita, Pratama, Fernández-Silva, and Reimer, 2023 [New Japanese name: Sesoko-hime-umishida-zoku] Nesometra integra n. sp. [New Japanese name: Kanzen-sesoko-hime-umishida] ( Figs 1 View Fig , 3–7 View Fig View Fig View Fig View Fig View Fig , 9A, B View Fig )

Antedon cf. iris View in CoL : Hemery et al. 2013: 513, 518, fig. 1, table 2. Nesometra sp. : Virgili et al. 2023: 13, 14, figs 1–3.

Material examined. Holotype: NSMT E-14402, Kedomi , Kakeroma-jima Island, Amami Islands, Kagoshima Prefecture, Japan, St. 10, 28°5.57′N, 129°14.55′E, 42–43.2 m depth, 29 May 2019, coll. T. Fujita.

Paratypes: NSMT E-13332, Seisui, Amami-Oshima Island, Amami Islands , Kagoshima Prefecture, Japan, St . 19, 28°7.71′N, 129°19.65′E, 28–33 m depth, 30 May 29, coll . T. Fujita; NSMT E-13603–13607, Sukomobanare Island , Amami Islands, Kagoshima Prefecture, Japan, 28°6.48′N, 129°10.17′E, 30 m depth, 8 October 2019, coll . R. Virgili; NSMT E-14401, 14403, 14404, 14406, Chichi-jima Island , Ogasawara Islands, Japan, St . 20, 27°4.23′N, 142°15.19′E, 52–54m depth, 29 October 2008, coll. T. Fujita, H. Namikawa, M. Okanishi; NSMT E-14405, Ikomo Bay , Kakeroma-jima Island , Amami Islands, Kagoshima Prefecture, Japan, St . 11, 28°5.44′N, 129°14.94′E, 45–75 m depth, 29 May 2019, coll. T. Fujita; NSMT E-14407, Shirahama Coast, Amami Islands , Kagoshima Prefecture, Japan, 28°11.87′N, 129°16.13′E, 17.5 m depth, 7 August 2015, coll . M . Arai; MV F160501 [as A. cf. iris in Hemery et al. (2013)], Northwest Shelf , Cunningham Island , Western Australia, St . 062, 17°35.71′S, 118°58.90′E, 110 m depth, 25 June 2008, coll . K . Naughton.

Comparative material. Antedon iris : holotype, ZMA. V.ECH. CR.1350, north of Pulau Damar (previously Salomakiëe) Island, Maluku Islands, Indonesia, Siboga Expedition, St. 144, 0°56.52′S, 128°19.45′E, 45 m depth, 7 to 9 August 1899.

Nesometra sesokonis : NSMT E-13579, Sesoko Island , Okinawa Prefecture, Japan, 26°39.30′N, 127°51.14′E, 13 m depth, 26 July 2019, coll . R. Virgili .

Diagnosis. A small-sized species with arm length of 11–41 mm. Centrodorsal flattened hemispherical, up to 1.5 mm in diameter; cavity opening rounded; radial impressions absent. Cirri present, IX–XXVIII, up to 7 mm long, up to 13 cirrals; cirrals laterally compressed; medially constrict- ed up to antepenultimate cirral in immature cirri, restrict- ed to cirrals before antepenultimate cirral in mature cirri; longest cirral L/W up to 4.2; cirral with perforate stereom, proximal spines present, distal end flared. Radials concealed by centrodorsal; proximal and distal facet trapezoidal with broadened aboral side; aboral side and circumoral nerve canal openings fused; adoral muscle fossae rounded; intermuscular ridge with shallow median furrow. IBr 2 united by synarthry; Ibr 2 ax pentagonal or rhombic with weak synarthrial tubercle. Ten arms. Distal intersyzygial interval 3. Proximal pinnulation complete; P 1 up to 4.5 mm long with up to 15 pinnulars; P 2 up to 3.1 mm long with up to 9 pinnulars; P 3 or P 4 first genital pinnule; pinnulars with spinous distal edge, not everted; comparative proximal pinnule length P 1> P 2> P 3 <P 4 <P 5 <PM <PD or P 1> P 2 ≤ P 3 <P 4 <P 5 <PM <PD. Disk globular, smooth; mouth central in fully developed individuals.

Description of the holotype NSMT E-14402 ( Fig. 1A–D, F View Fig ). Longest arm 25 mm in length. Centrodorsal flattened hemispherical, 1.5 mm across; polar area slightly convex, 0.4 times centrodorsal diameter, covered by obsolete cirrus sockets; cirrus sockets arranged in 2 alternating rows.

Cirri XXIV, slender, laterally compressed, cirri on upper rows slightly longer than lower rows, 4–7mm long; composed of 11–13 cirrals ( Fig. 1D View Fig ; see also NSMT E- 14405 in Fig. 4A, B View Fig ). Cirrals elongated except first cirral, smooth without aboral processes except opposing spine; distal end flared especially on oral side; proximal cirrals medially constricted; c1 L/ W 0.4 –0.7; c2 L/ W 1.0 –1.9; c3–5 longest, L/ W 2.3 –3; following cirrals gradually lose its medial constriction, shortened in length, only laterally compressed; antepenultimate L/ W 1.0 –1.2; penultimate L/ W 0.9 –1.3, with straight opposing spine on middle part; terminal claw curved beyond aboral margin of penultimate, approximately as long as antepenultimate.

Radials concealed by centrodorsal; IBr consists of 2 ossicles; Ibr 1 and Ibr 2 ax connected by synarthry; weak synarthrial tubercle present between Ibr 1 and Ibr 2 ax (see NSMT E- 14405 in Fig. 1E View Fig ); Ibr 1 broad oblong, distal side slightly incised (see also NSMT E- 14407 in Fig. 5C View Fig ), W/L 6.3–7.0; Ibr 2 ax pentagonal (see also NSMT E- 14405 in Fig. 5D View Fig ), W/L 1.5–1.8; each division series widely separated.

Ten arms, 2 arms broken, 8 arms almost equal in length, 8–25 mm long; 0.6 mm wide at first syzygy; 4.1 mm long from Ibr 1 to second syzygy. Brachials without aboral projection ( Fig. 1C View Fig ); br 1 broad oblong, not united interiorly, but slightly in contact with neighbor, width 0.669 mm, interior length 0.126 mm, exterior length 0.266 mm; br 2 irregular-shaped, completely separated from neighbor, width 0.681 mm, interior length 0.318 mm, exterior length 0.413 mm; weak synarthrial tubercle present between br 1 and br 2; br 3+4 oblong, interior length 0.629 mm, exterior length 0.439 mm; following brachials until br 8 wedge-shaped, with smooth, slightly everted distal edge; br 9+10 onward wedge-shaped, with smooth distal edge; brachials on distal half of arms elongated oblong, L/ W 1.1 –2.0. First syzygy at br 3+4; second at br 9+10; third at br 14+15; following at intervals of three muscular articulations.

Proximal pinnulation complete ( Fig. 1B View Fig ); pinnules slen- der except for P 1, tapering evenly to tip. P 1 on br 2, longest and stout, 4.5 mm long, composed of 14 pinnulars; p1 L/ W 0.6; p2 L/ W 1.4; p3 L/ W 1.7; following pinnulars elongated, L/W not more than 7 ( Fig. 1F View Fig ). P 2 2.2 mm long, 9 pinnulars; p1 L/ W 0.6; p2 L/ W 1.1; p3 L/ W 2.1; following pinnulars elongated, L/W not more than 3.5 ( Fig. 1F View Fig ). P 3 1.9 mm long, 9 pinnulars; p1 L/W 1; p2 L/ W 2.8; p3 L/ W 3.3; following pinnulars elongated, L/W not more than 4.5 ( Fig. 1F View Fig ). P 4 3 mm long, 11 pinnulars; p1 L/ W 0.6; p2 L/W 1; p3 L/ W 2.9; following pinnulars elongated, L/W not more than 8.5 ( Fig. 1F View Fig ). P 5 3.8 mm long, 11 pinnulars ( Fig. 1F View Fig ). PM 4–5 mm long, 12–13 pinnulars ( Fig. 1F View Fig ). PD 5.7 mm long, 11 pinnulars ( Fig. 1F View Fig ). Pa present, 2.42 mm long, 8 pinnulars ( Fig. 1F View Fig ). Genital pinnules not developed. All pinnules with distal edge of pinnulars spinous except on first and second pinnulars, not everted; P 2 /P 1 0.49, P 3 /P 2 0.86, comparative length P 1> P 2> P 3 <P 4 <P 5 <PM <PD.

Disk large, globular, 3.1 mm across, smooth without calcareous nodules, covering proximal ray until about br 1 ( Fig. 1A View Fig ); mouth central; anal papilla sub-central close to mouth, 1.7 mm high from adoral surface of centrodorsal.

Description of the Australian paratype MV F160501. Centrodorsal flattened hemispherical, 1.5 mm across; polar area slightly convex, 0.5 times centrodorsal diameter; cirrus sockets arranged in 2 alternating rows.

Cirri XXVIII, laterally compressed, 6 mm long, composed of 10–11 cirrals. Cirrals elongated except first cirral, smooth without aboral processes except opposing spine; distal end flared especially on oral side; proximal cirrals medially constricted; c1 L/ W 0.6; c2 L/ W 1.2; c3–4 longest, L/ W 2.2 –2.4; following cirrals gradually lose its medial constriction, shortened in length, only laterally compressed; antepenultimate L/ W 1.2; penultimate L/ W 0.9, with straight opposing spine on middle part; terminal claw curved beyond aboral margin of penultimate, approximately as long as antepenultimate.

Radials concealed by centrodorsal; IBr consists of 2 ossicles; Ibr 1 and Ibr 2 ax connected by synarthry; weak synarthrial tubercle present between Ibr 1 and Ibr 2 ax; Ibr 1 broad oblong, distal side slightly incised, W/L 4.5; Ibr 2 ax pentagonal, W/L 1.4; each division series widely separated.

Ten arms, broken, remaining arms 6–12 mm long; 0.8 mm wide at first syzygy; 5.3 mm long from Ibr 1 to second syzygy; brachials without aboral projection; br 1 broad oblong, slightly in contact interiorly, width 0.811 mm, interior length 0.179 mm, exterior length 0.449 mm; br 2 irregular-shaped, completely separated from neighbor, width 0.860 mm, interior length 0.540 mm, exterior length 0.367 mm; weak synarthrial tubercle present between br 1 and br 2; br 3+4 oblong, width 0.800 mm, interior length 0.734 mm, exterior length 0.613 mm; following brachials until br 8 oblong, with smooth, slightly everted distal edge; br 9+10 onward wedge-shaped, with smooth distal edge; distal part of arms lost. First syzygy at br 3+4; second at br 9+10; third at br 14+15; following at intervals of three muscular articulations.

Proximal pinnulation complete; pinnules slender, tapering evenly to tip. P 1 on br 2, broken. P 2 3.1 mm long, 9 pinnulars; p1 L/ W 1.0; p2 L/ W 2.0; p3 L/ W 3.4; following pinnulars elongated, L/W not more than 5.0. P 3 2.6 mm long, 9 pinnulars; p1 L/ W 0.5; p2 L/ W 1.0; p3 L/ W 2.3, following pinnulars elongated, L/W not more than 5. P 4 and P 5 mostly broken. PM 3 mm long, 10 pinnulars. PD lost. Pa present, broken. Genital pinnules not developed. All pinnules with distal edge of pinnulars finely spinous except on first and second pinnulars, not everted; P 3 /P 2 0.84.

Ossicle morphology of paratypes observed by SEM. Centrodorsal. Adoral surface concave, with five shallow interradial furrows, radial impressions absent ( Fig. 3A View Fig ). Cavity opening rounded, cavity diameter 0.4 times of ossicle diameter; interradial notch on cavity opening edge absent; overhanging rim at adoral edge absent, so internal diameter of cavity similar to opening. Aboral pole smooth without tuberculation, obsolete cirrus sockets present ( Fig. 3B View Fig ). Cirrus sockets with raised stereom surrounding the central lumen ( Fig. 3C View Fig ). Basal rosette located above cavity opening, pentagon, perforated centrally, with five radial processes ( Fig. 3D View Fig ); radial processes of basal rosette attached to proximal facet of radials, forming radial pentagon ( Fig. 3E, F View Fig ). Each corner of radial pentagon reaching centrodorsal margin.

Radial. Proximal facet trapezoidal with broadened aboral side, H/ W 0.74 –0.79, convex; ambulacral gutter not covered by spicules; aboral side and circumoral nerve canal openings fused ( Fig. 3G View Fig ). Distal facet trapezoidal with broadened aboral side, H/ W 0.77 –0.83, concave; adoral muscle fossae oblique to oral-aboral axis, interarticular ligament fossae deep, aboral ligament fossa deep; pairs of adoral muscle and interarticular ligament fossae laterally separated from adjacent ones ( Fig. 3H View Fig ). Adoral muscle fossae with solid and porous stereom, pores roughly arranged in concentric bands ( Fig. 3I View Fig ); rounded, H/ W 0.86 –1.30; oral border concave; each fossa diverges from neighbor. Intermuscular ridge broad, 0.07 mm wide, running from above central canal to ambulacral gutter, with shallow median furrow. Interarticular ligament fossae with galleried stereom ( Fig. 3J View Fig ); semicircular, broadened at outer edge, H/ W 1.53; separated from muscle fossae by narrow diagonal ridge; shallow furrow present on ridge. Aboral ligament fossa with galleried stereom ( Fig. 3K View Fig ); low semicircular, H/ W 0.30. Central canal oval. Ligament pit slightly smaller than central canal.

Cirral. Surface with perforate stereom, with round to elliptical pores ( Fig. 4C View Fig ); proximal spines present ( Fig. 4D View Fig ). Articulation not overlapped by preceding cirral.

First ossicle of division series (Ibr 1). Proximal facet trapezoidal with broadened aboral side, oral side with ambulacral gutter ( Fig. 5E View Fig ), H/ W 0.95; adoral muscle fossae perpendicular to oral-aboral axis, interarticular ligament fossae deep, aboral ligament fossa deep; pairs of adoral muscle and interarticular ligament fossae laterally separated from adjacent ones. Adoral muscle fossae with solid and porous stereom, pores roughly arranged in concentric bands; rounded, H/ W 0.86; oral border slightly rounded. Intermuscular ridge tapering from just above central canal to near ambulacral gutter, with shallow median furrow. Interarticular ligament fossae with galleried stereom; broadened, H/ W 1.45; separated from muscle fossae by oblique furrow. Aboral ligament fossa with galleried stereom; low semicircular, H/ W 0.27. Central canal oval. Ligament pit slightly smaller than central nerve canal. Distal facet rounded bifascial synarthry ( Fig. 5A View Fig ), H/ W 0.97. Fulcral ridge narrow, 0.04 mm wide. Synarthrial ligament fossae with galleried stereom; arch-shaped, gently sloping, H/ W 1.79. Central canal rounded.

Second ossicle of division series (Ibr 2 ax). Proximal facet trapezoidal bifascial synarthry, broadened oral side with shallow, wide ambulacral gutter ( Fig. 5F View Fig ), H/ W 0.81. Fulcral ridge narrow, 0.06 mm wide. Synarthrial ligament fossae with galleried stereom; arch-shaped, deep, H/ W 1.91. Central canal tall trapezoidal with broadened oral side. Distal facet 2 muscular articulations ( Fig. 5B View Fig ), H/ W 0.88; separated laterally by broad intermuscular ridge, 0.06–0.07 mm wide; running from transverse ridge on middle region to oral side, with shallow median furrow; horn-like protrusion present on oral end of intermuscular ridge. Adoral muscle fossae with solid and porous stereom, pores roughly arranged in concentric bands; oblique to oral-aboral axis; rounded, H/ W 0.79. Interarticular ligament fossae with galleried stereom; broadened, H/ W 0.96; separated from muscle fossae by narrow diagonal ridge. Aboral ligament fossa with galleried stereom, semicircular, H/ W 0.53. Central canals rounded. Ligament pits slightly larger than central canals.

Articulations on brachials. Synarthry between br 1 and br 2 with adambulacral portion of fulcral ridge slightly shorter than of abambulacral portion ( Fig. 6A View Fig ). Ligament fossae arch-shaped. Muscular articulation on proximal and middle brachials with two adoral muscular fossae, two interarticular ligament fossae, and an aboral ligament fossa ( Fig. 6B, C View Fig ). Adoral muscular fossae with solid and porous stereom, pores roughly arranged in concentric bands; roughly triangular on proximal brachials or pear-shaped on middle brachials. Intermuscular ridge tapering from just above central canal to near ambulacral gutter. Interaticular ligament fossae with galleried stereom; broadened; separated from muscle fossae by narrow diagonal ridge; slight impression may present on latero-aboral side of fossa on proximal brachials; rounded impression present on adoral portion of each fossa on middle brachials. Aboral ligament fossa with galleried stereom; semicircular. Central canal rounded. Ligament pit slightly larger than central canal. Syzygies on proximal and middle brachials with numerous narrow ridges ( Fig. 6D, E View Fig ); several ridges reaching central canal on proximal syzygies, but none reaching central canal on middle syzygies; articular area occupying almost entire ossicle facet. Pinnule sockets consist of two large fossae with labyrinthic stereom ( Fig. 6F View Fig ); separated by transverse ridge and central lumen.

Coloration. Two main coloration patterns: “solid” and “dotted”. In solid-colored specimens, disk red-orange-brown (holotype, Fig. 1A View Fig ; NSMT E-13603, Fig. 7A View Fig ), or purple (NSMT E-14401). Arm brachials uniformly colored, red-orange-brown with yellowish muscular articulation (holotype, Fig. 1A View Fig ; NSMT E-13603, Fig. 7A View Fig ), light brown with white muscular articulation (NSMT E-14406), or purple with pale purple muscular articulation (NSMT E-14401). Pinnules brown with yellowish distal ends (holotype) or purple with whitish distal ends (NSMT E-14401). Fleshy part of genital pinnules contain yellowish-brown gonad (NSMT E-14406) or purplish-brown gonad (NSMT E-14401). In dotted specimens ( Fig. 7B View Fig ), disk khaki (NSMT E-13604, 14403), and arm brachials irregularly alternating between dark brown and yellowish-khaki with white muscular articulation (NSMT E-13604, 14403). The same pattern is present on the pinnules. In both coloration types, cirri khaki (holotype) or light brown (NSMT E-14406).

These colorations were retained (holotype, NSMT E-14403, and NSMT E-14406) or faded (NSMT E-14401) in preserved specimens after fixation.

Habitat. Specimens observed in situ (NSMT E-13603– 13607) were found hiding below rocks during daytime at 30 m depth, and were exposed only after rock-flipping ( Fig. 7A, B View Fig ).

Distribution. Japan (Ogasawara Islands and Amami Islands) and Australia (Cunningham Island). Bathymetrical range: 17.5–110 m; 52–54 m in the Ogasawara Islands, 17.5– 75 m in the Amami Islands, 110 m in Cunningham Island.

Etymology. The specific name, integra (Latin adjective, feminine), means ‘complete’ after the adult complete proximal pinnulation of the new species compared to its congener, N. sesokonis .

The Japanese name of the genus Nesometra , Sesoko-hime-umishida-zoku (Ɩḽ¬LĮoìỳỵfi) is derived from Sesoko (ÊḂ), the type locality of the type species N. sesokonis , and hime-umishida-zoku (LĮoìỳỵfi), the Japanese name of antedonid comatulids and the Japanese word for genus, zoku (fi). The Japanese name of the new species, Kanzen-sesoko-hime-umishida (hžüžƖḽ¬LĮoìỳỵ), is derived from kanzen (Żḯ), means also ‘complete’ as a translation of the specific name, integra .

Molecular analysis. The Nesometra integra n. sp. COI sequences produced here had a 0.4 ± 0.1% mean genetic distance with Nesometra sp. (INSD accession numbers OP873135–OP873139) by Virgili et al. (2023), and 0.9 ± 0.4% with A. cf. iris (KC626511) of Hemery et al. (2013), suggesting conspecificity according to the mean COI intraspecific distances found in comatulids of 0 to 2.0 ± 0.7% (Summers et al. 2017; Virgili et al. 2023). The mean interspecific distance between N. integra n. sp. (including the A. cf. iris sequence) and N. sesokonis (OP873140–OP873152) was 3.3 ± 0.8% ( Table 1), which is slightly less than the usual mean interspecific distance in comatulids (ranging from 3.7 ± 0.9 to 6.7 ± 1.1%) as reported by Summers et al. (2017). Nevertheless, the multi-locus phylogenetic reconstruction and species delimitation analyses by Virgili et al. (2023) clearly separated these two clades, further suggesting that N. integra n. sp. is a separate species from N. sesokonis .

Remarks. Nesometra integra n. sp. and N. sesokonis share external characters of laterally compressed cirrals with flared distal edges, widely separated division series, rhombicshaped axillary, and weak synarthrial tubercles with the genus Antedon de Fréminville, 1811 . However, Virgili et al. (2023) showed that both N. sesokonis and N. integra n. sp. [= Nesometra sp. in Virgili et al. (2023)] were in a distinct molecular clade from the type species of Antedon , A. bifida (Pennant, 1777) . In addition, the present observation indicated that ossicle microstructures distinguish the two Nesometra species from A. bifida . Radial impressions on the adoral surface of centrodorsal are present for A. bifida (A. H. Clark 1915: figs 283, 593), but are absent for Nesometra ( Figs 3A View Fig , 8A View Fig ). Radials of A. bifida are rectangular in shape with aboral side and circumoral nerve canal openings separated on proximal facet (A. H. Clark 1915: fig. 600), while Nesometra has trapezoidal radials with aboral side and circumoral nerve canal openings fused on proximal facet ( Figs 3E View Fig , 8B View Fig ). These ossicle microstructures are shared by N. sesokonis and N. integra n. sp. and have not been reported in other Antedon species, suggesting that these characters are diagnostic of the genus Nesometra and clearly separate the genus from Antedon .

Some variations in external morphology and measurement were found in paratypes ( Table 2). At the species level, adult N. integra n. sp. is distinguishable from adult N. sesokonis based on the external morphology characters by having complete pinnulation of its proximal pinnules ( Fig. 1B View Fig ), larger centrodorsal diameter (1.0– 1.5 mm from 0.2–1.1 mm), longer cirri with more cirrals (3–7mm with 7–13 cirrals from 1.3–2.7 mm with 7–8 cirrals), and longer P 1 and P 2 with more pinnulars (P 1 1.6–4.5 mm with 7–15 pinnulars from 0.3–1.3 mm with 3–6 pinnulars; P 2 0.8–3.1 mm with 5–9 pinnulars from 0.2–0.8 mm with 3–5 pinnulars). Genital pinnules were not developed in the holotype; however, the first genital pinnule was found in paratypes at P 3 (NSMT E-14407) or P 4 (NSMT E-14406). Obuchi et al. (2010) and Virgili et al. (2023) suggested paedomorphic characteristics in cirral forms of N. sesokonis , and these clearly distinguish N. sesokonis from the new species. The cirrals of immature N. integra n. sp. have strong concavity on the oral and aboral side, creating a medially constricted appearance ( Fig. 9B View Fig ) from the second to the antepenultimate one, and this character is to the cirral before the antepenultimate one in the mature specimens ( Fig. 9A View Fig ). On the other hand, mature N. sesokonis retain strong medially constricted cirrals until the penultimate cirral (Obuchi et al. 2009: fig. 1D; Virgili et al. 2023: fig. 3H′) ( Fig. 9C View Fig ).

Obuchi et al. (2010) reported that N. sesokonis broods larvae on genital pinnules and is a simultaneous hermaphrodite, unique in Antedonidae . Hermaphroditism or brooding was not observed in the three mature N. integra n. sp. specimens (arm length 32–41 mm) studied here, but more observations on the life cycle of this novel species are required to confirm its reproduction mode.

Virgili et al. (2023) suggested, on the basis of molecular data, that A. cf. iris of Hemery et al. (2013) is conspecific to N. integra n. sp. Most of the characteristics and measurements of A. cf. iris (MV F160501) from the current study overlapped with those of N. integra n. sp. ( Table 2). On the other hand, A. iris has smaller width/length ratio of Ibr 1 (2.6–3.0), more numerous cirri (30) with longer length (8–9 mm), and longer P1 to P3 with more pinnulars (P 1 6.8–8.5mm, 16–18 pinnulars; P 2 4 mm, 11 pinnulars; P 3 3–4.5mm, 11 pinnulars) (A. H. Clark 1912; A. H. Clark and A. M. Clark 1967; this study; see Table 2) than both N. integra n. sp. and A. cf. iris . Considering these results, A. cf. iris of Hemery et al. (2013) is not A. iris but should be treat- ed as N. integra n. sp.