Phyllidiidae from the Red Sea 1986

JOURNAL OF NATURAL HISTORY, 1986, 20, 1401-1428

Red Sea Phyllidiidae (Mollusca, Nudibranchia), with descriptions of new species

NATHALIE YONOW

Marine Research Group, School of Biological Sciences, University College of Swansea, Swansea SA2 8PP, Wales, UK

(Accepted 29 January 1986)

All the known Red Sea nudibranchs of the family Phyllidiidae are described and their taxonomic positions discussed. The genus Fryeria Gray, 1853, is regarded as a junior subjective synonym of Phyllidia Cuvier, 1797, and Reyfria gen. nov. is introduced to describe those species differing from Phyllidia primarily in the position of the anus. Of 12 species described here from the Red Sea, four are new to science. Phyllidia arabica Ehrenberg, 1831, is distinguished from Phyllidia varicosa Lamarck, 1801, and Phyllidia sp. is described as the third species confused with this group. A subspecies of P. ocellata Cuvier, 1804, is described and illustrated, as are three new species: P. melanocera, P. monacha and P. multifaria.

KEYWORDS: Doridacea, Fryeria, gastropod, marine, mollusc, nudibranch, opisthobranch, Phyllidia, Phyllidiidae, Red Sea, Reyfria.

Introduction The Red Sea has attracted a number of studies on its opisthobranch fauna. The first

was published in 1828 by Riippell and Leuckart, who covered a vast number of opisthobranch species from the Gulf of Suez, but there was no further work for almost a century, until Vayssi6re (1906, 1912) described collections made in the Red Sea and the Gulf of Aden. Thereafter Eliot (1908) reported on a collection of nudibranchs from the Sudanese coast as far north as Suez; O'Donoghue (1928) compared the nudibranch fauna of the Red Sea and the Mediterranean; Eales (1938, 1970, 1979) studied the collections of the Murray Expedition, discussed the migration of tectibranchs from the Red Sea to the Mediterranean, and described the aplysiids from the Red Sea. (Barash and Danin (1980) relisted some Red Sea Aplysiidae.) White (1951) described the Bannwarth collection, made in 1914 from a number of Red Sea localities. Subsequently, Marcus and Marcus (1960) reported on opisthobranchs from the Red Sea and the Maldive Islands; Engel and van Eeken (1962) described some Israeli Red Sea opisthobranchs, and Fishelson (1971), in discussing the ecology and distribution of the benthic fauna of the Red Sea, included a few species of opisthobranchs. Most recently, Heller and Thompson (1983) described the non-nudibranch fauna and phyllidiids from a reef off the Sudanese coast; Rudman (1982, 1983, 1984, 1985) has described the chromodorids of the Indo-Pacific, including specimens from this latest collection.

On a circumnavigation of the Red Sea, nine species of the family Phyllidiidae were collected. Three species (Phyllidia sudanensis, Phyllidia varicosa and Phyllidia sp.) not present in this collection are also discussed. A total of 12 species of Phyllidiidae are thus

1402 N. Yonow

so far recorded for the Red Sea. Descriptions have been given for all species, with camera lucida drawings for the majority, and a detailed taxonomic discussion is provided for most species as the taxonomic status of many Phyllidiidae has been in a state of confusion for a long time.

Collection and methods The majority of the specimens were collected from various shallow reef localities

(maximum 20m; see Fig. 1), by scuba diving. All the sites were on reef faces, some with

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Red Sea Phyllidiidae 1403

shelves consisting of coral rubble. Nearly all the animals were photographed in situ. Descriptions and drawings were made before relaxation of the specimens in 7.5~ saline magnesium chloride and preservation in 10~ seawater formalin (4~o aqueous formaldehyde).

All specimens have been lodged in the British Museum (Natural History).

Stations and their positions: Daedalus Reef Whiskey Reef North of creek Creek South Tower Reef Abington Reef Sha'ab Rumi Shib Ammar Sha'ab Loka Wasaliyat Seghala Is. Zubair Islands Tongue Island Jezirat Seba

24°57'N, 35°55'E 30 km NNW of Jeddah, Saudi Arabia 23'5 km north of Jeddah, Saudi Arabia 21°44'N, 39°07'E 25 km WNW of Jeddah, Saudi Arabia 20°55'N, 37°26'E 19°57'N, 37°25% 19°54'N, 39°59'E 18°44'N, 38°33'E 16°45'N, 42°E 15°45'N, 40°47'E 15°2'N, 42°8'E 13°54'N, 42°42'E 12°28'N, 43°25'E

Phyllidia arabica Ehrenberg, 1831 (Fig. 10 A, B, C)

Phyllidia arabica Ehrenberg, 1831: 34. Phyllidia trilineata: Quoy and Gaimard, 1824: 419. Phyllidia varicosa: Lim and Chou, 1970: 135, pl. 16B; Wu and Romig, 1982: 149, figs 1, 2; Yonow,

1984: 227, figs C-E; not Bergh, 1905: 180, taf. 17, fig. 8; not Eliot, 1908: 120; not Faran, 1905: 345; not Guangyu, 1983: 149, pl. 1, fig. 9; not Heller and Thompson, 1983: 339, fig. 8A-G; not Quoy and Gaimard, 1832: pl. 21, fig. 25; not White, 1951: 252.

Phyllidia honloni: Guangyu, 1983: 149, pl. 1, fig. 6; Lim and Chou, 1970: 134, pl. 17B.

Material. (i) 5.7 x 3.0cm; 17.i.1983; North of creek; 9m depth. (ii) 5.0cm long; 16.vi.1983; Wasaliyat; 6m depth. (iii) 1"8 x0"8cm; 19.vi.1983; Tongue Island; 12m depth. (iv) 5.5 x 3.0 crn (preserved); 25.vi.1983; Zubair Island; 10m depth.

Description. This species has been well described and illustrated by numerous authors. The four Red Sea specimens exhibit the typical marking of Phyllidia arabica: four black longitudinal lines and three longitudinal lines or crests of tubercles on the dorsum, and an interrupted longitudinal black line on the foot. The first two specimens had orange-tipped oral tentacles. The smallest specimen (iii), despite its size, is a perfect example of P. arabica.

Remarks. Phyllidia arabica Ehrenberg is here regarded as the correct name for the well-known Indo-West Pacific species referred by most recent authors to P. varicosa. Lamarck (1801) introduced P. varicosa for a species described, but not named, by Cuvier (see Cuvier 1804). There was no illustration to accompany Lamarck's inadequate description, but Cuvier (1804) describing what seems to be the same

1404 N. Yonow

specimen, gave good figures of both dorsal and ventral surfaces, showing clearly that the ventral surface of the foot lacked a black line. Unfortunately, Cuvier introduced the spurious name P. trilineata for Lamarck's species. Ehrenberg (1831) founded his P. arabica on Red Sea material, and stated clearly that the foot bore a ventral longitudinal black line. His description fits most subsequent reports of P. 'varicosa'. Bergh (1869) recognized P. arabica and P. varicosa as distinct species, correctly placing P. trilineata Cuvier in the synonymy of the latter, but wrongly including P. varicosa Quoy and Gaimard (1824) in the synonomy of P. varicosa Lamarck. The Quoy and Gaimard (1824) species had a black line on the foot and Bergh (1869) wrongly incorporated this as a taxonomic character of P. varicosa Lamarck: this seems Io be the first reference to a black sole line in Lamarck's species and the source of all subsequent confusion surrounding the identity of these species.

Phyllidia arabica is a common Indo-West Pacific species (Wu and Romig 1982 as P. varicosa) which has been reported from the Red Sea by several authors. However, none of these records refers to P. arabica. Phyllidia varicosa Heller and Thompson (1983) is here referred to P. multifaria sp. nov. (p. 1408), and White's (195 l) specimen, which has a ventral anus, is assigned to Reyfria gen. nov. (p. 1417). I agree with Heller and Thompson (1983) that the photograph in Orr (1981:59) is probably not P. arabica. Sharabati (1981:82) figures purported P, 'varicosa' from the Red Sea and Bertsch and Johnson (1981:74-75) figure Hawaiian material, but in both cases there is no information on the ventral pigmentation of the animal, and as P. arabica is characterized by a black median line on the sole of the foot, the identity of these two records must remain in doubt.

In the latter part of the nineteenth century and first half of this century, the confusion over Phyllidia 'varicosa' was considerable. There seem to have been three species involved, two of which have not been found again:

(1) Phyllidia arabica as described above was recorded by a number of authors:

Phyllidia arabica Ehrenberg, 1831: 34; Bergh, 1869: 504. Phyllidia trilineata: Quoy and Gaimard, 1824:419. Phyllidia varicosa: Bergh, 1869: 499.

(2) The second species, Phyllidia varicosa Lamarck 1801 (see p. 1415), differs from the above in having no black line on the foot. It was for long confused with P. arabica but appears to be distinct from it. The following records may be referred to P. varicosa:

Phyllidia trilineata Cuvier, 1804: 267, figs 1-6. Phyllidia varicosa: Vasssi6re, 1912: 84; Gray, 1857: table 312, fig. 6 only.

The specimens of Riippell and Leuckart (1828), I ssel (1869), Sturany (1903), O'Donoghue (1928) (all from the Red Sea), and possibly those illustrated by Gray (1857: table 312, figs 3 and 4), may belong to either of the above, but without knowing whether or not the black line on the sole is present (if indeed they belong to this pair of species) they cannot be confidently assigned to either.

(3) Finally, a third species (see p. 1415) is distinguished by five rows of tubercles and in being much wider than both P. arabica and P. varicosa. The following descriptions clearly refer to this species:

Phyllidia caricosa: Eliot, 1908: 120; Gray, 1857: table 312. fig. 7 only: Quoy and Gaimard, t832: 292, p[. 21, fig. 25.


Neither Eliot (1908) not Gray (1857) described or illustrated the ventral side. Quoy and Gaimard (1832) had several specimens, and stated that there was not always a black line on the sole of the foot; it is possible that their account refers to more than one species, but the specimen illustrated definitely belongs to this specific group.

All three species of this complex have been recorded from the Red Sea, though only P. arabica has been recently collected again, despite the number of expeditions to this region in the past 50 years.

Phyllidia dautzenbergi Vayssi6re, 1912 (Fig. a0D, E)

Phyllidia dautzenberoi Vayssi6re, 1912: 85, pl. 1, fig. 14; Heller and Thompson, 1983: 341, fig. 8H, 1.

Material. (i) 1.9 x 0"5 cm; 23.vi.1983; Jezirat Seba; 1(~15 m depth. (ii) 0-6 x 0.4 cm; 6.vii.1983; Cousteau's habitat, Sha'ab Rumi; 10-15m depth. (iii) 0-6 x0 .3cm (preserved); 16.iii.1984; South Tower Reef; 15 m depth. Two colour slides of one specimen, Red Sea (D. Sharabati).

Description. Specimen (i) is of the general oval phyllidiid shape but the dorsal pattern is not of a typical phyllidiid. The usual large warts are absent, but under a binocular microscope one can see that the dorsum is covered with small pustules. The animal is rather flattened, but there are three longitudinal ridges: a median one (white and more prominent than the others) and one on either side of the median. A black elliptical pattern follows the edges of the lateral ridges, leaving a central oval of an off- white colour. From this black band radiate five pairs of black lines, the posterior pair forming a V shape. There is no single black band extending posteriorly as in the specimens described by Vayssi6re (1912) and Heller and Thompson (1983). Anteriorly, a black band projects forward, crossed by a black line anterior to the rhinophores. The rhinophores are long white finger-like projections, 1.5 mm long in the living animal, and arise from the white region. The anus is located on a raised tubercle slightly to the right of the midline.

Ventrally, the foot is much smaller than the mantle, and colourless. The gill lamellae are off-white and are visible on both sides along the whole length of the foot. The black markings on the dorsum are visible on the hyponotum. The latter has a very sculptured surface, better seen under the microscope: there are five to six ridges running concentrically around the gills. This part of the mantle is very thin and fragile.

The second specimen (ii) is similar to the first, except that there are only four pairs of black lines radiating from the central black ellipse, with the posterior pair again forming a V shape. The anterior black band, crossed by a perpendicular one, is also present (Fig. 10E).

The third specimen (iii) is again very small and has four lines radiating from the central ellipse. It is identical to specimen (ii), but with rudiments of a posterior longitudinal black line (Fig. 10D).

Remarks. This species is so distinctive in its markings from other species of Phyltidia that, despite the variations, all three specimens are clearly conspecific. The first (i) was found just north of the holotype locality (Vayssi~re, 1912). It differs from the holotype specimen only in that the single posterior line is replaced by two, connected apically. The second specimen (ii) was found just north of the Heller and Thompson (1983) locality, and differs from the holotype in having four, not five, symmetrical radiating lines and no posterior line. The third specimen (iii) was found offthe Jeddah coast, it is

1406 N. Yanow

the most northern record, and is the closest to the holotype in appearance. Phyllidia dautzenberoi shows no variation in the anterior mantle decoration, but the posterior black line may be present or absent, and the sides have between three and five paired bands. This species seems to have a very limited geographical range and is probably endemic to the Red Sea. It is illustrated in Sharabati's Red Sea Shells (1984: pl. 37, fig. 6).

Phyllidia melanocera sp. nov. (Figs 2, 10F-I)

Material. (i) 1.7x0-7cm; 17.i.1983; North of creek; 8m depth. (ii) 1.8cm long; 15.vi.1983; Shib Ammar; 17 m depth. (iii) 2.4 × 1.0 cm; 23.vi.1983; Jezirat Seba; 10-15 m depth. (iv) 2-3 x 1-0 cm; 30.vi.1983; Sha'ab Loka; 15 m depth. (v) 2.1 × 0.7 cm; 9.vii.1983; Daedalus Reef; 10 m depth. Six colour slides of two specimens, Red Sea (N. Yonow).

Deposition of types, HOLOTYPE. BM(NH) reg. no. 1985.206/1, specimen (iii) above. PARATYPE: BM(NH) reg. no. 1985.206/2, specimen (i) above.

Description. The preserved specimens slightly resemble preserved Phyllidia pustulosa Cuvier (1804), but the living animals differ considerably. The external character- istics are very uniform and can be seen in six colour slides in my possession. The tubercles are pink and cover most of the dorsal surface. The base colour, sometimes reduced to lines between groups of tubercles, is jet black. One specimen is noted to have been green under water, but unfortunately no photograph was taken.

There are three groups of tubercles: median, lateral, and marginal. The median ones are grouped into three large quadrangles between the rhinophores and anus, with one smaller tubercle group located posterior to the anus. Each quadrangle is composed of four multi-tuberculate pustules. The tubercles making up the four pustules are tiny and rather pointed, interconnected by little ridges; the whole structure, however, is low in profile and the animal itself is flattened (in contrast to P. pustulosa). The lateral tubercles are a little more variable: they range from isolated to interconnected groups, but generally form elongated clumps, alternating with the median clumps.

The pink mantle margin is very thin and there are tiny blister-like tubercles just inside the very edge; the pink coloration is continuous between the two in most cases. The smooth margin is distinct, although there may be one or two tubercles in it (Fig. 2A, B; Fig. 10F-I).

The rhinophores are long and pointed, black towards the tips and grey at the bases (Fig. 2D). They are located on the anterior-most quadrangle of median tubercles which is arranged in a diamond shape; the rhinophore pockets are found just behind and to the side of the anterior end of the diamond, on the white base (pink in living specimens) between the tubercles.

The anus is posterior to the last median quadrangle. In preservative, this region is black on one specimen, white on two specimens where tubercle bases have merged, and half black and half white on two animals.

The anterior edge of the foot is divided longitudinally in three specimens, and the oral tentacles are conical, joined at their bases (Fig. 2C).

Remarks. This species superficially resembles Phyllidia nobilis Bergh (1869) and P. elegans Bergh (1869). It differs in its coloration and the pattern of tubercles. Phyllidia elegans has yellow patches and yellow rhinophores, with a black line on the sole (see also p. 1410); P. nobilis has emerald green tubercles. It differes also from subsequent records of P. nobilis (e.g., Edmunds, 1972) in lacking concentric rings of black pigment and pale tubercles. It is also very similar to P. spectabilis Collingwood (1881). However, in


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FI~3. 2. Phyllidia melanocera sp. nov.: (a) dorsal view of specimen (iv); (b) dorsal view of specimen (iii) (HOLOTYPE); (C) ventral view of specimen (iii) (HOLOTYI'E); (d) Rhinophore of specimen (iv). All drawn with camera lucida.

P. spectabilis the tubercles are composed of up to 12 smaller ones, emerald green in colour, some with white tips; the dusters are irregularly shaped and dispersed, and the margin is also pale and very slightly tuberculate.

Phyllidia monacha sp. nov. (Figs 3, 11A)

Material. 1-4 × 0.75 cm (preserved); 15.xii.1983; Creek; 8 m depth. Deposition of types. HOLOTYPE: BM(NH) reg no. 1985.205. Description. This animal was sent to me with no colour notes or slides. The gills have

retained some pale creamy orange pigmentation, but the colour of the living animal remains unknown.

1408 N. Yonow

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Phyllidia monacha sp. nov.: (a) dorsal view; (b) ventral view. Both drawn with camera lucida.

There is a central black portion with five paired black radiating lines, rather indistinct on the left side, and a posterior ray. The semilunar areas thus created around the sides are white, and tuberculate; the mantle overlap is wide and very thin. In the central region of the black area is an irregularly shaped white tuberculate patch, elongated anteroposteriorly. Anteriorly, two white ellipses mark the rhinophore openings. The rhinophores are completely retracted, although one is just visible and is seen to be white. The anus is a small papilla located half on white, half on black along the midline (Fig. 3A, Fig. l lA).

Ventrally, the black dorsal pigmentation shows through the thin mantle; the hyponotum has a checkered sculpturing. The foot is divided anteriorly to form two rather thick laminae. The tentacles are simple conical structures (Fig. 3B). The gills are pale creamy orange. The tip of the dorsal surface of the foot has a black mark.

Remarks. There is nothing in the literature on Phyttidia that compares with this species. It is unique in its dorsal pattern, and the ventral tentacles and mouth region are different from any illustrated. Although the specimen is small, I have observed that tiny nudibranchs have the distinct adult pattern (e.g., P. arabica, p. 1403. R. ruppelii, p. 1418).

Phyllidia multifaria sp. nov. (Figs 4, 11B-H)

Phyllidia varicosa Heller and Thompson, 1983: 339, fig. 8A-G.

Material. (i) 3cm long; 14.i.1983; Whiskey Reef; 9m depth. (ii) 4.3× 1.9cm; 18.vi.1983; Zubair Islands; 10-12 m depth. (iii) 3.5 x 1-9 cm; 23.vi. 1983; Jezirat Seba; 10- 15m depth. (iv) 2.5 x 1-6cm; 23.vi.1983; Jezirat Seba; 10-15m depth. (v) 3.2 × 2.2 cm; 25.vi.1983; Zubair Island; 10 m depth. (vi) 3.6 × 1.0 cm (preserved); 26.vi. 1983; Seghala Is; 6m depth. (vii) 2.5 x 1.0 cm; 30.vi.1983; Sha'ab Loka; 10m depth. (viii) 3-5 x 1-3 cm; 7.vii. 1983; Abington Reef; 10-15 m depth. Twelve colour slides of five specimens, Red Sea (D. & I. Sharabati, N. Yonow).

Deposition of types. HOLOTYPE: BM(NH) reg. no. 1985.2071, specimen (iv) above. PARATVPE: BM(NH) reg. no. t985.207/2, specimen (viii) above.

Description. There is more variation in relative proportions of pigment in Ph.yllidia multifaria sp. nov. than in previously described species. However, all specimens can be placed in a series from least black to most black; the basic pattern of tubercles remains the same throughout, as does coloration.

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There is a row of median tubercles, extending from a single one just anterior to the rhinophores to, usually, a single tubercle posterior to the anus, which is located between two tubercles, on white ground. In specimens (iv) and (vii) the anus is on the last tubercle. In preservation, the rim around the anus has come to look like a tubercle. The bases of the median tubercles often merge; in some specimens, this median line merges with marginal tubercles posterior to it, and extends to the mantle edge. On each side of the median line, starting with a tubercle behind each rhinophore, is a row of similarly sized tubercles, sometimes merging with the median series posteriorly, sometimes extending posteriorly to the level of the anus.

The three central rows of tubercles are each bordered by black, which separates them from the lateral and marginal tubercles. Lateral to this central series of three rows the black pigmentation, rather than filling in between tubercles, determines and creates the pattern: it extends in tapering rays to the mantle margin, in elongated or angular crescents. The pattern of rays varies somewhat: usually a pair extends forwards from the rhinophores, sometimes with the addition of a median one, or even a pair. There is a series of rays running laterally: up to three major rays (paired in 50~ of the specimens) and any number of smaller dividing ones. There may be one, two or three caudal rays.

The lateral tubercles are smaller than the central ones, and the marginal tubercles smaller again. Between these lateral and marginal tubercles, there may be patches of black.

All specimens but one were pink and black when alive; (i) was recorded as grey. The central three rows of tubercles were tipped with brilliant yellow-orange in nearly all

1410 N. Yonow

cases; (vii) was pink and black only. A few lateral and marginal tubercles were also pigmented. The rhinophores were of the same bright yellow-orange colour: long lamellated curved processes (Fig. 4C). Specimen (viii) had orange tubercles with lemon yellow rhinophores.

Ventrally, all specimens bear a black line on the sole of the foot. They all have a black line of varying thickness on the top side of the foot on each side which fuses posteriorly to form a longitudinal line on the dorsal surface of the foot; in specimen (vii) this line was present, and incomplete, one one side only; in specimen (iv) the line did not extend onto the posterior dorsal surface of the foot. The gills of all specimens have some black pigmentation. Anteriorly, the rim of the foot is thickened with a medial concavity (Fig. 4B); in two specimens, (iv) and (vi), it is sharply notched. In all but two specimens, (ii) and (v), the foot seems to be bilaminate; the top lip is notched medially in four specimens, smooth in one, and lacking in two.

Twelve colour slides of five specimens, in my possession, can be ascribed with some confidence to this species.

Remarks. These specimens agree closely with the P. varicosa of Heller and Thompson (1983, Fig. 8A-G). P~yllidia arabica Ehrenberg (1831) and P. varicosa Lamarck (1801) are distinctive and consistently patterned species characterized by four longitudinal black lines on the dorsum, the first with a black line on the sole of the foot (see p. 1403). An anomalous specimen (2-4 × 1-2 cm; 15.vi. 1983; Shib Ammar; 11 m depth) (Fig. 11I) resembles P. multifaria sp. nov. in its tubercular pattern and general colour. It has orange tubercles and lemon yellow rhinophores, similar to specimen (viii), and the anteriorly divided foot and foot markings are all present. However, the colour pattern and texture are sufficiently different that I hesitate to assign it to P. multifaria sp. nov. It lacks the regularity of the other specimens, and ventrally the tentacles and mouth are very different: the foot is bilaminate anteriorly, and both laminae are divided longitudinally. The tentacles are long and slender. It is quite distinct even from the extreme form at the least black end of the colour scale, and is equally distinct from the extreme specimen figured by Heller and Thompson (1983) (Fig. 8D). More material is required before a definite identification can be made: as the other 15 specimens (including Heller and Thompson, 1983) are clearly similar, the hesitation with regard to this specimen may be well founded.

This species is closest to Phyllidia elegans Bergh (1869) in dorsal pattern: an elliptical black line with rays extending from it. However, P. elegans does not have the black ground colour in the ellipse, nor are the rays tapering, and it has yellow patches. Numerous authors have figured P. elegans from many localities, but there seem to be three forms under the name P. elegans, all different from the Red Sea specimens:

(1) The 'typical' P. elegans Bergh, 1869: 506, table 19, Far East.

(2) Edmunds, 1972: 82, Fig. 4B and Farran, 1905: (as P. nobilis) 345, pl. 3, West Pacific, which lacks the black line on the sole of the foot and the central black ring with rays as seen in specimens of form 1.

(3) Pruvot-Fol, 1956: (part) 64, Fig. IV, la, New Ireland, which is characterized by very little black in two more or less longitudinal rows, and (I assume) no black line on the foot. Of the two specimens she described, one was said to have a black line on the sole of the foot; I assume that this is conspecific with P. elegans sensu strictu (form 1, above) as the illustration (fig. IV, 1 b) of the dorsum agrees with form 1. The other, which is illustrated in fig. IV, 1 a, and which I assume lacks a black line, is distinctive and different from other P. elegans.


The Red Sea species, P. multifaria sp. nov., is distinct from all of these. The consistency of pattern in the 15 specimens discussed here and their consistent difference from all forms of P. eleoans are justification for the introduction of a new specific name. The high level of endemism in the Red Sea further strengthens the case for separating the Red Sea species from those of the Indo-Pacific region.

Phyilidia oceilata undula subsp, nov. (Figs 5, 12A, B)

Material. ( i )4x2 .3cm; 19.vi.1983; Tongue Island; 15m depth. (ii) 1.8 x 1.1 cm; 19.vi.1983; Tongue Island; 20m depth. (iii) 4.4 x 2"5 cm; 19.vi.1983; Tongue Island; 7 m depth. Eleven colour slides of two specimens, Red Sea (D. & I. Sharabati).

Deposition of types. HOLOrVPE: BM(NH) reg. no. 1985.208/1, specimen (ii) above. PARATYPE: BM(NH) reg. no. 1985.208/2, specimen (i) above.

Description. Specimen (ii) (Fig. 5A, 12B): the mantle is covered with globular warts. The pattern is nearly symmetrical about the midline and haphazard towards the edge of the mantle. The midline consists of nine spherical tubercles, two anterior to the rhinophores and the posterior in front of the anus, which is located slightly to the left of the midline. These tubercles were bright orange in the living specimen. There are a few scattered black patches, outlined by white, on each side. Behind each rhinophore a row of four to five tubercles converges towards the anus. The black pattern is approximately symmetrical on either side of these rows. Lateral to these rows on each side are three triangular groups of tubercles arranged around a central globular wart; within each group is a black blotch, surrounded by white, sometimes covering, completely or partially, one or more of the tubercles. These triangular groups of tubercles are partially surrounded, on the median side, by black crescent shapes. Medially, the crescents connect to form waves around the tubercles, again outlined in white. The rest of the mantle edge is covered by much smaller tubercles in no particular arrangement.

The long rhinophores (3 mm in life) were orange, with orange sheaths. The tubercle just posterior to each rhinophore with the midline tubercle and two lateral tubercles form a transverse row in a wide V shape (similar to Cuvier's (1804) anterior transverse row of five tubercles). Anterior to the rhinophores is a U-shaped area of black surrounding the two anterior-most median tubercles and open anteriorly. The base of this U partially overlaps the rhinophore sheaths, where the white outline is missing.

The anus is located on a tubercle which is in the black area. In life, the black was again surrounded by a white band.

Ventrally, the foot is elliptical, 5 mm at the widest part and 15 mm long in preserved material. It is notched at the anterior end, bilaminate, and the oral tentacles are conical structures connected at their bases (Fig. 5B). The foot is grey with a white margin. The gills are grey and limp, more or less triangular in outline. The dorsal pattern is visible through the rather thin mantle edge, and the hyponotum has a cracked checkered appearance produced by concentric lines crossed by transverse radiating lines.

Specimen (i): the pattern is less regular than in the previous specimen. In life, colour was bright orange with the black pattern enclosed in a white band. The anterior black U of specimen (ii) is a ring in this specimen, and the transverse line of tubercles is present. The crescent shapes are less regular, and in addition there are a few small black patches about the midline. The rhinophores were long finger-like projections, 4.5 mm long and orange in the living animal. The sheaths were also orange, touched anteriorly

1412 N. Yonow

T..,. i

/

i:

5 mm

. < . . . . . . . . . . . . . [

......... i : - ~ ']E

i" I r I

C '

B

FIc. 5. Phyllidiu ocellata undula subsp, nov.: (a) dorsal view of specimen (ii) (HOLOTYPE); (b) ventral view of specimen (ii) (HOLOTYPE); (C) rhinophore of specimen (iii). All drawn with camera tucida.

by the black ring, which is not surrounded by a white band. Ventrally, the mouth is partially visible as a flap between the conical oral tentacles and the notched foot. The gills are very flexible triangular flaps, the edges pigmented with black.

Specimen (iii) (Fig. 12A): this large specimen does not differ greatly from the other two; the black crescents are not all interconnected, although the basic pattern is present. Surrounding the black was a slate blue grey band; the tubercles again were bright orange and anteriorly there is a black ring. The oral tentacles are long conical structures, above a bilaminate foot, both laminae of which are notched. The rhinophores are rather fat and digitiform (Fig• 5C).

Remarks. There seems to be a fair amount of confusion over the identity of Phyllidia ocellata Cuvier. In my opinion the differences are consistent enough to enable me to divide them into five colour forms:

(1) Two paired black rings and one anterior.

Phyllidia ocellata Cuvier, 1804: 267, fig. 7 (Mer des Indes). Phyllidiopsis carinata Eliot, 1910: 435, pl. 25, figs 8, 9, 12 (Amirante, Seychelles)• Phyllidia japonica: Lim and Chou, 1970: 134, pl. 16C (Singapore).


(2) Five paired black spots or tubercles.

Phyllidia multituberculata Boettger, 1918: 129, tar. 8, fig. 4 (Aru Islands, Indonesia). Phyllidia baccata Pruvot-Fol, 1957:111, pl. 1, figs 1, 2 (Mauritius). Phyllidia ocellata: Edmunds, 1972: 77, fig. 2 (Seychelles).

(3) Three large paired black rings, as well as two small anterior and posterior ones.

Phyllidia japonica Baba, 1937; nom. nov. for tubereulata Baba 1930. Phyllidia tuberculata Baba, 1930:117, pl. 4, fig. la (Japan). Phyllidia ocellata Nishamura and Suzuki, 1971: pl. 51, fig. 5 (Japan) Rudman, 1981: calendar, November (Australia); Baba and Hamatani, 1975: 176, fig. 4

(Japan); K. Baba (personal communication): two original coloured drawings (Japan).

(4) Scattered small black spots and rings.

Phyllidia japonica: Baba, 1949: 157, pl. 29, fig. 107 (Japan).

(5) Anterior black ring or U-shape, scalloped black markings.

Phyllidia ocellata undula subsp, nov.: J. Heller (personal communication): postcard published by Palphot Ltd, Israel (Red Sea); D. Sharabati (personal communication): colour slides of two specimens (Red Sea).

The ringed forms above (1 and 3), as well as my specimens (5), all have white or slate grey around the black areas, whereas the tuberculate forms (2 and 4) do not. Photographs of two Red Sea specimens (D. Sharabati, personal communication) show this scalloped effect of black pigmentation surrounded by white. A postcard from Israel (published by Palphot Ltd) shows a specimen almost identical to specimen (ii) described above; a photograph of the same species is shown in Sharabati (1984: 87).

Phyllidia oeellata of Pruvot-Fot (1956) is impossible to assign to one of the five forms because there is no black pigment remaining on her material. O'Donoghue (1932) seems to have had two different species: of the larger he says, ' ... a number of blotches of. . . black,.., larger in the middle. . , and smaller at the edges. They are irregularly distributed... '; the smaller was mostly black with a yellow margin and yellow tubercles. Burn (1970) considers the larger to be similar to Pruvot-Fol's (1956) specimen as well as to Cuvier's (1804). However, the blotchy black pattern seems quite unlike the regular ringed pattern of Cuvier's (1804) specimen.

Phyllidia ocellata as presently recognized appears to be a variable entity with a broad geographical distribution. All of the specimens previously reported, despite the fact that they can be assembled into four perhaps taxonomically distinct groups, are alike in their vivid orange ground colour.

The Red Sea material described here represents a single taxonomic entity which differs consistently from all of the other groups detailed above. In view of its geographical separation from the main distribution centre of the 'ocellata complex', it may be regarded as subspecifically distinct.

Phyllidia pustulosa Cuvier, 1804 (Fig. 12C, D)

PhyUidia pustulosa Cuvier, 1804: 267, fig. 8; Heller and Thompson, 1983: 342, fig. 9. Phyllidia rotunda: Lim and Chou, 1970: 134, fig. 17A.

Material. O) 3.3 × 1.7cm; 19.vi.1983; Tongue Island; 12m depth. (ii) 4"4 x 1.7cm; 19.vi.1983; Tongue Island; 20m depth. (iii) 4.5 x 1.7cm; 23.vi.1983; Jezirat Seba; 10-15m depth. (iv) 3.5× 1.7cm; 26.vi.1983; Seghala Is; 6m depth. (v) 4-7× 1.6cm;

1414 N. Yonow

30.vi.1983; Sha'ab Loka; 15m depth. (vi) 5-0×2.5cm; 7.vii.1983; Abington Reef; 10-15m depth. Seven colour slides of four specimens, Red Sea (G. Bemert, D. Sharabati).

Description. All specimens have a dorsal pattern like those of Heller and Thompson (1983): median clusters of tubercles, and lateral and marginal tubercles. There is some variation, however, in the relative proportions of the black and white areas. Specimen (iv) has a greater amount of white: the black pigment is reduced to lines between clusters of tubercles and a few scattered spots within clusters. Specimen (i) (Fig. 12C) has a more equal proportion of the two, and the black areas are quite thick: the median clusters are composed of pale tubercles all separated from one another by black. The rest of the specimens form a gradation between the two extremes of (i) and (iv). The mantle edge does not have the black margin described by Lim and Chou (1970) in their material, but no other author discusses this point.

The anus is consistently located in a black area of the mantle, and the rhinophores each on a separate white area (contrary to Pruvot-Fol, 1957). In all but one specimen the long pointed rhinophores were completely black: in (iv) they were black with grey stalks.

In the preserved animals, the head and oral tentacles vary only slightly. Notes and drawings were made from life only for specimen (iv). The foot is angular anteriorly, and the oral tentacles rise separately from the head: they are tipped with black and have a longitudinal lateral groove. This is seen also in preserved specimens, although the foot of these has a concave edge, and in two cases is notched.

Colour seems to vary. Specimen (i) was noted to be 'pink, with a very pale green tinge'; (ii) was pink with a 'definite green tinge'; (iii) was 'green'; (iv) was 'greyish-pink'; (v) was 'pink'; (vi) was 'black with pink tubercles, green under water'. Preserved, all the tubercles, and a small area surrounding them, are grey white, and the ends of the tubercles themselves are creamy yellow. The black pigment is in dense bands between and around the tubercles, leaving a very narrow band of smooth unpigmented area round each tubercle or cluster.

Remarks. These specimens are most similar in external morphology to those of Cuvier (1804), HeUer and Thompson (1983), and to the Phyllidia rotunda of Lim and Chou (1970). All undoubtedly constitute a single species, and P. rotunda of Guangyu (1983) perhaps belongs also this species. Heller and Thompson (1983) suggest that other Pacific specimens of P. pustulosa represent a separate species, and I am inclined to agree with them. The specimens described in the following references are nearly identical in having groups of tubercles scattered on the dorsum (not divisible into median, lateral and marginal), smaller tubercles as well as smaller clusters, and a greater number of tubercles and clusters. They seem to constitute a single species distinct from Phyllidia pustulosa Cuvier:

Phyllidia pustulosa: Lim and Chou, 1970: 132, pl. 16 (Singapore); Bergh, 1869: 51(~511, table 24 (Timor Sea, Indonesia); Baba and Hamatani, 1975:175-176, fig. 2 (Japan); Guangyu, 1983: pl. 1, fig. 3 (China); Baba, 1936: pl. 3, fig. 1 (Ryukyu Is.); Bertsch and Johnson, 1981:76 (Hawaii).

Phyllidia (Phyllidiella) pustulosa: Baba, 1949: 56, pl. 29, fig. 107 (Japan). Phyllidia sp.: Orr, 1981:60 (Hong Kong).

The specimens of P. pustulosa described by Riippell and Leuckart (1828) and Eales (I 938), both from the Red Sea, have been mentioned by Edmunds (1972) and Heller and Thompson (1983). Their identity is discussed further below (p. 1418).


Phyllidia sudanensis Heiler and Thompson, 1983 (Fig. 6A)

Phyllidia sudanensis Heller and Thompson, 1983: 342, fig. 8J.

Remarks. This species has been described only by Heller and Thompson (1983) who gave no notes on coloration; it was 12 mm long. There is a central series of four groups of pale tubercles, the second and third with a small black patch centrally. These four groups are located on black ground, which is surrounded by a ring of pale tubercles. Surrounding this is a narrow bla~k band, and the margin of the mantle is pale. Nothing is known of the rhinophores or the ventral surface.

Although this species was not found in the present survey, it is included for the sake of completeness. Heller and Thompson (1983) had only one specimen, and no colour slides. Unfortunately, the specimen was sectioned and there is no holotype available for further study.

Phyllidia varicosa Lamarck, 1801 Phyllidia varicosa Lamarck, 1801: 66; Vayssirre, 1912: 84; not Risbec, 1929: 49, figs 10-16. Phyllidia trilineata Cuvier, 1804: 267, figs 1-6; Gray, 1857: table 312, fig. 7 only.

Remarks. This species was well described and illustrated by Cuvier (1804), and the illustration of the ventral surface quite clearly shows no black line; it is otherwise identical to Phyllidia arabica in dorsal pattern. Vayssirre (1912) describes the ventral surface in great detail, and nowhere does he state that there is a black line on the foot. The tortuous synonymy of these two species is discussed at length on p. 1403.

Phyllidia sp. (Fig. 6B)

Phyllidia varicosa: Eliot, 1908: 120; not Quoy and Gaimard, 1924: 419. Phyllidia trilineata: Gray, 1857: fig. 7; Quoy and Gaimard, 1832: 292, pl. 21, fig. 25.

Remarks. This species has been confused with both Phyllidia varicosa and P. arabica. However, from the available descriptions and illustrations it is quite recognizably distinct: it has five lines of whitish (or blueish) tubercles tipped with yellow or orange, and radiating lines of tubercles surrounding this central pattern. It is not known whether it has a line on the foot or not. Quoy and Gaimard (1832) describe a number of specimens which perhaps represented more than one species, but their illustration (1832: 292, pl. 21, Fig. 25) shows a species quite clearly different from P. varicosa or P. arabica. Their earlier reference to P. 'varicosa' (Quoy and Gaimard, 1824) correctly belongs in the synonymy of P. arabica, a species with.a black line on the ventral surface of the foot.

No original material is available for any of the three references above, and the species does not seem to have been recently collected or described. Consequently, the introduction of a specific name must await the collection of specimens suitable for designation as type-material.

Phyllidia ?varicosa var. quadrilineata Bergh, 1905 (Figs 7, 8, 12E)

Phyllidia varicosa var. quadrilineata Bergh, 1905: 181, taf. XII, fig. 41, 42.

Material. 4.1 x 1"7 cm; 7.vii.1983; Abington Reef; 10-15m depth. Description. Under water, the live animal was green with black lines, but at the

surface it became pink with black lines. The black is present around and between the

1416 N. Yonow

E E

A

FlG. 6. (a) Phyllidia sudanensis Heller and Thompson: (b) Phyllidia sp.

tubercles, which are clusters of smaller tubercles. One anterior black ray divides around the rhinophores past the central crest of tubercles. This median line of tubercles originates just anterior to the rhinophores, spreading to encompass them, and extends posteriorly to the anus. There are isolated clumps of tubercles on each side, sometimes touching, elongated anteroposteriorly and hence creating a longitudinal pattern. The black extends perpendicularly to the mantle edge in three paired rays and there are black marks in the resulting angular semi-lunar areas. Posteriorly, the black lines do not meet, stopping at the level of the anus to be diverted to the mantle edge as the third paired rays (Fig. 8).

The rhinophores are black with pink on the anterior face from approximately the middle down to the base (Fig. 7B). They were noted to be 5 mm long. A very thin raised rim surrounds the openings. The anus is situated on a tubercle resembling a volcano, inside of which there is a soft papilla; the hole in this structure is also visible.

Ventrally, some of the dorsal pigmentation is visible, and there is some black pigment. The hyponotum has a checkered appearance. The foot is simple, without markings, slightly angular anteriorly. The tentacles are fused around the mouth, very short, with a lateral groove (Fig. 7A). In life, the foot was white and the hyponotum pink. The gills have a tinge of black along the edges.

Remarks. This specimen seems to be closest in dorsal pattern to Phyllidia varicosa var. quadrilineata of Bergh (1905), from Indonesia. His illustration shows an animal too similar to be dismissed easily. More material is required to establish conclusively its taxonomic identity. It is distinct from both Phyllidia varicosa kamarck and P. arabica Ehrenberg, as well as Phvllidia sp.


5 mm A

1417

FIG. 7. Phyllidia ?varicosa var. quadrilineata Bergh: (a) ventral view; (b) rhinophore. Both drawn with camera lucida.

FIG. 8. (a) Phyllidia sudanensis Heller and Thompson; (b) Phyllidia sp.

1418 N. Yonow

Reyfria gen. nov. Genus. Reyfria gen. nov. Typical of the Phyllidiidae, Reyfria is dorsally tuberculate with black markings, has

no radula, and the gills are present~,aatrally along each side of the foot. This genus is characterized by a ventral anus located posteriorly between the foot and the mantle.

Type-species. Reyfria ruppelii (Bergh 1869). This genus is introduced for R@fria ruppelii (Bergh). Phyllidia pustulosa Cuvier was

described from the Red Sea by Rtippell and Leuckart (1828). Gray (1853) then introduced a new genus, Fryeria, for R/ippell and Leuckart's (1828) specimen of Phyllidia pustulosa Cuvier and one of his own, designating P. pustulosa Cuvier (1804) as the type-species. The most important character of Fryeria Gray is the ventral position of the anus. Bergh (1869) introduced Fryeria ruppelii nom. nov. for Rfippell and Leuckart's species, including Gray (1853, 1857) in its synonymy, implying correctly that this was distinct from Phyllidia pustulosa Cuvier. Howeveir, the type-species of the genus Fryeria Gray (1853) is Phyllidia pustulosa Cuvier, not the Rfippell and Leuckart specimen. Phyllidia pustulosa Cuvier has the anus in a dorsal position, and therefore does not differ from other species of Phyllidia. Fryeria Gray thus becomes a junior subjective synonym of Phyllidia, and a new genus is required for those Phyllidiidae with a ventral, anus.

Etymology. Reyfria is an anagram of Fryeria.

Reyfria ruppelii (Bergh, 1869) (Figs 9, 12F-J)

Fryeria pustulosa: Gray, 1853: 221; Gray 1857: table 312, fig. 3. Fryeria ruppelii Bergh, 1869:514. Fryeria ruppellii Bergh, 1875: 663, pl. 16, figs 5-10. Phyllidia pustulosa: Rfippell and Leuckart, 1828: 36, table XI, fig. la, b. Phyllidiella pustulosa: Eales, 1938: I10, pl. I, fig. 4. Phyllidia varicosa: White, 1951: 252.

Material. (i) 3-5 × 1-7cm; 17.i.1983; North of creek; 5m depth. (ii) 2-0x0-9cm (preserved); 18.vi.1983; Zubair Islands; 10-12m depth. (iii) 2.6 x 1.1 cm; 19.vi.1983; Tongue Island; 4 m depth. (iv) 0.7 x 0-5 cm; 25.vi.1983; Zubair Island; 10m depth. (v) 2,7 x 1.5 cm; 6.vii.1983; Cousteau's habitat, Sha'ab Rumi; 1(~15 m depth.

Type-material. I have been unable to trace original material of this species. In the absence of a recognized holotype, figs 1 a and 1 b in table Xl of R/ippell and Leuckart (1828) are designated as lectotype.

Description. This species is exquisitely patterned, very delicately and strikingly; the tubercle and colour patterns are consistent in all specimens. There are five tubercles down the centre (possibly six in specimen (i) (Figs 9A, 12F)), white with bright orange tips. The first is always just anterior to the rhinophores. The white spreads over an area larger than the tubercle base, and may have tiny tubercles. The white halos of the two posterior tubercles are usually confluent. The second tubercle rows begin with a small tubercle just behind each of the rhinophores; its white base spreads to encompass the rhinophore opening. These second row tubercles are three in number, the second two

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1420 N. Yonow

fitting into the gaps between median tubercles 2 and 3, and 3 and 4. There is no second row tubercle between median tubercles 4 and 5, which are usually connected by their white bases. The third row forms a ring around the first two rows, and it is between the second and third rows that the black pigment divides to form a scalloped pattern around the third row tubercles, with long black rays between them, which contain a few tubercles. At this third row, the white bases are very large, and the proportion of black to white is reversed, with more white than black. The fourth row is also a complete ring composed of smaller tubercles, and here the black again subdivides into more scallops and rays. It may become completely'separate, lacking, or only connected on one side of a loop. The tubercles are all white with bright orange ends. The mantle edge is thin and the margin is edged with orange. The black rays extend across this to the edge. The black regions are often tuberculate, with the tubercles continuous with, for example, the first and second pale tuberculate rows.

The rhinophores are curved finger-like projections, bright orange but of a darker tone than the tubercles, and located on white ground (Fig. 9D).

The anus is located on the midline, ventrally and posteriorly, tucked away between the foot and mantle. No papilla is visible (as in Baba and Hamatani, 1975: Fig. 5D), but it probably contracted upon preservation. The sole of the foot has no black line. The anterior edge of the foot is concave and bilaminate, and the oral tentacles are two simple fat conical structures (Fig. 9C).

Remarks. This species was first described by Rfippell and Leuckart (1828) as P. pustulosa Cuvier. Bergh (1869) introduced F. ruppelii as a nom. nov. for Rfippell and Leuckart's species, recognizing it as distinct from Cuvier's species, and including Gray's (1853, 1857) references in its synonymy. Gray (1853) described Fryeria pustulosa Cuvier from Cosseir (now known as Qussier) on the Egyptian coast of the Red Sea, referring to Rfippell and Leuckart (1828). He also discussed Cuvier's (1804) figure of P. pustulosa, considering it to be a bad illustration. In 1857 Gray listed five species of Phyltidiidae present in the collections of the British Museum (Natural History), illustrating all except his Phyllidia annulata Gray (1853). He illustrated two specimens as Fryeria pustulosa, neither of which fits his earlier description (Gray, 1853). Fig. 1 in table 312 is identical to Cuvier's (1804) illustration, and I am quite certain that it is P. pustulosa Cuvier. Gray's fig. 5 on the same plate is very similar indeed to Rfippell and Leuckart's (1828) figure, and one can detect the presence of a border around the margin. None of the specimens on which Gray's two accounts (1853, 1857) are based have survived. However, R. ruppelii (Bergh) must be founded upon the description of Rfippell and Leuckart (1828).

Examination of Eales' (1938) Phyllidiella pustulosa reveals no dorsal anus; similarly with White's (1951) P. varicosa. Unfortunately, both specimens (BM(NH)) reg. no. 1938. t 1.26.25 and 218; 1914.1.8) have been dissected and the posterior margin of both the foot and the mantle of Eales' specimen have been damaged; however, it may be safely inferred that this specimen belongs to Reyfria ruppelii. In White's specimen, the anus is visible ventrally as a hole.

The Red Sea specimens are consistent and distinct. All have similar dorsal patterns and pigmentation; the orange-yellow margin is just discernible in both the Eales and White specimens, and one can detect a border in Gray's specimen (1857: table 312, fig. 3). There are nine other references to Fryeria ruppelii and Fryeria pustulosa:

Bergh, 1889: 862, taf. 84, figs 19 22 (Mauritius). Eliot, 1906:563 (Maldives). Haas, 1920:140 (Red Seat.


i I

FIG. 10. A~C, P hyllidia arabica: A, specimen (ii); B, specimen (i); C, specimen (iii). D-E, Phyllidia dautzenbergi: D, specimen (iii); E, specimen (ii). F-I , Phyllidia melanocera sp. nov.: F, specimen (iii) HOLOTYPE; G, specimen (iv); H, specimen (v); I, specimen (i). Scale bar = 5 ram.

1422 N. Yonow

A I c BI ° r

D E F

ol FIG. 11. A, Phyllidia monacha sp. nov. B-H, Phyllidia multifaria sp. nov.: B, specimen (iii); C,

specimen (ii); D, specimen (v); E, specimen (viii); F, specimen (iv) HOLOTYPE; G, specimen (i); H, specimen (vii); I, cf. Phyllidia multifaria sp. nov. Scale ba r= 5 mm.


A BI c l 0

E F GI H

J

Fie. 12. A-B, Phyllidia ocellata undula subsp, nov.: A, specimen (iii); B, specimen (ii) HOLOTYPE. C-D, Phyllidia pustulosa: C, specimen (i); D, specimen (iii). E, Phyllidia ?varicosa vat. quadrilineata. F-J, Reyfria ruppelii: F, specimen (i); G, specimen (v); H, specimen Eales; I, specimen (ii); J, specimen (iv). Scale bar = 5 mm.

1424 N. Yonow

Risbec, 1929: 45, fig. 1 (Madagascar). Pruvot-Fol, 1957:114, pl. 1, figs. 3, 4 (Vietnam). Marcus, 1965:279 (Micronesia). Edmunds, 1972: 84, fig. 4A (Seychelles). Baba and Hamatani, 1975: 178, fig. 5 (Japan).

It is certain that these authors refer to a number of very similar but probably distinct species. Most similar to the Red Sea species are those described by Eliot (1906) and Baba and Hamatani (1975), of which only the latter is figured. Both lack the orange margin, but the dorsal pattern is otherwise within the range of variation found in the Red Sea specimens.

Another pair of similar specimens are those of Bergh (1889) and Edmunds (1972), from Mauritius and the Seychelles, respectively. Both are illustrated: ground colour is white, with black scalloped linear markings and scattered black markings around the edge.

A third pair are recorded by Pruvot-Fol (1957) and Marcus (1965), from Vietnam and Micronesia. According to Edmunds (1972), Burn examined the Marcus specimen and found it similar to that of Pruvot-Fol (1957). The dorsal pattern seems indistinct, with possibly three median rows of larger tubercles and scattered smaller tubercles surrounding them. Ground colour is black.

The specimen described by Risbec (1929) belongs to the genus Re3fria, but is quite different from Reyfria ruppelii (Bergh) from the Red Sea and from the three groups above. His illustrations at a first glance look just like Phyllidia arabica, but he states quite clearly that the anus is ventral, located posteriorly between the mantle and the foot.

The specimen of Haas (1920) is impossible to assign to any species, as he simply lists Fryeria pustulosa (Cuvier) from the Red Sea without any description or figure.

Thus, there have been 14 references to the genus Fryeria (excluding Bergh, 1869, 1875) who did not have any original specimens), encompassing about four different species. The Red Sea records may be assigned to Reyfria ruppelii (Bergh) but the others need to be redescribed, probably as distinct species. Fryeria variabilis Collingwood (1881) is probably a Phyllidia: he thought the genital opening was the anus (also considered by Risbec (1956), but as a distinct species). It seems very similar to Phyllidia albo-nigra Quoy and Gaimard 1832.

Discussion Species of the family Phyllidiidae are present in all tropical and subtropical seas and

are often conspicuous and common predators in reef talus habitats. There are approximately 50 described species; most of these are well characterized, but a number are poorly described, not, or only inadequately, illustrated, or have not been found since their original descriptions. Unfortunately, many of the specimens described by earlier authors have disappeared in the progress of time.

Table I lists the species recorded to date from the Red Sea, including the three new species and new subspecies described here. Of the remaining eight species, two (P. varicosa and Phyllidia sp.) are still known only from their original descriptions or illustrations. Their taxonomic status has been considered above and they seem to be valid species.

The phyltidiid fauna of the Red Sea is particularly interesting. Four of the species listed in Table 1 have broad geographical distributions within the Indo-West Pacific region, while P. ~'aric~a var. quadrilineata is described from Indonesia. Three of the


Table 1. Phyllidiidae recorded from the Red Sea.

1425

Species Author Distribution

Phyllidia arabica Phyllidia dautzenbergi

Phyllidia melanocera sp. nov. Phyllidia monacha sp. nov. Phyllidia multifaria sp. nov. Phyllidia ocellata undula

subsp, nov. PhyUidia pustuIosa

Phyllidia sudanensis Phyllidia varicosa Phyllidia sp. Phyllidia ?varicosa

vat. quadrilineata Reyfria ruppelii

Ehrenberg (1831) Vayssi6re (1912)

O'Donoghue (1928) Heller and Thompson (1983)

Heller and Thompson (1983)

Bergh (1873) Heller and Thompson (1983)

Heller and Thompson (1983) Vayssi6re (1912) Eliot (1903)

Rfippell and Leuckart (1828) Vassi+re (1912) O'Donoghue (1928) Eales (1938), White (1951)

Indo-West Pacific Red Sea

Red Sea

Indo-West Pacific

Red Sea Red Sea, Mer des Indes Red Sea, New Ireland Indonesia

Red Sea

previously described species (P. dautzenbergi, P. sudenensis and Reyfria ruppelii) seem to be endemic to the Red Sea, and P. ocellata undula subsp, nov. is clearly a geographical variant of a widely distributed Indo-West Pacific species. If the three newly described species prove to be similarly geographically limited in distribution, then the level of endemism suggested is particularly high. However, it must be emphasized that earlier systematic treatments of the Phyllidiidae are inadequate, that the often confusing variation in colour patterns and the form of mantle tubercles have tended to hinder recognition of conspecific forms, and that further taxonomic precision will only be achieved through careful study of the ecology of these sea slugs. Colour photography and field observations are particularly important in this respect. This preliminary survey of the Red Sea Phyllidiidae has enabled perhaps the majority of the species to be adequately characterized, but a number require further investigation.

Acknowledgements First, my thanks must go to Doreen and Issam Sharabati, who organized this trip

around the Red Sea on their yacht, the Ibn Batuta, and to my parents for their encouragement and support in my scientific endeavours. I am grateful to Professor J. S. Ryland, Marine Research Group, University College of Swansea, for the provision of facilities; to Mr G. Bemert and Mr and Mrs I. Sharabati for photographs; to Ms K. Way and Miss A. Thomson (British Museum (Natural History)), Miss A. Trew (National Museum of Wales), and Mr K. S. Williams (University of Bristol) for their generous assistance. Finally, I would like to thank Dr P. J. Hayward for his criticism and encouragement.

1426 N. Yonow

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Phyllidiidae from the Red Sea 1986

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