Upogebia Leach, 1814

Ngoc-Ho, Nguyen, 2003, European and Mediterranean Thalassinidea (Crustacea, Decapoda), Zoosystema 25 (3), pp. 439-555 : 508-536

Genus Upogebia Leach, 1814

TYPE SPECIES. — Cancer Astacus stellatus Montagu, 1808 , by original designation.

DIAGNOSIS. — Gastric region with lateral ridges, linea thalassinica depressed anteriorly, extending entirely or partly to posterior border of carapace; cervical groove well defined. Rostrum obtuse, bearing teeth or spines on lateral border, rarely unarmed. Anterolateral border of carapace unarmed or with spine. Telson often with faint inverted U-shaped carina dorsally, posterior bor- der straight or slightly convex, unarmed.

Eye peduncle cylindrical, with pigmented terminal cornea. Antennal scale reduced. Mxp1-3 with exopod; Mxp1 and Mxp3 with or without epipod, Mxp2 with epipod bent mesially; no spinous crest (crista dentata) on Mxp3 ischum. P1 equal, chelate or subchelate; P2- 4 simple, P5 subchelate. Plp1 absent in male, uniramous in female, Plp2-5 biramous, similar, lacking appendix interna. No suture on uropods.

Upogebia deltaura ( Leach, 1815) ( Figs 26 ; 27 )

Gebia deltaura Leach, 1815: 342 .

Gebia deltura Leach, 1816 : pl. 31, figs 9, 10. — Desmarest 1825: 204. — H. Milne Edwards 1837a: 314. — Bell 1846: 225, 1 fig. — White 1857: 97. — ° Fischer 1872: 428. — Van Beneden 1884: 647. — Carus 1885: 490. — Lovett 1885: 14. — Koehler 1886: 59. — Gourret 1888: 92, pl. 8, figs 16-29, pl. 9, figs 1-4. — Adensamer 1898: 621. — Sinel 1907: 217. — de Morgan 1910: 475, fig. 2. — RunnstrØm 1925: 33.

Gebia littoralis – * Sars 1884: 182, pls 3-5.

Gebia stellata – Lagerberg 1908: 54, pl. II fig. 10. — Schlegel 1912: 239. — Grieg 1927: 35 (part), 1 fig.

Gebiopsis deltaura – Balss 1926: 27, fig. 8.

Gebiopsis deltura – Stephensen 1910: 277.

Upogebia deltura – * Webb 1919: 85, pls 1-9, pl. 10, fig. 1, pl. 12 figs 1-4, 7, 8. — Gustafson 1934: 6. — Poulsen 1941: 216, 226, figs 6, 8.

Upogebia (Calliadne) deltaura – de Man 1927: 17, pl. 2, figs 8, 8b. — Balss 1936: 16.

Upogebia (Gebiopsis) deltaura – Borradaile 1903: 542. — Selbie 1914: 103. — Pesta 1918: 199, fig. 62. — Bouvier 1940: 106, fig. 70. — Gordon 1957: 249. — *Bourdillon-Casanova 1960: 109. — O’Céidigh 1962: 164. — Noël 1992: 82.

Upogebia deltaura – Schellenberg 1928: 77, fig. 58. — Miranda y Rivera 1933: 21. — ° Bertrand 1940: 30. — * Gurney 1942: 246, fig. 99. — Zariquiey Alvarez 1946: 107, 108; 1968: 231. — Wilson 1951: pl. 31, 1 fig. — * Rees 1955: 74, fig. 4.5 (part). — * Kurian 1956: 76. — Tambs-Lyche 1958: 12, fig. 2. — * Heegaard 1963: 458, pl. 1, fig. 7, text figs 27-33. —? Harmelin 1964: 94. — Picard 1965: 60. — Bourdon 1963: 429; 1965: 15. — Forest 1965: 347. — Holme 1966: 421, 452, fig. 14 (part). — Holthuis & Gottlieb 1958: 66. — de Gaillande 1970: 377. — de Saint Laurent 1971: 1261. — * Seridji 1971: 49. — Christiansen 1972: 41, fig. 48. — Naylor 1972: 69. — Lagardère 1973: 84. — Le Loeuff & Intès 1974: 56, figs 19, 19bis. — Neves 1974: 14. — Samuelsen 1974: 131. — ° Pastore 1976: 107. — * Thiriot 1976: 350, 367. — Tebble 1976: 85 (association with the bivalve mollusc Lepton squamosum (Montagu, 1803)) . — ° Glaçon 1977: 36. — Beaubrun 1979: 76. — de Saint Laurent & Le Loeuff 1979: 40, 93, fig. 3. — Monchartmont 1979: 73. — Domenech et al. 1981: 150. — Kocataș 1981: 162. — Ngoc-Ho 1981: 245; 1984: 511, fig. 6. — Adema et al. 1982: 28, fig. 8. — Manning & Froglia 1982: 324. — Cottiglia 1983: 78. — García Raso 1983: 318; 1990: 314. — Thessalou-Legaki & Zenetos 1985: 311. — Campbell & Nicholls 1986: 218, 1 fig. — Holthuis & Heerebout 1986: 62, fig. 81. — Lewinsohn & Holthuis 1986: 25. — Thessalou- Legaki 1986: 183. — Tunberg 1986: 753. — d’Udekem d’Acoz 1989: 176, fig. 5; 1995: 61, fig. 3; 1999: 156. — Moyse & Smaldon 1990: 520, fig. 10.13 (part). — Neves 1990: 670. — Števčić 1990: 215. — Dworschak 1992: 223. — ° García Raso et al. 1992: 258. — ° Koukouras et al. 1992: 223. — Noël 1992: 82. — ° Froglia 1995: 8. — Hayward et al. 1995: 433, fig. 8.52 (part). — Eneman 1996: 159, fig. 1. — Falciai & Minervini 1996: 149, 1 fig. — ° Brattegard & Christiansen 1997: 222. — Christiansen & Stene 1998: 76. — Astall et al. 1996:

821, tabls 1, 2; 1997a: 155, figs 1 (part), 3, 5, tabl. 1 (part); 1997b: 671, 675, fig. 1, tabls 1-5, pl. 2. — Hughes & Atkinson 1997: 640. — Lindahl & Baden 1997: 33. — Pinn et al. 1998a: 243, figs 2B, 2D; 1999a: 103, figs 2G, H, 3A, 4G, H; 1999b: 1461, figs 5, 6, tabls 1-6. — Hall-Spencer & Atkinson 1999: 871, figs 1-5. — Moen & Svensen 1999: 234, 2 figs. — Christiansen 2000: 233. — Taylor et al. 2000: 265, figs 1-3, tabls 1-3, 5, 6. — *González- Gordillo et al. 2001: 279. — Livory 2001: 33. — Markham 2001: tabls 1, 2. — * Martin 2001: 81, 1 fig. — ° Türkay 2001: 289.

Uppogebia deltaura –? Pérès & Picard 1964: 66.

TYPE MATERIAL. — Holotype: Kingsbridge Estuary , Devon, UK, English Channel , North East Atlantic, Leach collection:, posterior part of carapace damag- ed, left P1, P4, P5 present; left P2, right P1, P5 detached, merus missing ( NHML 259 e).

MATERIAL EXAMINED. — North Sea. Doggerbank Ost, 54°49.14’N, 3°3.30’E, 27.2-29.7 m, F. K. Senckenberg, 1 cl. 10.5 mm ( SMF 28334). — Spiekeroog, 27.VII.1959, 1 cl. 23 mm, 1 ovig. cl. 20 mm ( SMF 28335). — Near Helgoland, 1877, 1 ( RMNH D 4931). — NW Brandaris, in Mustelus mustelus (Linnaeus, 1758) , Zool. Station Den Helder coll., 20. V.1950, 1 ( RMNH D 7383). — Brown Banks, near Winterton Twenties, from stomach of Solea solea (Linnaeus, 1758) , 20-27.VIII.1960, leg. J. Kruuk, 3, poor condition ( RMNH D 16180). — 53°41’45”N, 3°55’E, 12.VIII.1971, leg. G. R. Heerebout, 2, 2 ( RMNH D 27524). — 53°43.5’N, 4°27.2’E, 33 m, 1. VI.1971, leg. Rijks. Instituut voor Visserij Onderzoek, 2, 2 ( RMNH D 27525). — 53°42’N, 3°55’E, 12.VIII.1971, Tridens , leg. G. R. Heerebout, 19 m, 7 ( RMNH D 27526). — 54°02.5’N, 4°07.9’E, 42 m, 30. VI.1971, leg. Rijks. Instituut voor Visserij Onderzoek, 1 ( RMNH D 28191 ). — 53°38’N, 4°31.6’E, 28-29 m, leg. Rijks. Instituut voor Visserij Onderzoek, 1 ( RMNH D 28192 ). — 53°48.7’N 4°22.5’E, 37 m, 1. VI.1971, leg. Rijks Instituut voor Visserij Onderzoek, 1 ( RMNH D 28193 ). — 53°38’N, 3°44’E, 34 m, 23. VI.1980, P. C. Goudswaard, stn 74, Rijks. Instituut voor Visserij Onderzoek, 4, 2 ( RMNH D 32960). — 53°48’N, 04°13’E, 24.IV.1972, Aurelia cruise, don. Netherlands Institute for Sea Research, 17, 4 ( RMNH D 33185). — 53°43’N, 04°17’E, 16.X- 1.XI.1972, Aurelia , stn 180, leg. F. Creutzberg, v.s., don. NIOZ, 3, 1 ( RMNH D 35738). — 53°38’N, 03°44’E, 34 m, 23. VI.1980, leg. P. C. Goudswaard, 2 ( RMNH D 35739). — 53°40’N, 04°22’E, 26.VI-7.VII.1972, Aurelia , stn 117, leg. F. Creutzberg, c.s. don. NIOZ, 2 ( RMNH D 35740). — North-Holland, 30 m, 21. VI.1958, don. NIOZ, 1 ovig. ( RMNH D 35741). — 53°39’N, 5°42’E, 20 m, 26.X.1977, leg. P. I. Van Leeuwen, don. R. Boddeke, RIVO, 2 ( RMNH D 35742). — 53°48’N, 04°07’E, 24. IV- 4. V.1972, Aurelia cruises, leg. F. Creutzberg, don. NIOZ, 3 ( RMNH D 35744). — 53°39’N, 04°06’E, 29.IX.1975, Aurelia , stn 333, don. NIOZ, 1 ( RMNH D 35745). — 53°52.5’N, 04°29.5’E, MS Aurelia , 2.II.1982, leg. P. C. Goudswaard, 1 ( RMNH D 35746). — 54°03’N, 02°50’E, MS Aurelia , 20.X.1976, leg. F. Creutzberg, 1 cheliped ( RMNH D 35747). — Surroundings Helgoland, 54°28’N, 06°13’E- 54°42’N, 06°35’E, 3-12. V.1995, leg. J. Verkuil, 2 ( RMNH D 46166 ).

Sweden. Prov. Gotebörg och Bohus, Fiskebackskil, c. 120 km N Göteborg, Kristineberg Marine Research Station, Pettke coll., IX.1976, 44 cl. 7- 20 mm, 41 cl. 7-17.5 mm ( SMF 28331) ; 4

cl. 17-21 mm, 1 cl. 12 mm ( SMF 28332) ; Kristineberg , IX.1975, 4 cl. 10.5-12.5 mm, 6 cl. 7.5- 15 mm ( SMF 28333). — Skagerrak, 20-30 m, exc. Leiden Biologists, stn 301, 3.VI.1971, 1 ( RMNH D 28194 ). — E of Bonden Island, 58°12’N, 11°19’E, 20-30 m, Amphioxus sand, 3.VI.1971, Exc. Studends Leiden, 1 ex. ( RMNH D 35734) .

Netherlands. Teakettle Hole, 50 miles WNW of Den Helder, in stomach of Egaleus galeus (Linnaeus), 24.XI.1945, don. Zool. Station Den Helder, 1 damag- ed ( RMNH D 6198). — NW of Brandaris, Texel, from stomach of Mustelus mustelus (Linnaeus, 1758) , near oystergrounds, 25.X.1950, don. Zoological Station Den Helder, 1 ( RMNH D 7382).

Great Britain. Firth of Clyde, off Lion Rock, R. B. Pike coll., 20.XII.1957, 3 ( NHML 1962.7.5.5.7). — Devon, English Chanel, Leach collection, 1 cl. 20 mm, tl. 65 mm approx. (figured) (holotype, NHML 259 e). — English Channel (see Holme 1961), 29 m, 3, 4 (2 ovig.) ( NHML 1999.257 - 264 ) ; 35 m, 1, 3 ( NHML 1999.247 - 256 ) ; 58 m, 4 ( NHML 1999.242 - 245 ) ; 37 m, 1 ( NHML 1999.241 ) ; 21 m, 1 carapace ( NHML 1999.229 ) ; 34 m, 1 broken ( NHML 1999.231 ) ; 18 m, 1 ( NHM 1999.228 ) ; 37 m, 2, 1 ( NHML 1999.239 - 249 ) ; 19 m, 1 ( NHML 1999.226 ) ; 36 m, 1 ( NHML 1999.233 ) ; 19 m, 1 abdomen ( NHM 1999.227 ) ; 37 m, 1 abdomen ( NHML 1999.232 ) ; 16 m, 1, 2 ( NHML 1999.230 ) ; 35 m, 1 ovig. ( NHML 1999.246 ) (see Holme 1966) ; 27 m, 3, 1 ( NHM 1999.269 - 272 ) ; 33 m, 1 ( NHML 1999 . 273) ; 36 m, 2, 3 ( NHML 1999.234 - 238 ) ; 35 m, 9.XI.1961, 4, 1 ( NHML 1999.306 - 310 ) ; 31 m, 1.II.1960, 4, 2 ( NHML 1999.278 - 285 ) ; 57 m, 11.VII.1961, 4, 3 ( NHML 1999.299 - 304 ) ; 22 m, 1.II.1960, 2 ( NHML 1999.286 - 289 ) ; 68 m, 1 carapace ( NHML 1999.321 ) ; 33 m, 2.II.1960, 1 exuvia ( NHML 1999.297 ) ; 27 m, 1 abdomen ( NHML 1999.294 ) ; 55 m, 31.I.1962, 5 (2 damaged) ( NHML 312-315 ) ; 38 m, 1.II.1960, 2 ( NHML 1999.276 - 277 ) ; 37 m, 1 abdomen ( NHML 1999.320 ) ; 52 m, 11.VII.1960, 1 ( NHML 305 ) ; 29 m, 1.II.1960, 2, 1 (without abdomen) ( NHML 1999.292 - 293 ) ; 46 m, 10.XI.1961, 1 ( NHML 1999.319 ) ; 64 m, 31.I.1960, 1 ( NHML 1999.275 ) ; 38 m, 1.II.1960, 2 ( NHML 1999.290 - 291 ) ; 26 m, 2 ( NHM 1999.295 - 296 ) ; 68 m, 3.II.1962, 1 ( NHML 1999.322 ) ; 44 m, 10.XI.1961, 1 ( NHML 1999.311 ) ; 51 m, 11.VII.1961, 1 (dried) ( NHML 1999.298 ) ; 68 m, 3.II.1962, 2, 1 ( NHML 1999.316 - 318 ) ; 31 m, 31.I.1960, 1 ( NHML 1999.274 ) ; 27 m, 30.I.1960, 3 (damaged) ( NHML 1999.265 - 268 ). — Plymouth, 1, 1 ( NHML 1910.7.1 -2) ; intertidal mud-flats, R. L. Rice coll., 8.IV.1971, 1 ( NHML 1971.21 ) ; II.1945, 1 ( NHML 1949.1.18.24) ; MBA purchased, VI.1950, 2 ( NHML 1950.8.22.18- 19) ; shell gravel, A. L. Rice coll., 18.III.1970, 1 ( NHML 1972.20 ). — Plymouth Sound, Montagu coll., 3 ( NHML 259 a, 259 b and 259 c). — Salcombe, Devon, 1 ( NHML 1901.4.16.1). — Salcombe Harbour, 1, 1 ovig. ( NHML 98.5.7.734). — Polperro, Cornwall, Norman collection, 1 ( NHML 1911.11.8.1034). — Weymouth Bay, B. Horton coll., 10.V.1990, 1 cl. 16 mm ( NHML 1999.995 ). — Plymouth, leg. Carlisle, V.1959, don. Zool. Station Napels, 1 ovig .

( RMNH D 13012).

Belgium. Ostende, E. Van Beneden coll., 1 ( NHML 98.5.7.731).

France. English Channel, Roscoff, collector unknown, 17.IX.1958, 1 cl. 18 mm, tl. 54 mm, 1 cl. 11 mm without abdomen (figured), 1 cl. 17.5 mm ( MNHN Th 1316). — Bricqueville, Normandy, low tides, B. Richer de Forges coll., III.1981, 1 cl. 21 mm, tl. 59 mm ( MNHN Th 643). — Baie de Seine, 20-30 m, Gentil coll., 1 cl. 15 mm ( MNHN Th 315). — Manche?, Baron of St-Joseph’s collection, No. 20, 1911, 2 cl. 14.5 and 21 mm ( MNHN Th 9), 2 cl. 8.5 and 11 mm (poor condition) ( MNHN Th 487). — Atlantic, Bay of Concarneau, Glémarec coll., 1 cl. 6.5 mm ( MNHN Th 8). — Coasts of France, 1 ovig. cl. 16.5 mm, 2 cl. 16-20 mm, 1 cl. 19 mm, tl. 58 mm (figured) ( MNHN Th 38). — St-Vaast-la-Hougue, A. Malard coll., 1903, 1 cl. 23 mm approx. (poor condition) ( MNHN Th 203). — Iroise sea, Brittany, Glémarec coll., 1 cl. 8 mm ( MNHN Th 313), 1 cl. 5.5 mm, 1 cl. 6 mm ( MNHN Th 314). — Bay of Morlaix, Brittany, J. H. Stock coll., 17.IX.1955, don. Zool. Museum Amsterdam, 1 ( RMNH D 10991). — Island of Glénoan, near Concarneau, Brittany, 10-30 m, 14.IX.1958, 1, 1 ( RMNH D 12533). — St-Servan, Brittany, IV.1924, 2 cl. 9 and 19 mm ( MNHN Th 554). — You Bridge?, Y. Gruet coll., 26.VIII.1972, 1 cl. 22 mm (poor condition) (MNHN-Th 612). — Island of Chausey (Pierre aux Vras), low tides, A. Crosnier coll., VIII.1976, 1 cl. 7.5 mm ( MNHN Th 440). — Mediterranean, Banyuls, sewage, P. Noël coll., 9. VI.1976, 3 cl. 4.5- 8 mm ( MNHN Th 1339). — Banyuls?, Thiriot coll., 1 cl. 7 mm ( MNHN Th 666). — Monaco, 22.VIII.1960, leg. C. Carpine, 1, 1 ( RMNH D 19889). — Fontaindreau, from the stomach of Raja clavata Linnaeus, 1758 , 110- 125 m, 3, 3 ovig.

( RMNH D 28198 ).

Spain. Cadaqués, VIII.1950, leg. R. Zariquiey Alvarez, 1 ( RMNH D 35743).

Italy. Porto Cesareo, Parenzan coll., 1 cl. 12 mm ( MNHN Th 543).

Mediterranean. Baleares, Ibiza, 5-6 m, 11.VIII.1954, 1 juv. cl. 3.5 mm (MNHN Th 1324).

Mauritania. Off Banc d’Arguin, 19°33’N, 16°55’W, depth 64 m, sticky grey mud, ophiurids, echinoids, bivalves, tubeworms, 1.2 m, Agassiz trawl, 15. VI.1988, 4, 3 (1 ovig.) ( RMNH D 48000 ).

Sierra Leone. 7°15’N, 13°00’W, 100 m, bottom with sandy mud, 18.IV.1956, A. R. Longhurst coll., 1 broken ( RMNH D 32139).

Ivory Coast (between Vridi and Jacqueville). 60 m, Le Loeuff coll., 23.XI.1966, 2 cl. 4.5 and 6 mm ( MNHN Th 283). — N Vridi, Reine Pokou Exped., 60 m, Le Loeuff coll., 22. VI.1966, 1 cl. 5 mm, 1 cl. 6 mm ( MNHN Th 663). — Jacqueville, Reine Pokou Exped., 60 m, Le Loeuff coll., 24.XI.1966, 1 cl. 8 mm ( MNHN Th 668). — East of Grand Bassam, 37 m, Le Loeuff coll., 25.II.1967, 1 juv. cl. 5 mm ( MNHN Th 670).

Togo. Togo coast, Ombango Exped., sandy mud, 59- 60 m, A. Crosnier coll., 6.X.1963, 2 cl. 6 and 6.5 mm ( MNHN Th 284).

DISTRIBUTION. — Eastern Atlantic: Sweden, Norway ( Samuelsen 1974; Christiansen 2000) to Togo, including the English Channel; North Sea (Poulsen 1940; Adema et al. 1982); Mediterranean including Adriatic Sea ( Pesta 1918; Števčić 1990; Dworschak 1992), Aegean sea ( Koukouras et al. 1992) and Cyprus ( Lewinsohn & Holthuis 1986).

DIAGNOSIS

Rostrum ( Fig. 26A, D ) obtuse distally, slightly longer than wide at base, with two or three teeth on lateral border; antero-lateral border of carapace unarmed ( Fig. 26B ); pleura of first abdominal segment ( Fig. 26G ) pointed postero-ventrally; telson ( Fig. 26C ) approximately 1.2 times as wide as long.

A1 peduncle ( Fig. 26B ) with large spine on lower margin of article 1, A2 peduncle with smaller spine on lower margin of article 3. Md with large mesio-anterior pointed tooth. Mx1 and Mx2 ( Fig. 27D, E ) as figured. Mxp1 ( Fig. 27H ) without epipod, Mxp3 ( Fig. 27G ) with epipod.

P1 slightly stouter in male ( Fig. 27A ) than in female ( Fig. 26E, F ); merus with upper subdistal spine and two to five lower spines or spinules; carpus with large upper mesial distal spine, small lower distal spine and (sometimes) small median spines near mesial distal margin ( Fig. 26F ); propodus with small upper subdistal spine, both lateral and mesial surface with one or two distal spines near base of fixed finger; fixed finger as long as dactylus in young specimen, often shorter than dactylus in adult, with three to four triangular teeth on proximal half of cutting edge; dactylus with longitudinal row of round tubercules on external and mesial surface, small round teeth on cutting edge. P2 ( Fig. 27B ) with upper subdistal spine on merus and carpus. P3 ( Fig. 27E ) with spinules and tubercles on lower margin of merus. Large coxal spines on P1 and P2. Arthrobranch with double row of fine lamellae on either side of the rachis.

Plp1 absent in male, uniramous in female ( Fig. 26G ); Plp2-5, biramous, foliaceous. Exopod of uropod with proximal spinule, basipod also with spinule ( Fig. 26C ).

Colour

Bright orange, some specimens whitish (d’Udekem d’Acoz pers. comm.); grey yellow tinged green ( Campbell & Nicholls 1986); dirty yellow, tinged white or red ( Moyse & Smaldon 1990).

Size

Holotype of cl. 21 mm, tl. 65 mm approx. Largest specimens in material examined: cl. 18- 21 mm, tl. 54-59 mm. Largest size reported: tl. up to 80 mm or exceptionally 150 mm ( Moyse & Smaldon 1990).

ECOLOGY AND BIOLOGY

The population ecology of the species in western Sweden was studied by Tunberg (1986). In western Norway, it occurs in shell and mineral sand with some mud, at 3 m depth ( Samuelsen 1974), in shallow water in southern part of the boreal region ( Christiansen 2000). It was found in muddy-gravel association off Plymouth, English Channel ( Holme 1966), and was occasionally dredged from mud or muddy sand ( Gordon 1957). It is common in Brittany, France, in coarse or muddy sand, with or without seagrass (d’Udekem d’Acoz 1989). Specimens were collected from shore, in Ireland ( Selbie 1914), near the water edge on the beach ( Southern 1915), between 20-40 m, in Portugal ( Neves 1990), and were present in a variety of substrates from sand with high percentage of pebbles to muddy sand, in Patras Gulf, Greece, at 5-15 m depth (Thessalou-Legaki & Zenetos 1985).

Reproduction occurs in February-April in Roscoff, France ( Bourdon 1965) in spring and summer in Plymouth, UK ( Gordon 1957). Larvae are present in Roscoff during February- November, abundant in July-August, mainly near the bottom (i.e. at 15 m depth in a bottom of 20 m) during day time, in surface waters at night ( Thiriot 1976), common in the Kilkieran Bay plankton, Ireland in April-October, plentiful in August ( O’Céidigh 1962).

Aspects of biology studied

Functional anatomy of foregut (Ngoc-Ho 1984); branchial parasitic isopods ( Astall et al. 1996); burrow morphology and burrow-dwelling lifestyle ( Astall et al. 1997a); branchial morphology, gill area and gill ultrastructure ( Astall et al. 1997b); particle size selectivity and resource partitioning ( Pinn et al. 1998b); burrows in scottish mearl beds (Hall-Spencer & Atkinson 1999); gut morphology and gut microflora ( Pinn et al. 1999a); mouthpart morphology and mouthpart setal fringes ( Pinn et al. 1999b); functional response in filter feeding ( Lindahl & Baden 1997); oxygen transporting properties of heamocyanin ( Taylor et al. 2000).

VARIATIONS

There are variations in: 1) the rostrum often bears two, three or (rarely) four lateral rostral teeth; 2) the large upper distal spine on P1 carpus is sometimes accompanied mesially by one or two spinules ( Fig. 26F ); 3) the lower border of P1 propodus has two to five spinules on the proximal half, three on the left and eight (rare) on the right in the holotype ( Fig. 26E ); 4) the distal lateral spine, as well as distal mesial spine near the base of P1 fixed finger, is larger in female than in male, and impaired, as on left P1 of holotype (rarely paired as on right P1 of holotype, Fig. 26E, F ); 5) P1 fixed finger is about as long as the dactylus in small specimens (cl. 7-8 mm, tl. c. 20-22 mm), shorter than the dactylus in most large specimens, shortest in large males; and 6) P3 merus is sometimes unarmed.

REMARKS

Upogebia deltaura is close to U. mediterranea Noël, 1992 which is often associated with Posidonia oceanica seagrass. Some materials assigned to U. deltaura and reported as associated with Posidonia ( Harmelin 1964; Pérès & Picard 1964) could actually belong to U. mediterranea . Differentiation of U. deltaura and U. mediterranea is discussed under the latter species.

U. deltaura is similar to U. demani de Saint Laurent & Le Loeuff , from the Gulf of Guinea, in the shape and spinulation of the rostrum (de Saint Laurent & Le Loeuff 1979: 41, fig. 4b), also in having a lower spine on article 3 of A2 peduncle. De Saint Laurent & Le Loeuff (1979: 42, fig. 4a) fail to mention and figure the latter character, but give in detail the differences between the two species.

The pointed postero-ventral pleura of abdominal segment 1 is the most important characteristic of U. deltaura . Along with the morphology of the rostrum and the unarmed antero-lateral border of the carapace, it permits a quick identification even of small specimens (e.g., MNHN Th 1324; cl. 3.5 mm). The carapace is thin and weakly calcified in some specimens.

Genital openings are visible in young male and female of cl. 5-5.5 mm (tl. c. 15-17 mm) but Plp1 appear at a later stage in female of cl. 6.5- 8 mm (tl. c. 19-22 mm).

Upogebia deltaura , widely distributed in Europe, was also found in Western Africa ( Ivory coast and Togo) by Le Loeuff and Crosnier ( MNHN Th 283, 284, 663, 668, 670). Specimens from these areas are apparently smaller than in Europe, but the adult size is not known as no ovigerous females were available for study.

Upogebia mediterranea Noël, 1992 ( Figs 28 ; 29 )

Upogebia n. sp. “ mediterranea ” Noël, 1992: 82.

Upogebia deltaura –? Harmelin 1964: 94.

Uppogebia deltaura –? Pérès & Picard 1964: 66.

Upogebia cf. deltaura – Kocataș 1981: 162. — Dworschak 1992: 223.

Upogebia mediterranea – Dworschak 1983: 40 (appendix) nomen nudum.

Upogebia mediterranea – ° Froglia 1995: 8. — Asgaard et al. 1997: 23. — Abed-Navandi & Dworschak 1998: 609. — d’Udekem d’Acoz 1999: 157. — ° Türkay 2001: 289.

Upogebia nitida – Le Loeuff & Intès 1974: 58 (part), fig. 20.

Upogebia cf. nitida – de Saint Laurent & Le Loeuff 1979: 93.

Upogebia nitida mediterranea – d’Udekem d’Acoz 1995: 60, 61, fig. 2.

Upogebia n. sp. – Thessalou-Legaki 1986: 184.

TYPE MATERIAL. — Lectotype: from Tamaris-sur- Mer , near Toulon, France ( MNHN Th 1372) by present designation.

Paralectotypes: Tamaris-sur-Mer , same data as lectotype: 4, 3 ( MNHN Th 1369) ; Banyuls , France, 1 ( MNHN Th 665), 1 (posterior part of abdomen missing) ( MNHN Th 667) ; Ibiza , Baleares, Spain, 1 ( MNHN Th 661) ; Gulf of Gabès, Tunisia, 1 ( MNHN Th 921), 1 ( MNHN Th 922), 1 ( MNHN Th 923) and 1 ( MNHN Th 924) ; Gulf of Oran, Algeria, 2, 2 ( MNHN Th 486) .

TYPE LOCALITY. — Tamaris-sur-Mer, France.

MATERIAL EXAMINED. — France. Tamaris-sur-Mer, near Toulon, collection Sollaud, No. ES 38, 25.XII.1932, lectotype cl. 15 mm, tl. 48 mm (figured) ( MNHN Th 1372); same data, 1 paralectotype cl. 12 mm, tl. 36 mm (figured) and 3 paralectotypes cl. 13-14.5 mm, tl. 38-43 mm, 3 paralectotypes cl. 13-18 mm, tl. 42-56 mm ( MNHN Th 1369). — Banyuls, France, in Posidonia rhizome, 7 m, P. Noël coll., 12.VIII.1976, 1 paralectotype cl. 6 mm, tl. 17 mm ( MNHN Th 665); in Herbier of Racou, 5-7 m, P. Noël coll., 27.III.1979, 1 paralectotype (posterior part of abdomen missing) cl. 9 mm ( MNHN Th 667). — Banyuls?, Thiriot coll., 1 ovig., broken rostrum, approx. cl. 8 mm ( MNHN Th 1312); Banyuls, in bed of Posidonia , 5 m, P. Noël coll., 3. VI.1977, 1 (just moulted, poor condition) approx. cl. 11 mm ( MNHN Th 664). — Cap d’Ail, near Villefranche-sur-Mer, night catch, C. Vadon coll., 28. V.1980, 1 cl. 11.5 mm ( MNHN Th 1323). — Marseille, Port Cros Island, 1.8 m, A. Willsie coll., 18. V.1983, 1 cl. 10 mm (poor condition) ( SMF 28328). — Marseille, in sediments with Posidonia, A. Willsie coll., 1982, 1 cl. 12.5 mm, tl. 38 mm, ( MNHN Th 1325); 1988, 1 cl. 11 mm ( SMF 28329). — Ponteau, W of Marseille, H. Zibrowius coll., 13.II.1978, 1 cl. 12 mm, tl. 37 mm ( MNHN Th 1326), 1 cl. 11.5 mm ( SMF 28330). — Villefranche-sur-Mer, Alpes- Maritimes, W. G. N. v. d. Sleen coll., 26.IV.1920, 1, 1 ovig. ( RMNH D 16179). — Monaco, H. Nouvel coll., 15 m, 15.IV.1952, 4, cl. 9-11 mm, 3 cl. 10.5- 11 mm ( RMNH D 34028); 1974, 3 cl. 5.5-9.5 mm, 3 cl. 10-11 mm ( RMNH D 34029); 10. V.1938, 1 cl. 10 mm ( RMNH D 34030); 2.VII.1937, 2 cl. 6.5 and 9.5 mm ( RMNH D 34031); IV.1938, 2 cl. 6.5 and 8.5 mm ( RMNH D 34033); 15-25. VI.1937, 1 cl. 6 mm ( RMNH D 34034); 1936, 10 cl. 5.5- 7.5 mm, 7 (4 ovig.) cl. 7-11 mm ( RMNH D 34035); 2.IV.1953, 1 cl. 7.5 mm ( RMNH D 34036); VI.1937, 24 cl. 5.5- 12 mm, 4

(3 ovig.) cl. 8-11 mm ( RMNH D 34037). — Cap d’Ail, H. Nouvel coll., 9.IV.1953, 1 cl. 7 mm ( RMNH D 34032).

Spain. Baleares, Ibiza, 11.VIII.1954, 5- 6 m, 1 paralectotype cl. 8 mm, tl. 23.5 mm ( MNHN Th 661).

Italy. Ischia, Castello A., in Posidonia rhizome, W. Tertschnig coll., 14. V.1981, 4 cl. 8-13 mm and 1 cl. 13 mm ( NHMW 6763). — Ischia, Lacco Ameno, in Posidonia rhizome, 2 m, K. Wittmann coll., 1. VI.1987, 1 cl. 12.5 mm, 2 ovig. cl. 13 and 14 mm ( NHMW 6764). — Naples, Adensamer leg. 1898, 1 cl. 12 mm, 1 cl. 13.5 mm ( NHMW 10959). — Naples, J. G. de Man, 1876, 1 cl. 10.5 mm ( RMNH D 993); don. Zoöl. Station Naples, V.1959, 19 cl. 5-16mm, 9 (3 ovig.) cl. 6-16 mm ( RMNH D 13069).

Greece. Rhodos, R. Bromley coll., 2.VIII.1998, 1 cl. 11.5 mm ( NHMW 17936). — Tavros, Chios Islands, G. Potts coll., 1 cl. 5 mm ( NHML 1968: 607). — Kreta (Stavros), given by fishermen, C. d’Udekem d’Acoz coll., 15.VII.1987, 1 cl. 3.5 mm (d’Udekem d’Acoz). — Aegean Sea, 40 m, 18.VIII.1985, Strimonikos G., 1 (A.U.TH P 4529).

Malta. M’xiokk Bay, P. J. Schembri coll., 1 cl. 12.5 mm, 1 cl. 13 mm ( NHML 1977: 158). — Market, Norman collection, 4 cl. 14-15.5 mm, 5

(3 ovig.) cl. 9-15 mm ( NHML.1911.11.B.1024- 1033).

Israel. Haifa Bay , 19 m, E. Gottlieb coll., 7.VI.1955, 1 juv. cl. 1.5 mm ( RMNH D 13327) ; 18 m, 7.IX.1954, 1 cl. 5 mm ( RMNH D 13328) ; 20 m, 18.X.1955, 1 juv. cl. 2 mm ( RMNH D 13329) .

Tunisia. Gulf of Gabès, Calypso , muddy sand with detritus, 25 m, 2.V.1965, 1 cl. 9 mm, tl. 27 mm ( MNHN Th 921) ; 1 paralectotype cl. 11 mm, tl. 31 mm ( MNHN Th 922) ; muddy and chalky sand, 25 m, 19.IV.1965, 2 paralectotypes cl. 8 mm, tl. 24 mm ( MNHN Th 923), cl. 9 mm, tl. 26 mm ( MNHN Th 924) .

Algeria. Gulf of Oran, P. Pallary coll., 1900, 1 paralectotype cl. 9.5 mm, tl. 29 mm (figured), 1 paralectotype cl. 10 mm, tl. 30 mm and 2 paralectotypes, cl. 11 and 12 mm, tl. 32 and 34 mm ( MNHN Th 486).

Morocco. Melilla, J. Ruttlant, 1 cl. 8 mm ( RMNH D 5254).

Congo. Near Pointe Noire, trawling, 18 m, A. Crosnier coll., Ombango, 25.XI.1966, 1 cl. 5.5 mm ( MNHN Th 231).

DISTRIBUTION. — Mediterranean and Congo (west African coast).

DIAGNOSIS. — Rostrum elongated triangular, slightly longer than wide at base, with two teeth on each lateral border; anterolateral border of carapace unarmed; pleura of first abdominal segment rounded posteroventrally; telson subquadrate. A1 peduncle with large lower spine on article 1; A2 peduncle unarmed. Md with large mesioanterior pointed tooth. P1 merus with upper subdistal spine and lower spinules or tubercles; carpus with large upper mesial distal spine and small lower distal spine; propodus with upper subdistal spine and lateral distal spinule near base of fixed finger, latter with three or four teeth on proximal half of cutting edge; dactylus as long as fixed finger with mesial proximal round tooth on cutting edge. P2 with upper subdistal spine on both merus and carpus. P3 merus unarmed. Large acute coxal spine on P1 and P 2 in male and female. Uropods about as long as telson.

DESCRIPTION

Rostrum ( Fig. 28A, B ) elongated triangular, slightly longer than wide at base, projecting beyond eyes, with two large spiniform teeth on each lateral border. Faint median dorsal groove on rostrum and anterior part of gastric region. Lateral groove narrow and shallow, lateral ridge with 10 to 12 teeth, larger distally. Linea thalassinica distinct, extending to posterior border of carapace. Anterolateral border of carapace unarm- ed. Cervical groove deep, bearing minute denticles and tubercles on either side. Epistome rounded distally or with minute distal spinule.

First abdominal segment slightly shorter than second, proximal part of pleura rounded ventrally ( Fig. 28E ). Telson ( Fig. 28F, G ) approximately quadrate, lateral and posterior borders weakly convex, an U-shaped carina and a faint median groove on upper surface.

Eyestalk ( Fig. 28B ) moderately long, cornea pigmented in preserved specimens. A1 ( Fig. 29A ) with large lower distal spine on peduncular article 1; peduncular article 3 about as long as second and first together; flagella not longer than peduncle. A2 peduncle unarmed ( Fig. 29B ), scale large, terminating in a spinule.

Md ( Fig. 29C ) with large mesio-anterior pointed tooth. Mxp1 without epipod, Mxp3 with small epipod. Arthrobranch with double or single (rare) row of fine lamellae on either side of rachis.

P1 ( Figs 28C, D ; 29F ) cheliform, slightly stouter in male. Ischium with one or two lower spinules or tubercles. Merus over twice as long as wide, bearing upper subdistal spine and eight to 12 lower spinules and tubercles. Carpus with slight longitudinal groove on upper part of lateral surface, large mesial upper distal spine sometimes accompanied by one or two mesial distal spinules; smaller lower distal spine. Propodus about twice as long as wide in female, slightly stouter in male, with two to six spinules or denticles on proximal third of lower border, upper subdistal spine, and lateral distal spinule or tubercle near base of fixed finger; fixed finger with three or four teeth on proximal half of cutting edge. Dactylus approximately as long as fixed finger, with rounded tubercles on upper margin, proximal one largest and slightly pointed; longitudinal row of tubercles on both lateral and mesial surfaces, proximal mesial round tooth on cutting edge followed distally by faint teeth; corneous tip. P2 ( Fig. 29D ) merus over 4.5 times as long as wide with upper subdistal spine; carpus half as long as merus with small upper subdistal spine. P3 ( Fig. 29E ) and P4 unarmed, propodus and dactylus with lateral longitudinal row of corneous tubercles; dactylus pectinate on lower margin. Large coxal spine on P1 and P2.

Uropod ( Figs 28F ; 29G ) exopod with proximal spinule and denticles on slightly convex posterior margin; posterior margin of endopod almost straight; protopod with spinule hanging over endopod.

Coloration of a live specimen (MNHN Th 667) (Noël pers. comm.)

Gastric region with uniform orange colour on upper part, whitish laterally, with punctiform red chromatophores. Abdominal tergites and pleurites orange, with chromatophores, the latter a shade of light gray. Sternites transparent. Eye peduncle orange, with red chromatophores; cornea dark-wine. A1 peduncle transparent with a few red chromatophores. A2 peduncle with red chromatophores on articles 4 and 5; other articles whitish; flagella transparent.

P1: merus to dactylus orange, with a few red chromatophores on upper surface, lower surface lighter in color. Tip of dactylus dark brown and transparent. P2: distal part of merus to dactylus orange, lighter than on P1, with a few red chromatophores; proximal and lower part of merus opalescent white. P3-5 transparent, P3 and P4 with a few chromatophores from merus forwards. Pleopods transparent, with a few chromatophores on basis.

Size

Largest specimens in material examined: cl. 12- 15 mm, tl. 36-48 mm.

ECOLOGY

Upogebia mediterranea occurs in muddy sand or sediments with Posidonia , 0- 25 m. Burrows in firmground were studied by Asgaard et al. (1997).

VARIATIONS

1) The upper mesial distal spine of P1 carpus is accompanied laterally by one to three spinules; 2) the longitudinal row of tubercles on the lateral and mesial sufaces of the dactylus, as well as the proximal mesial round tooth on cutting edge is sometimes faint or absent (rare); and 3) arthrobranchs are composed of a single row of lamellae on either side of the rachis in specimens from the Gulf of Gabès, Tunisia, and a double row of lamellae in the rest of the material examined.

D

REMARKS

This species, similar to both U. deltaura and U. nitida , was first mentioned by de Saint Laurent & Le Loeuff (1979: 93) as an undescribed species related to U. nitida .

According to Dworschak (1992: 223), it was cited again, under the present name mediterranea , as a taxon related to U. deltaura , by de Saint Laurent in a key to western Atlantic and Mediterranean species of Upogebia (“Tableau de détermination des Upogebia de l’Atlantique nordoriental et de Méditerranée”). This key was given to participants of the IInd Colloquium Crustacea Mediterranea in Ancona, Italy (1979) but was never validly published. The species was subsequently recognized by Kocata ș (1981), Thessalou-Legaki & Zenetos (1985) and Thessalou-Legaki (1986). It was mentioned by Noël (1992) in an identification key to decapod crustaceans from France and common European species (“Clé préliminaire d’identification des Crustacea Decapoda de France et des principales autres espèces d’Europe”). As Noël’s key was published and contained diagnostic details, Noël is now the species’ author according to the International Code of Zoological Nomenclature ( ICZN 1999). The type series includes all specimens examined by him or de Saint Laurent before the species name was published. They were considered as syntypes and a lectotype was selected from them.

Upogebia mediterranea resembles U. deltaura in: 1) the shape and armature of the rostrum; and 2) the anterolateral border of carapace unarmed. It differs in the following characters: 1) the pleura of the first abdominal segment is rounded postero-ventrally in U. mediterranea (pointed in U. deltaura ); 2) A2 peduncle is unarmed in U. mediterranea (with a lower distal spine on article 3 in U. deltaura ); 3) P1 merus bears spinules and/or tubercles on its lower border in U. mediterranea (with spines and spinules in U. deltaura ); 4) P1 propodus has no mesial distal spine near the base of the fixed finger in U. mediterranea (spine present in U. deltaura ); 5) P1 dactylus bears a proximal mesial round tooth on the cutting edge in U. mediterranea (tooth often absent in U. deltaura ); 6) P1 dactylus and fixed finger are of about the same length in U. mediterranea (dactylus often longer than fixed finger in U. deltaura ); 7) P3 merus is unarmed in U. mediterranea (with spinules and tubercles on the lower border in U. deltaura ); and 8) the telson is approximately quadrate in U. mediterranea (about 1.2 times as wide as long in U. deltaura ).

Upogebia mediterranea was considered closely related to U. nitida (A. Milne-Edwards, 1868) by de Saint Laurent & Le Loeuff (1979: 39, 93) and d’Udekem d’Acoz (1999: 157) or a subspecies of the latter (d’Udekem d’Acoz 1995). It was actually confused with the U. nitida in Le Loeuff & Intès (1974: 58, fig. 20). The small female these authors reported from Congo, of 5.5 mm in carapace length, and now deposited in the MNHN ( MNHN Th 231), actually belongs to U. mediterranea . The excellent figures given agree perfectly with those of the latter species ( Figs 28 ; 29 ), but with neither U. nitida ( Figs 30 ; 31 ) nor U. furcata (Aurivillius, 1898) (see Le Loeuff & Intès 1974: fig. 18), a common species in the area. This female, bearing a large coxal spine on both P1 and P2, represents the only record of U. mediterranea outside the Mediterranean.

More likeness can be found between U. mediterranea and a newly established, non-european species, U. senegalensis Ngoc-Ho, 2001 from Senegal. The latter resembles both U. mediterranea and U. nitida and the three are sometimes hard to differentiate.

Similarities and differences between U. mediterranea and U. senegalensis are presented below with some details and summerised in Table 2. U . mediterranea and U. senegalensis are compared with U. nitida under the latter species and summerized in the same table.

U. mediterranea resembles U. senegalensis (see Ngoc-Ho 2001a: figs 1-3) in: 1) the shape of rostrum; 2) the shape and spinulation of P1 except for a subdistal tooth absent on dactylus cutting edge; and 3) an acute coxal spine on both P1 and P2.

It differs from U. senegalensis in: 1) the large A2 scale with pointed tip (small A2 scale with blunt tip in U. senegalensis ); 2) absence of subdistal tooth on the cutting edge of P1 dactylus (tooth present in U. senegalensis ); 3) P2 bears a dorsal subdistal spine on both merus and carpus (P2 unarmed in U. senegalensis ); and 4) the telson is subquadrate (telson slightly wider than long in U. senegalensis ).

Upogebia nitida (A. Milne-Edwards, 1868) ( Figs 30 ; 31 )

Gebiopsis nitidus A. Milne-Edwards, 1868: 63 , pl. 18, figs 4-7. — ° Bouvier 1917: 118.

Gebia (Gebiopsis) nitida – Ortmann 1893: 50, pl. 4, fig. 2.

Upogebia (Calliadne) nitida – de Man 1928 b: 24, 50 (key).

Upogebia nitida – de Saint Laurent & Le Loeuff 1979: 38, fig. 2. — d’Udekem d’Acoz 1999: 157.

Non Upogebia nitida – Le Loeuff & Intès 1974: 58 (part), fig. 20a-o (= Upogebia mediterranea Noël, 1992 ).

TYPE MATERIAL. — Lectotype:; paralectotypes, 2, from Cape Verde Islands , Cape of St Vincent, Atlantic, subsequent designation by de Saint Laurent & Le Loeuff (1979) ( MNHN Th 22).

MATERIAL EXAMINED. — France. Tamaris-sur-Mer, near Toulon, Sollaud coll., No. ES 38, 25.XII.1932, 2 cl. 16 and 16.5 mm, tl. 51 and 52 mm (figured) ( MNHN Th 1393). — Banyuls, P. Noël coll., 4.VII.1975, 1 ovig. cl. 10 mm, tl. 31 mm, without P1 ( MNHN Th 660).

Italy. Toscana, Livorno, G. Herweck leg., 1 cl. 15.5 mm, partly dried, without pereopods ( SMF 28327).

Algeria. Gulf of Oran, P. Pallary coll., 1900, 1 cl. 13 mm, tl. 38 mm ( MNHN Th 1392).

St Vincent. Cape Verde Islands, Atlantic, de Folin coll., lectotype cl. 8 mm, tl. 21 mm, 2 paralectotypes tl. 16 and 17 mm (specimens broken, poor condition, measurements by de Saint Laurent & Le Loeuff 1979) (figured) ( MNHN Th 22). — Cape Verde Islands, Talisman Exp. , dredge, 26.VII.1883, 1 cl. 6 mm, 1 ovig. cl. 8 mm, 4 juv. ( MNHN Th 23).

Mauritanie. G. Bolloré coll., 1965, 2 broken, cl. 12 and 14 mm (measurements by de St Laurent & Le Loeuff 1979) ( MNHN Th 383).

Senegal. Gorée, in lobsters net, Cadenat coll., 10.X.1950, 1 ovig. cl. 11.5 mm, tl. 35 mm (figured) ( MNHN Th 278) ; Paraïso coll., 27. V.1948, 1 cl. 5 mm ( MNHN Th 279). — Between Takona and Bel Air , 14-15 m, I. M. Marchad coll., 19.III.1954, 1 cl. 11 mm ( MNHN Th 282) ; in gallery of Toredo molluscs, 19.XI.1950, 1 cl. 2.75 mm ( MNHN Th 281). — Dakar, R. Mauny coll., 25.IX.1949, 1 cl. 11 mm, 1 cl. 10 mm ( MNHN Th 276) ; R. Mauny coll., 14.IX.1949, 1 cl. 11 mm ( MNHN Th 277) ; 25.IX.1949, 3 cl. 6-11mm, 1 cl. 6 mm ( MNHN Th 273) .

Gulf of Guinea. Sao Tomé , Calypso , stn 18, in front of Praia Lagarto, 11.VI.1956, 1 ovig. cl. 7 mm ( MNHN Th 270) ; stn 19, 4- 5 m, 12.VI.1956, 2 ovig. cl. 4 and 5.5 mm ( MNHN Th 271). — Principe, stn 94, 31 m, 27.VI.1956, 1 ovig. cl. 5.5 mm ( MNHN Th 272). — Annobon Island , Ombango, dredge, A. Crosnier coll., 11.XII.1965, 1 cl. 6 mm ( MNHN Th 229), 50-60 m, 1 cl. 5 mm ( MNHN Th 230) .

DIAGNOSIS

Rostrum ( Figs 30A, B ; 31A, B ) approximately triangular, slightly overreaching eyestalks, about as long as wide at base, with two teeth on either lateral border; anterolateral border of carapace unarmed; telson ( Figs 30C ; 31G ) 1.1-1.2 times as wide as long. A1 ( Figs 30B ; 31B ) peduncle with lower spine on article 1. P1 slightly stouter in male ( Fig. 31D, E ) than in female ( Figs 30D, E ; 31C ) merus with lower border unarmed or denticulate; carpus with upper mesial distal spine; fixed finger with three or four round teeth on proximal half of cutting edge; dactylus about as long as fixed finger with proximal and (often) subdistal round tooth on cutting edge ( Fig. 31E, J ). P2 ( Figs 30F ; 31F ) unarmed. Small coxal spine on either P1 or P2, seldom on both (and in male only).

Colour

Unknown.

Size

Types and material from northwest Africa: cl. 8- 13 mm, tl. 21-38 mm.

Specimens examined from Europe: cl. 15- 16.5 mm, tl. 51-52 mm.

VARIATIONS

1) rostral spines, often small, are larger in the lectotype ( Fig. 31A ); 2) P1 carpus bears a very small lower distal spine in a few specimens ( Fig. 31I ), likewise for lateral spinule near the base of the fixed finger ( Fig. 30D ); the proximal and/or subdistal tooth on the dactylus cutting edge is small or absent ( Fig. 30D, E ); and 3) the telson is subquadrate in small specimens of cl. <6-6.5 mm.

REMARKS

Upogebia nitida differs from U. mediterranea in many features: 1) the rostrum is triangular in U. nitida (elongated triangular in U. mediterra-

elongated nea); 2) P1 merus has no upper subdistal spine, the lower border is unarmed or with denticles, P1 carpus bears a small upper mesial distal spine and no lower distal spine in U. nitida (P1 merus with upper subdistal spine, lower spinules and tubercles; P1 carpus with large upper mesial distal spine, small lower distal spine in U. mediterranea ); 3) P1 propodus with the upper border unarmed and no lateral spinule near the base of fixed finger in U. nitida (P1 propodus with upper subdistal spine and a lateral spinule near the base of fixed finger in U. mediterranea ); 4) P1 dactylus with a mesial proximal and subdistal tooth on the cutting edge in U. nitida (no subdistal tooth on the cutting edge in U. mediterranea ); this character varies; 5) P2 is unarmed in U. nitida (both P2 merus and carpus bear an upper subdistal spine in U. mediterranea ); and 6) small coxal spine on either P1 or P2, seldom on both in U. nitida (long and acute coxal spine on both P1 and P 2 in U. mediterranea ).

Similarities between U. nitida and U. senegalensis are (see Ngoc-Ho 2001a: figs 1-3): 1) A2 scale is small, not acute at tip; 2) a subdistal round tooth is present on the cutting edge of P1 dactylus; 3) P2 is unarmed; and 4) the similar shape of the telson.

Differences between U. nitida and U. senegalensis are: 1) the shape of the rostrum; 2) the spinulation of P1, the same as for U. mediterranea ; and 3) small coxal spine is present on either P1 or P2, seldom on both in U. nitida (coxal spine present on both P1 and P 2 in U. senegalensis ).

Distinguishing characters between U. nitida , U. mediterranea and U. senegalensis are presented in Table 2.

Upogebia pusilla ( Petagna, 1792) ( Figs 32 ; 33 )

Astacus pusillus Petagna, 1792: 418 , pl. 5, fig. 5.

Thalassina littoralis Risso, 1816: 76 , pl. 3, fig. 2.

Gebia venetiarum Nardo, 1869: 314 , pl. 13, fig. 3.

Gebia litoralis – Stalio 1877: 107. — Carus 1885: 490. Gebia littoralis – Desmarest 1825: 234. — H. Milne Edwards 1837a: 313; 1837b: pl. 49, figs 1-11. — Lucas 1840: 475. — Heller 1863: 205, pl. 5, figs 12- 15. — Czerniavsky 1868: 126. — Lafont 1868: 522. — ° Fischer 1872: 429. — de Folin & Périer 1879: 211. — Stossich 1880: 206. — Sars 1883: 44. — Claus 1884: 746. — ° Graeffe 1902: 69. —?Gibert i Olivé 1920: 51, 1 fig.

Gebios littoralis – Risso 1827: 51.

Upogebia (Upogebia) littoralis – Borradaile 1903: 543. — de Man 1927: 29, figs 11-11b; 1928b: 23, 41 (key). — Monod 1937: 2.

Upogebia (Upogebia) litoralis – Pesta 1918: 197 (part), fig. 61a.

Upogebia littoralis –?* Sars 1884: 182, pls 3-5. — Balss 1926: 26 (key). — Tucker 1930: 1. — Monod 1937: 2. — Makarov 1938: 52. — Bouvier 1940: 107, fig. 71. — Popovici 1940: 421, 485. — Zariquiey Alvarez 1946: 107, 108, pl. 5 fig. b, c. — ° Gottlieb 1953: 441. — * Heegaard 1963: 460, figs 34-39. —? Luther & Fiedler 1965: 50, 175, fig. on pl. 23. — Daguerre de Hureaux 1971: 67, pls 1-4. — Ngoc-Ho 1981: 245.

Upogebia litoralis – Ott et al. 1976: 62, figs 1-3, tabls 1, 2, pls 1, 2. — Balss 1936: 14.

Upogebia pusilla – Holthuis 1947: 321, fig. 1(5); 1956: 325; 1961: 32; 1977: 57. — * Bourdillon- Casanova 1960: 109. — Lewinsohn & Holthuis 1964: 54. — Pérès & Picard 1964: 55. — Picard 1965: 60. — Băcescu 1967: 220, figs 101-103. — de Gaillande 1968: 382; 1970: 377. — Zariquiey Alvarez 1968: 231, fig. 94a. — Le Gall 1969: 400. — Števčić 1969: 128; 1971: 529; Števčić 1990: 215. — Ktari- Chakroun & Azouz 1971: 21. — de Saint Laurent 1971: 1261 (key). — ° Pastore 1976: 107. — Kattoulas & Koukouras 1974: 346. — Neves 1974: 15, fig. 5; 1987: 256. — * Thiriot 1976: 350. — Holthuis 1977: 57. — Merker Pocek 1977: 110. — Ngoc-Ho 1977: 313. — Beaubrun 1979: 77, fig. 49. — de Saint Laurent & Le Loeuff 1979: 43, fig. 5a. — Monchartmont 1979: 73. — Bourdon 1980: 1. — Domenech et al. 1981: 150, figs 37, 38. — Dworschak 1981: 25, figs 1-8; 1983: 20, figs 1-14; 1987a: 338, figs 1-7; 1987b: 421, fig. 1; 1988: 52, figs 1-10; 1992: 224, fig. 18a, c, g, h. — Manning & Števčić 1982: 296. — Cottiglia 1983: 74, figs 27b, 28, 29. — García Raso 1983: 320, fig. 1; 1985: 21. — Riedl 1983: 483, pl. 177, 2 figs. — Chaud 1984a: 1- 176; 1984b: 194. — d’Udekem d’Acoz 1986: 103, figs 3-5; 1989: 176, fig. 6; 1995: 60; 1999: 157. — Thessalou-Legaki & Zenetos 1985: 311. — Müller 1986: 117. — Thessalou-Legaki 1986: 182. — Türkay et al. 1987: 92. — Geiss 1990: 208, 1 col. fig. — Moyse & Smaldon 1990: 520, fig. 10.13 (part). — Pérez Sánchez & Moreno Batet 1991: 140, 1 fig. — de Vaugelas 1991: 56, photo 2; 1998: 3, figs 1-3. — ° García Raso et al. 1992: 258. — ° Koukouras et al. 1992: 223. — Noël 1992: 82. — ° Mayoral et al. 1994: 236. — Froglia 1995: 8. — Gonzalez Pérez 1995: 136. — Hayward et al. 1995: 434, fig. 8.52 (part). — Moosleitner & Patzner 1995: 120, 1 fig. — Falciai & Minervini 1996: 149, 4 figs. — Astall et al. 1997b: 671, fig. 1, tabls 1-5. — Ingle 1997: 80, fig. 7.4. — Kevrekidis et al. 1997: 799, figs 1-7, tabls 1- 3. — Pinn et al. 1999a: 103, figs 2E, F; 1999b: 1461, figs 1F-H, 7, 8, tabls 1-6. — *González-Gordillo et al. 2001: 279. — Markham 2001: 197, tabls 1, 2. — * Martin 2001: 82, 1 fig. — ° Türkay 2001: 289. — Lopez de Rosa et al. 2002: 88. — Noël 2002: 241.

Upogebia (Upogebia) pusilla – Noël 1992: 82 (key).

Upogebbia pusilla – Pérès & Picard 1964: 56.

TYPE MATERIAL. — Whereabouts unknown.

MATERIAL EXAMINED. — Great Britain. W English Channel, Holme’s collection, stn 114, 50°19.1’N, 4°27.2’W, 35-37 m, 1 cl. 13 mm ( NHML 2002.839).

France. Atlantic, Trébeurden, 1987, 4 cl. 10- 16 mm, 2 cl. 15.5 and 16 mm ( MNHN Th 1329) ; F. Meunier coll., III.1983, 1 cl. 7.5 mm, 1 cl. 14 mm ( MNHN Th 1330). — Île Grande, clean sand, F. Meunier coll., 21.IX.1984, 3 cl. 10-17 mm ( MNHN Th 1331) ; muddy sand, 2.X.1978, 1, 1, cl. 16 mm ( MNHN Th 1340). — Brest, muddy sand, M.-N. Helleouet coll., 6.V.1997, 2 cl. 13.5 and 16 mm ( MNHN Th 1332). — Bassin of Marennes-Oléron, in sediments of “Claires de Sendre”, D. Fouché coll., II.1994, 1 cl. 18 mm, tl. 50 mm (figured) ( MNHN Th 1317), 1 cl. 13 mm ( MNHN Th 1410). — Bay of Chingoudy ( western Pyrénées ), A. Chaud coll., V.1982, 14 cl. 9- 17 mm, 3 (1 ovig.) cl. 13-14.5 mm ( MNHN Th 652). — Bassin of Arcachon , port de la Hume, intertidal (0.8-1.2 m), A. Chaud coll., 3 cl. 6.5- 18 mm, 2 cl. 16 and 17 mm, tl. 46 and 48 mm (figured) and 3 cl. 9-16 mm ( MNHN Th 645). — Auray , Brittany, R. Bourdon coll., 1 cl. 14 mm ( MNHN Th 541). — Concarneau , Baron de St- Joseph’s coll., VIII.1892, 1 cl. 15 mm ( MNHN Th 39). — Carry-le-Rouet, A. Vayssière coll., 1912, 1 cl. 13 mm, 1 cl. 10 mm ( MNHN Th 36). — Coast of France , 1939, 1 cl. 12 mm ( MNHN Th 35). — Concarneau , in burrows of muddy sand, under rocks, excursion RMNH, 12.IX.1958, 1, 1 exuvia ( RMNH D 12438). — Bay of Arcachon , R. Bourdon, III.1964, 2 ( RMNH D 21559) ; A. W. Lacourt, VII.1976, 1 dry ( RMNH D 45673). — Baie of Chingoudy , Hendaye, H. Nouvel coll., 1.VIII.1953, 145, 128 (93 ovig.) ( RMNH D 34027) ; A. Chaud coll., 2 cl. 13 and 15 mm, 1 cl. 14 mm ( MNHN Th 646) .

Mediterranean, Bay St-Trinité, 5 km from Porto Vecchio, Corsica, W. F. Roodenburg, 21.VII.1968, 1 ovig. ( RMNH D 26474). — Banyuls, meadows of Grosse Island , P. Noël coll., 16.V.1978, 2 cl. 8 and 9 mm, 1 ovig. cl. 15.5 mm, tl. 47 mm (figured) and 2 cl. 12.5 and 14 mm ( MNHN Th 621) ; 12.V.1978, 1 cl. 14 mm, tl. 45 mm (figured), 1 cl. 14 mm ( MNHN Th 622) ; 22.V.1976, 1

cl. 6 mm (MNHN Th 673) ; Y. Bhaud coll., II.1976, 1 cl. 12 mm (MNHN Th 672) ; M. de Saint Laurent coll., V.1959, 2 cl. 7 and 13 mm, 1 juv. cl. 4 mm (MNHN Th 318) ; J. Forest coll., 14.VII.1950, 1 cl. 8.5 mm (MNHN Th 219) ; under a rock, J. M. Amouroux coll., 24. V.1976, 1 cl. 14 mm (MNHN Th 659) ; excursion Leiden Biologists, 25.IX.1937, 9 juv. ( RMNH D 11728). — Nice, Travailleur, VII.1881, 1 cl. 11mm (MNHN Th 37) ; 1

cl. 12 mm ( MNHN Th 204). — Monaco, Prince de Monaco coll., 6.I.1905, 1 just moutled and exuvia, approx. cl. 13 mm ( MNHN Th 1328). — Marseille , Ponteau , 3 cl. 4.5- 12 mm, 1 cl. 15 mm (poor condition) ( SMF 28322). — Golfe de Fos , near Electricity plant at Ponteau, 2 m, H. Zibrowius coll., 5.IV.1977, 5 cl. 8-11 mm, 21 9-11.5 mm (poor condition) ( MNHN Th 598). — Anse des Laurons, near golfe de Fos, sand, 3 m, H. Zibrowius coll., 21.IV.1974, 1 cl. 10.5 mm ( MNHN Th 366). — Le Brusc, Var, muddy sand, 0.5 m, P. Noël coll., 14.XI.1997, 1 cl. 12.5 mm ( MNHN Th 1334). — Languedoc, lagune de Manguis, J. L. Bouchereau coll., 1992, 1 cl. 14 mm ( MNHN Th 1335). — Corsica, near Porto Vecchio, C. Monniot coll., VII.1997, exuviae only, 2 rostrums, several pereiopods ( MNHN Th 1333). — Cap Martin, Monaco, 10-15 m, dredge, 13.XII.1912, don. H. Nouvel, 11, 9 ( RMNH D 34038). — Monaco, H. Nouvel, 2.VII.1937, 2, 3 ( RMNH D 34039) ; 1936, 2, 1 ovig. ( RMNH D 34040) ; 1 ( RMNH D 34041) ; IV.1938, 1, 1 ( RMNH D 34042) ; 19.IV.1912, Prince de Monaco coll., H. Nouvel don., 1 ( RMNH D 34043) ; 1936, collection H. Nouvel , 1 ( RMNH D 34044) .

Portugal. Olhão, C. d’Udekem d’Acoz coll., 19.VII.1980, 5 cl. 10.5-17.5 mm, 2 ovig.

cl. 14.5 mm (d’Udekem d’Acoz). — Cabo de Arnel, near Faro, excursion RMNH, 29.VII.1962, 1 exuvia ( RMNH D 19026). — Praia do Faro, Algarve, excursion RMNH, 6. V.1971, 4, 2 (1 ovig.) ( RMNH D 27490). — Caldeira do Mainho, Costado do Pedro José, Algarve, burrows in dry mud, at low tides, excursion RMNH, 5. V.1971, 3, 8

(4 ovig.) ( RMNH D 27491). — Caldeira do Moinhoc Costado do Pedro José, Algarve , burrows in dry mud, at low tide, excursion RMNH, 3.V.1971, 1 ( RMNH D 28197 ). — Praia de Faro, Algarve, mudflat at low tide, excursion RMNH, 2.XI.1974, 3, 3 ( RMNH D 36259) ; 3.XI.1974, 9, 12 ( RMNH D 36260). — SPMOPO1974, stn 080, Algarve , slightly W of Faro, mudflats near Praia de Faro, low tide, excursion RMNH 1974 , 3.XI.1974, 2 ( RMNH D 38477) .

Spain. Port Lligat, 1-5 m, G. Fischer coll., 14

cl. 12.5- 15 mm, 16 (6 ovig.) cl. 9-15 mm ( SMF 28974). — Port Lligat, U. Pettke coll., 1.VIII.1985, 6 cl. 12.5-13.5 mm, 8 (3 ovig.) cl. 8.5- 13.5 mm ( SMF 28975). — Gerona, Cadaqués , M. Türkay coll., 30.III.1985, 8 spec. badly damaged, cl. c. 10-12 mm ( SMF 28976). — Cadaqués , I. Gordon coll., 2 exuviae ( NHML 1955.2.28.91) ; Zariquiey Alvarez pres., 1, 1 ( NHML 1954.12.30.96- 97). — Mouth of Ebre river, Calypso Exped., 1977 , stn 4, 1 cl. 12 mm (MNHN Th 623). — Canary islands, Lanzarote, Santaella coll., IX.1973, 2 cl. 8 and 10 mm, 1 ovig. cl. 9 mm (MNHN Th 317) ; A. K. Totten coll., 10, 6 (3 ovig.) ( NHML.1956.5.2.92-101). — Cadaqués near Barcelona, R. Zariquiey Alvarez , 1940, 2 broken spec. ( RMNH D 5253 ). — Playa Jonquet , N of Cadaqués, in c. 50 cm water, in holes in the ground, L. B. Holthuis, 13.VIII.1950, 3, 1, 1 juv. ( RMNH D 6751 ). — Rio Fluvia, in front of mouth of Golfo de Rosas , brackish water, L. B. Holthuis, 16.VIII.1950, 1 ovig. ( RMNH D 6752 ). — Cadaqués , sand and stones, in c. 75 cm water, in holes, L. B. Holthuis, 6.VIII.1950, 3, 2 ( RMNH D 6753 ) ; from fishermen, 4-16.VIII.1949, 1 ( RMNH D 6200 ) ; in c. 20 cm, 14.VIII.1959, 1 ( RMNH D 15229 ). — Cala Junquet, N of Port Lligat, burrows in sand with stones, c. 20 cm, L. B. Holthuis, 20.VIII.1959, 1, 2 (1 ovig.), 6 juv. ( RMNH D 15230 ). — Playa d’en Pere Fet, Cadaqués, sand, L. B. Holthuis, 23.VIII.1959, 1 ( RMNH D 15231 ). — Cala Jugadora, S of Cabo de Creus, in sand, 0-0.2 m, L. B. Holthuis, 31.VII.1961, 7, 1 ( RMNH D 16182). — Ria de Arosa, N of Cabo Cruz , excursion Leiden Biologists , 23.VII.1962, 7, 8 ovig .

( RMNH D 17914). — Rio Unia, SE of Ria de Arosa, washed ashore, excursion RMNH, 17.VII.1962, 1 exuvia ( RMNH D 19027). — Mouth of bay, SE of El Grove, from mud at low water mark, excursion RMNH, 24.VII.1962, 1 ( RMNH D 19028). — Playa Barraña, Peninsula Chazo, in holes, sandy beach with some rocks, intertidal, excursion RMNH, 7.VII.1963, 22, 28 ovig. ( RMNH D 19541). — Punta San Vicente, Peninsula del Grove, in sand, excursion RMNH, 1.VII.1963, 2 ( RMNH D 19900). — Peninsula del Grove, Ria de Arosa, 0- 0.5 m, excursion RMNH, 10.VII.1963, 1 exuvia ( RMNH D 19901). — Playa de Barraña, Ria de Arosa, excursion RMNH, 10.VIII.1964, 6 (4 ovig.) ( RMNH D 23797 ). — Playa Cerantes, Vigo, R. Margalef, II.1954, 1 ( RMNH D 28185 ). — Playa Cerantes, Vigo, A. Figueras, 6.III.1954, 3 ( RMNH D 28186 ). — Cabo de Creus, N of Cadaqués, 50 fms, J. Fortuny, VI.1947, 1 ovig.

( RMNH D 28189 ); 27.VII.1945, 1 ( RMNH D 28190 ). — Bahia de Cadiz , 5.5 m, sand, gravels and algae, large dredge, excursion RMNH, 13.V.1971, 1 exuvia ( RMNH D 28196 ) ; 12-18 m, sand and algae, excursion RMNH, small dredge, 11.V.1971, exuviae of 3- 4 spec. ( RMNH D 28199 ). — Baleares, Ibiza, San Antonia, R. Zariquiey Cenarro, VII.1934, 3, 1 juv. ( RMNH D 28200 ) ; 1 damaged spec. ( RMNH D 35722). — SPMOPO1974, stn 049, Bay of Cadiz, coast of Punta de la Cruz, washed ashore, excursion RMNH, 26.X.1974, 1, 1 ( RMNH D 38479). — East bank of Guadalquivir river near Bonanza , mud, H. O. von Hagen, summer 1960, 1 ( RMNH D 23607 ). Italy. Gulf of Gaeta , Licola , 1966/1968, Doerjes leg., 1 cl. 13 mm ( SMF 28326). — Sicilia, K. Koltar coll., 2 (damaged), larger specimen cl. 12 mm ( ZMUC 2706 ). — Grado, N Adriatic, intertidal, C. d’Udekem d’Acoz coll., IX.1997, 1 cl. 18 mm, tl. 50 mm (figured) and 2 cl. 15 and 16.5 mm, 3

(1 ovig.) cl. 15.5-17.5 mm (MNHN Th 1318). — Grado lagoon, 0.3-0.4 m, P. Dworschak leg., 1979, 6 cl. 16-20 mm, 3 (1 ovig.) cl. 11-18 mm (MNHN Th 630). — Lido di Starangano, 0.2-0.3 m, P. Dworschak leg., 1979, 5 cl. 7-15 mm, 3

cl. 8-15 mm (MNHN Th 631). — Entrance of Venice Lagoon, P. Sabbioni, in Zostera noltii meadow, intertidal, C. d’Udekem d’Acoz coll., 1.IX.1997, 1

cl. 6 mm, 1 ovig. cl. 17 mm (d’Udekem d’Acoz). — Naples, R. Gurney presented, 2, 2 (1 ovig.) ( NHM. 1947.3.18. 756-757). — Naples, 2, 1 ( RMNH D 937); G. Stiasny coll., VIII.1924, 3 ( RMNH D 2421). — Mergellina, Naples, 3-4 m, L. B. Holthuis coll., 29.IV.1950, 54, 44

(21 ovig.) ( RMNH D 6516); E. Caroli coll., V.1959, 14, 16 (2 ovig.) ( RMNH D 13014) ; J. H. Stock coll., V.1957, 1, 3 ( RMNH D 15228 ). — Porto Cesaro, Strea Bay, in sand, 0.2- 0.5 m, II.1967, 3 cl. 11-14 mm, 2 cl. 11 and 11.5 mm (MNHN Th 318). — Porto Cesaro , 1 ovig. cl. 15 mm (MNHN Th 319) .

Croatia. Rovinj, Kuvi, seagrass, Excursion University Frankfurt , 18.VIII.1989, 2 cl. 8.5 and 14 mm, 3 cl. 9-15 mm ( SMF 28323). — Rovinj, 1987, 2 cl. 8.5 and 10 mm, 2 cl. 11.5 and 12.5 mm ( SMF 28324). — Istria, Rovinj, Val Salina , F. B. Burin coll., 21.VIII.1989, 3 cl. 10-14 mm ( SMF 28325). — Rovinj, Števčić coll., 9.XI.1977, 2 cl. 11 and 13 mm ( MNHN Th 613) ; P. Dworschak leg. 1979, 2 cl. 9 and 11.5 mm, 2 cl. 8 and 13 mm ( MNHN Th 632) .

Romania. Near Constantine, Calypso Exped. 1977 , stn 3, 6.X.1977, 1 juv. cl. 4 mm ( MNHN Th 628).

Greece. Thessalonique, 25.IX.1930, 1 cl. 19 mm ( MNHN Th 209). — Lesbos Island , C. d’Udekem d’Acoz coll., 10.VII.1992, 5 cl. 5-7 mm ( d’Udekem d’Acoz ). — Aegean Sea , near harbour of Porto Lago , 0-2 m, excursion Leiden Biologists, 29.VI.1959, 16, 38 ( RMNH D 13073) ; leg. Cardo Godo, don. R. Zariquiey Alvarez, 2, 1 ovig. ( RMNH D 28195 ) ; Kolovrechtis, Evvoia I., 0-2 m, 2.VI.1971, 2, 2 (A.U. TH. G1 767). — Chalkidiki, Kassandra, Nea Fokia, G. Gorgiadis, 15.VIII.1971, 1 ( RMNH D 28724). — S of Euboea Gulf , c. 38°30’N, 24°E, A. Koukouras, 3.II.1972, 3 ( RMNH D 30380 ). — Chochlakos, Rhodes, SLM 4012, 16.X. 1970, 26 m, 11 (all young specimens), 6 juv. ( RMNH D 35724) .

Turkey. Izmir, A. Kocataș coll., 2 cl. 13 and 14 mm, 2 cl. 13 and 15 mm ( MNHN Th 619). — Tuzla near Izmir, A. Kocataș, 10.X.1966, 8, 1 ( RMNH D 23345 ).

Malta. C. G. Thorogood pres., 14, 3 ovig.

( NHML.1945. VI.15.1-15); 12, 10 (7 ovig.) ( NHML.1945.VII.12.11-20).

Mediterranean coast of Israel. El Akhziv (= Gesher- Hasiv), N of Nahariya , together with Synapta spec. , L. Fishelson, 25.II.1960, 1 ( RMNH D 19217) ; 24.VI.1962, 1 ( RMNH D 19218) .

Suez. Suez Canal, A. Gruvel coll. 27.II.1904, 4

cl. 10-11.5 mm, 2 cl. 9 mm (MNHN Th 34).

Algeria. Bône, H. Lucas coll., 1849, 4 cl. 9- 12.5mm, 2 cl. 8 and 10 mm ( MNHN Th 210). — Castiglione, R. Dieuzeide, VIII.1947, 3, 5 ( RMNH D 6199).

Morocco. IV.1965, 2 cl. 10 mm (poor condition), 1 cl. 10 mm ( MNHN Th 40) ; X.1964, 1 cl. 14.5 mm, 2 cl. 9 and 10 mm ( MNHN Th 43). — Fedhala, J. Lionville coll., VIII.1923, 2 cl. 5.5 and 6 mm, 2 cl. 5.5 and 8 mm ( MNHN Th 546) ; 1 cl. 10 mm ( MNHN Th 544) ; Lionville, Lepiney & Bernaudot coll., 5.II.1935, 1 cl. 11 mm ( MNHN Th 44) ; Lionville, Lepiney & Mineur coll., 5.III.1955, 1 cl. 11.5 mm ( MNHN Th 41). — Bon Znina, Lepiney coll. 30.X.1940, 2 cl. 7 mm ( MNHN Th 547). — Tanger, H. Charnier coll., 1, 1 cl. 10 mm ( MNHN Th 545). — SPMOPO 1974, stn 34, S of Rabat, 1 km S of Oued Yquem, rocky shore with sandy lagoon, excursion RMNH, 19.X.1974, 1 ( RMNH D 38476). — SPMOPO 1974, stn 38, atlantic coast, 23 km S of Rabat, 0- 0.2 m, rock pools, excursion RMNH, 20.X.1974, 1 ( RMNH D 38478). — SPMOPO 1974, stn 40, atlantic coast, 16 km S of Rabat, Temara, sandy beach with rocks, in rock pools, excursion RMNH 1974 , 21.X.1974, 1 ( RMNH D 38480 ) .

Tunisia. Tunis, El Biban, Tidevandzonen, 17. V.1938, Nordisk Insulin-Lab’s Exp., 1 cl. 10 mm ( ZMUC 2707). — Gulf of Tunis, between Sidi Rais and A in Oktor, Manning, Forest, de Saint Laurent and Jones coll., 16.VIII.1973, 1 cl. 10.5 mm, tl. 30 mm, 10 cl. 8.5-16.5 mm, 16 (14 ovig.) cl. 8-16 mm ( MNHN Th 713). — Salammbô, R. Manning & R. Ingle coll., 21.II.1974, 1, 1 ( NHML.1974.182).

Mauritania. G. Bolloré coll., 1965, 1 cl. 13 mm ( MNHN Th 384).

?São Thomé (West Africa). M. Nobre coll., 1889, 1 cl. 14 mm (MNHN Th 798).

DISTRIBUTION. — South of the British Isles, northwestern Brittany, France in the North, Mediterranean including Corsica (de Vaugelas 1991, 1998) to Mauritania and São Thomé? in the South. According to Pesta (1918), Bouvier (1940), Zariquiey Alvarez (1968), Upogebia pusilla can be found as far north as Norway, the North Sea while according to Poulsen (1941), the species was absent from Norway. More recently, Moyse & Smaldon (1990) considered the distribution of Upogebia pusilla to include the British Isles, probably only the South. In this work, a male from the English coast of the English Channel has been found ( NHML 2002.839).

The southernmost localities of the species distribution in Eastern Atlantic ( Mauritania and Sao Thomé) were given by a single female of cl. 13 mm (tl. 43 mm) collected in 1965 and a male of cl. 14 mm (tl. 45 mm), collected in 1889. Both are in fair condition, but it is questionable whether the label accompanying the specimen from Sao Thomé is authentic.

The species is common around the Mediterranean and is also found in the Black Sea ( Makarov 1938) and the Suez Canal ( Monod 1937).

DIAGNOSIS

Rostrum ( Fig. 32A, B ) obtuse distaly, about 1.2- 1.3 times as long as wide at base, with four or five teeth on lateral border; anterolateral border of carapace with spine; telson ( Fig. 32H ) approximately 1.2-1.3 times as wide as long, distal margin narrower than proximal, posterior border straight or slightly concave medially.

A1 and A2 peduncle ( Fig. 32B ) with spine on lower margin of article 1 and 3 respectively; A2 scale with distal spine. Md with large mesio-anterior pointed tooth. Mxp1 without epipod, Mxp3 with epipod.

P1 stouter in male ( Fig. 32C, D ) than in female ( Fig. 32F, G ). Ischium with two to four spines on lower border. Merus with upper subdistal spine, three to six spines and spinules on proximal half of lower border and a double or single row of denticles distally. Carpus with large upper mesial distal spine and lower distal spine. Propodus more slender in female, dilated distally in large male (cl.> 11.5- 12 mm); upper border with lateral longitudinal smooth crest and mesial crest bearing tubercles or denticles ( Fig. 32E ); small mesial upper subdistal spine, large mesial upper distal spine; both lateral and mesial surfaces with distal spine near cutting edge of dactylus; fixed finger shorter than dactylus, cutting edge smooth or with small teeth on proximal half. Dactylus 2.5-4 times as long as fixed finger, cutting edge with round teeth.

P2 ( Fig. 33A ) with upper subdistal spine and one to three lower spines on merus; carpus with upper and lower distal spine. P3 ( Fig. 33B ) with three or four lower spines on merus, carpus with lower distal spine. Small coxal spine on P1 and P2. Basipod of uropod with spinule hanging over endopod; exopod ( Fig. 32G, H ) triangular, about as long as distal width, with proximal spinule.

Colour

Usually dull green, sometimes brown (d’Udekem d’Acoz pers. comm.).

Variable, blueish to greenish-brownish (Dworschak pers. comm.); a coloured photograph is presented in Geiss (1990).

Size

Large specimens in material examined: cl. 15.5- 18 mm, tl. 47-50 mm. In Northern Adriatic: males are of cl. 20-22 mm, tl. 55-61 mm, females of cl. 16-18 mm, tl. 46-52 mm, with a maximum size of 66 mm in males and 60 mm in females; males of tl.> 50 mm have a longer carapace than females of the same size ( Dworschak 1988). Largest size reported, from North Aegean Sea: 82 mm for females, 106 mm for males ( Kevrekidis et al. 1997).

BIOLOGY AND ECOLOGY

The biology of U. pusilla in the Bay of Chingoudy (Pays Basque), France was reported by Chaud (1984a, b). Its biology in Northern Adriatic, including burrow construction, environment, growth and production, feeding, pumping rate, was studied in detail by Dworschak (1981, 1983, 1987a, b, 1988).

In the Patra Gulf and Ionian Sea ( Greece), this species was found living in a wide range of substrates with little or no mud, in shallow water; it was absent from Posidonia meadows (Thessalou- Legaki & Zenetos 1985). In Trébeurden, Brittany, many specimens were found at 20- 25 cm depth, in an area with muddy sand and small Zostera (d’Udekem d’Acoz 1986) , and also in coarse sand. They were dark green or brown coloured and may carry the parasitic isopod Gyge branchialis Cornalia & Panceri, 1861 ( Tucker 1930; d’Udekem d’Acoz 1989). In Marseille, France, U. pusilla was regarded as the characteristic species of biocoenoses with muddy sand, in shallow and calm water ( Pérès & Picard 1964; de Gaillande 1968).

Other aspects of biology studied

Branchial morphology, gill area and gill ultrastructure ( Astall et al. 1997b); population dynamics, reproduction and growth ( Kevrekidis et al. 1997); comparison of gut morphology and gut microflora ( Pinn et al. 1999a); mouthpart morphology and mouthpart setal fringes ( Pinn et al. 1999b); abundance of population in Corsica (de Vaugelas 1991); relation between burrow architecture, feeding mode and structure of the sediments (de Vaugelas 1998).

VARIATIONS AND REMARKS

Upogebia pusilla is the most common European upogebiid species but varies morphologically. Variation is greatest among Mediterranean populations and bring them near an allied species in the area, U. tipica ( Nardo 1869) . The two are sometimes so similar or hardly distinguishable that García Raso (1983), studying material from southern Spain, thought they might be identical. Examination of the present material reveals however that both species are valid, and their differentiation is discussed under U. tipica .

Most characteristics given to U. pusilla by authors such as de Man (1927), Bouvier (1940), de Saint Laurent (1971) concern P1 propodus, e.g., the distal dilation, the length/width ratio or the spinulation of its upper margin, but these are variable, as reported by García Raso (1983), and Dworschak (1992).

Characteristics of typical U. pusilla as listed in the diagnosis fit best specimens from the Atlantic, and some from the western Mediterranean.

Certain variations mentioned by García Raso (1983), affecting few specimens, can be consider- ed as usual within large populations; they are: 1) paired or bifid postocular spine on anterolateral border of carapace (usually single); 2) bifid A2 scale (usually simple); and 3) lateral distal border of P1 carpus with two or three spinules (one usually).

Other variations occur more often, mainly in the rostrum and P1:

The rostrum is between 1.2-1.4 times as long as wide at base.

P1 propodus: 1) the length/width ratio varies between 1.7-2.65 in male, 2.15-2.8 in female, 2.8-3.7 in juvenile ( García Raso 1983); 2) the mesial longitudinal crest on the upper margin is sometimes armed with spines ( Fig. 33E, G ); 3) both the lateral and mesial crests on the upper margin are sometimes very faint; 4) the distal dilation at level of the fixed finger is nearly absent in female, weak in young male, pronounced in large male; 5) the median spine on the mesiodistal margin (near the cutting edge of the dactylus) is large in female ( Fig. 32G ), much smaller or absent in male ( Fig. 32D ); 6) the fixed finger is distal in female, slightly or strongly subdistal in male; 7) the fixed finger is about one-fourth to half as long as the dactylus; and 8) the dactylus and fixed finger cutting edge is unarmed or with small teeth.

The following characters which contrast with U. tipica , are more constant (the last one to a lesser extent): 1) the rostrum is less than 1.5 times as long as wide at base with four or five teeth on each lateral border; 2) P1 merus bears three to six spines and spinules on the proximal half of the lower margin, and denticles distally; 3) P1 propodus has a longitudinal upper smooth crest on the lateral surface, an upper longitudinal tuberculate or spinous crest on the mesial surface, an upper subdistal spine, an upper distal spine and a median distal spine near the cutting edge of dactylus (in females especially); and 4) the telson is wider than long, the uropodal exopod is triangular and about as long as its distal width, this appendage is slightly longer in juvenile.

Upogebia stellata ( Montagu, 1808) ( Figs 34 ; 35 )

Cancer Astacus stellatus Montagu, 1808: 89 , tabl. III, fig. 5.

Gebia stellata – Leach 1816: tabl. 31, figs 1-8. — Desmarest 1825: 204. — Bell 1846: 223, 1 fig. — White 1857: 97, pl. 7, fig. 3. —? Bonnier 1887: 248. — Stephensen 1910: 277. — Lagerberg 1908: 55, pl. 2, fig. 10. — Schlegel 1912: 239, 250. — RunnstrØm 1925: 28, pl. 3. — Grieg, 1927: 35 (part), not fig. — ° Vilela 1936: 226.

Upogebia stellata – Stebbing 1893: 185. — Norman & Scott 1906: 12. — ° Norman 1907: 357. — de Morgan 1910: 475, fig. 1. — Selbie 1914: 104. — * Webb 1919: 102, pl. 10, figs 2-11, pl. 11 figs 2-6, pl. 12 figs 5, 6, 9, 10. — Schellenberg 1928: 75, fig. 57. — Gustafson 1934: 12. — Bouvier 1940: 108, fig. 72. — Poulsen 1941: 216, 224, figs 5, 7, 9. — * Gurney 1942: 249, fig. 100. — Holthuis 1950: 111, fig. 39; 1958: 6, fig. 9. — Gordon 1957: 250. — Tambs-Lyche 1958: 14. — Holme 1961: 415, 416; 1966: 433, 352. — Bourdon 1965: 16. — Allen 1967: 18. — Christiansen 1972: 41, fig. 47. — Naylor 1972: 73. — Samuelsen 1974: 132. — * Thiriot 1976: 350, 367. — Tebble 1976: 85 (association with the bivalve mollusc Lepton squamosum (Montagu)) . — Lacourt 1977: 246. — Adema et al. 1982: 28, fig. 5, tabl. 6. — Holthuis & Heerebout 1986: 62, fig. 80. — Thessalou-Legaki 1986: 183. — Moyse & Smaldon 1990: 520, fig. 10.13 (part). — ° Koukouras et al. 1992: 223. — ° Froglia 1995: 8. — Hayward et al. 1995: 434, fig. 8.52 (part). — Nickell & Atkinson 1995: 181, fig. 2D, E, tabl. 3. — d’Udekem d’Acoz 1996: 60; 1999: 157. — Astall et al. 1996: 821, tabls 1, 2; 1997a: 155, fig. 1 (part), 2, 4, 6, tabl. 1 (part); 1997b: 669, fig. 1, tabls 1-5, pls 2, 3, 5. — ° Brattegard & Christiansen 1997: 222. — Pinn et al. 1997: 1083, figs 1, 2, tabls 1-3. — Christiansen & Stene 1998: 76. — Nickell et al. 1998: 745, 752, figs 6, 7. — Pinn et al. 1998a: 243, fig. 2B, D; 1998b: 211, figs 1A, 2A; 1999a: 103, figs 3B-F, 5H; 1999b: 1461, tabls 1-6. — °Pancucci-Papadopoulou et al. 1999: 393. — Christiansen 2000: 233. — Taylor et al. 2000: 265, figs 1, 2, tabls 1-3, 5, 6. — Markham 2001: tabls 1, 2. — * Martin 2001: 83, 1 fig. — ° Türkay 2001: 289. Upogebia (Upogebia) stellata – Borradaile 1903: 543. — de Man 1927: 36, figs 14-14d. — Noël 1992: 83.

Non Upogebia stellata Grieg, 1927: 35 (part), fig. (= U. deltaura ( Leach, 1815)) . — Nobre 1931: 195, fig. 109; 1936: 122, fig. 102 (= U. pusilla ( Petagna, 1792)) .

TYPE MATERIAL. — Two specimens of this species from Plymouth Sound, previously belonging to Montagu’s collection are in the Natural History Museum, London ( NHML 258a and 258b) but the type, from the estuary of Kingsbridge, is missing, probably lost.

MATERIAL EXAMINED. — North Sea. NW of Texel, 53°36’N, 3°45’E, 34 m, P. C. Goudswaard, 23.IV.1980, 1 ovig. ( RMNH D 35729); 53°52.5’N, 4°29.5’E, 42 m, MS Aurelia , leg. P. C. Goudswaard, 2.II.1982, 1 juv. ( RMNH D 35730). — Off the English coast, 51°50’- 52°20’N, 1°20’- 2°00’E, NIOZ, leg. D. Eisma, 6-18.IX.1965, 1 spec. ( RMNH D 35731). — C. 53°37’N, 3°35’E, Aurelia cruise, 6.X.1975, leg. F. Creutzberg, don. NIOZ, 1 ( RMNH D 35732). — 53°43’N, 4°14’E, Aurelia , 24. IV- 5. V.1972, leg. F. Creutzberg, don. NIOZ, 1 ovig. ( RMNH D 35733).

Great Britain. Millport, Firth of Clyde, 20 m, R. B. Pike coll., 4, 2 (1 ovig.) ( NHML 1962.7.5.1-4). — Port Erin , Isle of Man , Hartnoll coll., 1 ( NHML 1960.VII.25:150). — Plymouth Sound , Montagu collection (dried, rehydrated), 1 cl. 11.5 mm ( NHML 258 a), 1 broken ( NHML 258 b). — Plymouth, 1, 1 ( NHML 1910.7.1.3- 4) ; D. B. Carlisle, V.1959, 1, 1 ( RMNH D 15227 ). — Salcombe Harbour, 1875, Norman coll., 1 ( NHML.98.7.735). — English Channel , 55 m, 17.VII.1959 (see Holme 1966), 1 ( NHML 325 ) ; 35 m, 1 ( NHML 324 ) ; 37 m, 1 ( NHML 323 ) .

France. Roscoff , Brittany, R. Bourdon coll., 1 cl. 14 mm, tl. 41 mm (figured) and 2 cl. 11.5 and 13 mm ; 2 (1 ovig.) cl. 12.5 mm ( MNHN Th 324) ; a dozen of pereopods with 2 P1 ( MNHN Th 322). — Meadow opposite Beuvan Bras, M. Cantacuzène coll., 20.XI.1957, 1 cl. 10 mm, 1 cl. 12 mm ( MNHN Th 323). — Port de Primel , R. Bourdon coll., 18.II.1965, 1 cl. 12.5 mm ; 1 cl. 16 mm, tl. 49 mm (figured) and 1 ovig. cl. 14 mm ( MNHN Th 320) ; 16.II.1965, 1 cl. 14.5 mm ( MNHN Th 321). — Callot island, near Roscoff, sandy beach at low tide, excursion RMNH, 17.IX.1958, 2 ( RMNH D 12437). — Rade de Brest , Glémarec coll., 1 cl. 12 mm ( MNHN Th 49) ; 1 cl. 7 mm ( MNHN Th 420) .

Greece. Orei Channel, stn DIO5, 21.XI.1991, 1 cl. 9 mm. — Rhodes, stn R. 48, XI.1983, 1 damaged spec. with 2 P1 (collection Thessalou-Legaki).

DISTRIBUTION. — European coasts of eastern Atlantic: Swedish west coast ( Gustafson 1934), western Norway (Samuelson 1974), Denmark (Poulson 1941), Firth of Clyde, west Scotland ( Allen 1967), Netherlands ( Holthuis 1950), southern North Sea ( Adema et al. 1982), coasts of Ireland ( Selbie 1914), western English Channel ( Holme 1961), Brittany, France; Greece (Thessalou-Legaki 1986).

DIAGNOSIS

Rostrum ( Fig. 34A, B ) obtuse distally, about twice as long as wide at base, six or seven lateral teeth on either border; antero-lateral border of carapace with one or two spines; lateral groove of gastric region well defined in anterior half, faint posteriorly. Telson ( Fig. 34C ) approximately 1.4-1.5 times as wide as long, lateral borders regularly curved, posterior border slightly concave medially.

A1 and A2 peduncle ( Fig. 34B ) with spine on lower margin of article 1 and 3 respectively. Md with large mesio-anterior pointed tooth. Mxp1 without epipod, Mxp3 with epipod.

P1 slightly stouter in male ( Fig. 34D, F ) than in female ( Figs 34G ; 35B ). Ischium with one or two spines on lower margin. Merus with upper subdistal spine, five to eight spines on proximal twothirds of lower margin and up to eight spinules distally. Carpus with upper, lower distal spine and median distal spine on mesial surface. Propodus with upper subdistal spine followed posteriorly by one to four pointed tubercles; mesial surface with upper distal spine near articulation with dactylus; fixed finger about half as long as dactylus, cutting edge smooth or with round teeth proximally. Dactylus with proximal upper conical tooth often small in female, cutting edge with mesial proximal round tooth, small or absent in female.

P2 ( Fig. 34E ) with upper subdistal spine and two or three lower spinules on merus; carpus with upper and lower subdistal spine.

P3 ( Fig. 35C ) with two to four spines on lower margin of merus, carpus with lower subdistal spine.

Small coxal spine on P1 and P2, single pleurobranch on P5 ( Fig. 35A ) (absent in a young male of cl. 5.5 mm from North Sea ( RMNH D 35730) and a female from Orei Channel of cl. 9 mm (collection Thessalou-Legaki).

Uropods ( Fig. 34C ) about as long as telson; basipod with spinule hanging over endopod; exopod about 1.5 as long as distal width, with proximal spinule.

Colour

Yellowish white, with minute stellated orange spots ( Montagu 1808; White 1857; Moyse & Smaldon 1990).

Size

Type: nearly 2 inches long (tl. about 50 mm) ( Montagu 1808). Largest specimens in material examined: cl. 14-16 mm, tl. 41-49 mm.

ECOLOGY AND BIOLOGY

In Gullmarfjord, Sweden, the species was found in a bottom of clay, sand and gravel, at 25-30 m depth ( Gustafson 1934). It was less common in the North Sea than U. deltaura but was found in nearly the same localities ( Adema et al. 1982). In western Norway, it was collected on the littoral, at low water, in mineral sand with some mud, or in shell sand, at 7-13 m; one specimen was found in mud at 120 m ( Samuelsen 1974). It was recorded in shallow water in southern part of the boreal region ( Christiansen 2000), collected between 25-36 m in Danish waters ( Poulsen 1941) and was rare in the Clyde sea area ( Allen 1967). It is common on all coasts of the British Isles ( Moyse & Smaldon 1990); fairly common around Roscoff, France, in the same intertidal areas as U. deltaura , but less abundant ( Bourdon 1965). According to Holme (1966), it lives in a muddy-gravel substrate, off Plymouth, English Channel. Reproduction occurs in February- August in Roscoff ( Bourdon 1965). Larvae are abundant in Roscoff from February to July, with the same distribution as U. deltaura ( Thiriot 1976) .

Aspects of biology studied

Burrow morphology and feeding behaviour ( Nickell & Atkinson 1995); branchial parasitic isopods ( Astall et al. 1996); burrow morphology and burrow-dwelling lifestyle ( Astall et al. 1997a); branchial morphology, gill area and gill ultrastructure ( Astall et al. 1997b); microbial flora associated with digestive system ( Pinn et al. 1997); morphology of mouthparts and pereiopods in relation to feeding, ecology and grooming ( Nickell et al. 1998); diet ( Pinn et al. 1998a); particle size selectivity and resource partitioning ( Pinn et al. 1998b); gut morphology and gut microflora ( Pinn et al. 1999a); mouthpart morphology and mouthpart setal fringes ( Pinn et al. 1999b); oxygen transporting properties of heamocyanin ( Taylor et al. 2000).

REMARKS

With an exception, the southernmost distribution of this species in the material examined, is Brittany, northwest France. Several samples collected in southwest France (Arcachon), Spain or Portugal and previously assigned to U. stellata , are actually U. pusilla . The possible exception is the material from Greece reported by Thessalou- Legaki (1986). It includes a small female (cl. 9 mm) from Orei Channel and two P1 of a very small and badly damaged specimen (not examin- ed) from Rhodes. The female from Orei differs from typical U. stellata by having no pleurobranchs on P5 of both sides, and by the uropods distinctly longer than the telson. It could possibly belong to U. stellata in virtue of the shape of the rostrum, the posterior border of telson medially concave and a median distal spine on mesial surface of the P1 carpus. This is the only possibly correct record of this species (with variations) in the Mediterranean. Nobre (1931, 1936) reported U. stellata from Portugal, with a figure representing, with little doubt, a specimen of U. pusilla . Bonnier (1887) and Vilela (1936) reported U. stellata from the Bay of Concarneau, France and from Faro, Portugal respectively; their materials have not been checked. More recently, Lacourt (1977) considered U. stellata as part of the marine fauna of the Bay of Concarneau. Specimens he studied were deposited (dried) in the Rijksmuseum of Natural History, Leiden, and one of these ( RMNH D 45673) has been examined. It actually belongs to U. pusilla .

U. stellata was often confused with U. deltaura and a part of the material studied by Grieg as U. stellata (1927, with a figure) is likely U. deltaura . De Morgan (1910) revealed the distinction between the two species in the anterolateral border of the carapace, with a spine in U. stellata , unarmed in U. deltaura . Also the P1 fixed finger is half as long as the dactylus in the former species, as long or nearly as long as the dactylus in the latter.

Upogebia stellata has also been confused with U. pusilla and U. tipica , for all three have a spine on the antero-lateral border of the carapace, a subcheliform P1 and like P2 and P3.

It can readily be differentiated from U. pusilla and U. tipica , as well as all other European Upogebia , in having a pleurobranch on the P5. Additional distinguishing features are: 1) P1 carpus has a median spine on its mesial distal border and the dactylus bears an upper proximal conical tooth (spine and tooth absent in U. pusilla and U. tipica ); and 2) the telson is about 1.5 times as wide as long with lateral borders regularly curved and the posterior border slightly concave medially (it is 1.3 times as wide as long or approximately quadrate in U. pusilla and U. tipica respectively with lateral borders convex and the posterior bor- der straight).

Figures

FIG. 26. — Upogebia deltaura (Leach, 1815); A-C, from Roscoff, France (MNHN Th 1316); D-G, holotype, from Devon, UK (NHML 259 e); A, D, anterior part of carapace and rostrum, dorsal view; B, anterior part of carapace, lateral view; C, telson and uropods; E, F, distal articles of pereopod 1, lateral and mesial view; G, abdominal segments 1 and 2, lateral view. Scale bars: A-F, 2 mm; G, 4 mm.

FIG. 27. — Upogebia deltaura (Leach, 1815); A, from coasts of France (MNHN Th 38); B-H, from Roscoff, France (MNHN Th 1316); B, C,; D-H,; A-C, pereopod 1-3; D, E, maxillule and maxilla; F-H, maxilliped 2, 3 and 1 respectively. Scale bars: A-C, 2 mm; D-H, 1 mm.

FIG. 28. — Upogebia mediterranea Noël, 1992; A-E, lectotype, from Tamaris-sur-Mer, France (MNHN Th 1372); A, B, anterior part of carapace, dorsal and lateral view; C, D, pereopod 1 and distal articles in mesial view; E, abdominal segments 1 and 2, lateral view; F, paralectotype,, same locality (MNHN Th 1369), telson and uropods. Scale bars: 2 mm.

FIG. 29. — Upogebia mediterranea Noël, 1992; A-C, paralectotype, from Tamaris-sur-Mer, France (MNHN Th 1369); D, E, lectotype,, same locality (MNHN Th 1372); F, paralectotype, from Baleares (MNHN Th 661); G, paralectotype, from Gulf of Oran, Algeria (MNHN Th 486); A, B, antennule and antenna; C, mandible; D-F, pereopod 2, 3 and 1 respectively; G, telson and uropods. Scale bars: A, B, D-G, 1 mm; C, 0.5 mm.

FIG. 30. — Upogebia nitida (A. Milne-Edwards, 1868), from Tamaris-sur-Mer, France (MNHN Th 1393); A, B, anterior part of carapace, dorsal and lateral view; C, telson and uropods; D, E, pereopod 1 and distal articles in mesial view; F, pereopod 2. Scale bars: 2 mm.

FIG. 31. — Upogebia nitida (A. Milne-Edwards, 1868); A-C, F, lectotype, from Cape Verde Islands (MNHN Th 22); D, E, paralectotype, from Cape Verde Islands (MNHN Th 22); G-J, from Gorée, Senegal (MNHN Th 278); A, B, anterior part of carapace, dorsal and lateral view; C, D, I, pereopod 1; E, J, distal articles of pereopod 1, mesial view; F, P2; G, telson and uropods; H, rostrum. Scale bars: 1 mm.

FIG. 32. — Upogebia pusilla (Petagna, 1792); A-E, from Oléron, France (MNHN Th 1317); F-I, from Arcachon, France (MNHN Th 645); A, B, anterior part of carapace, dorsal and lateral view; C, F, pereopod 1; D, G, distal articles of pereopod 1, mesial view; E, pereopod 1 propodus, upper view; H, telson and uropods; I, uropods. Scale bars: 2 mm.

FIG. 33. — Upogebia pusilla (Petagna, 1792); A, B, from Oléron, France (MNHN Th 1317), pereopod 2 and 3; C, from Italy (MNHN Th 1318); D, E, H, from Banyuls, France (MNHN Th 622); F, G, from Banyuls (MNHN Th 621); C, D, F, pereopod 1; E, G, distal articles of pereopod 1, mesial view; H, anterior part of carapace. Scale bars: 2 mm.

FIG. 34. — Upogebia stellata (Montagu, 1808); A-F, from Roscoff, France (MNHN Th 324); G, from Port de Primel, France (MNHN Th 320); A, B, anterior part of carapace, dorsal and lateral view; C, telson and uropods; D, pereopod 1; E, pereopod 2; F, G, distal articles of pereopod 1, mesial view. Scale bars: 2 mm.

FIG. 35. — Upogebia stellata (Montagu, 1808); A, C, from Roscoff, France (MNHN Th 324); B, from Port de Primel, France (MNHN Th 320); A, abdominal segments 1, 2 and gills of pereopod 4 and 5 in lateral view, posterior arthrobranch of pereopod 4 omitted; B, C, pereopod 1 and 3. Scale bars: 1 mm.