Callianassa Leach, 1814

Ngoc-Ho, Nguyen, 2003, European and Mediterranean Thalassinidea (Crustacea, Decapoda), Zoosystema 25 (3), pp. 439-555 : 465-475

Genus Callianassa Leach, 1814

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

DIAGNOSIS. — Carapace with dorsal oval, lacking rostral carina and posterodorsal cardiac prominence. Eyestalk short, flattened dorsoventrally; cornea subdistal, dorsal or dorsolateral. Mxp1 with anteriorly truncate epipod, endopod minute. Mxp2 with small epipod. Mxp3 lacking exopod, subpediform or operculiform; propodus not expanded ventrally, less than three times as wide as dactylus, latter digitiform, at least twice as long as wide.

P1 unequal, usually with meral hook, P2 chelate, P3 and P4 simple, P5 subchelate. P3 lacking or bearing reduced lower proximal heel on propodus.

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

Plp1 and Plp2 often absent or rudimentary in male, larger in female; female Plp2 biramous. Plp3-5 with stubby, projecting appendix interna. Uropodal exopod with dorsal plate.

REMARKS

The genus Callianassa is provisionally considered a polyphyletic group, with the species included as listed in Tudge et al. (2000). Sakai (1999a) proposed a much broader concept of the genus as he synonymised several genera erected by Manning & Felder (1991) with Callianassa . Although his action is not considered fully justified, the genera established by Manning & Felder (1991), focusing on American callianassids, probably need to be reconsidered, especially in view of their global use. Examination of the abundant, highly diverse material from the Indo-Pacific, together with a detailed phylogenetic analysis is desirable to show the true relationships between species and provide a solid base for their grouping. This however is beyond the scope of this work.

Nevertheless a number of additional characters, besides those given by Tudge et al. (2000), are suggested to be useful in the taxonomic study of this genus. Taking the three European Callianassa species as an example, it can be noted that, though they all fit the diagnosis of the genus as given above, C. subterranea has a slender minor P1 ( Fig. 9C ) not compressed laterally, with the carpus four or five times as long as wide; the P3 ( Fig. 9J ) propodus not ovate, with the lower border dentate; the third segment of Md palp ( Fig. 9M ) of same width throughout and the basal endite of Mx1 ( Fig. 10A ) approximately rounded distally.

By contrast, C. acanthura and C. truncata have the minor P1 ( Fig. 8D ) slightly compressed laterally, with the carpus less than twice as long as wide; the P3 propodus ovate ( Fig. 8F ) with the lower border regularly curved; the third segment of Md palp ( Fig. 8K ) narrowing distally and the basal endite of Mx1 ( Fig. 8E ) of inverted-triangle shape with an expanded distal border bearing spiniform setae.

These characters were not considered by Tudge et al. (2000) in their analysis but are probably of phylogenetic significance; those of C. acanthura and C. truncata are presumably apomorphic.

Callianassa acanthura Caroli, 1946 ( Fig. 8 )

Callianassa acanthura Caroli, 1946: 66 , figs 1a, 2. — Holthuis 1953: 94, fig. 5. — Monchartmont 1979: 71. — de Saint Laurent & Božić 1976: 21, figs 3, 11, 19, 25, 30. — Türkay 1982: 225; ° 2001: 289. — ° Türkay et al. 1987: 92. — ° Koukouras et al. 1992: 223. — Noël 1992: 81. — ° Froglia 1995: 7. — Falciai & Minervini 1996: 147, 3 figs. — d’Udekem d’Acoz 1996: 54, 56. — Abed-Navandi & Dworschak 1998: 605, figs 1-8. — Sakai 1999a: 14, 128 (addendum). — ° Türkay 2001: 289.

Callianassa (Trypaea) acanthura – Zariquiey Alvarez 1968: 229.

Necallianassa acanthura – Heard & Manning 1998: 883. — d’Udekem d’Acoz 1999: 155.

TYPE MATERIAL. — Lectotype:, Naples, Italy, E. Caroli coll., received V.1959, don. Zoological Station Naples ( RMNH D 15212 a); paralectotype: ( RMNH D 15212 b), by present designation.

MATERIAL EXAMINED. — Italy. Naples, E. Caroli coll., 2 lectotype cl. 11 mm, tl. 43 mm (figured), and paralectotype cl. 8 mm, tl. 36 mm ( RMNH D 15212 a and b) ; Gulf of Naples , E. Caroli don., 1 ( RMNH D 6588 ) .

Croatia. Adriatic, Kornati, Zirje Island, 3 m, D. Abed- Navandi coll., 17.IX.1997, 1, 1 cl. 10 mm (figured) ( NHMW 15372).

Greece. NW Pogonia, 1.5 m, C. d’Udekem d’Acoz coll., 12.VII.1993, 1 cl. 8.5 mm, tl. 36.5 mm (figured) ( MNHN Th 1400) ; Lesbos (Aegean sea), C. d’Udekem d’Acoz coll., 12.VII.1992, 1 cl. 10.5 mm ( MNHN Th 1401), 1 cl. 10 mm (d’Udekem d’Acoz) .

OTHER MATERIAL EXAMINED. — Necallianassa berylae Heard & Manning, 1998, South Carolina , 1 cl. 2.5 mm, 1 cl. 4.5 mm (paratypes, USNM 260885 ) ; Georgia, 1 cl. 3.2 mm (paratype, USNM 260888 ). DISTRIBUTION. — Mediterranean: Adriatic (Abed- Navandi & Dworschak 1998), Naples, Aegean, Ionian sea (d’Udekem d’Acoz 1996) .

DIAGNOSIS

Carapace ( Fig. 8A ) with dorsal oval; rostrum approximately triangular with blunt tip, rostral spine absent. Telson ( Fig. 8G ) about as long as proximal width, a little narrower distally, lateral border weakly convex with long curved subdistal spine, posterior border weakly rounded with median spinule.

Eyestalk ( Fig. 8A ) short; cornea dorsal, subterminal, disk-shaped. A1 peduncle slightly shorter than that of A2. Md ( Fig. 8K ) with threesegmented palp, last segment pointing distally. Mx1 ( Fig. 8E ) with basal endite of invertedtriangle shape, bearing spiniform setae. Mx2 with large endopod. Mxp1 ( Fig. 8H ) with small endopod, epipod without anterior lobe. Mxp3 ( Fig. 8I, J ) operculiform, ischium-merus length about 1.5 times merus width, ischium with crista dentata of 14-16 spines on mesial surface; propodus 1.5 times as long as wide; dactylus digitiform, 1.5 times to twice as long as wide.

P1 unequal in adult male, subequal in young male and female with both chelipeds similar to minor cheliped of adult male. Male major P1 ( Fig. 8B ) with lower border of merus convex on distal half ( Fig. 8C ) bearing small rounded teeth; meral hook pointed distally and denticulate on proximal border; carpus and propodus unarmed; dactylus with curved tip and densely setose near base. Male minor P1 ( Fig. 8D ) slender with carpus 1.7-1.8 times as long as wide and twice as long as propodus; both fixed finger and dactylus with denticulate cutting edge. P3 ( Fig. 8F ) propodus with lower border rounded, no proximal heel. P4 propodus slender, over three times as long as wide.

Male Plp1 ( Fig. 8M ) small, two-articulated; male Plp2 absent. Female Plp1 ( Fig. 8N ) uniramous, two-articulated; female Plp2 ( Fig. 8O ) biramous, exopod longer than endopod. Plp3-5 biramous, foliaceous, appendix interna ( Fig. 8E ) small, narrowing distally, partly embedded, the rest projecting from inner border of endopod.

Uropod ( Fig. 8G ) as long as telson; exopod with long proximal spine, dorsal plate with setal distal row at short distance from posterior border; endopod with long lateral subdistal spine.

Colour

Body whitish in living specimens, white P1; tailfan mottled with whitish or dull-yellowish dots and spots that are characteristic of the species (d’Udekem d’Acoz 1986).

Tailfan and eyestalk orange, ripe ovaries red (Dworschak pers. comm.).

Size

Medium: cl. 8.5- 11 mm, tl. 36-43 mm.

ECOLOGY

The species lives in muddy sand, at 2-3 m depth ( Caroli 1946), or in a mixture of mud and fine or coarse sand, at 1-1.5 m, together with C. truncata or U. pusilla (d’Udekem d’Acoz 1996) , or in pure sand, at 0.5 m depth ( Türkay 1982). It is abundant on sublittoral sandy bottoms of the Kornati Archipelago and near Rovinj, in fine or medium sand, at 3-6 m depth, associated with the fish Trachinus sp. and Bothus sp. (Abed-Navandi & Dworschak 1998).

REMARKS

This species, together with Necallianassa berylae Heard & Manning, 1998, can be distinguished from all other callianassids by the presence of large acute spines on the lateroposterior border of the telson and uropodal endopod, as well as proximally on the uropodal exopod. Spines are present on the lateroposterior border of the telson in a few other species, e.g., C. truncata or C. intermedia de Man, 1905 (see Sakai 1999a: 128) but in none of these are they so large (about onefifth length of telson and uropodal endopod in C. acanthura ) and acute.

Among its European and Mediterranean congeners, C. acanthura is close to C. truncata but differs essentially in: 1) the large acute spines on the telson and uropods, as mentioned above; and 2) the absence of an appendix interna on female Plp2.

Callianassa subterranea ( Montagu, 1808) ( Figs 9 ; 10 )

Cancer Astacus subterraneus Montagu, 1808: 89 , pl. 3

figs 1, 2. Callianassa helgolandica Lutze, 1938: 174 , figs 52-61.

Callianassa pestae – Lutze 1937: 6, figs 1-6; 1938: 167, figs 10-21.

Callianassa subterranea – Leach 1815: 343; 1816: pl. 32. — Desmarest 1825: 205, pl. 36 fig. 2. — White 1847: 70; 1857: 94, pl. 7 fig. 2. — Bell 1846: 219, 1 fig. — ° Lafont 1868: 522. — A. Mi1ne-Edwards 1870: 80, 101. — ° Fischer 1872: 428. — Stalio 1877: 106. — °de Folin & Périer 1879: 211. — Stossich 1880: 206. — Van Beneden 1884: 647. — Carus 1885: 489. — Koehler 1886: 59. — Bonnier 1887: 247. — Ortmann 1891: 55, pl. 1, fig. 10. — Adensamer 1898: 620. — ° Graeffe 1902: 69. — ° Norman 1907: 357. — Sinel 1907: 216. — Lagerberg 1908: 53, pl. 1, fig. 15. — * Lo Bianco 1909: 603. — Schlegel 1912: 238, 250. — * Boraschi 1921: 6, pl. 1 fig. 2. — * Webb 1921: 403, pl. 3, figs 2, 5. — Miranda y Rivera 1933: 21. — Gustafson 1934: 14. — Lutze 1938: 170, figs 28-51. — Poulsen 1941: 229, figs 10-12. — * Gurney 1942: 244, fig. 98. — Zariquiey Alvarez 1946: 106. — * Rees 1955: 74, fig. 4.5 (part). — Holme 1966: 433, fig. 23 (part). — Allen 1967: 57, 90 (fig.). — Naylor 1972: 69. — ° Pastore 1976: 107. — ° Christiansen 1972: 41, fig. 49; 2000: 234. — de Saint Laurent & Božić 1976: 17, figs 1, 9, 17, 28. — * Thiriot 1976: 350, 367. — de Saint Laurent & Le Loeuff 1979: 52, fig. 9c, e. — Moncharmont 1979: 71. — Domenech et al. 1981: 150. — Adema et al. 1982: 23, fig. 6a-c, map 4. — Christiansen & Greve 1982: 213. — Atkinson 1986: 358, fig. 1M. — Campbell & Nicholls 1986: 218, 1 fig. — Holthuis & Heerebout 1986: 62, fig. 83. — Moore 1986: 32. — Thessalou-Legaki 1986: 182; 1987: 457. — Witbaard & Duineveld 1989: 209-219, fig. 1. — Atkinson & Nash 1990: 403, figs 1, 2, tabls 1, 2. — Moyse & Smaldon 1990: 520, fig. 10.13 (part). — Števčić 1990: 217. — Manning & Felder 1991: 768, fig. 8. — Dworschak 1992: 203. — ° Koukouras et al. 1992: 223. — Noël 1992: 81. — Lindley et al. 1993: 53. — ° Diez et al. 1994: 47. — Rowden & Jones 1994: 623, figs 1-5, tabls 1-3; 1995: 1155, figs 1-4, tabl. 1; 1997: 153, figs 1-3. — ° Froglia 1995: 7. — Hayward et al. 1995: 432, fig. 8.52 (part). — Nickell & Atkinson 1995: 181, fig. 1, tabl. 1. — Falciai & Minervini 1996: 147, 4 figs. — Astall et al. 1997b: 669, 674, 675, fig. 1, tabls 1-5, pls 2, 4, 5. — Brattegard & Christiansen 1997: 220. — Hughes & Atkinson 1997: 639. — Johns et al. 1997: 127, tabls 1, 2. — Rowden et al. 1998: 1365, figs 1-4, pls 1-2; tabl. 1. — Christiansen & Stene 1998: 75. — Nickell et al. 1998: 735, 749, 754, figs 1-3. — Pinn et al. 1998b: 243, fig. 3; 1999a: 103, figs 1, 3, 4; 1999b: 1461, tabls 1-6. — Stamhuis et al. 1997: 156; 1998a: 43; 1998b: 197. — Stamhuis & Videler 1998: 2152. — d’Udekem d’Acoz 1999: 155. — Sakai 1999a: 19. — Hughes et al. 2000: 189, figs 1-5. — Öksnebjerg 2000: 78. — Taylor et al. 2000: 265, figs 1-3, tabls 1-3, 5, 6. — *González-Gordillo et al. 2001: 279. — Livory 2001: 33 C. — Markham 2001: 196, tabls 1, 2. — * Martin 2001: 77, 1 fig. — ° Türkay 2001: 289.

Callianassa (Callianassa) subterranea – Zariquiey Alvarez 1968: 229.

Callianassa (Cheramus) subterranea – Borradaile 1903: 545. — de Man 1928a: 6, pl. 1 figs 1-1h; 1928b: 27, 91, 92, 94, 97. — Makarov 1938 (English edition, 1962): 63, fig. 21. — Bouvier 1940: 101, fig. 67. — Gurney 1944: 82, figs 1A, 14, 15. — Gordon 1957: 249. — * O’Céidigh 1962: 163.

Callianassa tyrrhena – Holthuis & Gottlieb 1958: 62, fig. 13.

Cheramus subterraneus – Colosi 1923: 6.

Non Callianassa subterranea – H. Milne Edwards 1837a: 309; 1837b: 130, pl. 48 figs 3-3e (= Pestarella tyrrhena ( Petagna, 1792) n. comb.). — Heller 1863: 202, pl. 6 figs 9-11 (= P. tyrrhena n. comb.). — Ortmann 1891: 55, pl. 1, fig. 10 (= P. tyrrhena n. comb.). — Giard & Bonnier 1890: 362, figs 1, 3 (= P. candida ( Olivi, 1792) n. comb.).

TYPE MATERIAL. — Lectotype:, Kingsbridge Estuary, S Devon, Great Britain ( NHML 1939.2.20.1); paralectotype: 1 ovig. ( NHML 1939.2.20.2) by present designation.

MATERIAL EXAMINED. — North Sea. Kettle Hole, southern North Sea, 23.III.1955, leg. et don. P. Smit, 1 broken spec. ( RMNH D 10985 ). — Near Pit bouy, 17.III.1955, depth 150 ft, leg. et don. P. Smit, 1 chela ( RMNH D 10986). — Bruine Bank near Winterton Twenties, c. 53°N, 3°E, 20-27.VIII.1960, leg. et don. J. Kruuk, 1 broken spec. ( RMNH D 16614). — 53°41’45”N, 3°55’E, ms Tridens , haul 44, 12.VIII.1971, leg. G. R. Heerebout, 1 ( RMNH D 29227).

Great Britain. S Devon, 1 lectotype cl. 13 mm, tl. 52 mm ( NHML 1939.2.20.1), 1 paralectotype cl. 12 mm, tl. 47.5 mm ( NHML 1939.2.20.2) ; 2 spec., dried ( NHML 260 a, b). — Plymouth Sound, 1 cl. 11 mm, tl. 47 mm (figured), 1 cl. 9 mm, 2 cl. 10 mm and 10.5 mm, tl. 42.5 and 44.5 mm (figured), 3 (1 ovig.) cl. 10-10.5 mm ( MNHN Th 211). — Lion Rock, Millport , Firth of Clyde, sandy mud, R. B. Pike coll., 2.IX.1959, 1, 1 juv. ( NHML 1962.7.5.8-9). — English Channel, stn 107, MBA coll., 10, 6, 3 juv. (damaged) ( NHML 1999.81 - 101 ) ; stn 44, 29 m, 2, 1 ( NHML1999.72 - 73 ) ; stn 152, 42 m, 1 without abdomen ( NHML 1999.126 ) ; stn 103, 37 m, 1 ovig. ( NHML 1999.74 ) ; stn 110, 51 m, MBA coll., 7, 1 ovig. (poor condition) ( NHML 1999.102 - 114 ) ; stn 173, 54 m, MBA coll., 7, 1 (poor condition) ( NHML 1999.127 - 138 ) ; stn 152, 42 m, 2, 1 abdomen of (poor condition) ( NHML 1999.115 - 116 ). — Western Channel, MBA coll., 17.III.1970, 3, 2 (damaged) ( NHML 1971.24 ). — Salcombe , Devon, spring low tides, P. Gibb coll., 26.III.1970, 1 ovig. ( NHML 1971.23 ) ; R. Gurney coll., 21.III.1935, 3, 3 (damaged) ( NHML 1947.3.18.723-725). — 1 mile S of Knight Errant muddy grounds, 22 fms, 8.III.1970 ,

Ngoc-Ho N.

1, 1 ovig. ( NHML not registered). — Jersey, 3 dried spec. ( NHML 260d).

France. Roscoff , Thalassa, 1970 , 1 juv. cl. 3 mm ( MNHN Th 103), 1 cl. 5.5 mm, 1 juv. ( MNHN Th 104), 1 juv. cl. 2 mm ( MNHN Th 105), 1 damaged spec. ( MNHN Th 106). — Grande Vasière, near Concarneau, Glémarec coll., 1 cl. 6 mm, 1 cl. 5 mm ( MNHN Th 102) ; Glémarec coll., IV.1972, 2 cl. 4 and 8 mm (MNHN-Th 108), 1 just moulted cl. 10 mm ( MNHN Th 109), 1 cl. 5.5 mm, 1 cl. 5 mm, 1 juv. ( MNHN Th 110), 1 cl. 5.5 mm, 3 cl. 5-7 mm, 2 juv. ( MNHN Th 111). — Bay of Biscay , 120-180 m, leg. Lagardère, 2 cl. 4.5 and 6 mm, 1 ovig. cl. 5.5 mm ( MNHN Th 107). — Banyuls, mud with Turritella , 30 m, P. Noël coll., 3.VIII.1976, 1 juv. cl. 3 mm ( MNHN Th 1349) ; VI.1977, 1 cl. 4.5 mm, 1 ovig. cl. 5 mm ( MNHN Th 1350) ; mud, 90 m, 14. VI.1977, 1 cl. 4 mm ( MNHN Th 1351) .

Portugal. 42°9,4’N, 8°58,3’W, Thalassa , stn 819, 103 m, 21.X.1968, 1 juv. cl. 4 mm ( MNHN Th 635).

Spain. Alboran sea, Calypso , 35°50’N, 3°19’W, muddy sand, 500 m, 3.IX.1958, 1 cl. 8 mm ( MNHN Th 367).

Greece. Calypso , stn 1, dredge, 94 m, 27.IX.1977, 1, 1 ovig. cl. 4.5 mm ( MNHN Th 625). — Aegean Sea, Strymonikos G., 25 m, 5.II.1977, 2 (A.U.TH P 1529).

Israel. Ascalon , bottomgrab, 54 m, 27.VII.1947, leg. A. Wirszubski, No. 590A, 1 juv. ( RMNH D 13796) ; 30.IX.1947, leg. A. Wirszubski, No. 652, 2 juv. ( RMNH D 13797). — Haifa Bay , stn 7, bottomgrab, 38 m, 7.VI.1955, leg. E. Gottlieb, No. 225, 1 juv. ( RMNH D 13798) ; stn 9, bottomgrab, 42 m, 11.X.1954, leg. E. Gottlieb, No. 148, 1 ( RMNH D 13799). — Nabi Junis, bottomgrab, 54 m, 18.VIII.1949, leg. A. Wirszubski, No. 787, 1 ( RMNH D 13801). — Nahariya, bottomgrab, 70 m, 18.IX.1947, leg. A. Wirszubski, No. 638, 2 juv. ( RMNH D 13802). — Tel Aviv, bottomgrab, 54 m, 12.XII.1949, leg. A. Wirszubski, No. 838, 1 juv. ( RMNH D 13803). — Nabi Rubin, bottomgrab, 56 m, 24.V.1949, leg. A. Wirszubski, No. 741, 1 juv. ( RMNH D 13804). — Rafan, bottomgrab, 36 m, 22.VI.1947, leg. A. Wirszubski, No. 545, 1 juv., broken ( RMNH D 13805) .

Dahomey. Ombango, dredge, 40 m, A. Crosnier coll., 9.X.1963, 1 juv. cl. 3.5 mm ( MNHN Th 265).

DISTRIBUTION. — Eastern Atlantic: S Scandinavia ( Gustafson 1934; Poulsen 1941; Christiansen & Greve 1982; Christiansen & Stene 1998), SW of Scotland ( Allen 1967; Nickell et al. 1998), S of North Sea, English Channel, W coast of France, Dahomey.

Mediterranean: France, Israel, Greece, Alboran Sea, Adriatic Sea.

C. subterranea is recorded on the Coast of Norway to 60°N from 5-7 m to 60-100 m ( Christiansen 2000). According to de Saint Laurent & Božić (1976), the species is common in the northern part of its distribution range ( Great Britain, Denmark, Norway), scarce along the atlantic coast of France and in the Mediterranean with smaller specimens living in deeper waters (35-500 m).

DIAGNOSIS

Carapace ( Fig. 9A ) with dorsal oval; rostrum approximately triangular with blunt tip, rostral spine absent. Telson ( Fig. 9F ) about as long as proximal width, narrowing distally, lateral border convex proximally, posterior border with median spinule. Eyestalk ( Fig. 9A ) short, cornea dorsal, indistinct. A1 peduncle shorter than that of A2. Md ( Fig. 9M ) with three-segmented palp, last segment of same width throughout. Mx1 and Mx2 ( Fig. 10A, B ) as figured. Mxp1 ( Fig. 10C ) with small rounded endopod, epipod without anterior lobe. Mxp2 ( Fig. 10D ) with short exopod. Mxp3 ( Fig. 9D, E ) subpediform; row of 13-15 spines on mesial surface of ischium, smaller proximally; propodus about 1.5 as long as wide; dactylus 2.5- 3 times as long as wide.

Major P1 ( Fig. 9B ) stouter in male than in female, strong meral hook, curved anteriorly, smooth or with denticles on proximal border; carpus about as long as distal width, propodus slightly longer, cutting edge of fixed finger and dactylus unarmed or with small round teeth. Minor P1 ( Fig. 9C ) slender with carpus approximately four times as long as distal width. P2 ( Fig. 9I ) as figured. P3 ( Fig. 9J ) propodus with lower border dentate bearing separated tufts of setae, no proximal heel. P4 ( Fig. 9K ) propodus slender, nearly three times as long as wide. P5 ( Fig. 9L ) subchelate.

Male Plp1 ( Fig. 9G ) uniramous, male Plp2 ( Fig. 9H ) small, sometimes absent. Female Plp1 ( Fig. 9O ) uniramous, female Plp2 ( Fig. 9P ) biramous, exopod overreaching endopod.

Plp3-5 ( Fig. 9N ) biramous, foliaceous, appendix interna approximately cylindrical, partly embedd- ed, distal half projecting from inner border of endopod. Uropodal endopod ( Fig. 9F ) about as long as telson; exopod longer, setal row of dorsal plate at short distance from posterior border.

Colour

Colour during life more or less orange, sometimes yellow on the sides and on the tail, the arms usually pink ( White 1857); dorsal abdomen pale pink (Dworschak pers. comm.); white to light pink or light blue ( Campbell & Nicholls 1986).

Size lectotype of cl. 13 mm, tl. 52 mm; paralectotype of cl. 12 mm, tl. 47.5 mm.

In the present material examined, specimens are larger in northern Atlantic, of cl. 10-13 mm, tl. 42-52 mm, smaller in southern Atlantic (e.g., France, Spain) and in the Mediterranean, of cl. 4-8 mm, ovigerous of cl. 4.5-5.5 mm.

ECOLOGY AND BIOLOGY

This species shows all the characteristics of a deposit feeder, according to Stamhuis et al. (1997). It is common in subtidal (10-80 m) muddy fine sands and muds ( Adema et al. 1982; Dworschak 1992; Moyse & Smaldon 1990). In Plymouth, UK, larvae are very common in the plankton in summer and early autumn ( Gordon 1957).

Its ecology and general biology including burrow structure in the North Sea were dealt with by Witbaard & Duinevelt (1989), Atkinson & Nash (1990), Rowden & Jones (1994, 1995).

Other aspects of biology studied

Burrow morphology and feeding behaviour ( Nickell & Atkinson 1995); burrows and bioturbation ( Rowden & Jones 1997); burrow architecture and burrow ventilation ( Stamhuis et al. 1997; Stamhuis & Videler 1998); branchial morphology, gill area and gill ultrastructure ( Astall et al. 1997b); sulphide metabolism ( Johns et al. 1997); morphology of mouthparts and pereopods in relation to feeding, ecology and grooming ( Nickell et al. 1998); morphology, motion and function of appendages ( Stamhuis et al. 1998a); size selectivity and resource partitioning ( Pinn et al. 1998b); optimal foraging and grain size selection ( Stamhuis et al. 1998b); role in sediment turnover resuspension ( Rowden et al. 1998); gut morphology and gut microflora ( Pinn et al. 1999a); effects of burrows on sedimentswater solute fluxes ( Hughes et al. 2000); oxygen transporting properties of haemocyanin ( Taylor et al. 2000).

REMARKS

As reported by de Saint Laurent & Božić (1976), the specimen from Helgoland assigned by Lutze (1938) to Callianassa pestae (p. 167, figs 10-21) as well as his new species Callianassa helgolandica (p. 174, figs 52-61) actually belong to C. subterranea . This is confirmed by the figures of their Mxp3, both subpediform (fig. 11, also part of fig. 26a, b for C. pestae and fig. 53 for C. helgolandica ). On the other hand, Dworschak (1992: 205) contends that the specimen collected by the “Pola Expedition” ( NHMW 6613) and mention- ed by Adensamer (1898: 620) is in fact a juvenile of the present species, not of Gourretia denticulata ( Lutze 1937) .

There is variation in the length of major P1 carpus and propodus, also in the length and shape of the dactylus and fixed finger. This appears to be not linked with sex ( Nickell et al. 1998).

Callianassa truncata Giard & Bonnier, 1890 ( Fig. 11 )

Callianassa truncata Giard & Bonnier, 1890: 362 , figs 2, 4. — Caroli 1940: 73; 1946: 66, figs 1b, 3. — Holthuis 1953: 91, fig. 4. — * Bourdillon-Casanova 1960: 108. — Picard 1965: 38. — Lagardère 1966: 195, pls 2-5; 1973: 84. — ° Faure 1970: 773. — de Saint Laurent & Božić 1976: 19, figs 2, 10, 18, 29. — Beaubrun 1979: 84, figs 56, 57, 62-64. — Monchartmont 1979: 72. — ° Kocataș 1981: 162. — Mikashavidze 1981: 1417. — Thessalou-Legaki & Zenetos 1985: 309. — Thessalou-Legaki 1986: 183. — ° Koukouras et al. 1992: 223. — Noël 1992: 81. — ° Froglia 1995: 7. — Falciai & Minervini 1996: 147, 4 figs. — d’Udekem d’Acoz 1996: 54. — Ziebis et al. 1996: 619, fig. 1. — Abed-Navandi & Dworschak 1997: 565, figs 1-8. — Sakai 1999a: 20. — *González-Gordillo et al. 2001: 279. — Markham 2001: 197. — ° Türkay 2001: 289.

Callianassa italica Parisi, 1915: 64 , figs 1, 2.

Callianassa (Trypaea) italica – de Man 1928a: 11, figs 5- 5h; 1928b: 27, 101.

Callianassa (Trypaea) truncata – Borradaile 1903: 546. — de Man 1928 b: 28, 101. — Bouvier 1940: 102, fig. 68. — Gurney 1944: 82, figs 1E, 3, 4. — Zariquiey Alvarez 1950: 81, fig. 1, pl. 6 figs 1-6, pl. 7 fig. 2; 1968: 229. — *Dolgopolskaia 1954: 186, figs 5-9; *1969: 316, pls 35-38.

Callianassa (Trypaeta) truncata . —* Băcescu 1967: 229, fig. 104.

Necallianassa truncata – Heard & Manning 1998: 883. — d’Udekem d’Acoz 1999: 156. — Dworschak et al. 2000: 302. — * Lindley et al. 2001: 46. — Markham 2001: tabls 1, 2.

Pestaina truncata – Öksnebjerg 2000: 78 nomen nudum.

TYPE MATERIAL. — Whereabouts unknown.

MATERIAL EXAMINED. — France. Bay of Chingoudy (eastern Pyrénées), A. Chaud coll., V.1982, 2 cl. 9.5 and 10 mm, tl. 40 and 41 mm (figured), 6 cl. 10-11.5 mm, 2 (1 ovig.) cl. 9 mm ( MNHN Th 653) ; III.1983, 1 cl. 9.5 mm ( MNHN Th 658). — Noirmoutiers Island , Y. Gruet coll., 3.V.1973, 1 ovig. cl. 7 mm ( MNHN Th 604) ; 2.VII.1973, 1 ovig. cl. 9 mm ( MNHN Th 605) .

Italy. Gulf of Naples, donated by E. Caroli, 3, 2 ( RMNH D 6586). — South of La Gaiola, 100 m, in mud, 11.VIII.1963, don. J. A. W. Lucas, 1 small ( RMNH D 9070). — Naples, collection E. Caroli, received III.1959 from Zoological Station Naples, 47, 23, 24 ovig. ( RMNH D 12988).

Greece. Kreta, W of Souda Bay , C. d’Udekem d’Acoz coll., 14.VII.1987, 1 cl. 6 mm, 3 ovig. cl. 6- 6.5 mm ( MNHN Th 1354) ; 15.VII.1987, 1 cl. 7.5 mm, 6 (3 ovig.) cl. 5.5-6.5 mm (d’Udekem d’Acoz). — Kalives Harbour, 11.VII.1987, 2 ovig. cl. 4.5 and 5 mm ( d’Udekem d’Acoz ). — Tersanas (west of Stavros ), 15.VII.1987, 22 cl. 3.5- 6 mm, 25 (2 ovig.) cl. 3-6.5 mm ( d’Udekem d’Acoz ). — Pogonia , 0.5-1.5m, C. d’Udekem d’Acoz coll., 12.VII.1993, 1 cl. 5 mm, 8 (7 ovig.) cl. 4-5.5 mm ( MNHN Th 1415). — Georgiopolis, C. d’Udekem d’Acoz coll., 10.VII.1987, 7 cl. 5- 9 mm, 12 (6 ovig.) cl. 5-8 mm (d’Udekem d’Acoz) . Morocco. Leg. Institut scientifique chérifien, III.1966, 3 cl. 5-6.5 mm, 1 cl. 4.5 mm ( MNHN Th 112). — Melilla, VI.1946, leg. J. Rutlant, R. Zariquiey Alvarez coll., 1 ( RMNH D 35798a) .

DISTRIBUTION. — Atlantic coast of France (Noirmoutiers Island, Bay of Biscay) to Atlantic coast of Morocco (de Saint Laurent & Božić 1976). Mediterranean: Greece, Italy, Melilla. Adriatic Sea (Abed-Navandi & Dworschak 1997), Ionian Sea (Thessalou-Legaki 1986; d’Udekem d’Acoz 1996), Aegean Sea ( Kocataș 1981; Koukouras et al. 1992), southeast of Black Sea ( Mikashavidze 1981).

DIAGNOSIS

Carapace ( Fig. 11A ) with dorsal oval, rostrum approximately triangular with blunt or slightly acute tip, rostral spine absent. Telson ( Fig. 11D ) about as long as proximal width, a little narrower distally; lateral borders weakly convex, lateroposterior corner with two transparent minute spines, posterior border with median spinule.

Eyestalk short ( Fig. 11A ), cornea dorsal, subterminal, disk-shaped. A1 peduncle approximately as long as that of A2. Md ( Fig. 11F ) with three-segmented palp, last segment tapering distally. Mx1 ( Fig. 11G ) with basal endite of inverted triangle, distal border widened bearing spiniform setae. Mxp3 ( Fig. 11J, K ) operculiform, length about 1.6-1.7 times merus width, ischium with row of 15-16 spinules on mesial surface; propodus about 1.6-1.7 times as long as wide; dactylus 1.8-2 times as long as wide.

P1 unequal in adult male and female, subequal in certain small specimens. Major P1 of male ( Fig. 11C ) with lower border of merus convex on distal half bearing small rounded teeth; meral hook pointed distally, denticulate on both proximal and distal border; carpus unarmed; propodus dentate on lower border; dactylus with curved tip and dense setae on proximal two-thirds and near base of fixed finger. Major P1 of smaller male and of female similar to that of male but more slen- der, with fewer or very few setae (in female) on dactylus. Minor P1 ( Fig. 11B ) slightly compressed laterally, merus with median spine on lower border, carpus about 1.4-1.5 times as long as wide and 1.5 as long as propodus, lower border regularly curved. P3 ( Fig. 11E ) propodus approximately oval, with lower border rounded, no proximal heel. P4 propodus slender, over three times as long as wide.

Male Plp1 ( Fig. 11L ) small, male Plp2 absent. Female Plp1 ( Fig. 11H ) uniramous, female Plp2 ( Fig. 11I ) biramous, exopod about as long or slightly longer than endopod, latter with distal appendix interna. Pleopods 3-5 ( Fig. 11M ) foliaceous, appendix interna approximately cylindrical, slightly narrowing distally, partly embedded, partly projecting from inner border of endopod.

Uropod ( Fig. 11D ) as long as telson; exopod with distal setal row of dorsal plate at short distance from posterior border; endopod with small lateral distal spine.

Colour

Body yellowish, abdomen dull yellow, pereopods whitish (d’Udekem d’Acoz 1996).

Creamy to yellowish on dorsal abdomen, chelipeds white, sometimes yellow, hepatopancreas orange-brown, ripe ovaries bright orange-red (Dworschak pers. comm.). A coloured photograph is presented in Ziebis et al. (1996).

Size

Medium: cl. 5.5-11.5 mm, largest specimens of cl. 9.5-11.5 mm, tl. 40-43 mm approx.

ECOLOGY

The species occurs in fine sand with or without mud, sometimes with pebbles. It seems to prefer fine sand and can be locally abundant (d’Udekem d’Acoz 1996). According to Picard (1965), C. truncata is characteristic of muddy bottoms along the coast of France. In the Bay of Biscay and nearby areas, specimens were captured in the littoral and at 44-57 m depth (Lagardère 1965).

REMARKS

De Saint Laurent & Božić (1976) cited a pair of fine spinules on each lateroposterior angle of the telson and a small laterodistal spine on the uropodal endopod as diagnostic characters for this species. It must be noted, however, that the spinules on the lateroposterior angle of the telson (figured in this work) are transparent and indistinct in most specimens. By contrast, the laterodistal spinule on the uropodal endopod is obvious. It makes a good distinguishing feature for the species in Europe, together with the lower meral spinule on the minor P1.

In certain populations of small size, e.g., MNHN Th 1415 from Pogonia, Greece, all females (eight specimens with seven ovigerous, cl. 4-5.5 mm) have their P1 subequal and similar to the minor P1 of males. This does not occur in larger specimens, e.g., those of cl. 8-9 mm.

Figures

FIG. 9. — Callianassa subterranea (Montagu, 1808); A, from Plymouth, UK, anterior part of carapace (from de Saint Laurent & Božić 1976); B-P, from Plymouth, UK (MNHN Th 211); B-E, I-P,; F-H,; B, C, major and minor pereopod 1; D, E, maxilliped 3 and mesial view of ischium, merus; F, telson and uropods; G, H, male pleopod 1 and 2; I-L, pereopod 1- 5; M, mandible; N, pleopod 3; O, P, female pleopod 1 and 2. Scale bars: A-E, G-M, O, P, 1 mm; F, N, 2 mm.

FIG. 10. — Callianassa subterranea (Montagu, 1808), from Plymouth, UK (MNHN Th 211); A, B, maxillule and maxilla; C, D, maxilliped 1 and 2. Scale bars: 1mm.

FIG. 8. — Callianassa acanthura Caroli, 1946; A-D, F, G, M, lectotype, from Naples, Italy (RMNH D 15212 a); E, H-L, from Pogonia, Greece (MNHN Th 1400); N, O, from Zirje island, Croatia (NHMW 15372); A, anterior part of carapace; B, C, major pereopod 1 and merus in mesial view; D, minor pereopod 1; E, appendix interna on mesial border of pleopod 3; F, pereopod 3; G, telson and uropods; H, maxilliped 1; I, J, maxilliped 3, mesial and lateral view; K, mandible; L, maxillule; M, male pleopod 1; N, O, female pleopod 1 and 2. Scale bars: A-J, M-O, 2 mm; K, L, 1 mm.

FIG. 11. — Callianassa truncata Giard & Bonnier, 1890; A, from Morocco, anterior part of carapace (from de Saint Laurent & Božić 1976); B-M, from Chingoudy Bay, France (MNHN Th 653); B-G, J-M,; H, I; B, C, minor and major pereopod 1; D, telson and uropods; E, pereopod 3; F, mandible; G, maxillule; H, I, female pleopod 1 and 2; J, K, maxilliped 3 and mesial view of ischium, merus; L, male pleopod 1; M, pleopod 3. Scale bars: 1 mm.