Hippolyte varians Leach, 1814

Cédric, 2007, New records of Atlantic Hippolyte, with the description of two new species, and a key to all Atlantic and Mediterranean species (Crustacea, Decapoda, Caridea), Zoosystema 29 (1), pp. 183-207 : 200-205

Hippolyte varians Leach, 1814

Hippolyte varians Leach, 1814: 431 . — Zariquiey Álvarez 1953: 105, figs 1, 2, 5, 7. — Ledoyer 1969: 342, pl. 1a. — D’Udekem d’Acoz 1995: 497, figs 2a-q, 4a-m; 1996: 90, in part, figs 44-48, not figs 49, 50 (based on H. holthuisi ). — García Raso et al. 1998: 455, fig. 2.

MATERIAL EXAMINED. — Norway. TromsØ area, SommarØy , north coast, 69°38’03”N, 18°01’21”E, lower shore, 0.5 m depth, amongst the brown alga Halidrys siliquosa (L.) Lyngb., 11.VIII.2002, coll. Cédric d’Udekem d’Acoz, 4 specimens (including 2 ovigerous ♀♀) ( TMU, TSZCr 12177). — Same locality, same depth, also amongst Halidrys siliquosa , 30.IV.2006, coll. Cédric d’Udekem d’Acoz, 8 specimens (all with uniform brown colour; no ovigerous specimens) (MNHN-Na16270). — Between 2002 and 2004, a few other specimens including ovigerous females have been observed at the same station, almost always amongst Halidrys , but they have not been preserved .

France. Bay of Biscay, INTRIGAS cruise, sample BB 1- TS01N-F1 (night-time sampling), 46°57’N, 3°28’W, 104 m, 2. VI.2001, coll. Jean-Claude Sorbe, 1 specimen (MNHN-Na16271).

DISTRIBUTION. — East Atlantic: from Norway and Faeroe Islands to Southern Spain, Azores, Madeira, Canary Islands.

REMARKS

D’Udekem d’Acoz (1996) stated that the deepest reliable records of H. varians were at 60 m depth and that the deeper record (200 m) by Kemp (1910) could be based on misidentified H. leptometrae , a species which was not described in 1910. Hippolyte varians is here recorded at 104 m.

In Norway, Dons (1915: 128) stated that H. varians only occurs in the area of Bergen and that it is not present in the northern part of the country. Later on, Sivertsen (1925) recorded it well above Bergen, at his station VII in the Foldenfjord. According to Soot-Ryen (1925), this station is close to RØsvik, and its approximate coordinates are 67°30’N, 15°22’E. Dons (1933) and Brattegard & Holthe (1997) gave no new records farther north. The species is here recorded at 69°38’N, i.e. for the first time above the Lofoten Islands.

The occurrence of a breeding population of H. varians above the Arctic Circle is not necessarily a recent range extension. Indeed in the northernmost part of its range, the distribution of a species is likely to be patchy and a small shrimp like H. varians could be easily overlooked. Furthermore the species is common in the cold waters surrounding the Faeroe Islands ( Berggren 1994), indicating that it is able to maintain populations in subarctic conditions. The collecting station, SommarØy, is situated on the outer coast which is bathed by the Gulf Stream and where the winter temperature is higher than in adjoining fjords.

Few other true temperate European decapods (i.e. species with breeding populations in the Iberian Peninsula or farther south) are common in the area of TromsØ: Sergestes arcticus KrØyer, 1855 , Pasiphaea multidentata Esmark, 1866 , Dichelopandalus bonnieri Caullery, 1896 , Pandalina brevirostris (Rathke, 1843) , Pandalina profunda Holthuis, 1946 , Crangon crangon (Linnaeus, 1758) , Philocheras bispinosus (Hailstone, 1835) , Pontophilus norvegicus (M. Sars, 1861) , Calocaris macandreae Bell, 1846 , Calocarides coronatus (Trybom, 1904) , Galathea nexa Embleton, 1834 , Munida sarsi Huus, 1935 , Munida tenuimana G. O. Sars, 1872 , Pagurus bernhardus (Linnaeus, 1758), and Carcinus maenas (Linnaeus, 1758) . All these species have been collected by the author in 2002-2004. Other decapods recorded as far north are either northern species or erratic individuals of temperate species, which do not form breeding populations above the Lofoten Islands (e.g., Cancer pagurus Linnaeus, 1758 and Geryon trispinosus (Herbst, 1803)) .

KEY TO MATURE FEMALE HIPPOLYTE LEACH, 1914 OF THE ATLANTIC, THE MEDITERRANEAN AND NEIGHBOURING SEAS

The current knowledge of the Hippolyte species of the Atlantic Ocean and the Mediterranean basin is summarized by the present identification key and the checklist given after it. Since the key is intended to be a practical tool only, some species appear twice in it. Species like H. proteus and H. kraussiana key out by geographical area only since morphological accounts on them are insufficient. Only mature females are considered since males are not known for some species, and because males are often much more difficult to identify than females.

1. First article of peduncle of antennula with 1-3 teeth on outer distal corner ................ 2

— First article of peduncle of antennula without teeth on outer distal corner ................. 4

2. First article of peduncle of antennula with 1 tooth on outer distal corner ................... 3

— First article of peduncle of antennula with 2 or 3 teeth on outer distal corner ............... ........................................................................................................... H. obliquimanus

3. Fourth pleonite without pair of dorsolateral styliform teeth ............ H. group ventricosa

— Fourth pleonite with a dorsolateral styliform tooth on each side ............. H. coerulescens

4. No postrostral tooth ................................................................................................... 5

— Postrostral tooth/teeth present .................................................................................. 21

5. Distal article of third maxilliped densely setose (paintbrush-like) ............................... 6

— Distal article of third maxilliped not densely setose .................................................... 7

6. Rostrum with two widely separated dorsal teeth, one on the middle of the rostrum and one close to tip of rostrum. European species ......................................... H. leptometrae

— Rostrum with one dorsal tooth, on the middle of rostrum. South African species .......... ...................................................................................................... H. catagrapha n. sp.

7. Merus of P3-P4 not presenting the combination lateral row of spines + ventral row of spines (there can be a lateral row of spines, or a single distal spine, or no spines at all) ............ 8

— Merus of P3-P4 with lateral row of spines + ventral row of spines (rostrum long [often overreaching scaphocerite], either with no dorsal tooth or with 1 proximal dorsal tooth. Extended forward, P3 falls slightly short of midlength of scaphocerite. Large slender species which sometimes overreaches 40 mm) .............................................. H. inermis

8. Tip of dactylus of P3-P4 with one single very strong distal spine and 2 or 3 spinules on flexor border ............................................................................................................... 9

— Tip of dactylus of P3-P4 with 2 or 3 strong spines (which can be different-sized) and a row of 3-14 well developed spines on flexor border .................................................. 10

9. Dactylus of P3 about 0.85 times as long as carpus. Last two spines of flexor border of P3-P5 about as long as or longer than width of dactylus at the level of insertion of distal unguis. Merus of P3-P4 without spine. Rostrum with 1-3 dorsal teeth, at least one of them being far beyond half of rostrum. East Atlantic species ........... H. longiallex n. sp.

— Dactylus of P3 about 0.50 times as long as carpus. Last two spines of flexor border of P3-P5 no more than 0.25 as long as width of dactylus at the level of insertion of distal unguis. Merus of P3 (and sometimes of P4) with one subdistal outer spine. Rostrum with 0 or 1 (rarely 2) dorsal teeth; if one tooth present, it is on the middle of rostrum. West Atlantic species ......................................................................................... H. nicholsoni

10. Tip of dactylus of P3-P5 with 3 spines, the anterior being significantly shorter than the next two ones. West Atlantic .................................................................................... 11

— Tip of dactylus of P3-P5 with 2 spines, the anterior being longer or subequal to the next one. East Atlantic, Mediterranean, South Africa, Suez Canal .................................... 12

11. Rostrum not overreaching antennular peduncle ................................. H. pleuracanthus

— Rostrum distinctly overreaching antennular peduncle .............................. H. zostericola

12. Rostrum with no dorsal teeth ................................................................................... 13

— Rostrum with at least one dorsal tooth ..................................................................... 16

13. Dactylus of P3-P5 with 8-14 spines (flexor and apical spines considered). Most anterior spine of tip of dactylus of P3-P4 1.0-1.5 times as long as next spine ......................... 14

— Dactylus of P3-P5 with 5 or 6 (sometimes 7) spines (flexor and apical spines considered). Most anterior spine of tip of dactylus of P3-P4 2.2-3.3 times as long as next spine. (First article of carpus of P2 significantly longer than two distal ones combined. P3-P4 merus with a single tooth; adult females with a colour pattern made of transverse red, yellow and white stripes) ................................................................................ H. prideauxiana

14. Stout species. Scaphocerite 2.2-3.0 times as long as wide. Rostrum far from reaching tip of scaphocerite. Extended forward, P3 reaches or nearly reaches tip of scaphocerite ........ 15

— Slender species. Scaphocerite 4.7-5.0 times as long as wide. Rostrum very long (and narrow), almost reaching or overreaching tip of scaphocerite. Extended forward, P3 falls slightly short of midlength of scaphocerite ........................................ H. niezabitowskii

15. Rostrum shorter than eyes (hence considerably shorter than antennular peduncle). P3 merus with 0 or 1 lateral spine ..................................................................... H. palliola

— Rostrum considerably longer than eyes, nearly reaching or slightly overreaching tip of antennular peduncle. P3 merus with several lateral spines ........................ H. lagarderei

16. Rostrum longer than eye .......................................................................................... 17

— Rostrum shorter than eye (very stout species with no more than one dorsal tooth on the rostrum and no ventral tooth) ...................................................................... H. palliola

17. Extended forward, P3 reaches or nearly reaches tip of scaphocerite. Scaphocerite 3.0-3.8 times as long as wide ................................................................................................ 18

— Extended forward, P3 falls slightly short of midlength of scaphocerite. Scaphocerite 4.7- 5.0 times as long as wide (rostrum very long and slender, with 0-4 dorsal teeth at various positions) .......................................................................................... H. niezabitowskii

18. North Atlantic and Mediterranean species ................................................................ 19 — Red Sea species entering the Suez Canal ....................................................... H. proteus — South African species .............................................................................. H. kraussiana — Equatorial West African species ................................................................ Hippolyte sp.

19. Rostrum measuring at least 0.92 of carapace length; large proximal dorsal tooth always present (rostrum dorsally with one subdistal and one proximal teeth). Pereiopods fairly robust or slender. First article of P2 at least 3.1 times as long as wide. Dorsal length of sixth pleonite 1.9-2.6 times longer than its height. Spines of dactylus of P3-P5 long or medium-sized ........................................................................................................... 20

— Rostrum measuring 0.67-0.78 of carapace length; small proximal tooth present or absent (rostrum dorsally with one subdistal tooth, and with or without proximal tooth).Pereiopods very robust. First article of P2 about 2.4-2.6 times as long as wide. Dorsal length of sixth pleonite 1.7-1.8 times longer than its height. Spines (particularly the terminal ones) of dactylus of P3-P5 short ............................................................................ H. lagarderei

20. Robust outline. Stylocerite reaching 0.8-1.0 of basal segment of antennular peduncle. Scaphocerite 3.0-3.5 times as long as wide. Mandibular incisor process with 5 teeth (rarely 4). Ratio between dorsal length and height of sixth pleonite 1.9-2.1. North East Atlantic ........................................................................................................ H. varians

— Slender outline. Stylocerite reaching 0.6-0.7 of basal segment of antennular peduncle. Scaphocerite 3.6-3.8 times as long as wide. Mandibular incisor process with 6 or 7 teeth. Ratio between dorsal length and height of sixth pleonite 2.5-2.6. Mediterranean .......... ................................................................................................................... H. holthuisi

21. P 3 in mature females reaching or slightly overreaching midlength of scaphocerite when extended forward. Outer antennular flagellum shorter than inner antennular flagellum. Scaphocerite 3.8-4.2 times as long as wide. Extremity of hepatic spine very distant of anterior margin of carapace. Dorsal length of sixth pleonite 2 times as long as its height. Eggs large ................................................................................................................. 22 — P3 almost extending to or overreaching apex of scaphocerite. Outer antennular flagellum longer than inner. Scaphocerite 2.5-3.1 times as long as wide. Extremity of hepatic spine almost reaching or slightly overreaching anterior margin of carapace. Dorsal length of sixth pleonite 1.6-1.8 times as long as its height. Eggs small ..................................... 23

22. Rostrum very long, almost reaching extremity of scaphocerites. One postrostral tooth (very rarely 2 teeth) and 2 (rarely 1 or 3) proximal dorsal rostral teeth present; 2 or 3 (rarely 1 or 4) ventral rostral teeth present ............................................... H. sapphica forma A

— Rostrum very short, sometimes almost indistinct, always shorter than eyestalks.One postrostral tooth, no dorsal rostral tooth, and no ventral tooth present .............. H. sapphica forma B

23. Lateral subdistal spine of merus of P5 almost always absent. Rostrum long or short (in Atlantic Ocean and Alboran sea usually shorter than antennular peduncle, often much longer in many Mediterranean populations); when longer than eyestalk, usually narrow or very narrow in Atlantic Ocean and Alboran sea, but sometimes very high in some Mediterranean populations. 1-6 (usually 3 or 4) dorsal teeth: first dorsal tooth and sometimes all dorsal teeth may be erected. Proximal teeth usually narrowly spaced. When at least 3 dorsal teeth, base of second dorsal tooth almost never distinctly overreaching extremity of supraorbital spine (second dorsal tooth and supraorbital spine often approximately at same level). Space between ultimate and penultimate dorsal tooth can be longer, equal or shorter than space between penultimate and antepenultimate dorsal tooth. Ventral rostral tooth or teeth absent or present. In many but not all populations, second pair of dorsolateral pair of dorsolateral spines of telson very frequently closer to first pair than to apex of telson; second pair can be more than 2 times closer to first pair than to telson apex. Incisor process of mandible with 4 (rarely 5) teeth .......................... H. leptocerus

— Lateral subdistal spine present on merus of P 5 in almost all Mediterranean specimens and in 25% of specimens of Arcachon Bay. Rostrum always long (always reaching or overreaching tip of antennular peduncle), high or rather high; 3, occasionally 4 dorsal teeth (rarely 2 teeth in Arcachon Bay), not erected and separated by large space. Base of second dorsal tooth distinctly overreaching extremity of supraorbital spine. Space between ultimate and penultimate dorsal tooth can be equal or shorter (but never distinctly longer) than space between penultimate and antepenultimate teeth. Ventral rostral tooth or teeth always present. Second pair of dorsolateral spines of telson very frequently closer to apex of telson than to first pair; second pair at most 1.4 times closer to first pair than to apex of telson. Incisor process of mandible with 5 or 6 (rarely 4) teeth ............ H. garciarasoi

CHECKLIST OF HIPPOLYTE SPECIES KNOWN IN THE ATLANTIC,

THE MEDITERRANEAN

AND NEIGHBOURING SEAS

H. catagrapha n. sp. West coast of South Africa, on the crinoid Tropiometra carinata (Lamarck, 1816) . Figures: present paper.

H. coerulescens (Fabricius, 1775) . Tropical and subtropical East and West Atlantic in offshore waters; lives on drifting substrates, especially Sargassum . Figures: Chace (1972); Crosnier & Forest (1973); d’Udekem d’Acoz (1996).

H. garciarasoi d’Udekem d’Acoz, 1996 . South of the Bay of Biscay to Mauritania, Mediterranean. Often confused with long-rostred forms of H. leptocerus (Heller, 1863) . Figures: d’Udekem d’Acoz (1996).

H. holthuisi Zariquiey Álvarez, 1953 . Mediterranean Sea. Hippolyte holthuisi is the Mediterranean sister species of the Atlantic H. varians . D’Udekem d’Acoz (1996) synonymized both forms with hesitation. However there is today a consensus for considering them as separate species. Figures: Zariquiey Álvarez (1953); Ledoyer (1969); d’Udekem d’Acoz (1995, 1996, as H. varians , part, Mediterranean specimens: fig. 29b, 49, 50); García Raso et al. (1998).

H. inermis Leach, 1815 . Northeast Atlantic: Ireland to Morocco, Mediterranean. Usually found amongst seagrasses of various species, including Posidonia oceanica (L.) Delile (see d’Udekem d’Acoz 1996). Also found on the red alga Gelidium sesquipetale (Turner) Thur. in the South of the Bay of Biscay (d’Udekem d’Acoz 1999b) and amongst the green alga Caulerpa prolifera (Forssk.) J.V.Lamour. (see López De La Rosa et al. 2006). Figures: Lagardère (1971); d’Udekem d’Acoz (1993, 1996); García Raso et al. (1998).

H. kraussiana (Stimpson, 1860) . South Africa and Madagascar. Figures: Lenz & Strunck (1914 as Virbius capensis Lenz & Strunck, 1914 ); Barnard (1950); Ledoyer (1968); Kensley (1972). Species not adequately described but close to Hippolyte sp. , H. proteus , H. varians and H. holthuisi .

H. lagarderei d’Udekem d’Acoz, 1995 . Northeast Atlantic: Morocco, amongst algae, intertidal on exposed coasts. Figures: Lagardère (1971 as Hippolyte sp. ); d’Udekem d’Acoz (1995, 1996).

H. leptocerus (Heller, 1863) (senior subjective synonym of H. longirostris (Czerniavsky, 1868)) . Northeast Atlantic:Western Ireland and Normandy to Mauritania, Madeira, Cape Verde Islands, Mediterranean, Black Sea. Very variable species. Figures: d’Udekem d’Acoz (1996).

H. leptometrae Ledoyer, 1969 . Northeast Atlantic: Bay of Biscay, Mediterranean, on crinoids of the genus Leptometra . Figures: Ledoyer (1969); Noël (1983); d’Udekem d’Acoz (1996); present paper.

H. longiallex n. sp. Equatorial East Atlantic, on gorgonians. Figures: present paper.

H. nicholsoni Chace, 1972 . Tropical and subtropical West Atlantic, on gorgonians. Figures: Chace (1972); present paper.

H. niezabitowskii d’Udekem d’Acoz, 1996 . Mediterranean. Beds of small seagrasses, at least Cymodocea nodosa (Ucria) Ascherson (see García Raso et al. 2006), but never amongst Posidonia oceanica (L.) Delile. In non-lagunar conditions. Superficially similar to H. inermis but has only one row of spines on the merus of P3-P4 (instead of two), shorter stylocerites, and a more slender rostrum (with a more variable number of dorsal teeth in the populations of the central Mediterranean). Figures: d’Udekem d’Acoz (1996); García Raso et al. (1998).

H. obliquimanus Dana, 1852 (senior subjective synonym of H. curacaoensis Schmitt, 1924 ). West Atlantic: North Carolina to Brazil. Figures: Chace (1972 as H. curacaoensis Schmitt, 1924 ); d’Udekem d’Acoz (1997).

H. palliola Kensley, 1970 . West and South Africa. Figures: Kensley (1970); Crosnier (1972); d’Udekem d’Acoz (1996).

H. pleuracanthus (Stimpson, 1871) . Northwest Atlantic: Connecticut to North Carolina. Very close to H. zostericola . Figures: Chace (1972).

H. prideauxiana Leach, 1817 (senior subjective synonym of H. huntii (Gosse, 1877)) . Northeast Atlantic: Scotland to Canary Islands, Mediterranean, on crinoids of the genus Antedon . Figures: Nouvel (1953, as H. hunti ); d’Udekem d’Acoz (1996).

H. proteus ( Paulson, 1875) . Red Sea, Suez Canal. Figures: Paulson (1875, in part, as Virbius proteus group A I); Gurney (1927). Species not adequately described, close to H. kraussiana , H. varians and H. holthuisi . According to drawings of literature, the first article of the carpus of P2 is much stouter in H. proteus than in H. holthuisi .

H. sapphica d’Udekem d’Acoz, 1993 forma A. Mediterranean basin: Adriatic, Ionian, Aegean and Black seas. Amongst small seagrasses in lagoons, sheltered bays and tidal mud flats. Figures: Băcescu (1967 as H. inermis ); d’Udekem d’Acoz (1993, 1996).

H. sapphica d’Udekem d’Acoz, 1993 forma B. Mediterranean basin: Adriatic and Ionian seas. Amongst small seagrasses in lagoons, sheltered bays and tidal mud flats. No intermediates between the formae A and B of Hippolyte sapphica have been recorded so far. Figures: d’Udekem d’Acoz (1996).

Hippolyte sp. Equatorial West Africa. Figures: present paper.

H. varians Leach, 1814 . Northeast Atlantic from Northern Norway and Faeroe to Portugal, Azores, Madeira, Canary Islands. Figures: Zariquiey Álvarez (1953); Ledoyer (1969); d’Udekem d’Acoz (1995, 1996, as H. varians , part, Atlantic specimens: figs 44-48); García Raso et al. (1998).

H. group ventricosa H. Milne Edwards, 1837 . Indo-Pacific, Red Sea, Suez canal, East of South Africa. Figures of syntypes of H. ventricosa and of the species of the group living in the Red Sea ( H. orientalis Heller, 1862 ): d’Udekem d’Acoz (1999a). Different related species (some presumably not yet named) have been and are still confused under the name H. ventricosa (see d’Udekem d’Acoz 1996, 1999a, 2001).

H. zostericola (S. I. Smith, 1873) . Northwest Atlantic: Massachusetts to Mexico, Caribbean, Bermuda. Figures: Chace (1972).