Pestarella, Ngoc-Ho, 2003

Ngoc-Ho, Nguyen, 2003, European and Mediterranean Thalassinidea (Crustacea, Decapoda), Zoosystema 25 (3), pp. 439-555 : 475-486

Genus Pestarella n. gen.

Astacus Petagna, 1792: 418 (part).

Cancer Olivi, 1792: 51 (part).

Callianassa Leach, 1814: 400 (part).

TYPE SPECIES. — Astacus tyrrhenus Petagna, 1792 , by present designation.

SPECIES INCLUDED. — Astacus tyrrhenus Petagna, 1792 , Cancer candidus Olivi, 1792 , Callianassa rotundicaudata Stebbing, 1902 , Callianassa convexa de Saint Laurent & Le Loeuff, 1979 , Callianassa whitei Sakai, 1999 .

ETYMOLOGY. — The genus is named in honour of Dr Otto Pesta. The gender is feminine.

DIAGNOSIS

Carapace with dorsal oval, rostrum approximately triangular, rostral spine absent. Abdominal somite 2 about equal in length with abdominal somite 6; abdominal somites 3-5 with lateral tufts of setae. Telson rounded in posterior half.

Eyestalk short, flattened dorsoventrally; cornea distinct, disk-shaped. A1 peduncle as long or long- er than that of A2. Mxp1 epipod truncate anteriorly; Mxp3 operculiform, a row of numerous spinules on mesial surface of ischium, dactylus digitiform; exopod absent.

Exopod on Mxp1-2; single arthrobranch on Mxp2, paired arthrobranch on Mxp3 and P1-5.

P1 unequal in male and female, major with meral hook. P3 propodus oval, no proximal heel. P4 propodus narrow; P5 subchelate. Male Plp1 and Plp2 absent; female Plp1 uniramous, female Plp2 biramous, Plp3-5 biramous, foliaceous in both sexes, appendix interna small, projecting from mesial border of endopod. Uropodal exopod with dorsal plate terminating in posterior setal row near or apart from posterior border.

REMARKS

The late Dr Ray Manning had intended to separate the two European species Callianassa candida and C. tyrrhena into a genus dedicaced to Dr Otto Pesta. The present generic name fulfills that intention. The genus nevertheless has not hitherto been erected and the names Pestaina truncata and Pestaina candida used by Öksnebjerg (2000: 78) are nomina nuda.

In the subfamily Callianassinae of the Callianassidae , the new taxon is related to Callianassa . It differs by the telson rounded in posterior half, the operculiform Mxp3, and the absence of Plp1 and Plp 2 in males.

Apart from Pestarella whitei n. comb., the four other species of this genus were placed in a small group within Callianassa by de Saint Laurent & Le Loeuff (1979: 96). They inhabit eastern Atlantic: three European species, P. convexa n. comb. from Senegal and P. rotundicaudata n. comb. from South Africa.

Pestarella candida ( Olivi, 1792) n. comb. ( Fig. 12 )

Cancer candidus Olivi, 1792: 51 , pl. 3 fig. 3.

Gebios Davyanus Risso, 1822: 243 .

Callianassa subterranea forma pontica Czerniavsky, 1884: 81 (part).

Callianassa (Callichirus) Pestae de Man, 1928a: 34 , pl. 9 figs 16-16E; 1928b: 29, 111.

Callianassa algerica Lutze, 1938: 168 , figs 22-26a, b, 27.

Callianassa candida – Giordani Soika 1943: 83 (part); 1946: 944. — Lewinsohn & Holthuis 1986: 20. — ° Koukouras et al. 1992: 223. — Dworschak 1992: 194; 1998a: 1535, figs 1, 3, 5-9 (part), tabl. 1; 1998b: 143. — Noël 1992: 81. — Falciai & Minervini 1996: 147, 4 figs. — d’Udekem d’Acoz 1996: 54; 1999: 154. — Sakai 1999a: 14, fig. 2. — *González- Gordillo et al. 2001: 279. —?* Lindley et al. 2001: 46. — ° Türkay 2001: 289. — Dworschak 2002: 64, figs 1a, b; fig. 2; tabl. 1 (part).

Callianassa laticauda – Heller 1863: 203.

Callianassa (Callichirus) laticauda – Pesta 1918: 204. — Bouvier 1940: 103, fig. 69.

Callianassa pestae – Manning & Števčić 1982: 295. — García-Raso 1985: 21. — Froglia & Grippa 1986: 261.

Callianassa pestai – Holthuis 1953: 95, fig. 3. — *Dolgopolskaia 1969: 316, pls 32-34. — Mikashavidze 1981: 1417, fig. 2. — Türkay et al. 1987: 92.

Callianassa (Callichirus) pestai – Zariquiey Alvarez 1968: 230. — Kattoulas & Koukouras 1974: 344, fig. 1.

Callianassa pontica – *Dolgopolskaia 1954: 179, figs 1-4. — Števčić 1972: 102. — de Saint Laurent & Božić 1976: 24, figs 5, 13, 21, 32. — Beaubrun 1979: 84, figs 58, 59, 68-70. — Monchartmont 1979: 71. — García Raso 1983: 323, fig. 3. — Thessalou- Legaki & Zenetos 1985: 311. — °Thessalou-Legaki 1986: 182.

Callianassa (Callichirus) pontica – Caroli 1946: 71; 1950: 190. — Makarov 1938: 73, 297, figs 27, 28. — Băcescu 1967: 231, fig. 105.

Callianassa (Callichirus) stebbingi – Pesta 1918: 101 (part), fig. 63.

Callianassa subterranea – Giard & Bonnier 1890: 362, figs 1, 3.

Callianassa tyrrhena – Forest & Guinot 1956: 31.

Pestaina candida – Öksnebjerg 2000: 78 nomen nudum. Non Callianassa pestae – Lutze 1937: 6, figs 10-21; 1938: 167, figs 10-21 (= C. subterranea ( Montagu, 1808)) .

TYPE MATERIAL. — Neotype:, from Kuvi Bay , south of Rovinj, Istria, Croatia, selected by Sakai 1999a ( SMF 23572).

MATERIAL EXAMINED. — France. Banyuls, ex. collection Sollaud, No. ES 30, 1932, 2 ovig. cl. 14 and 16 mm, tl. 52 and 58 mm (figured) ( MNHN Th 1358). — Villefranche, Old Harbour, Travailleur, 1886, 1 cl. 13 mm ( MNHN Th 123). — Îles Cerbicales, Corsica, C. Monniot coll., VII.1994, 1 cl. 10.5 mm ( MNHN Th 1348).

Portugal. Algarve, mudflat near Praia de Faro, just W of Faro, along lagoon, behind dunes, 2.XI.1974, excursion RMNH, stn 79, 1 ( RMNH D 36281 as Callianassa pestae ).

Spain. Cadaqués, NE Spain, VI.1950, leg. J. Fortuny, collection R. Zariquiey Alvarez, 2 ( RMNH D 35799b as Callianassa pestae ).

Italy. Naples, collection E. Caroli, received V.1959 from Zoological Station Naples, 7, 6 ( RMNH D 13005, as Callianassa pestae ). — Gulf of Naples, Italy, donated by E. Caroli, 2 ( RMNH D 6587 as Callianassa pontica ). — Lido di Staranzano, intertidal, P. Dworschak coll., 9.X.1984, 1, 2 ( NHMW 6788). — Punta Sabbioni, Venice Lagoon, mud flat, P. Dworschak coll., 25.III.1989, 2, 1 ( MNHW 6761). — Porto Cesareo, P. Parenzan coll., 1969, 1 cl. 10 mm ( MNHN Th 214).

Adriatic. Norman coll., 1 ( NHML 1997.322). — Kuvi, Rovinj, P. Dworschak coll., 11.VII.1997, 1, 1 ovig. ( NHMW 15369).

Slovenia. Strunjan (near Piran), intertidal, P. Dworschak coll., IX.1985, 1, 1 ( NHMW 6789).

Greece. Kreta, Stavros, C. d’Udekem d’Acoz coll., 4.VII.1987, 1, 1, 1 juv. (d’Udekem d’Acoz), 12.VII.1987, 2, 1 (d’Udekem d’Acoz) ; Kato Zakro, 21.VII.1987, 1 (d’Udekem d’Acoz). — Amvrakikos Kolpos, SE Koronissia, 2.5- 3 m, C. d’Udekem d’Acoz coll., 16.VII.1993, 1 (d’Udekem d’Acoz). — SE Peloponesos, Epidavros Limera, C. d’Udekem d’Acoz coll., 3.VII.1986, 1 (d’Udekem d’Acoz). — Khoklakos, Rhodos, don. Zoological Museum, Tel Aviv University, Israel, 13.X.1970, 2 ( RMNH D 35802, as Callianassa pestae ) .

Israel. Haifa Bay, 11. VI.1963, leg. C. Lewinsohn, 1, 1 ( RMNH D 35803 as Callianassa pestae ).

Algeria. Castiglione, leg. R. Dieuzeide , V.1939 (material previously identified by Lutze as Callianassa algerica ), 3 cl. 7.5- 9 mm, 1 cl. 9 mm ( MNHN Th 96) ; 21.VI.1931, 1 cl. 6.5 mm ( MNHN Th 97) ; 23.IV.1929, 1 cl. 5 mm, 1 cl. 6 mm, 1 juv. cl. 4 mm ( MNHN Th 98) .

Morocco. Near Tanger, G. Buchet coll., 1901, 3

cl. 7.5- 10 mm, 2 (1 ovig.) cl. 8 and 12 mm (MNHN Th 804).

Tunisia. Gulf of Tunis, R. B. Manning coll., 16.VIII.1973, 1, 1 cl. 9 mm, tl. 32 mm (figured) and 56, 37 cl. 6-9 mm, 40 juv. ( MNHN Th 633) ; 1978, 1 cl. 12 mm ( MNHN Th 626). — Les Bibans, Cherbonnier coll., 18. V.1955, 1 cl. 7 mm (assigned to C. tyrrhena by Forest & Guinot 1956) ( MNHN Th 122). — Salammbô, Ingle & Manning coll., 18.II.1974, 1 ( NHML 1997.320 ) ; on sand flat, 1 m or less, R. B. and B. A. Manning coll., 23.XI.1972, 7 juv. cl. 2.5- 5 mm ( MNHN Th 316). — Punic Port, Salammbô, M. de St Laurent coll., IX.1973, 7 cl. 6- 13 mm, 2 cl. 7 mm, 5 juv. ( MNHN Th 419) ; northern Punic Port , sandy mud, 5-20 cm, Manning, Forest and de Saint Laurent coll., 7.VIII.1973, 12

cl. 6-10 mm, 3 cl. 6.5- 9 mm (MNHN Th 712).

DISTRIBUTION. — This species is mainly confined to the Mediterranean with a few specimens in nearby areas, Cadix Gulf in southern Spain ( García Raso 1983, 1985) and Algarve, in southern Portugal.

DIAGNOSIS

Rostrum ( Fig. 12A ) with blunt tip. Telson ( Fig. 12H ) with proximal width about 1.3-1.5 times median length.

Eyestalk ( Fig. 12A ) with blunt tip or slightly pointed mesially, cornea disk-shaped, dorsolateral or dorsal. Peduncles of A1 and A2 of approximately same length. Mxp1 ( Fig. 12E ) as figured. Mxp3 ( Fig. 12F, G ) propodus about 1.5 times as long as wide, lower border convex; ischium-merus length about 1.4-1.5 times merus width, dactylus digitiform, 2.5 to 3 times as long as wide.

Major P1 ( Fig. 12D ) merus with lower border straight or slightly convex; meral hook pointed distally, with denticules on lower border, posterior to distal point. Minor P1 ( Fig. 12C ) unarmed on lower border, carpus slightly narrower proximally than distally, about as wide as propodus and 1.6-2 times its length. Plp3-5 ( Fig. 12I ) with small appendix interna. Uropodal exopod ( Fig.12H ) longer than distal width, dorsal plate with distal row of setae well apart from posterior border; endopod oblong, about 1.3-1.5 times length of telson, posterior border rounded.

Colour

Body creamy, tailfan sometimes yellowish, chelipeds always white (Dworschak pers. comm.).

Size

Medium, of cl. 6-16 mm, ovigerous of cl. 12- 16 mm, tl. 50-58 mm.

ECOLOGY

Pestarella candida n. comb. was found in the Gulf of Tunis, in muddy sand with or without Zostera , at 20-30 m depth, together with P. tyrrhena n. comb. and Upogebia pusilla (de Saint Laurent & Manning 1982) ; but it occurs higher up in relation to intertidal level and in more muddy sediments ( Dworschak 1992). It is also common in coarse sand or mud in the intertidal and shallow subtidal, at 7-9 m depth ( García Raso 1983; Thessalou-Legaki & Zenetos 1985). Its burrow and functional morphology are reported in Dworschak (2002).

REMARKS

This species has been known in the past under Callianassa pontica or Callianassa pestae ( pestai ) and the priority of these names was discussed by Holthuis (1953) and de Saint Laurent & Božić (1976). In 1986, Lewinsohn & Holthuis pointed out the priority of the name candida . Although both Dworschak (1992) and d’Udekem d’Acoz (1999) found Lewinsohn and Holthuis’ arguments (based on the white chelipeds of the species) not quite convincing, especially given the rudimentary work by Olivi (1792), the name candida was used by both, and also in this work, for the sake of nomenclature stability. Sakai (1999a) selected a neotype ( SMF 23572) for the species.

As reported by de Saint Laurent & Božić (1979: 26), a part of the material assigned by Lutze (1938) to Callianassa algerica and now deposited in MNHN (MNHN Th 96-98), belongs to the present species. The identity of the two forms is confirmed.

Variations occur in: 1) the eyestalk distal tip is blunt or mesiodistally produced; 2) the lower border of P1 merus is straight or convex; 3) the length ratios of carpus/propodus/fingers vary in both P1; and 4) the meral hook of the major cheliped is often relatively large with the lower bor- der straight and a distinct pointed distal tip ( Fig. 12B ) or smaller with the lower border convex and the distal tip more discrete ( Fig. 12D ).

Besides characters given in the key, additional details differentiating P. candida n. comb. and its two congeners are presented under these latter species.

Pestarella tyrrhena ( Petagna, 1792) n. comb. ( Figs 13 ; 14 )

Astacus tyrrhenus Petagna, 1792: 418 , pl. 5, fig. 3.

Callianassa laticauda Otto, 1821: 11 ; 1828: 345, pl. 21 fig. 3. — Stalio 1877: 107. — Stossich 1880: 206. — Carus 1885: 489. — Giard & Bonnier 1890: 365. — Caroli 1940: 73; 1946: 71; 1950: 189. — Zariquiey Alvarez 1946: 106. — Holthuis 1953: 91, fig. 2. — Picard 1957: 48. — ° Faure 1970: 773.

Callianassa sp. – Stebbing 1893: 184.

Callianassa (Callichirus) Stebbingi Borradaile, 1903: 547 . — Selbie 1914: 100, pl. 14, figs 8-10.

Callianassa candida – Giordani Soika 1943: 83 (part). Callianassa (Callichirus) laticauda – Borradaile 1903: 547. — Pesta 1918: 204. — de Man 1928a: 33, pl. 8 fig. 15-15d; 1928b: 28, 91, 92, 111. — Bouvier 1940: 102, fig. 69. — Gurney 1944: 82, figs 1D, 10, 11. — O’ Céidigh 1962: 164.

Callianassa Stebbingi – Southern 1915: 35. — Steinitz 1933: 147.

Callianassa stebbingi – Balss 1926: 27. — Schellenberg 1928: 78, figs 59, 60. — ° Bodenheimer 1937: 281. — Lutze 1937: 6, figs 1-6; 1938: 165, figs 1-9. — Ott et al. 1976: 61, fig. 2 (part), tabl. 2, pls 1, 2 (part).

Callianassa (Callichirus) stebbingi – Pesta 1918: 201 (part).

Callianassa subterranea – H. Milne Edwards 1837a: 309; 1837b: 130, pl. 48 fig. 3-3 e. — Heller 1863: 202, pl. 6 figs 9-11. — Ortmann 1891: 55, pl. 1 fig. 10.

Callianassa tyrrhena – Holthuis 1947: 320, fig. 1; 1953: 91, 93, fig. 1; 1958: 8, fig. 4; 1977: 57; 1991: 252, 264, figs 457, 458. — Holthuis & Gottlieb 1958: 62 (part). — *Bourdillon-Casanova 1960: 108. — Forest & Gantès 1960: 348. — Pérès & Picard 1964: 54. — Picard 1965: 59. — Bourdon 1965: 15. — Forest 1967: 6. — Le Gall 1969: 398, figs 1-4, tabls 1-5. — ° Pastore 1976: 107. — * Thiriot 1976: 350, 367. — de Saint Laurent & Božić 1976: 22, figs 4, 12, 20, 31. — ° Glaçon 1977: 36. — Beaubrun 1979: 84, 90, figs 55, 60, 61, 66, 67. — de Saint Laurent & Le Loeuff 1979: 53. —? Domenech et al. 1981: 149, figs 35, 36a, b. — ° Kocataș 1981: 162. — Adema et al. 1982: 26, fig. 7ac. — Manning & Števčić 1982: 295. — García Raso 1983: 323, fig. 2. — Riedl 1983: 483, pl. 177, 2 figs. — Holthuis & Heerebout 1986: 62, fig. 83a. — d’Udekem d’Acoz 1986: 101, fig. 2; 1989: 176; 1996: 54; 1999: 155. — Thessalou-Legaki 1986: 182; *1990: 659, figs 1-5, tabls 1-4. — Dworschak 1987a: 421, fig. 1 (part); 1992: 206; 1998a: 1535, figs 1, 2, 4, 4-9 (part), tabl. 1; 1998b: 143, figs 1-4; 2000: 155, figs 1- 6. — Thaker & Haritos 1989a: 63, figs 1-8; 1989b: 199, figs 1-7. — Moyse & Smaldon 1990: 520, fig. 10.13 (part). — Števčić 1990: 217. — Pérez Sánchez & Moreno Batet 1991: 189, 1 fig. — de Vaugelas 1991: 56, photo 1; 1998: 3, figs 1-3. — ° García Raso et al. 1992: 258. — ° Koukouras et al. 1992: 223; 1993: 195. — Noël 1992: 81. — Gruner 1993: 997, fig. 619. — ° Mayoral et al. 1994: 236. — ° Froglia 1995: 7. — Gonzalez Pérez 1995: 136, photo 78. — Hayward et al. 1995: 432, fig. 8.52 (part). — Falciai & Minervini 1996: 147, 4 figs. — Ingle 1997: 78, fig. 7.3. — Thessalou-Legaki & Kiortsis 1997: 435, figs 1-4. — Thessalou-Legaki et al. 1997: 439, figs 1-3, tabls 1-3. — Sakai 1999a: 21. — Dworschak et al. 2000: 302. — *González-Gordillo et al. 2001: 279. — Livory 2001: 33. — * Martin 2001: 78, 1 fig. — * Lindley et al. 2001: 46. — Markham 2001: tabl. 2. — ° Türkay 2001: 289. Callianassa (Callichirus) tyrrhena – Zariquiey Alvarez 1968: 230. — Monchartmont 1979: 72. — Neves 1974: 13, fig. 4.

Pestaina tyrrhena – Öksnebjerg 2000: 78 nomen nudum.

Non Alpheus Tyrhenus – Risso 1816: 94, pl. 2 fig. 2 (= Pontonia pinnophylax ( Otto, 1821)) .

Callianassa tyrrhena – Risso 1827: 54 (= Pontonia pinnophylax ( Otto, 1821)) . — Forest & Guinot 1956: 31; 1958: 10 (= Pestarella candida ( Olivi, 1792) n. comb.). — Holthuis & Gotlieb 1958, fig. 13 (= Callianassa subterranea ( Montagu, 1808)) .

Callianassa stebbingi – Gottlieb 1953: 440 (= Gourretia denticulata ( Lutze, 1937)) .

Callianassa (Callichirus) stebbingi – Pesta 1918, fig. 63 (part) (= Pestarella candida ( Olivi, 1792) n. comb.).

Callianassa (Callichirus) tyrrhena – Cottiglia 1983: 80, figs 27a, 30, 31 (= Pestarella candida ( Olivi, 1792) n. comb.).

TYPE MATERIAL. — Whereabouts unknown.

MATERIAL EXAMINED. — North Sea. 53°10’N, 2°15’E, depth 20-22 fms, 16.IV.1963, don. Netherlands Institute of Sea Research, received VII.1973, 2 ( RMNH D 29676). — Southern North Sea, 52°11’N, 4°21’E, 6-18.IX.1965, leg. D. Eisma, don. Netherlands Institute of Sea Research, 1 (poor condition, RMNH D 35804).

The Netherlands. Diepe Gat, 17 fms, in stomach Raja clavata (L.), don. Netherlands Institute of Sea Research, 1 poor condition ( RMNH D 6203 ). — 30 miles NW of Ijmuiden, 7.VII.1950, leg. et don. Zoological Station Den Halder , 1 ( RMNH D 7381 ). — Province of Zuid Holland, off Scheveningen , sand pumped from quadrant, 52°10’00.46”N, 4°02’28.81”E- 52°12’00.02”N, 4°06’00.00”E- 52°12’49.50”N, 4°04’52.08”E- 52°10’59.90”N, 4°01’20.86”E, 2 broken, several P1 ( RMNH D 40764 ) ; c. 30 km off Scheveningen, sand pumped on the beach, H. L. Strack coll., 31.III.1991, 2 chela ( RMNH D 40769 ). — Diepe Gat near Den Helder, 17 fms, 16.II.1951, from stomach of Mustelus mustelus (L.), leg. et don. Zoological Station Den Halder, 1 spec. ( RMNH D 7634 ) .

Great Britain. Jersey, Norman coll., 2, 2 ( NHML 98.5.7.751.52). — Jersey, purchased 30. V.1950, 3, 1 ovig. ( NHML 1950.8.22.13- 17). — St Clement, Jersey, J. Sinel coll. (types of Callianassa stebbingi ), 1, 1 ( NHML 84.18). — St-Aubius Fort, Channel Islands, Jersey, R. Pike coll., 1 ( NHML 1951.9.4.1-2).

France. Brittany, Pointe du Château, N of Tréguier , P. Noël coll., 28.II.1994, 2 cl. 9 and 15 mm, tl. 36.5 and 55.5 mm ; 1 cl. 14 mm, tl. 57 mm (figured) ( MNHN Th 1346). — St-Malo Bridge, N. Rabet coll., 1.VIII.1992, 1 cl. 13 mm, tl. 54.5 mm (figured) ( MNHN Th 1345). — Réville, Normandie, J. I. Legrand coll., VII.1974, 1 cl. 12.5 mm ( MNHN Th 422). — Wimereux, leg. R. P. Dolfus, 1905, 2 cl. 12.5 mm, 1 cl. 10 mm, and 1 cl. 12.5 mm, tl. 53 mm (figured) ( MNHN Th 677). — Noirmoutiers, fine sand, Y. Gruet coll., 1.VII.1973, 1 cl. 15 mm ( MNHN Th 600) ; 2.VII.1973, 1 ovig. cl. 13.5 mm ( MNHN Th 601) ; 13.VIII.1972, 1 cl. 6 mm ( MNHN Th 602) ; 1.VII.1973, 1 ovig. cl. 14 mm ( MNHN Th 603). — Concarneau, Baron of St-Joseph coll., 1911, 1 ovig. cl. 13 mm ( MNHN Th 227) ; Cabellou Beach, J. Forest coll., VIII. 1957, 1 cl. 10 mm ( MNHN Th 781). — Rade of Brest, Point of Bindy, P. Noël coll., 23.I.1992, 2 cl. 5.5 and 8.5 mm ( MNHN Th 1347). — Coutainville, Lessertisseur coll., IX.1962, 1 cl. 10.5 mm ( MNHN Th 135). — Roscoff, F. Gentil coll., 1972, 1 cl. 12 mm, 1 cl. 9 mm ( MNHN Th 221). — St-Servan , Brittany, Monot coll., 1 cl. 12.5 mm ( MNHN Th 133) ; unknown coll., IV.1924, 1 cl. 10 mm ( MNHN Th 555) ; E. Fischer coll., IX.1929, ex. coll. Sollaud, No. ES 33, 2 cl. 13 and 14.5 mm, 1 cl. 13.5 mm ( MNHN Th 1364). — St-Malo, Vaillant coll., 2 cl. 7.5 and 8.5 mm ( MNHN Th 132). — Dinard ?, about 4.88 gr. W, 54.5 gr. N, 1930 (?), ex coll. Sollaud, No. ES 28, 1 cl. 12.5 mm ( MNHN Th 1356). — Lancieux , about 4.99 gr. W, 52.2 gr. N, V.1927, ex coll. Sollaud, No. ES 36, 1, infested by isopod, with left Plp 1 present ( MNHN Th 1367). — Audierne , Finistère , Dr Piton don., No. 10.1924, 3 (2 ovig.) cl. 14-15 mm, 1 broken ( MNHN Th 220). — Chausey Island , western Atlantic , northern area, flats exposed at low tides, A. Crosnier and P. Clark coll., IX.1992, 1, 4 ( NHML 1992.1086 ). — Rocheneuf, between St-Malo and Mont-St-Michel, Ille-et-Vilaine, 20.VIII.1958, collection H. Nouvel, 1 ( RMNH D 35806). — Mediterranean, 1 cl. 13.5 mm ( MNHN Th 224) .

Portugal. Olhão, C. d’Udekem d’Acoz coll., 18.VII.1988, 2 cl. 11 and 12 mm (d’Udekem d’Acoz). — Algarve, lagoon of Manta Rota, C. d’Udekem d’Acoz coll., 20.VII.1988, 1 cl. 8 mm, 3

cl. 9-11.5 mm (d’Udekem d’Acoz). — Algarve , mudflat near Praia de Faro, just W of Faro, 2.XI.1974, excursion RMNH, stn 79, 1, 3 ( RMNH D 36278) ; 1, 3 ( RMNH D 36279) ; 3.XI.1974, excursion RMNH, stn 80, 3 ( RMNH D 36280) .

Spain. Playa Gentils in Cala de Cutip , N of Cabo de Creus, NE Spain, 12.VIII.1950, leg. et don. L. B. Holthuis, No. 127, 1 juv. ( RMNH D 6735 ). — Playa Jonquet, N of Cadaqués, NE Spain, c. 0.5 m, sand and stones, 13.VIII.1950, leg. et don. L. B. Holthuis, No. 128, 1 juv. ( RMNH D 6736 ). — Cala Junques, N of Port Lligat near Cadaqués , sand and stones, c. 0.2 m, 20.VIII.1959, leg. L. B. Holthuis, 1 juv. ( RMNH D 15214 ). — Ria de Arosa, N of Cabo Cruz, stn 0.32, 23.VII.1962, low water mark, excursion Rijksmuseum van Natuurlijke Historie, 2 ( RMNH D 18544). — Ria de Arosa, Peninsula Chazo, Playa Barraña, sandy beach with some rocks, in burrows in sand between low and highwater marks, 7.VII.1963, excursion RMNH, stn 0.50, 2, 1 ( RMNH D 20539) ; Ria de Arosa , Playa de Barrana, 10.VIII.1964, excursion RMNH, stn o.126, 1, 1 ovig. ( RMNH D 23659 ) ; mouth of Ria de Arosa , Peninsula del Grove , Punta San Vicente, littoral, burrowed in sand, 1.VI.1963, excursion RMNH, stn 0.44, 1 ( RMNH D 20540). — Bay between Cadaqués and Cabo de Creus, Rierca de Junquet, 19.VIII.1950, leg. R. Zariquiey Alvarez, No. 1331, 3 ( RMNH D 35805). — VIII.1951, leg. R. Zariquiey Alvarez, No. 1330, 1, 1 juv. ( RMNH D 35807) .

Italy. Porto Cesareo, P. Parenzan coll., 1969, 1 ovig. cl. 8.5 mm ( MNHN Th 207). — Naples, 1920, C. Gravier, 1 cl. 13.5 mm ( MNHN Th 125); Norman coll., 2, 2 ( NHML 98.5.7.748-50); 1 ( NHML 1910.2.4.91); 1, 1 ovig. (bought I.1925) ( NHML 1950.1.2.113-114); 1876, leg. J. G. de Man, 3, 1, 2 ovig. ( RMNH D 938); collection E. Caroli, received V.1959 from Zoological Station Naples, 23, 12, 4 ovig.

( RMNH D 13007). — Gulf of Naples ; V.1924, leg. et don. G. Stiasny, 1 ( RMNH D 2391 as Callianassa laticauda ) ; V.1957, leg. et don. J. H. Stock, Zoologisch Museum, Amsterdam, 4 ( RMNH D 15213 ) .

Greece. Kreta, Kato Zakro, C. d’Udekem d’Acoz coll., 21.VII.1987, 1 cl. 6 mm, 1 ovig. cl. 9 mm (d’Udekem d’Acoz). — Tersanas, W of Stavros, 2.VII.1987, 1 cl. 7 mm, 1 cl. 6 mm (d’Udekem d’Acoz). — Stavros, 4.VII.1987, 1 ovig. cl. 8.5 mm, 1 juv. (d’Udekem d’Acoz); 11.VII.1987, 2 cl. 6 and 9 mm, 1 juv. (d’Udekem d’Acoz). — Ormos Livadi, 9.VII.1987, 3 cl. 6-10 mm, 1 ovig. cl. 9 mm (d’Udekem d’Acoz). — W of Goudouras, 19.VII.1987, 4 cl. 6-11 mm (d’Udekem d’Acoz). — Island of Lesbos, Gavatas, C. d’Udekem d’Acoz coll., 8.VII.1992, 1 cl. 6.5 mm (d’Udekem d’Acoz). — Aegean Sea, Calypso , stn 779, NE of Salamina, 90-100 m, 21.IX.1955, Saronikos G., 1 (A.U.TH P 3505).

Israel. Tantura, 6.VII.1952, don. Tel Aviv Institute of Natural Sciences, 1 ( RMNH D 13800). — Mediterranean coast near Tantura, S of Haifa, in rockpools, 0-2 m, with Pro-nox fish poison, 3. V.1962, leg. A. Ben-Tuvia, E. Gilat and L. B. Holthuis, 2 ( RMNH D 18545).

Morocco. 1 km S of Oued Yquem, 22 km S of Rabat, along highway Rabat-Casablanca, rocky coast with sandy lagoon, 19.X.1974, excursion RMNH, No. 39, 9, 2 ( RMNH D 36277). — SPMOPO 1974, stn 034, Morocco, atlantic coast, on highway Rabat-Casablanca, south of Rabat, 19.X.1974, Exc. RMNH 1974, 1 ( RMNH D 38482). — SPMOPO 1974, stn 038, 0-0.2 m deep, rock pools. 20.X.1974, Exc. RMNH 1974, 1 ( RMNH D 38483).

Tunisia. R. B. Manning coll., 12.VIII.1973, 13 juv. ( MNHN Th 1344). — Salammbô, grass flats, in sand, Manning, Forest and de Saint Laurent coll., 19.VIII.1973, 3 cl. 7-12.5 mm, 3 cl. 12- 14 mm ( MNHN Th 711).

Guinea. Témara, îles de Los, H. Gantès coll., No. 852, 14.III.1952, 1 cl. 11 mm ( MNHN Th 131); No. 1696, 14. V.1953, 1 cl. 9.2 mm, 1 cl. 10 mm ( MNHN Th 128); No. 2312, 24.II.1955, 1 moult, broken ( MNHN Th 129).

OTHER MATERIAL EXAMINED. — Pestarella convexa n. comb., Gambia, south of Cape of Bald, 18 m, unknown collector, 31.III.1954, 1 ovig. cl. 5 mm, tl. 24 mm (holotype), 1 tl. 24 mm, 1 tl. 18 mm (paratypes) ( MNHN Th 617). — Mauritania N’Diago, stn 61, in boulders, B. Richer de Forge coll., 18.IV.1982, 3 ovig. cl. 8-9.5 mm ( MNHN Th 723).

Pestarella rotundicaudata n. comb., South Africa, Kowie, Port Alfred, A. Penther coll., V.1896, 1 cl. 8.5 mm ( NHMW 6619).

DISTRIBUTION. — From Southern North Sea, Eastern Atlantic: Ireland ( Selbie 1914), English Channel, Atlantic coasts of France, Spain, Portugal, Morocco to Guinea. Mediterranean including Adriatic ( Števčić 1990; Dworschak 1992) and Corsica (de Vaugelas 1991, 1998).

DIAGNOSIS

Carapace ( Fig. 13A ) with blunt tip, no rostral spine. Abdominal segments ( Fig. 13C ) unarmed, second largest, segments 3-5 with lateral tufts of setae.Telson ( Fig. 13F ) about as long as proximal width.

Eyestalk with blunt distal tip; cornea distinct, dorso-lateral or dorsal. A1 peduncle slightly long- er than A2 peduncle ( Fig. 13A ). Md ( Fig. 13B ), Mxp1, Mxp2 ( Fig. 14F, G ) as figured; Mxp3 operculiform ( Fig. 13G, H ), ischium-merus length about 1.3-1.4 times merus width, propodus 1.5- 1.7 times as long as wide, lower border convex; dactylus digitiform.

Major P1 ( Fig. 13D ) slightly stouter in male; meral lower border straight or slightly convex; meral hook with rounded, denticulated lower border. Minor P1 ( Fig. 13C ) unarmed, carpus slightly wider proximally than distally, wider than propodus.

P2-5 ( Fig. 14 B-E) as figured, P3 propodus with lower border slightly dentate.

Female Plp1 ( Fig. 13I ) uniramous, two-articulated; female Plp2 ( Fig. 13J ) biramous, endopod twoarticulated. Uropodal exopod ( Fig. 13F ) about as long as proximal width, dorsal plate with distal row of setae at short distance and indistinct from posterior border; endopod slightly longer than telson.

Colour

General colour of body similar to that of P. candida n. comb., creamy with a slight tint of pink, tailfan sometimes yellowish, chelipeds usually slightly to bright pink (but sometimes completely white), ripe ovaries yellow to orange (Dworschak pers. comm.; d’Udekem d’Acoz 1999). A colour- ed photograph is presented in Perez Sanchez & Moreno Batet (1991) and González Pérez (1995).

Size

Largest specimens in material examined of cl. 12.5- 15 mm, tl. 53-57.5 mm; average size of cl. 6-15 mm, ovigerous of cl. 8-15 mm, tl. 60 mm ( Balss 1926).

Body length up to 67 mm in the British Isles ( Moyse & Smaldon 1990).

ECOLOGY AND BIOLOGY

Pestarella tyrrhena n. comb. occurs in intertidal and shallow subtidal zones ( Dworschak 1998a), in fine sand (d’Udekem d’Acoz 1986; Dworschak 1992), in muddy sand with or without Zostera seagrass (d’Udekem d’Acoz 1989; Števčić 1990) and in mud ( Le Gall 1969), at 5-20 m or deeper ( Moyse & Smaldon 1990). Picard & Pérès (1964) and Picard (1965) considered this species as characteristic of a relatively undisturbed muddy subtrate.

Aspects of biology studied

Feeding behaviour ( Dworschak 1987a); cadmium, mercury bioaccumulation and their effects ( Thaker & Haritos 1989a, b); reproductive output (percentage of mother weight devoted to egg production) (Thessalou-Legaki & Kiortsis 1997); physiological aspects in relation to pollution (Thessalou-Legaki et al. 1997); biometric analysis ( Dworschak 1998a); abundance of population in Corsica (de Vaugelas 1991); relation between burrow architecture, feeding mode and structure of the sediments (de Vaugelas 1998); burrow and burrow construction with role of tegumental glands ( Dworschak 1998 b, 2000).

REMARKS AND VARIATIONS

This is the most common callianassid species along the French Atlantic coast and the European coast of the Mediterranean, with much confusion in nomenclature.

It was described by Otto (1821, 1828) under the name Callianassa laticauda but was often confused with C. subterranea ( Montagu, 1808) . In 1893, Stebbing reported some Callianassa material from Jersey which he considered distinct from subterranea and which was named later Callianassa Stebbingi by Borradaile (1903). De Man (1928a) showed its identity with C. laticauda .

Giordani Soika (1943) stated that Cancer candidus Olivi, 1792 was identical with Callianassa laticauda Otto, 1821 , and that Olivi’s name, being the older of the two, should be employed. On the other hand, Holthuis (1947) considered that the three names Astacus tyrrhenus Petagna, 1792 , Cancer candidus and Callianassa laticauda actually referred to the same species. He ( Holthuis 1953) definitely settled the question and presented the main reason why Petagna’s name should be preferred over Olivi’s (Petagna’s figure is far superior). The species name tyrrhena ( Petagna, 1792) has been used therafter.

Variations occur in: 1) the lower border of major P1 merus which is straight or slightly convex; 2) the length ratio of carpus/propodus/fingers in major P1, with fingers often shorter in male than in female; and 3) the Mxp3 ischium ( Fig. 13G ) is often, not always, narrower proximally than distally and narrower than the merus. The colour pattern is also variable: whilst most specimens are more or less tinged with pink, specimens with completely white pereopods are occasionally found (d’Udekem d’Acoz pers. comm.).

Characters given for this species in the key are reliable but its European congeners can also be differentiated by the dorsal plate of the uropodal exopod: its setal distal row is at short distance and indistinct from the posterior border in P. tyrrhena n. comb. and well apart from the posterior border in P. candida n. comb. and P. whitei n. comb.

Pestarella whitei (Sakai, 1999) n. comb. ( Fig. 15 )

Callianassa Davyana White, 1847: 70 . — de Man 1928a: 37.

Callianassa whitei Sakai, 1999a: 23 , fig. 4C. — ° Türkay 2001: 289. — Dworschak 2002: 64, fig. 1c, d, tabl. 1 (part).

Callianassa candida – Dworschak 1992: 194 (part).

TYPE MATERIAL. — Holotype:, from Kap Monsena, S Val Salina , Istria near Rovinj, Croatia ( SMF 14033); paratype: same locality, ( SMF 24574).

MATERIAL EXAMINED. — Croatia. Rovinj , leg. U. Pettke, VII.1982, 1 holotype cl. 11 mm, tl. 49 mm (data given by Sakai; no data label accompagnying specimen) (figured, SMF 14033). — S Palu, Rovinj, intertidal under stones, Dworschak coll., 2.VIII.1982, 1 (just moulted) ( NHMW 6902 ). — Montauro, Rovinj, boulder field in 3 m, sand under stones, J. Ott coll., VII.1988, 1 cl. 12 mm ( NHMW 6778 ) .

DISTRIBUTION. — Currently known only from Croatia.

DIAGNOSIS

Rostrum ( Fig. 15A ) with blunt tip, no rostral spine. Telson ( Fig. 15D ) with proximal width about 1.5 times median length.

Eyestalk short, distal tip blunt or slightly pointed mesially; cornea disk-shaped, dorso-lateral or dorsal. Peduncle of A1 longer and stouter than that of A2. Mxp3 ( Fig. 13F ) operculiform, ischium-merus length about 1.5-1.6 times merus width, propodus slender, about twice as long as wide, lower border nearly straight, dactylus digitiform, 2.5-3 times as long as wide.

Major P1 ( Fig. 15B ) merus with lower border straight or slightly convex, lower border of meral hook denticulated, pointed distally. Minor P1 unarmed ( Fig. 15C ), carpus with similar proximal and distal width, 2-2.5 times length of propodus. Plp3-5 ( Fig. 15G ) with small appendix interna, projecting from inner border of endopod. Uropodal exopod ( Fig. 15D ) a little longer than distal width, dorsal plate with distal row of setae well apart from posterior border; endopod elongated, nearly twice as long as telson, posterior border rounded.

Colour

Similar to P. candida n. comb. (Dworschak pers. comm.).

Size

Holotype: cl. 11-12 mm, tl. 49 mm.

ECOLOGY

Living in mud, under stones or in seagrass meadows, 0-4 m ( Sakai 1999a; Dworschak 2002).

REMARKS

The chelipeds of the holotype ( Sakai 1999a: fig. 4c, d) were probably figured in situ and certain features are inaccurate, e.g., the shape of the major P1 meral hook, or the length ratio of the carpus and propodus. The figures are also presented as mirror images, as if the major cheliped was on the left and the minor cheliped was on the right while it is actually the opposite. The two appendages are here detached from the body and laid flat for drawing ( Fig. 15B, C ). Their morphology is very similar to that of P. candida n. comb.

In his work on the fauna of the Adriatic, Pesta (1918: fig. 63, 63a, 63b) reported some material he assigned to Callianassa stebbingi (= P. tyrrhena n. comb.). Later, de Man (1928a: 34) and de Saint Laurent & Božić (1976: 24) placed it in C. pestae or C. pontica (= P. candida n. comb.), while Sakai (1999a: 23) considered it to belong to P. whitei n. comb.

The material examined by Pesta was probably a mixture of P. candida n. comb. and P. tyrrhena n. comb. (and also P. whitei n. comb.). Figure 63 of Pesta (1918) is likely to represent a specimen of the former species (meral hook of major P1 with pointed distal tip, uropodal exopod with setal row of dorsal plate well apart from posterior bor- der). Figure 63a (an operculiform Mxp3) could be of either species, while figure 63b is clearly of P. tyrrhena n. comb. (telson about as long as wide, uropodal exopod with setal row of dorsal plate near posterior border). This material (or part of it) is still extant and housed at the Natur- historisches Museum Vienna. It includes specimens of P. candida n. comb. (Dworschak pers. comm. and sketch) with an exception (NHMW 319) that could be a variation of P. whitei n. comb. (peduncle of A1 slightly longer than that of A 2 in one side, about as long in the other; Mxp3 propodus slender).

Pestarella whitei n. comb. is similar to P. candida n. comb. in: 1) the eyestalk is short with the distal tip blunt or slightly pointed mesially; 2) the morphology of both P1; 3) the telson is wider than long; and 4) the uropodal exopod and endopod greatly exceed the telson. Distinguishing features are listed in the key and this species can be differentiated from its European congeners especially by the shape of its Mxp3 propodus ( Fig. 15E, F ). The latter is slender, approximately twice as long as wide, with the lower border nearly straight, while it is 1.5 times as long as wide, with the lower border convex in P. candida n. comb. and P. tyrrhena n. comb.

Pestarella whitei n. comb. is also similar to a non- European congener, but with a close distribution range, which is P. convexa n. comb. from Senegal (de Saint Laurent & Le Loeuff 1979: 73, fig. 10). It differs in: 1) the telson is about 1.5 times as wide as long in P. whitei n. comb. (1.3-1.4 times in P convexa n. comb.); 2) uropods are more elongated, endopods nearly twice as long as telson in P. whitei n. comb. (uropodal endopod about 1.5 times as long as telson in P. convexa n. comb.); and 3) the uropodal exopod with the setal row of dorsal plate well apart from posterior border in P. whitei n. comb. (setal row of dorsal plate near posterior border in P. convexa n. comb.).

Figures

FIG. 12. — Pestarella candida (Olivi, 1792) n. comb.; A, from Castiglione, Algeria, anterior part of carapace (from de Saint Laurent & Božić 1976); B, C, F-H, from Banyuls, France (MNHN Th 1358); D, E, I, from Gulf of Tunis, Tunisia (MNHN Th 633); B, D, major pereopod 1; C, minor pereopod 1; E, maxilliped 1; F, G, maxilliped 3, lateral and mesial view; H, telson and uropods; I, pleopod 3. Scale bars: 2 mm.

FIG. 13. — Pestarella tyrrhena (Petagna,1792) n. comb.; A, from Morocco, anterior part of carapace (from de Saint Laurent & Božić 1976); B, G-J, from Wimereux, France (MNHN Th 677); C, from Pointe du Château, France (MNHN Th 1346); D-F, from St- Malo, France (MNHN Th 1345); B, Mandible; C, lateral view, pereopod 1 omitted; D, E, major and minor pereopod 1; F, telson and uropods; G, H, maxilliped 3 and mesial view of ischium; I, J, female pleopod 1 and 2. Scale bars: A, B, D-J 2 mm; C, 4 mm.

FIG. 14. — Pestarella tyrrhena (Petagna,1792) n. comb.; A, from Pointe du Château,France (MNHN Th 1346), pereopod 1; B-E, from St-Malo, France (MNHN Th 1345), pereopod 2-5; F-H, from Wimereux, France (MNHN Th 677); F, G, maxilliped 1 and 2; H, pleopod 3. Scale bars: 2 mm.

FIG. 15. — Pestarella whitei (Sakai, 1999) n. comb., holotype from Rovinj, Croatia (SMF 14033); A, D, after Sakai 1999a; A, anterior part of carapace; B, C, major and minor pereopod 1; D, telson and uropods; E, F, maxilliped 3, lateral and mesial view; G, pleopod 3. Scale bars: 2 mm.