Lyphira heterograna
Creators: name: Rahayu, Dwi Listyo
name: Ng, Peter K. L.
Publisher: name: Zenodo
Year: 2024
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Malacostraca
Decapoda
Leucosiidae
Lyphira
Lyphira heterograna

description: Lyphira heterograna (Ortmann, 1892)
(Figs. 1A‒D, 2, 3A‒C, 6A, 7A‒C, 9A, 10A, B, 11A, B, 13A, 14A, 15A–C, 16A, B, 17A‒G, 21A)
Philyra heterograna Ortmann, 1892: 582, pl. 26, fig. 17; Ihle 1918: 315; Balss 1922: 128; Urita 1926: 38; T. Sakai 1937: 158, text fig. 33; Uchida 1949: 717, fig. 2075; Miyake, 1961 b: 171; Miyake et al. 1962: 127; T. Sakai, 1965: 48; 1976: 111, pl. 32, fig. 3, text fig. 63a; Serène 1968: 46; Takeda & Miyake 1970: 230; Kim 1973: 306, pl. 77, fig. 67a, b, text fig. 99; T. Sakai 1976: 111, text fig. 63a, pl. 32 fig. 3; Takeda 1982: 101, fig. 297; Dai et al. 1986: 78, fig. 39.2, pl. 9.7; Chen 1987: fig. 2; Dai & Yang 1991: 86, fig. 39.2, pl. 9.7; Takeda 1993: 56; Muraoka 1998: 19; Komai 1999: 82; K. Sakai 1999: 18, fig. 1b, pl. 7C; Ng et al. 2001: 9, fig. 3a; Chen & Sun 2002: 379, text fig. 169I, pl. 14.4; Takeda et al. 2006: 191; Ng et al. 2008: 93; Galil 2009: 300 (in part).
Philyra pisum. — Nobili 1903: 11. (not Philyra pisum De Haan, 1841 = Pyrhila pisum (De Haan, 1841)).
Philyra peitahoensis Shen, 1932: 18, pl. 1.1-2, text figs. 10–12, 16b; Serène 1968: 46.
Philyra anatum.— Rathbun 1910: 312. (not Cancer anatum Herbst, 1783).
Lyphira heterograna.— Ng et al. 2017: 48.
Type material. Lectotype: male (12.4 × 12.2 mm), MZS Cru 3191 (specimen number Ortmann 658), Tokyo Bay, Japan, coll. L. Döderlein, 1880–1881.
Other material examined. 1 male (16.5 × 16.9 mm), KPM NH 0130526, 1 male (15.2 × 15.9 mm) KPM NH 0130527, Mikawa-Isshiki, Aichi, Japan, ca. 50 km south of Nagoya City, central Japan, T. Odawara collection, coll. 20 November 1958. 14 males (16.4 × 16.3 mm, 16.4 × 16.6 mm, 16.5 × 17.6 mm, 16.6 × 17.0 mm, 16.6 × 17.2 mm, 17.1 × 17.6 mm, 17.2 × 18.1 mm, 17.5 × 17.9 mm, 17.6 × 17.5 mm, 17.7 × 18.7 mm, 18.1 × 18.4 mm, 18.4 × 18.9 mm, 18.9 × 20.0 mm, 1 broken), 5 females (14.9 × 15.0 mm, 16.8 × 17.3 mm, 17.4 × 17.6 mm, 17.5 × 17.6 mm, 17.5 × 17.6 mm), KPM NH 0103043, Isshiki, Aichi, central Japan, coll. T. Sakai, 1967; 2 males (8.8 × 9.5 mm, 8.8 × 9.3 mm), NHM 1930.11.14.5–6 (paratypes of Philyra peitahoensis Shen, 1932), Pei-Tai-Ho, Peichihli Bay, northern China, coll. S.S. King, summer 1926; 2 males (14.2 × 14.4 mm, 14.0 × 14.4 mm), 2 females (13.5 × 13.7 mm, 11.7 × 11.7 mm), NHM 2006.558–561 (ex 1912.12.15.31–43), Chusan, South Korea, 10–14 fathoms (18–25 m), coll. P. Basset-Smith, no date.
Diagnosis. Carapace approximately as broad as long, or slightly longer than broad, dorsal surface covered with small and minute granules, large, more prominent granules on cardiac, branchial and intestinal regions, sparse granules on hepatic region (Figs. 1A, 2A, B, 3A–C, 6A); frontal margin granulated; postfrontal lobe slightly convex with slight concave medially (Figs. 1B, 11A, B); endostomial spine low, barely visible from dorsal view (Figs. 1A, B, 2A, B, 13A, 14A); lateral margin of carapace with row of small granules interspersed by larger granules; posterior margin gently convex, with large granule medially and on each posterolateral border (Figs. 1A, 2A, B, 3A–C, 6A); posterior carapace margin sinuous, with large granule medially and on each posterolateral corner, with margin between them gently concave to almost straight (Figs. 1A, 2A, B, 3A–C, 6A, 10A‒B). Third maxilliped exopod 2.9 times as long as broad, forming petaliform structure with convex outer margin (Figs. 1C, 2D, 11A, B, 17A). Chelipeds subequal; merus, carpus and chela surfaces smooth or covered with low, small rounded granules (Figs. 1A, 2A–C, 3A–C); chela stout, palm surface smooth, large granules on lower outer surface; fingers longer than length of palm; pollex slightly bent, cutting edge lined with denticles; dactylus 1.4 times as long as palm, row of small granules on upper margin, cutting edge with median large tooth, remaining of edge with small denticles (Fig. 15A–C). P2–P5 slender, short (Figs. 1A, 2A‒C, 3A, B); merus distinctly longer than carpus and propodus, dactylus longer than propodus, lanceolate; fourth leg shortest, with merus 4.3 times as long as broad, margins lined with small granules, more prominent on ventral margin (Fig. 16A, B). Thoracic sternites transversely narrow (Figs. 2D, 7A–C). Pleon narrow, slender, long; somites 2–6 fused, forming elongate trapezoidal plate, shallow suture just visible between somites 5 and 6, surface smooth; somite 6 longitudinally subrectangular, surface with one granule medially; telson triangular, longer than wide (Figs. 7A–C, 17B). G1 elongate, slender, slightly dilated on distal area; apical process long, very slightly bent, with blunt tip (Fig. 17C–G). Female pleon broad, convex, smooth, somites 2‒6 fused, suture between somites 2 and 3; telson shorter than its basal width (Fig. 9A); vulvae spaced far apart on sternite 6, each opening weakly crescent-shaped, relatively short, without opercular cover (Fig. 21A).
Description. Carapace approximately as broad as long, or slightly longer than broad (Figs. 1A, 2A, B, 3A‒C, 6A); most regions indistinct, branchio-cardiac grooves just visible, hepatic, cardiac and intestinal regions distinct, swollen; dorsal surface of carapace covered with minute and small granules, large, more prominent granules on cardiac, branchial, and intestinal regions, sparse granules on hepatic region. Frontal margin granulated, postfrontal lobe slightly convex with slight concave medially (Fig. 1B, 11A, B).Antennules folding transversely.Antennae short, longitudinally inserted between antennular fossa and base of ocular peduncle. Orbits small, rounded, upper orbital margin entire. Eyes short, retractable. Endostomial spine low, barely visible from dorsal view (Figs. 13A, 14A). Hepatic facet indistinct, rows of granules on lower margin continuing to distal part, almost reaching frontal margin; upper margin with sparse low granules. Lateral margin of carapace with row of small granules interspersed by larger granules; posterior margin gently convex, with large granule medially and on each posterolateral border (Figs. 1A, 2A, B, 3A–C, 6A). Posterior carapace margin sinuous, with large granule medially and on each posterolateral corner, with margin between them gently concave to almost straight (Figs. 1A, 2A, B, 3A–C, 6A, 10A, B). Pterygostomial region granulated; sub-branchial region almost smooth.
Third maxilliped exopod 2.9 times as long as broad, approximately as broad as basal part of endopod, forming petaliform structure with convex outer margin, with submarginal row of low to very low granules along inner margin; endopod with completely fused basis-ischium, distinctly longer than merus, with shallow submarginal sulcus near inner margin; merus triangular with blunt tip, margins unevenly granulated; palp on inner surface, dactylus longest (Figs. 1C, 2D, 11A, B, 17A). Coxa prominent, forming curved plate.
Chelipeds subequal; merus subcylindrical, surface granulated, larger granules on proximal area and along margins; carpus smooth but with granules along margin (Figs. 1A, 2A‒C, 3A‒C). Chela stout, palm surface smooth, upper and lower margins with row of granules, longitudinal row of relatively large granules on lower outer surface of palm; fingers longer than length of palm, surface with longitudinal sulcus flanked by row of small granules; pollex slightly bent, cutting edge lined with denticles; dactylus 1.4 times as long as palm, row of small granules on upper margin, cutting edge with median large tooth, remaining of edge with small denticles (Figs. 15A‒C).
P2–P5 slender, short (Figs. 1A, 2A‒C, 3A, C); merus distinctly longer than carpus and propodus; dactylus longer than propodus, lanceolate, terminating in corneous tips; second and third legs longest, fourth leg shortest, with merus 4.3 times as long as broad, margins lined with small granules (more prominent on ventral margin) (Fig. 16A, B).
Thoracic sternites transversely narrow (Fig. 7A‒C), surface finely granulated, larger granules laterally; sternites 1–3 completely fused without trace of sutures; sternite 3 separated from sternite 4 by shallow groove. Sternites 4–7 progressively narrower sternite 8 narrowest.
Male sternopleonal cavity deep, reaching to proximal distance between fused thoracic sternites 1–3; pleonal locking mechanism formed by lateral emargination of thoracic somite 5 and pleon (Fig. 7A–C).
Male pleon narrow, slender, long; somite 1 longitudinally narrow, wide; somites 2–6 fused, forming elongate trapezoidal plate, shallow suture just visible between somites 5 and 6, surface smooth; somite 6 longitudinally subrectangular, surface with one granule medially; telson triangular, longer than wide (Figs. 7A‒C, 17B).
G1 elongate, slender, slightly dilated on distal area; apical process long, very slightly bent, with blunt tip (Fig. 17C‒F); G2 small, apical process blunt (Fig. 17G).
Female sternopleonal cavity reaches distal part of sternite 1 (Fig. 9A). Female pleon broad, convex, smooth, somites 2‒6 fused, suture between somites 2 and 3; telson shorter than its basal width (Fig. 9A); vulvae spaced far apart on sternite 6, each opening weakly crescent-shaped, relatively short, narrow, without opercular cover (Fig. 21A).
Colour. Pale orange on dorsal surfaces (Sakai 1976: pl. 32 fig. 3).
Remarks. Komai (1999: 82) first reported on Ortmann’s (1892) type material in MZS from Tokyo Bay, listing five males and eight females as syntypes. Galil’s (2009: 300) comments on the type material are as follow: “ Syntypes: Japan. Tokyo Bay, 1880–1881, id. Ortmann, 1 ♂ cl 12.2 mm, 5 ♀♀ cl 10.8–17.0 mm (Musée zoologique, Strasbourg, reg. no. 658).—Same data, 4 ♂♂ cl 18.4–20.0 mm (Musée zoologique, Strasbourg, reg. no. 149) (the ♂ cl 12.2 mm is here designated the lectotype and the remaining specimens are now considered to be paralectotypes).” She did not state that she examined this material, and no figures of the types were provided. In the paper, she also synonymised Philyra peitahoensis Shen, 1932, and Philyra acutidens Chen, 1987, under L. heterograna. We, however, regard Philyra acutidens Chen, 1987, as a good species (see remarks for the species later).
There are two specimens of “ Philyra heterograna ” in the NHM (catalogue number NHM 1930.11.14.5–6) had been identified as “ Philyra tuberculata Stimpson ” by C. J. Shen and appear to have been deposited there before 1932. All his original labels are there, and they agree what he later describes as P. peitahoensis Shen, 1932. The data matches the material he listed as types of P. peitahoensis and the specimens are clearly paratypes. The holotype of P. peitahoensis was stated to a male from with a catalogue number of 2032 (Shen 1932: 22). These specimens agree very well with what is here defined as L. heterograna s. str. and we agree with Galil’s (2009) action. As for P. acutidens, we prefer to recognise it as a separate species (see discussion for next taxon).
Galil’s (2009) concept of the species is somewhat confusing. She had material from Japan, China, East China Sea, Korea, Philippines, Malaysia (Pontian and Malacca) and Thailand, and she figured the overall dorsal and ventral habitus of one male specimen (NHM 1900.10.22.340–345) (Galil 2009: fig. 14) and its G1 (Galil 2009: fig. 12B); noting that it was from Indonesia. The specimen with this catalogue number, however, is from Malaysia; she actually did not list any material from Indonesia. We have examined her Malaysian material but not been able to find her specimens from the East China Sea and Philippines (they have not been returned at the time of writing). In her diagnosis of L. heterograna as well as her key to the species in the genus, she states that the endostomial spine of the species is not prominent (Galil 2009: 301 314). It is indeed the case for all the specimens from Japan, northern China and Korea (Figs. 13A–D, 14A–B) as well as some specimens we have examined from Singapore and Malaysia (Figs. 3E‒G, 13D, E). But this is not the case for specimens we have from southern China and Hong Kong, as well as most of the material from Thailand and Singapore, all of which have well developed endostomial spines (Figs. 13F‒L). Galil (2009: 301, 314) also states that L. heterograna has the exopod of the third maxilliped relatively narrow and “lozenge-like” or “paddle-shaped” but most of the specimens from southern China, Hong Kong, Malaysia and Singapore actually have proportionately much wider and expanded exopods (Figs. 11D‒J, 12). Only those from Japan, northern China and Korea, and some from Malaysia and Singapore have proportionately narrower exopods (Figs. 11A‒C, 17A, H). Detailed comparisons of all the material we have on hand shows that East and Southeast Asia has six different species under the name “ Lyphira heterograna ”, four of which are undescribed. They differ not just in the strength of the endostomial spines and third maxilliped exopod proportions but also in other characters.
Lyphira heterograna s. str. is distinct and is known for certain from Japan, Korea, Taiwan and northern China; all the specimens from this region, including the lectotype, have weak endostomial spines, the carapace is often longer than broad, the posterior carapace margin is usually sinuous and divided into two concave lobes by three large granules, the exopod of the third maxilliped is more slender, the chela are relatively longer, the cutting edges of the fingers of the chela are lined with low teeth and denticles, the ambulatory merus is relatively slender, the distal process of the G1 is relatively longer and gently curved, and the vulvae are spaced apart with the opening small (cf. diagnosis). Sakai (1976: text fig. 63a) depicts the distal part of the G1 of a specimen from Japan; it agrees very well with the present material from that country (Fig. 17C–F).
The carapace shape of L. heterograna s. str. varies slightly, with most of them longer than broad, although this is more pronounced in males. The proportions of the fingers on the chelae and ambulatory legs, however, do not vary. The carapace shape is a useful character to distinguish the various species in the L. heterograna species-group, but it is not always reliable as it does vary in L. heterograna s. str. to some degree. It is nevertheless important to note that for the other taxa in the species-group, the carapace shape is always more rounded and are never longer than broad.
Nobili (1903: 11–12) had three male specimens from Yokohama in Japan which he identified with P. heterograna but left it under his heading for Philyra pisum, presumably because he regarded the two species as very close and not easily separated. Ng et al. (2001: fig. 3a) shows a dried specimen ostensibly from Kaohsiung in Taiwan that has all the features of L. heterograna s. str., with the carapace proportionately more elongate and the chelae with the fingers as long as the palm, not inflated and without large cutting teeth. No other specimens from Taiwan are known.
The two males from South Korea (NHM 2006.558–561) (Figs. 3C, 7C, 11B, 16B) are probably L. heterograna s. str. They differ from specimens from Japan in that the dorsal surface of the carapace is distinctly smoother and is slightly higher and more convex in frontal view (Figs. 3C, 11B), and the ambulatory merus is also relatively stouter and shorter (Fig. 16B). The endostomial spines, chelae, G1s and vulvae of the Korean specimens, however, agree with the Japanese material. In one male, the granules on pleonal somite 6 are rather low while in the other it is distinctly higher; in the larger male, pleonal somites 1 and 2 are clearly fused but in the smaller one, somite 2 appears more distinctly demarcated with the suture deeper, although it is still functionally fused.
Type locality. Tokyo Bay, Japan.
Distribution. Japan, Korea, northern China and Taiwan, at 0–30 m depth.
description: Published as part of Rahayu, Dwi Listyo & Ng, Peter K. L., 2024, On the genus Lyphira Galil, 2009 (Crustacea: Brachyura: Leucosiidae), with descriptions of four new species from the West Pacific, pp. 358-392 in Zootaxa 5476 (1) on pages 360-364, DOI: 10.11646/zootaxa.5476.1.29, http://zenodo.org/record/12681921