thecaprellidea(crustacea:amphipoda)fromceuta,northafrica ... ·...

Journal of Natural History, 2002, 36, 675–713

The Caprellidea (Crustacea: Amphipoda) from Ceuta, North Africa,with the description of three species of Caprella, a key to the species

of Caprella, and biogeographical discussion

J. M. GUERRA-GARCIÂ A² and I. TAKEUCHI³ 1

² Laboratorio de Biologõ Âa Marina, Departamento de Fisiologõ Âa y Biologõ ÂaAnimal, Facultad de Biologõ Âa, Universidad de Sevilla, Apdo. 1095, 41080

Sevilla, Spain, e-mail: [email protected]

³ Otsuchi Marine Research Center, Ocean Research Institute, the

University of Tokyo, Akahama, Otsuchi, Iwate 028-1102, Japan

(Accepted 16 November 2000)

The caprellidean fauna of Ceuta, located on the North African side of the Straitof Gibraltar, was studied with regard to species composition and biogeographicalcharacteristics. A new species, Caprella ceutae n. sp. is described and comparedwith the closest species of this genus, C. equilibra and C. fretensis. Furthermore,two species, C. tuberculata and C. liparotensis are redescribed in detail. Ofthe nineteen species collected, three are recorded for the ® rst time in theMediterranean Sea, i.e. C. fretensis, C. erethizon and C. tuberculata. With regardto biogeographical distribution, the Strait of Gibraltar has a very high proportionof endemic species of the Caprellidea, contributing 30.8% of the Mediterraneanendemisms. A key to the 13 species of Caprella from the region of Ceuta is alsoprovided.

Keywords: Amphipoda, Caprellidea, Caprella, taxonomy, biogeography, Ceuta.

Introduction

Although the caprellidean amphipods are considered to be very important as

secondary and tertiary producers in benthic marine communities, they have not been

su� ciently studied in the Mediterranean Sea. Most general amphipod studies havebeen focused mainly on gammarids and only little supplementary information is

given about caprellids (Nicolaidou and Karakiri, 1989; Conradi et al., 1997;

Procaccini and Scipione, 1992; Conradi and LoÂ pez-GonzaÂ lez, 1999). Moreover the

amphipod fauna of the North African coast has still been relatively unexplored,

except for scarce and very fragmented data. Following research on North African

1Present address: Departament of Life Environment Conservation, Faculty ofAgriculture, Ehime University, 3-5-7 Tarumi, Matsuyama, Ehime 790-8566, Japan,e-mail: [email protected]

Journal of Natural HistoryISSN 0022-2933 print/ISSN 1464-526 2 online Ñ 2002 Taylor & Francis Ltd

http://www.tandf.co.uk/journalsDOI: 10.1080/00222930010025923

J. M. Guerra-Garc Ì a and I. Takeuchi676

amphipods conducted up to the beginning of the twentieth century (Lucas, 1849;

Chevreux, 1888, 1910, 1911; Schellenberg, 1928, 1936) and Mayer’s monographs(Mayer, 1890, 1903), a long interval ensued until the recent work of Bakalem and

Dauvin (1995), carried out on the Argelian coast, and our previous work on Ceuta

(Guerra-Garcõ Â a and Takeuchi, 2000).

Material and methods

This study was conducted at Ceuta (Northern Africa), during the summers of1998 and 1999. Caprellids, together with their natural substrata, mainly algae and

hydroids, were collected by SCUBA diving at 23 localities along diŒerent depths

(A, 5± 15m; B, 20 ± 25m; C, 30± 35m; D, 40 ± 45m) (® gure 1). These substrata were

collected in plastic bags containing seawater. In addition to SCUBA diving collec-

tion, soft bottom samples were taken with box-core and Van veen grabs at 10

localities (® gure 1). The sediment samples were sieved using a 0.5 mm mesh.Caprellids were separated, sorted, preserved in a 10% seawater-formol solution and

then identi® ed to species level and counted.

Results

A total of 1051 specimens was examined and grouped into 19 diŒerent speciesof caprellids (table 1).

The dominant species collected from the hard bottom samples were

Fig. 1. Location of Ceuta in the Strait of Gibraltar. Arabic numerals correspond to localitiesof Ceuta where caprellids were collected by SCUBA diving (Depth: A, 5± 15 m; B,20± 25 m; C, 30 ± 35 m; D, 40± 45 m). Sediment samples are indicated in Roman numerals(grabs were always used between 5 and 10 meters in each locality).

The Caprellidea from Ceuta, North Africa 677

Table 1. Species composition of the Caprellidea from Ceuta and number of specimensstudied.

Hard bottom† Soft bottom

Species Number % Number %

Caprella acanthifera Leach, 1814 91 9.7 ± ±C. cavediniae Krapp-Shickel and Vader, 1998 5 0.6 ± ±C. ceutae n. sp (present study) 5 0.6 ± ±C. danilevskii Czerniavskii, 1868 136 14.5 ± ±C. dilatata KroÈ yer, 1843 1 0.1 ± ±C. erethizon Mayer, 1901 29 3.1 ± ±C. fretensis Stebbing, 1878 2 0.2 ± ±C. grandimana Mayer, 1882 3 0.3 ± ±C. hirsuta Mayer, 1890 2 0.2 ± ±C. liparotensis Haller, 1879 11 1.2 ± ±C. penantis Leach, 1814 72 7.7 ± ±C. santosrosai SaÂ nchez-Moyano et al., 1995b 103 11 ± ±C. tuberculata Bate and Westwood, 1868 27 2.9 ± ±Caprella sp (armata-group) 50 5.3 ± ±Pariambus typicus KroÈ yer, 1844 ± ± 100 86.9Pedoculina garciagomezi SaÂ nchez-Moyano et al., 2 0.2 ± ±

1995aPhtisica marina Slabber, 1769 172 18.4 15 13.1Pseudoprotella inermis Chevreux, 1927 13 1.4 ± ±P. phasma (Montagu, 1804) 212 22.6 ± ±

† Mainly composed of the seaweeds Cystoseira usneoides, Asparagopsis armata,Cladostephus verticillatus, Halopteris scoparia and the hydroid Sertularella gayi.

Pseudoprotella phasma Montagu, with 22.6% of total individuals, followed by

Phtisica marina Slabber (18.4%), Caprella danilevskii Czerniavski (14.5%), Caprella

santosrosai SaÂ nchez-Moyano et al. (11%) and Caprella acanthifera Leach (9.7%).

The soft bottoms provided only two species of Caprellidea, i.e. Pariambus typicus

KroÈ yer (86.9%) and Phtisica marina Slabber (13.1%).

Systematics

Caprella acanthifera Leach, 1814

Caprella acanthifera Leach, 1814: 404; Mayer, 1882: 39± 43; Mayer, 1903: 77± 78; Stephensen,1929: 181, ® gure 43; Harrison, 1944: 13± 15, ® gure 5; Krapp-Schickel and Vader, 1998:952± 957, ® gures 2± 3.

Caprella hystrix: KroÈ yer, 1843: 585, pl. 8, ® gures 20± 26.Caprella calva: Bate, 1856: 60, ® gure 57.Caprella aspera: Heller, 1866: 55, pl. 4, ® gures 20± 21.Caprella elongata: Haller, 1880: 409, ® gure 45.Caprella acanthifera forma typica: Mayer, 1890: 44; Chevreux and Fage, 1925: 446 ± 449,

® gures 427 ± 428.Caprella acanthifera intermedia: Monterosso, 1915, ® gures 1± 2.

Material examined. 1A: two males, two females, two ovigerous females; 2A: one

male; 2B: one male, two females, two ovigerous females; 3A: six males, four females,

four ovigerous females, four juveniles; 4A: one male; 5A: three males, one ovigerous

female; 6A: one male; 7A: two males, two females; 11A: one female; 12B: one female;13A: four males, two females, one juvenile; 15A: three males, one female; 15B: three


males, two females, one juvenile; 18A: one male, one ovigerous female; 18C: three

female, one ovigerous female; 19A: nine males, 13 females, four juveniles.

Caprella cavediniae Krapp-Schickel and Vader, 1998

Caprella cavediniae Krapp-Schickel and Vader, 1998: 961 ± 963, ® gure 7.Caprella acanthifera var. acanthifera: Cavedini, 1982: 494, ® gures 1± 2; Krapp-Schickel, 1993:

774± 779, ® gures 528 ± 529.

Material examined. 1C: one female; 23A: three males, one female.

Caprella ceutae n. sp.

(® gures 2± 5)

Material examined. 15D: ® ve males.

Diagnosis

Male. Head with a developed rostrum. Body smooth, except for a pair of

dorsolateral projections on pereonite 6. Inferior surface of the third peduncular

segment of antenna 1 bearing ® ne setae. Basis of gnathopod 2 one-quarter ofpereonite 2 length and carrying small dorsal teeth. Propodus oblong with dorsal

setae.

Etymology

The speci® c name ceutae was chosen to dedicate this species to Ceuta, the locality

where specimens were collected.

Type material

Holotype male from hydroid Sertularella gayi Lamouroux. Depth 40 metres,

August 1999, coll. Juan Rodrõ Â guez. Paratypes: four adult males found together with

holotype. Holotype and two paratypes are deposited in the Museo Nacional de

Ciencias Naturales de Madrid, Spain (No. MNCN 20.06/4647). The other twoparatypes are in the Laboratorio de Biologõ Â a Marina, Universidad de Sevilla, Spain.

Type locality

Ceuta, North Africa, Mediterranean (35ß 54 ¾ N, 5 ß 16 ¾ 35 ² W).

DescriptionHolotype male

Body length. 8.9 mm

Lateral view. Body slender. Head with a well-developed rostrum. Pereonite 2

elongated. Peronites 3, 4 and 5 subequal. Pereonite 6 with a pair of acute dorsolateral

projections close to the coxae.

Gills. Oval, length ca twice width.Antennae. Antenna 1 about half of body length, with 14 articles in the ¯ agellum;

the third peduncular article bearing dense, plumose setae on its inferior surface.

Antenna 2 about half length of antenna 1; peduncular articles 3 and 4 carrying nine

pairs of long serrated setae; ¯ agellum two-articulate, setose.

Mouthparts. Upper lip symmetrically bilobed, pubescent apically. Inner lobe of

the lower lip round, outer lobe round apically; both inner and outer lobe pubescentapically. Mandibular process strong; incisor and lacinia mobilis of both left and


Fig. 2. Caprella ceutae, new species. Male lateral view. Scale bar: 1 mm.


Fig. 3. Caprella ceutae, new species. Male. a, lower lip; b, upper lip; c, maxilliped; d, leftmandible; e, right mandible; f, maxilla 1; g, maxilla 2. Scale bars: 0.1 mm.

right mandibles ® ve-toothed. Left mandible with a row of three pectinated setae,

right with two. Maxilla 1 outer lobe carrying seven serrated spines on its end; article2 of palp with ® ve robust apical setae and ® ve lateral slender setae. Maxilla 2 inner


Fig. 4. Caprella ceutae, new species. Male. a, antenna 1; b, antenna 2; c, gnathopod 1;d, gnathopod 2. Scale bars: a, b: 1 mm; c: 0.5 mm; d: 1 mm .

lobe rounded, a little shorter than outer, which is slender; both inner and outer lobewith 15 marginal setae. Maxilliped inner plate rectangular, carrying two teeth, and

one simple and seven plumose setae on the apical margin; outer plate, twice as large

as inner, with four teeth and ® ve simple setae; palp four-articulate, articles 2 and 3

with long setae on inner margin; article 4 with two rows of setulae on grasping

margin.

Gnathopods. Gnathopod 1 setose; ischium round, merus and carpus each withmarginal row of seven setae posteriorly; propodus posterior margin with two prox-

imal grasping spines. Gnathopod 2 inserted near to the distal end of pereonite 2;

basis one-quarter length of pereonite 2, with a strong lateral ridge which bears

several small tubercles; ischium round; merus length ca 1.5 times ischium, with an

acute projection on its ventral margin; carpus very short; propodus palm slightly

convex, proximally with a robust tooth and one spine, distally with a largerectangular process; dorsal surface profusely setose.


Fig. 5. Caprella ceutae, new species. Male. a, pereopod 5; b, pereopod 6; c, pereopod 7;d, abdomen (ventral view). Scale bars: 0.5 mm.

Pereopods. Pereopods 5, 6 and 7 increasing in length; pereopods 5± 7 basis with

a developed carina; carpus and merus with a dorsal row of four setae; palm of

propodus concave with two proximal grasping spines.

Abdomen. With a pair of large appendages, a pair of lateral lobes and a singledorsal lobe. Abdominal appendage two-articulate; basal article about twice longer

than distal one, length ca 1.5 times width; distal segment round, carrying a lateral

seta and a distal one or two. Dorsal lobe with two plumose setae. Penes medial,

length ca twice width.

RemarksThe genus Caprella is the largest genus in the suborder Caprellidea. The species

closest to Caprella ceutae, n. sp. are C. equilibra Say 1818 and C. fretensis Stebbing,

1878. Specimens of C. equilibra and C. fretensis belonging to the private collection

of the senior author (JMGG) collected along the Atlantic and Mediterranean coasts

of southern Spain, were examined with the purpose of comparison. The character-

istics of C. equilibra from Southern Spain are in complete agreement with thoseincluded in the description of Krapp-Schickel (1993). Caprella ceutae is distinguished


from C. equilibra as follows: (1) in C. ceutae the prominent ventral spine between

the second gnathopod, present in C. equilibra, is lacking; (2) C. ceutae lacks a pairof tubercles anteriorly on pereonite 5, however these tubercles are characteristic of

C. equilibra; (3) the inferior margin of the third peduncular article of antenna 1 and

the dorsal surface of gnathopod 2 have ® ne and dense setae in C. ceutae. These

setae are lacking in C. equilibra.

Caprella ceutae is also close to the Atlantic C. fretensis, also found in Ceuta

(although only two damaged specimens). A detailed comparison betweeen C. ceutaefrom Ceuta and undamaged specimens of C. fretensis collected by JMGG from

Huelva (Atlantic coast of southern Spain) re¯ ected several constant diŒerences

between them. In C. fretensis the inferior surface of the second and third peduncular

articles of antenna 1 bears rows of short ® ne setae and the propodus of gnathopod

2 is entirely covered with dense setae. These characteristics, present in specimens

from Southern Spain, are in agreement with descriptions and ® gures of C. fretensisfrom the coasts of England and Ireland, included in Chevreux and Fage (1925) and

Harrison (1944). Together with these diŒerences in antenna 1 and gnathopod 2

propodus, we have observed that in C. fretensis from Southern Spain the tubercles

on the lateral ridge of gnathopod 2 basis and the acute dorsolateral projections on

pereonite 6 are lacking. Furthermore, the abdominal appendages are diŒerent; inC. fretensis the proximal article is ca one-quarter as long as the distal one and half

as long as wide. Moreover, the distal article is clearly serrated distally whereas it is

smooth in C. ceutae.

The present specimens of C. ceutae were collected from hydroids in an area of

high transparency; visibility is ca 10 ± 30 m. depending on the weather. The area,

which is located near the top of the Ceuta Peninsula, is exposed to the strong eŒectsof local currents. Caprella ceutae lives together with Pseudoprotella inermis Chevreux,

1927, Caprella santosrosai SaÂ nchez-Moyano et al., 1995b, and the Atlantic species

C. erethizon Mayer, 1901 and C. tuberculata Bate and Westwood, 1868.

Caprella danilevskii Czerniavski, 1868

(® gure 6)

Caprella Danilevskii Czerniavski, 1868: 92, pl. 6, ® gures 21± 34; Mayer, 1890: 58, pl. 5,® gure 44; pl. 7, ® gures 12 ± 13.

Caprella Danilewskii: Chevreux and Fage, 1925: 454, ® gure 432.Caprella danilevskii: McCain, 1968: 22, ® gures 10± 11; McCain and Steinberg, 1970: 16;

Cavedini, 1982: 499; Krapp-Schickel, 1993: 779, ® gure 531.

Material examined. 1A: 53 males, 34 females, four ovigerous females, 36 juveniles;

3A: one male, one female; 4A: one male, one ovigerous female; 6A: two males; 15A:

two males, one female.

Caprella dilatata Kroyer, 1843

Caprella dilatata Kroyer, 1843: 585, pl. 8, ® gures. 1± 9; McCain, 1968: 38; McCain andSteinberg, 1970: 17; Cavedini, 1982: 499.

Caprella acutifrons: Mayer, 1882: 48, pl. 1, ® gure 9; pl. 2, ® gures 12± 22; pl. 4, ® gures 26± 28;pl. 5, ® gures 15, 22± 23.

Caprella acutifrons forma typica: Mayer, 1890: 54, pl. 2, ® gure 34; pl. 4, ® gures 62± 63; pl. 7,® gures 16± 17; Chevreux and Fage, 1925: 452.

Caprella acutifrons forma minor: Mayer 1890: 54, pl. 2, ® gure 35; pl. 4, ® gures 54, 64.Caprella penantis forma typica: Schellenberg, 1928: 678; Schellenberg, 1936: 24.Caprella acutifrons forma typica: Fischetti, 1937: 13.


Fig. 6. Caprella danilevskii Czerniavski, 1868. Lateral view: a, male; b, female. Scale bar:1 mm.

Material examined. 17A: one male.

Caprella erethizon Mayer, 1901

(® gure 7)

Caprella erethizon Mayer, 1901: 239± 245, ® gures 1± 3; Mayer, 1903: 101, pl. 4, ® gures 12± 13;pl. 7, ® gure 77; Chevreux and Fage, 1925: 459 ± 460, ® gure 436; Bertrand, 1941: 9, 17,20± 23, ® gure 1; Bocquet and Peltier, 1963: 152± 165, ® gures 1± 10, pls. 1 ± 2.


Fig. 7. Caprella erethizon Mayer, 1901. Lateral view: a, male; b, female. Scale bar: 1 mm.

Material examined. 2B: three males; 7D: one male; 15D: 12 males, 12 females,

one juvenile.

Remarks

Caprella erethizon Mayer has been considered, as well as C. tuberculata, a rare

species (Chevreux and Fage, 1925). The original description of Mayer (1901) waspoor, with only the female described. Chevreux and Fage (1925) redescribed the


female and described the juvenile male of C. erethizon with very schematic ® gures.

Bertrand (1941) found an adult male of C. erethizon but he added little informationabout its morphological characteristics. Bocquet and Peltier (1963) ® nally gave a

precise redescription of this species. The specimens of C. erethizon found in Ceuta,

North Africa are identical to those described by these authors from RoscoŒ(Atlantic

French coasts). This species was previously recorded on Atlantic coasts of England

and France and only one specimen has been found on the Atlantic Portuguese coast

(Marques and Bellan-Santini, 1985).

Caprella fretensis Stebbing, 1878

Caprella fretensis Stebbing, 1878: 31± 4, pls. 5, ® gure 1; Stebbing, 1879: 521; Mayer, 1882: 58;Mayer, 1890: 62± 63, pl. 4, ® gures 38± 39; pl. 5, ® gures 41± 42; Chevreux and Fage, 1925:457± 459, ® gure 435; Toulmond and Truchot, 1964: 36.

Material examined. 2A: one male; 5A: one male.

Caprella grandimana Mayer, 1882

Caprella grandimana Mayer, 1882: 43, pl. 1, ® gure 5; pl. 2, ® gures 23± 29; pl. 4, ® gures 29± 31;Cavedini, 1982: 501, ® gure 2.

Caprella acanthifera var. grandimana : Mayer, 1890: 47.Caprella hirsuta f. longimana: Chevreux, 1913: 5, ® gure 1; McCain and Steinberg, 1970: 23.Caprella acanthifera ssp. grandimana : Carausu and Carausu, 1959: 409.

Material examined. 2B: two males, one juvenile.

Caprella hirsuta Mayer, 1890

Caprella hirsuta Mayer, 1890: 77, pl. 2, ® gure 19; pl. 4, ® gures 26 ± 29; Chevreux and Fage,1925: 449, ® gure 429; McCain and Steinberg, 1970: 23; Cavedini, 1982: 503.

Material examined. 15A: one male; 17A: one male.

Caprella liparotensis Haller, 1879(® gures 8± 11)

Caprella liparotensis Haller, 1879: 232; Haller, 1880: 404, pl. 23, ® gures 41 ± 42; Mayer, 1890:57; Mayer, 1903: 114, pl. 8, ® gure 23; Chevreux and Fage, 1925: 452, ® gure 431; Fiorencis,1940: 15, ® gure 6; McCain and Steinberg, 1970: 29; Cavedini, 1982: 504.

Caprella dentata: Haller, 1880: 744, ® gures 4± 9; Mayer, 1882: 50, pl. 1, ® gure 8; pl. 3, ® gures1± 9; pl. 4, ® gure 33.

Material examined. 17A: 10 males, one female.

Diagnosis

Head with developed rostrum. A pair of lateral spines anteriorly on pereonites

2, 3 and 4 in male; in female only on pereonite 3. In male two pairs of dorsal

tubercles on pereonite 5, one pair on pereonite 6 and two pairs of small tubercles

on pereonite 7. These two pairs on pereonite 7 are lacking in female. Peduncular

articles 2± 3 of antenna 1 with dense plumose setae on ventral margin. Basis ofgnathopod 2 very short, about one-® fth of pereonite 2, with carina.


Fig. 8. Caprella liparotensis Haller, 1879. Lateral view: a, male; b, female. Scale bar: 1 mm.


Fig. 9. Caprella liparotensis Haller, 1879. Male. a, lower lip; b, upper lip; c, maxilliped;d, left mandible; e, right mandible; f, maxilla 1; g, maxilla 2. Scale bars: 0.1 mm.

Material examined

Male `a’ from the seaweed Cystoseira tamariscifolia Papenfuss. Depth 10 metres,June 1999, coll. JoseÂ Manuel Guerra-Garc õ Â a. Female `b’ found together with male


Fig. 10. Caprella liparotensis Haller, 1879. a± d, male. a, antenna 1; b, antenna 2;c, gnathopod 1; d, gnathopod 2. e, female gnathopod 2. Scale bars: 0.5 mm.


Fig. 11. Caprella liparotensis Haller, 1879. a± d, male. a, pereopod 5; b, pereopod 6;c, pereopod 7; d, abdomen (ventral view). e, female abdomen (ventral view). Scalebars: a± c: 0.5 mm; d, e: 0.1 mm.

à’ . Other specimens studied: nine males collected together with male à’ and female

`b’ . Male à’ , female `b’ and three more specimens are deposited in the MuseoNacional de Ciencias Naturales de Madrid, Spain (No. MNCN 20.06/4648 ). The


other six specimens are in the Laboratorio de Biologõ Â a Marina, Universidad de

Sevilla, Spain.

Locality

Ceuta, North Africa, Mediterranean (35ß 53 ¾ 30 ² N, 5 ß 17¾ 00 ² W ).

Redescription

Male `a’

Body length. 8.2 mm

Lateral view. Head with a well developed rostrum. A pair of lateral spines

anteriorly on pereonites 2± 4. Two pairs of dorsal tubercles on pereonite 5 and 7,

One pair on pereonite 6.

Gills. Rounded. Length ca 1.3 times width.

Antennae. Antenna 1 shorter than cephalon (head1 pereonite 1) and pereonite

2 together; peduncular articles 1 and 2 slightly widened; articles 2 and 3 carrying

dense plumose setae on ventral margin and very short ® ne setae on dorsal surface;

¯ agellum 14-articulate. Antenna 2 a little longer than antenna 1 peduncle; peduncular

articles 3 and 4 carrying ten pairs of long serrated setae; ¯ agellum two-articulate;

basal article with seven pairs of plumose setae.

Mouthparts. Upper and lower lip similar to those in Caprella ceutae n. sp.

Mandibular molar process strong, bordered by teeth. Pars incisiva and lacinia

mobilis with ® ve teeth on each; left mandible with three pectinated setae, right with

only two; a row of short and ® ne setae between these pectinated setae and the molar

process. Maxilla 1 outer lobe carrying seven serrated spines apically; article 2 of the

palp with 11 marginal setae and 13 lateral ones disposed along the surface. Maxilla

2 inner lobe with 12 apical setae, outer lobe with nine setae. Inner plate of maxilliped

nearly as large as the outer plate; inner plate with nine plumose setae and three

teeth; outer plate with long simple seta and ® ve teeth; articles 2 and 3 of palp

carrying many long setae on inner margin; article 4 shorter than article 3, with rows

of setulae on its grasping margin.

Gnathopods. Gnathopod 1 setose; ischium to merus with many ventral setae;

propodus with two proximal grasping spines, palm shallowly convex. Gnathopod 2

inserted near distal end of pereonite 2, as in Caprella ceutae n. sp.; basis short, about

one-® fth of pereonite 2, with a very developed carina; ischium round; merus length

ca twice ischium, with an acute projection on ventral margin as in Caprella ceutae

n. sp.; propodus longer than basis to ischium combined; length ca 2.5 times width;

palm concave with one projection bearing a tooth and spine halfway from proximal

end, another projection one-quarter length from distal end, followed by `U’ notch

distally.

Pereopods. Pereopods 5, 6 and 7 increasing in length; basis with carina on

posterior margin; merus and carpus carrying three setae on outer margin; carpus

with several pairs of strong setae on the anterior margin; palm of propodus concave

with two proximal grasping spines and two anterior rows of short and wide setae.

Abdomen. With a pair of two-articulate appendages, a pair of lateral lobes and

a single dorsal lobe which carries two plumose setae. Penes short, length about 1.2

times width.


Female `b’ .

Body length 6.8 mm. A pair of lateral spines anteriorly on pereonite 3. Pereonite7 without dorsal tubercles. Gnathopod 2 inserted on the anterior half of pereonite

2; length of propodus ca 1.5 times width. Brood lamellae of pereonite 3 setose on

the posterior margin; brood lamellae on pereonite 4 not setose. Abdomen with a

pair of lateral lobes and a dorsal lobe carrying 2 plumose setae.

Remarks

Although Chevreux and Fage (1925) and Krapp-Schickel (1993) provided

detailed descriptions of Caprella liparotensis from Cette (France) and Napoli (Italy)respectively, we have included complete drawings of antennae, mouthparts and

abdomens based on the newly collected specimens from Ceuta. The comparison

reveals several diŒerences between the previous descriptions and present specimens

from Ceuta: (1) female from Ceuta carries a pair of lateral spines on pereonite 3,

while in C. liparotensis from Napoli the female lacks lateral spines on pereonites 2and 3 (Krapp-Schickel, 1993); (2) males from Ceuta bear two pairs of tubercles on

pereonite 7 while those from Napoli lack these tubercles (Krapp-Schickel, 1993);

(3) the specimens of C. liparotensis from Ceuta possess three teeth on the inner

plate of the maxilliped and four to six teeth on the outer plate. Chevreux and Fage

(1925) reported that the maxilliped of C. liparotensis was similar to that in

C. acutifrons. They described for C. acutifrons (old complex of C. penantis,C. andreae, C. dilatata and other close species) an inner plate with two large teeth

and an outer plate with seven to eigth teeth. Nevertheless these diŒerences are not

enough to consider these specimens from Ceuta as a new species.

Caprella penantis Leach, 1814

(® gure 12)

Caprella Penantis Leach, 1814: 404.Caprella acutifrons: Mayer, 1882: 48; Mayer, 1890: 50, pl. 2, ® gures 36± 37, 39± 41; pl. 4,

® gures 52± 53, 55, 57± 61, 65± 69; Mayer, 1903: 79, pl. 3, ® gures 4± 28; pl. 7, ® gures 62± 65.Caprella acutifrons f. testudo: Chevreux and Fage, 1925: 452, ® gure 430.Caprella penantis: McCain, 1968: 33, ® gures 15± 16; McCain and Steinberg, 1970: 33; Cavedini,

1982: 508.

Material examined. 2A: one female; 3A: three females, two juveniles; 6A: ® ve

males, two females, one ovigerous female, two juveniles; 13A: one male; 15A: three

males, three females, three juveniles; 17A: 14 males, 14 females, three ovigerous

females, 15 juveniles.

Caprella santosrosai SaÂ nchez-Moyano et al., 1995(® gures 13 ± 16)

Caprella santosrosai SaÂ nchez-Moyano et al., 1995b: 197 ± 204, ® gures 1± 5.

Material examined. 7D: two males, two females; 14A: six males, two females,

one ovigerous female, three juveniles; 15A: one male; 15D: 42 males, 22 females,

® ve ovigerous females, 17 juveniles.

Remarks

The record of Caprella santosrosai on the coast of Ceuta represents the ® rst forthis species since its original description (SaÂ nchez-Moyano et al., 1995b). Although


Fig. 12. Caprella penantis Leach, 1814. Lateral view: a, male; b, female. Scale bar: 1 mm.


Fig. 13. Caprella santosrosai SaÂ nchez-Moyano et al., 1995. Lateral view: a, male; b, female.Scale bar: 1 mm.


Fig. 14. Caprella santosrosai SaÂ nchez-Moyano et al., 1995. Male. a, lower lip; b, upper lip;c, maxilliped; d, left mandible; e, right mandible; f, maxilla 1; g, maxilla 2. Scale bars: 0.05 mm.

this species was fully described, we have included complete ® gures because several

diŒerences have been found between specimens from southern Spain and northernAfrica. The general lateral view is very similar. However the mouthparts and


Fig. 15. Caprella santosrosai SaÂ nchez-Moyano et al., 1995. a± d, male. a, antenna 1;b, antenna 2; c, gnathopod 1; d, gnathopod 2. e, female gnathopod 2. Scale bars:a: 0.5 mm; b: 0.3 mm; c: 0.5 mm; d: 0.2 mm; e: 0.3 mm.


Fig. 16. Caprella santosrosai SaÂ nchez-Moyano et al., 1995. a± d, male. a, pereopod 5;b, pereopod 6; c, pereopod 7; d, abdomen (ventral view). e, female abdomen(ventral view). Scale bars: a± c: 0.5 mm; d, e: 0.1 mm.


abdomen are diŒerent: (1) in specimens collected from Ceuta the outer lobe of the

maxilliped has four teeth and the inner lobe three, while in the specimens fromsouthern Spain it has three and two respectively; (2) specimens from Ceuta have no

plumose seta on the outer lobe of maxilla 1, while this seta, which is rare among

species of Caprella, is present in specimens from southern Spain; (3) the most

remarkable diŒerences are found in the male abdomen; while specimens from Ceuta

have the ® rst pair of pleopods clearly biarticulate, no distinct articulation was found

in the specimens from southern Spain (SaÂ nchez-Moyano et al., 1995b). Furthermore,the distal article of the ® rst pleopods is serrated on the apical margin in male

specimens from Ceuta, whereas it is smooth in specimens from southern Spain.

Caprella sp (belonging to the armata-group, see Krapp-Schickel and Vader, 1998)

Material examined. 15A: three males, four females, one juvenile; 16A: four males,

one ovigerous female; 17A: four males, three females, three juveniles; 17B: three

males, one female, one ovigerous female, three juveniles; 20A: two males, one female;

22A: ® ve males, three females, two ovigerous females, three juveniles; 23A: two

males, one female.

Remarks

The specimens studied are very similar to those described as Caprella acanthiferadiscrepans (Krapp-Schickel, 1993). They are characterized by the presence of an

axillary spine near the coxa of gnathopod 2. The taxa with this spine, belonging to

the armata-group, are still awaiting revision (Krapp-Schickel and Vader, 1998)

Caprella tuberculata Bate and Westwood, 1868

(® gures 17 ± 20)

Caprella tuberculata GueÂ rin, 1836, pl. 28, ® gure 1; Goodsir, 1842: 188 ± 189, pl. 3, ® gures 6± 7;Bate and Westwood, 1868: 68± 70; Mayer, 1882: 56 ± 57, ® gures 15 ± 16; Chevreux and Fage,1925: 460± 461, ® gure 437; Schellenberg, 1942: 230± 239, ® gure 199.

Caprella acanthifera: Bate, 1862: 366, pl. 57, ® gure 2.

Material examined. 2B: ® ve males, six females; 15D: four males, six females, four

ovigerous females, two juveniles.

Diagnosis

Head with a short and acute rostrum in males, and small round protuberance

in females. Except the ® rst, all body pereonites with strong and numerous dorsal

tubercles. Males with a pair of lateral tubercles on pereonite 5. Pereonites 1 and 2

elongated in male, pereonite 2 with plumose setae in males. Palm of the propodusof gnathopod 2 in males with large thumb-like protuberance de® ning the palm, and

with long and dense, plumose setae. Anterior surface of basis and propodus also

with plumose setae. Pereopods 5 and 6 with a pair of proximal grasping spines.

Pereopod 7 with four grasping spines in males (in clusters of 2-1-1) and three in

females (2-1).

Material examined

Male `a’ from the hydroid Sertularella gayi Lamouroux (35 ß 55 ¾ 20 ² N, 5ß 22 ¾ W ).Depth 28 metres, July 1999, coll. Jose Manuel Guerra-Garc õ Â a. Female `b’ found


Fig. 17. Caprella tuberculata Bate and Westwood, 1868. Lateral view: a, male; b, female.Scale bar: 1 mm.

together with male à’ . Other specimens: four males, ® ve females collected together

with male à’ ; four males, ten females and two juveniles on seaweed Dilophus spiralisHamel (35 ß 54 ¾ N, 5 ß 16 ¾ 35 ² W ), August 1999, coll. Juan Rodrõ Â guez. Male à’ , female


Fig. 18. Caprella tuberculata Bate and Westwood, 1868. Male. a, lower lip; b, upper lip;c, maxilliped; d, left mandible; e, right mandible; f, maxilla 1; g, maxilla 2. Scalebars: 0.05 mm.

`b’ and another three specimens are deposited in the Museo Nacional de Ciencias

Naturales de Madrid, Spain (No. MNCN 20.06/4649), the remaining specimens arein the Laboratorio de Biologõ Â a Marina, Universidad de Sevilla, Spain.


Fig. 19. Caprella tuberculata Bate and Westwood, 1868. a± d, male. a, antenna 1; b, antenna2; c, gnathopod 1; d, gnathopod 2. e, female gnathopod 2. Scale bars: a,b: 0.5 mm;c: 0.2 mm; d: 0.5 mm; e: 0.2 mm.


Fig. 20. Caprella tuberculata Bate and Westwood, 1868. a± d, male. a, pereopod 5; b, pereo-pod 6; c, pereopod 7; d, abdomen (ventral view). e± f, female. e, pereopod VII;f, abdomen (ventral view). Scale bars: a± c, e: 0.5 mm; d,f: 0.05 mm.


Locality

Ceuta, North Africa, Mediterranean (35ß 55 ¾ 20² N, 5 ß 22 ¾ W; 35ß 54 ¾ N, 5 ß 16¾ 35 ² W ).

Redescription.

Male `a’

Body length. 5.9 mm.

Lateral view. Head with a short, acute and curved rostrum. Except the ® rst one,

all pereonites with tubercles following the formula 0-2.1-2.1-1.2.1- 2( lateral ).2.2-4-2.2. Pereonites 1 and 2 elongated; pereonite 2 bearing long plumose setae dorsally

and ventrally.

Gills. Oval, length ca twice width.

Antennae. Antenna 1 with the same length as cephalon and pereonite 2 together;

peduncular articles setose; ¯ agellum with 11 articles. Antenna 2 about half length

of antenna 1; peduncular articles 3 and 4 carrying six pairs of long, serrated setae;¯ agellum two-articulate, setose.

Mouthparts. Upper and lower lip similar to those in Caprella ceutae n. sp. Pars

incisiva and lacinia mobilis with ® ve teeth on each; left mandible with three pectinate

setae and the right with two; a row of short and ® ne setae between the pectinate

setae and the molar process. Maxilla 1 outer lobe with seven serrated spines distally;article 2 of palp with eight distal setae and a row of four lateral setae. Inner lobe

of maxilla 2 shorter than outer, with 12 apical setae; outer lobe with nine apical

setae. Inner plate of maxilliped shorter and a little wider than outer plate, with seven

plumose setae and two teeth; outer plate with long simple setae and three teeth;

article 2 and 3 of palp with long setae; article 4 with rows of setulae on grasping

margin.Gnathopods . Merus and carpus of gnathopod 1 with row of four setae on posterior

margin; propodus with a pair of proximal grasping spines; palm of propodus with

a few setae; dactylus partially serrated on the inner margin. Gnathopod 2 inserted

on the posterior part of the pereonite; basis one-third length of pereonite 2, with

carina and several plumose setae on anterior surface; ishium, merus and carpus

rounded; merus about 1.5 times larger than ischium; carpus very short, length caone-® fth of merus; palm of propodus de® ned proximally by a large, thumb-like

protuberance and distally by a round projection; between these two projections, the

palm of the propodus is very concave, covered with a mixture of dense ® ne plumose

and simple setae; dorsal surface of propodus also with plumose setae; dactylus with

medial protuberance and plumose setae.Pereopods. Pereopods 5, 6 and 7 increasing in length respectively; palm of

propodus concave; a pair of proximal grasping spines on pereopods 5, 6 and 7; on

pereopod 7, two additional grasping spines, four in total.

Abdomen. With a pair of appendages, a pair of lateral lobes and a single dorsal

lobe; appendage two-articulate, the two articles of each appendage rounded and

approximately the same size; both articles carrying three setae; distal article serratedon the apical margin; lateral lobes carrying four simple setae and dorsal lobe with

two plumose setae. Penes median, slender, length ca three times width.

Female `b’

Body length. 4.1 mm. Head with a short dorsal protuberance in the middle.

Pereonite 1 and 2 not elongated; tubercle formula on pereonites 0-1.2.1-2-1.2.1 -2-4-2.2, pereonite 5 lacking lateral tubercles; pereonite 2 without setae. Peduncular


articles of antenna 1 scarcely setose; ¯ agellum of antenna 1 with eight articles.

Gnathopod 2 inserted on anterior half of pereonite 2; palm de® ned by two smallteeth instead of thumb-like protuberance, without distal projection at the dactylar

hinge; basis, propodus and dactylus without ® ne, plumose setae. Pereopod 7 pro-

podus with three grasping spines. Anterior brood lamellae setose all round, posterior

pair not setose. Abdomen with a pair of lateral lobes which carry two setae and a

dorsal lobe with two plumose setae.

Remarks

As McCain and Steinberg (1970) pointed out, the nomenclature of this speciesis one of the most confused among the caprellid species. Indeed, Mayer (1882)

considered that the specimens described by GueÂ rin (1836) and Goodsir (1842) were

not C. tuberculata. Stebbing (1888) mentioned that GueÂ rin’s C. tuberculata could

be a synonym of C. scaura. The description of C. tuberculata given by GueÂ rin (1836)

is quite brief.The specimens found in Ceuta are in agreement with the description of

C. tuberculata given by Bate and Westwood (1868) for Atlantic specimens. However,

we have observed several diŒerences. Following the descriptions of Bate and

Westwood (1868), Chevreux and Fage (1925) and Harrison (1944) the most striking

diŒerence between C. tuberculata of Bate and Westwood and C. tuberculata found

at Ceuta is the size; males of Caprella tuberculata from Ceuta do not exced 5.9 mm(although we have observed only nine adult males), while specimens from British

and French coasts (Bate and Westwood, 1868) usually reach 15 mm at adulthood.

Caprella tuberculata from Ceuta is also distinguished from Atlantic specimens as

follows: (1) males from Ceuta have a short and round rostrum on the head while

Atlantic specimens have a very short, central, dorsal spine; (2) male `a’ and female

`b’ from Ceuta have tubercles following the formula (0-2.1-2.1-1.2.1- 2( lateral ).2.2-4-2.2 in male, and 0-1.2.1-2-1.2.1-2-0-2. 2 in female). These tubercles are always

present in the other specimens examined but some small tubercles may occur

depending on the individuals. There is no speci® c formula for tubercles in descrip-

tions of Atlantic specimens; (3) in C. tuberculata from Ceuta the propodus of

pereopod 7 has four grasping spines in males and three in females, while there is

only a single pair of grasping spines in Atlantic specimens (both male and females);(4) the posterior brood lamella are not setose on the posterior edges in females from

North Africa; a setose margin is present in Atlantic specimens.

Before this study, Caprella tuberculata had been found on the British and French

coast but also at Santander, northern Spain (Chevreux and Fage, 1925). Marques

and Bellan-Santini (1985) also reported 27 specimens of C. tuberculata from betweenPeniche and Arrabida (Portuguese Atlantic coast). An extensive comparative study

between Atlantic individuals of C. tuberculata and the specimens of the Strait of

Gibraltar would be necessary in order to elucidate whether the above diŒerences are

intra-speci ® c or inter-specic. Recently, a detailed comparison has been successfully

conducted for the species complex Caprella acanthifera ( Krapp-Schickel and Vader,

1998) collected from Atlantic and Mediterranean waters.

Pariambus typicus Kroyer, 1844

Podalirius typicus KroÈ yer, 1844: 283, pl. 3, ® gure 1; Mayer, 1882: 75, ® gures 30± 31; pl. 4,® gure 14; Mayer, 1890: 92, pl. 4, ® gures 7± 8; pl. 5, ® gures 62± 64; pl. 6, ® gure 17; Mayer,1903: 63, pl. 7, ® gures 46± 47; pl. 10, ® gures 4± 9.


Pariambus typicus : Chevreux and Fage, 1925: 441, ® gure 425; McCain and Steinberg, 1970: 62.

Material examined. I: four males, eight females; III: one male; IV: 13 males, 36

females, 12 juveniles, V: three males, three females, VI: three males, two females;VII: two males, ® ve females, one juvenile; VIII: one male, one female, one ovigerous

female; IX: one male; X: one male, two females.

Pedoculina garciagomezi SaÂ nchez-Moyano et al., 1995

Pedoculina garciagomezi SaÂ nchez-Moyano et al., 1995a: 419 ± 427, ® gures 2 ± 6.

Material examined. 16A: one male; 18D: one male.

Phtisica marina Slabber, 1769

Phtisica marina Slabber, 1769: 77, pl. 10; Chevreux and Fage, 1925: 434, ® gure 422; Fiorencis,1940: 11, ® gure 1, pl. 1, ® gures 1± 2; Costa, 1960: 99; McCain, 1968: 91, ® gures 46± 47;McCain and Steinberg, 1970: 65.

Proto brunneovittata: Heller, 1879: 231; Haller, 1880: 339, pl. 22, ® gures 19± 22.Proto pedata: Haller, 1879: 230; Haller, 1880: 398.Proto ventricosa: Mayer, 1882: 22, pl. 1, ® gure 1; pl. 3, ® gures 16± 29; pl. 4: ® gures 12± 13; pl.

5: ® gures 1 ± 5; Mayer, 1890: 12, pl.3, ® gures 4± 6; pl. 5, ® gures 3± 6; pl. 6, ® gure 1; pl. 7,® gure 1; Mayer, 1903: 20, pl. 6, ® gure 23; Monterosso, 1915: 3.

Material examined. 1A: nine males, one female; 1C: two males, ® ve females; 2A:

two males, one female, one juvenile; 2B: ® ve males, seven females, two ovigerous

females, three juveniles; 3A: ® ve males, three females, one ovigerous female, seven

juveniles; 5A: two males, one female, one ovigerous female; 7A: two males; 7B: one

male, two females; 8A: one ovigerous female; 9A: four males, three females, oneovigerous female, two juveniles; 10A: one male; 11A: ten males, four females, three

juveniles; 12A: one male, two ovigerous females, one juvenile; 12B: four males; 13A:

one juvenile; 14A: two males, one female; 15A: one male; 15B: four males, one

female, one ovigerous female, one juvenile; 15D: two males, one female, three

juveniles; 16A: one male, one female, one juvenile; 17B: ® ve males, one female; 17C:ten males, four females, four ovigerous females, four juveniles; 18C: two males, one

female; 19A: two males, one female, one juvenile; 20A: two males, two females; 21A:

two males; 22A: ® ve males, three females, one ovigerous female, one juvenile; IV:

one male; VI: three males, three females, three juveniles; VII: four males, one female.

Pseudoprotella inermis Chevreux, 1927

Pseudoprotella inermis Chevreux, 1927: 135, pl. 14, ® gures 3± 5; McCain and Steinberg, 1970:72; Guerra-Garcõ Â a and Takeuchi, 2000: 980 ± 988, ® gures 2± 5.

Material examined. 15D: seven males, two females, four juveniles.

Pseudoprotella phasma (Montagu, 1804)

Cancer Phasma Montagu, 1804: 66, pl. 6, ® gure 3.Caprella quadrispinis: Grube, 1864: 63.Protella phasma: Mayer, 1882: 29, pl. 1, ® gure 2; pl. 4, ® gures 1± 8, ® gures 34 ± 37; pl. 5,

® gures 19± 21.Pseudoprotella phasma: Chevreux and Fage, 1925: 437 ± 439, ® gure 423; Cavedini, 1982: 527.

Material examined. 1A: three males, three females; 1C: one female; 1D: four

males, three females, two ovigerous females, one juvenile; 2B: 26 males, 20 females,seven ovigerous females, 22 juveniles; 5A: eight males, ® ve females, one ovigerous


female, ten juveniles; 7A: one male, one female, one ovigerous female, one juvenile;

7B: two males, one female, one juvenile; 7C: one male, one female, one juvenile;12B: one male, one female, one juvenile; 14A: one ovigerous female; 15B: one male,

one female, one juvenile; 15D: seven males, six females, one juvenile; 17B: three

males, two females, one female carrying juveniles, 17C: nine males, nine females,

two ovigerous females, eight juveniles; 18C: six males, two females; 19D: ® ve males,

® ve females, one ovigerous female, four juveniles; 20A: two males; 21A: one male;

22A: one male; 23A: one male, two females carrying juveniles.

Key to species of Caprella from Ceuta

The key is based primarily on gnathopod 2 of adult males, following Takeuchi

(1995 ).

1 Basis of gnathopod 2 clearly shorter than one-third of pereonite 2 . . . . . 2± Basis of gnathopod 2 longer than one-third of pereonite 2 . . . . . . . 5

2 Propodus of gnathopod 2 profusely setose . . . . . . . . . . . . 3± Propodus of gnathopod 2 scarcely setose . . . . . . . . . . . . 4

3 Propodus of gnathopod 2 entirely covered with ® ne plumose setae. Articles 2 and 3of antenna 1 with dense plumose setae on the posterior margin . . . C. fretensis

± Propodus of gnathopod 2 profusely setose only proximally. Only article 3 of antenna1 with dense setae posteriorly . . . . . . . . C. ceutae n. sp (® gures 2± 5)

4 Head without rostrum. Pereopods without grasping spines . C. danilevskii (® gure 6)± Head with rostrum. Pereopods with grasping spines . . C. liparotensis (® gures 8± 11 )

5 Antennae 2 with short setae posteriorly . . . . . . . . . . . . 6± Antennae 2 with long setae posteriorly . . . . . . . . . . . . . 9

6 Body with dorsal tubercles . . . . . . . . . . . . . C. acanthifera± Body without dorsal tubercles . . . . . . . . . . . . . . . 7

7 Body with a few rounded humps . . . . . . . . . . . C. cavediniae± Body smooth . . . . . . . . . . . . . . . . . . . 8

8 Propodus of gnathopod 2 round with dense, long setae dorsally; basis about half ofpereonite 2. . . . . . . . . . . . . . . . . . . C. hirsuta

± Propodus of gnathopod 2 with sparse, short setae dorsally; basis as long as pereonite 2. . . . . . . . . . . . . . . . . . . . C. grandimana

9 Pereonite 1 shorter than head . . . . . . . . . . . . . . . 10± Pereonite 1 longer than head . . . . . . . . . . . . . . . 11

10 Gnathopod 2 with distal poison tooth . . . . . . . . . . C. dilatata± Gnathopod 2 with proximal poison tooth . . . . . . C. penantis (® gure 12)

11 All the body pereonites with dorsal acute projections . . . C. erethizon (® gure 7)± Body pereonites with tubercles . . . . . . . . . . . . . . . 12

12 Head with a central dorsal projection. Propodus of gnathopod 2 setose on posteriorsurface. Acute ventral projections on pereonites 2± 4 . C. santosrosai (® gures 13± 16)

± Head with a short acute rostrum. Propodus of gnathopod 2 setose on anterior andposterior surface. Acute ventral projections lacking . C. tuberculata (® gures 17 ± 20)

Discussion

Ceuta is located on the North African side of the Strait of Gibraltar. The Strait

of Gibraltar is a very important geographical-geologica l formation formed in the

® nal phases of the Pliocene period. It is the boundary for the Mediterranean region(to the east), the Lusitanian region (to the northeast ) and the Mauretanian region


(to the southeast). The Strait is a zoogeographical area, especially from the point

of view of the interchange of the species between the Atlantic and the Mediterranean,and the European and African faunas. Thus, the Strait has attracted the attention

of marine taxonomists in latter years and several biogeographical studies have been

published (e.g. bryozoans: LoÂ pez de la Cuadra and Garcõ Â a-GoÂ mez, 1995; sponges:

Carballo et al., 1997; ascidians: Naranjo et al., 1998; molluscs: Gofas, 1998).

A recent cladistic analysis by Bellan-Santini and RuŒo (1998) indicates that the

Mediterranean and Atlantic fauna of amphipods are very close and that a largeproportion of Mediterranean amphipods could be of Atlantic origin. Recently, the

benthic Gammaridea fauna of Algeciras Bay (Iberian side of the Strait of Gibraltar)

has been studied (Conradi and LoÂ pez-GonzaÂ lez, 1999). In this case, all species

collected, except Urothoe hesperiae which was described as a new species (Conradi

et al. 1995), had been found in Mediterranean waters in previous studies. Thus,

Algeciras Bay can be considered a typical Mediterranean locality despite beingsituated within the Strait of Gibraltar.

In the Mediterranean Sea, 33 species of caprellids have been recorded so far

including the present study (table 2, ® gure 21). Of these species, 13 (39.4%) have

been collected only in Mediterranean waters and they can be considered as

Mediterranean endemic. On the coast of Ceuta, which is only about 20 kilometreslong we have collected 19 of these 33 species; four of them (Caprella ceutae, Caprella

santosrosai, Pedoculina garciagomezi and Pseudoprotella inermis) are, so far, endemic

to the Strait of Gibraltar. Thus, 30.8% of Mediterranean endemic species are endemic

to the Strait of Gibraltar. Caprella fretensis, C. erethizon and C. tuberculata had

previously been collected only on Atlantic coasts (Chevreux and Fage, 1925,

Harrison, 1944, Marques and Bellan Santini, 1985). With the present work thedistributions of these species are extended to Mediterranean waters. The species

Caprella santosrosai and Pedoculina garciagomezi are recorded for the ® rst time on

the coast of north Africa, demostrating that these species inhabit both sides of the

Strait of Gibraltar.

The percentage of caprellid endemism in the Mediterranean (39.4%) is only a

little higher than the 37% reported by Bellan-Santini and RuŒo (1998) for caprellidsand gammarids combined. Both values are higher than the 26.6% calculated by

Fredj et al. (1992) for all of the Mediterranean fauna taken together. As indicated

by Bellan-Santini and RuŒo (1998), further researches along the Atlantic coast of

North Africa and the Iberian Peninsula would likely reduce the number of amphipods

that are considered endemic to the Mediterranean. For example, Caprella hirsutaand C. grandimana, considered as Mediterranean endemics in Krapp-Schickel (1993),

have been recently reported on the Atlantic african coasts from Cape Spartel to

Cape Blanc (Bellan-Santini and RuŒo, 1998). A similar change in biogeographic

distribution was found for Caprella rapax Mayer, 1890, which was collected recently

on the Atlantic Iberian coast (Bellan-Santini and RuŒo, 1998). Although Parvipalpus

major Carausu, 1941 was considered as an endemic species by Bellan-Santini andRuŒo (1998), specimens of this species were collected in the Bay of Biscay (Laubitz

and Sorbe, 1996) so this species also extends to Atlantic waters and is not a

Mediterranean endemic.

Nevertheless, more extensive researches are necessary to expand our understand-

ing of the biogeography of the caprellid fauna of African coasts, not only northern

Africa but also Atlantic coasts of Africa, and especially the African coasts of theIndian Ocean. The caprellid fauna on the Indian coast has a special taxonomic


Table 2. Caprellids cited in Mediterranean waters.

Name of species ME SGE Presence on North African coast

Caprella acanthifera* Algeria: Cherchell, Bou Ismail (Bakalem andDauvin, 1995); Morocco (Bellan-Santini andRuŒo, 1998); Ceuta (Present study)

Caprella andreae Algeria (McCain, 1968)Caprella cavediniae* Ceuta (Present study) (see Krapp-Schickel and

Vader, 1998)Caprella ceutae, n. sp.* 1 1 Ceuta (present study)Caprella danilevskii* Algeria: Alger, Cherchell (Chevreux and Fage,

1925); Algeria: Alger, Bou Ismail (Bakalemand Dauvin, 1995); Morocco (Bellan-Santiniand RuŒo, 1998), Ceuta (present study)

Caprella dilatata* Algeria (McCain and Steinberg, 1970),Morocco (Bellan-Santini and RuŒo, 1998),Ceuta (present study)

Caprella equilibra Egypt: Port Said (Schellenberg, 1928); Algeria:Cherchell, Bou Ismail (Bakalem and Dauvin,1995 )

Caprella erethizon* Ceuta (present study)Caprella fretensis* Ceuta (present study)Caprella grandimana* Morocco (Bellan-Santini and RuŒo, 1998);

Ceuta (present study)Caprella hirsuta* Tunisia: La Galite; Algeria: Cherchell

(Chevreux and Fage, 1925); Morocco (Bellan-Santini and RuŒo, 1998); Ceuta (presentstudy)

Caprella lilliput 1 Not collected on North African coastCaprella linearis Algeria: Bou Ismail (Bakalem and Dauvin,

1995 )Caprella liparotensis* Egypt: Abu Qir (Schellenberg, 1936); Algeria:

Cherchell (Chevreux and Fage, 1925);Morocco (Bellan-Santini and RuŒo, 1998);Ceuta (present study)

Caprella mitis 1 Algeria (Bakelem and Dauvin, 1995)Caprella penantis* Egypt: Abu Qir (Schellenberg, 1936); Algeria:

Alger (Mayer, 1890); Morocco (Bellan-Santiniand RuŒo, 1998); Ceuta (present study)

Caprella rapax Not collected at North African coastCaprella santosrosai* 1 1 Ceuta (present study)Caprella sp (armata-group)* Ceuta (present study)Caprella telarpax 1 Not collected at North African coastCaprella tuberculata* Ceuta (present study)Deutella shieckei 1 Not collected at North African coastLiropus elongatus 1 Algeria: Alger (Bakalem and Dauvin, 1995)Liropus minimus 1 Not collected at North African coastPariambus typicus* Algeria: Alger (Chevreux and Fage, 1925);

Algeria: Bou Ismail, Alger (Bakalem andDauvin, 1995); Morocco (Bellan-Santini andRuŒo, 1998); Ceuta (present study)

Parvipalpus linea 1 Not collected at North African coastParvipalpus major Not collected at North African coastPedoculina bacescui 1 Not collected at North African coastPedoculina garciagomezi* 1 1 Ceuta (present study)Phtisica marina* Algeria: Bou Ismail, Alger, Bejaia (Bakalem

and Dauvin, 1995); Ceuta (present study)


Table 2. (Continued ).

Name of species ME SGE Presence on North African coast

Pseudolirius kroyeri 1 Algeria: Bou Ismail, Alger, Bejaia, Annaba(Bakalem and Dauvin, 1995)

Pseudoprotella inermis* 1 1 Spain: CaÂ diz (Chevreux, 1927); Ceuta (Guerra-Garcõ Â a and Takeuchi, 2000; present study)

Pseudoprotella phasma* Egypt: Abu Qir (Schellenberg, 1936); Algeria:Bou Ismail (Bakalem and Dauvin, 1995);Morocco (Bellan-Santini and RuŒo, 1998);Ceuta (present study)

Sources: Krapp-Schickel (1993) for all species except Caprella cavediniae (Krapp-Schickeland Vader, 1998), Caprella linearis (Bakalem and Dauvin, 1995), Caprella santosrosai(SaÂ nchez-Moyano et al., 1995b ), Pedoculina garciagomezi (SaÂ nchez-Moyano et al., 1995a),Pseudoprotella inermis (Chevreux, 1927; Guerra-Garcõ Â a and Takeuchi, 2000), Caprella ceutae,n. sp. (present study) and the atlantic species C. erethizon, C. fretensis and C. tuberculata(present study).

ME: Mediterranean endemic, SGE: Strait of Gibraltar endemic. Presence on NorthAfrican coast is indicated. The asterisk (*) after species names indicates that specimensof these species have been found on the coast of Ceuta. More detailed information aboutbiogeographical distribution of the species is included in Bellan-Santini and RuŒo (1998).

Fig. 21. Number of species of the Caprellidea in North Atlantic and Mediterranean areas.

interest since it is situated between distribution centres of the Atlantic and Paci® c

(Takeuchi, 1993).

Among the species of Caprella from Ceuta we can diŒerentiate a group of speciesby the short length of the gnathopod 2 basis, i.e., C. danilevskii, C. liparotensis,

C. fretensis and C. ceutae. Clinging behaviour has been studied in C. danilevskii by

Takeuchi and Hirano (1995), who report that the short basis in this species allows

the parallel posture to be adopted more easily.

The species with a longer gnathopod 2 basis can be separated into two subgroups.

One is composed of species which have no long setae on antenna 2, i.e. C. acanthifera,C. cavediniae, C. hirsuta and C. grandimana. The other two species, Caprella penantis


and C. dilatata, develop a setose ventral surface of antenna 2. These two species can

be easily distinguished from the remaining species of Caprella from Ceuta in thatthey have a very short pereonite 1 of the cephalon. Caprella penantis has been

recorded under several species or subspecies names from the temperate regions of

the world (McCain and Steinberg, 1970) and there is need of further studies to

determine its nomenclatural status at each locality (McCain, 1968; Laubitz, 1972;

Takeuchi and Hirano, 1995). As well as C. danilevskii, C. penantis also displays the

`parallel posture’ (Takeuchi and Hirano, 1995). The remaining species of Caprellafrom Ceuta, C. erethizon, C. tuberculata and C. santosrosai can be easily diŒerentiated

by ornamental characteristics (tubercles and acute projections) along their bodies.

These three species seem to be very close, sharing morphological characteristics.

Probably C. santosrosai known only in the Strait of Gibraltar so far, will soon be

found on Atlantic coasts living together with C. erethizon and C. tuberculata. We

also consider it possible that this species could be the Mediterranean vicariant ofthe Atlantic species Caprella septentrionalis KroÈ yer.

Caprella tuberculata, C. erethizon, C. santosrosai and C. ceutae n. sp. have been

found together on similar substrata, and at localities on the littoral of Ceuta with

similar ecological characteristics. How can they coexist and avoid competition?

Perhaps they have diŒerent life-cycles through the year. It would be interesting toundertake rearing experiments in the laboratory with these species using Takeuchi’s

methodology (Takeuchi and Hirano, 1991, 1992) and to study the clinging behaviour

on the substratum. The short gnathopod basis in C. ceutae could enable it to adopt

a more parallel posture, categorized by Takeuchi and Hirano (1995). These four

species have, until now, a restricted distribution. Caprella ceutae could have evolved

from the ancestor of the cosmopolitan C. equilibra, adapting to particular ecologicalcharacteristics present in the Strait of Gibraltar.

AcknowledgementsWe express our thanks to CompanÄ õ Â a del Mar and Club Calypso for assistance

in the ® eld. The senior author (JMGG) thanks Dr. JoseÂ Carlos Garcõ Â a-GoÂ mez,

director of the research project `Biodiversidad y Medio Ambiente Ceutõ Â ’ for enabling

the study in Ceuta and Asamblea de Ceuta and a grant, `Programa de FormacioÂ n

de Profesorado Universitario y Personal Investigador’ , from the Ministry ofEducation and Culture of Spain for ® nancial support. The work was completed at

the Otsuchi Marine Research Center. The work was partially supported by the

Cooperative International Research on Marine and Coastal Environment by the

United Nations University, Ocean Research Institute, The University of Tokyo and

Iwate Prefecture Government.

References

Bakalem, A. and Dauvin, J. C., 1995, Inventaire des crustaceÂ s Amphiphodes (Gammaridea,Caprellideaa, Hyperiidea) des coÃ tes d’AlgeÂ rie: essai de syntheÁ se, MeÂsogeÂ e (Bulletin duMuseum d’Histoire Naturelle de Marseille), 54, 49± 61.

Bate, C. S., 1856, On the British Edriophthalma. Part I. The Amphipoda, Report of theBritish Association for the Advancement of Science, 25, 18± 62.

Bate, C. S., 1862, Catalogue of the Specimens of Amphipodou s Crustacea in the Collection ofthe British Museum (London: British Museum), 399 pp.

Bate, C. S. and Westwood, J. O., 1868, Caprellidae, A History of the British Sessile-eyedCrustacea, 2, 68± 70.

Bellan-Santini, D. and Ruffo, S., 1998, Faunistics and Zoogeography, in S. RuŒo (ed.)


The Amphipoda of the Mediterranean, MeÂmoires de l’Institut Oceanographique , Monaco,13 (4), 895± 911.

Bertrand, H., 1941, Les CrustaceÂ s MalacostraceÂ s de la reÂ gion Dinardaise, Bulletin duLaboratoire Maritime de Dinard, 23, 20± 23.

Bocquet, C. and Peltier, A. M., 1963, Redescription de l’Amphipode Caprella erethizonMayer, Bulletin de la SocieÂ teÂ LinneÂ de la Normandie, 10 (4), 152± 165.

Carausu, A. and Carausu, S., 1959, Contribution aÁ l’eÂ tude des Caprellidae (CrustaceÂ sAmphipodes aberrants) de la mer Noire. Caprella acanthifera Leach (morphologie,eÂ cologie, variabiliteÂ ), in Lucrarile sesiunii stiintiWce (15± 27 septembrie 1956) a Statiuniizoologice marine `Prof. Ioan Borcea’ , Agigea, 353± 420.

Carballo, J. L., Naranjo, S. and GarciÁ a-GoÁ mez, J. C., 1997, Where does the MediterraneanSea begin? Zoogeographical a� nities of the littoral sponges of the Straits of Gibraltar,Journal of Biogeography, 24, 223± 232.

Cavedini, P., 1982, Contributo alla conoscenza dei Caprellidi del Mediterraneo (Crustacea,Amphipoda), Bollettino del Museo civico di storia naturale di Verona, 8, 493± 531.

Chevreux, E., 1888, Sur quelques CrustaceÂ s Amphipodes recueillis aux environs de Cherchell,Association francË aise pour l’avancement des sciences, 17 (2), 343 ± 353

Chevreux, E., 1910, Note sur les CrustaceÂ s Amphipodes d’AlgeÂ rie et de Tunisie, Bulletin dela SocieÂ teÂ d’histoire naturelle de l’Afrique du Nord, 2(9), 135± 137.

Chevreux, E., 1911, Campagnes de la Melita. Les Amphipods d’AlgeÂ rie et de Tunisie,MeÂmoires de la SocieÂ teÂ zoologique de France, 23 (3± 4), 145 ± 285.

Chevreux, E., 1913, Sur quelques inteÂ ressantes espeÁ ces d’Amphipodes provenant des paragesde Monaco et des peÃ ches peÂ lagiques de la Princesse-Alice et de l’Hirondelle II enMeÂ diterraneÂ e, Bulletin de l’Institut oceÂ anographique , Monaco, 262, 26 pp.

Chevreux, E., 1927, CrustaceÂ s Amphipodes. ExpeÂditions ScientiWques du `Travailleur’ et de`Talisman’ pendant les anneÂ s 1880, 1881, 1882, 1883, 9 MalacostraceÂ s (suite), 3, 41± 152.

Chevreux, E. and Fage, L., 1925, Amphipodes, Faune de France, 9, 1± 488.Condradi, M. and LoÁ pez-GonzaÁ lez, P. J., 1999, The benthic Gammaridea (Crustacea,

Amphipoda) fauna of Algeciras Bay (Strait of Gibraltar): distributional ecology andsome biogeographical considerations, Helgolander Marine Research, 53, 2± 8.

Conradi, M., LoÁ pez-GonzaÁ lez, P. J. and Bellan-Santini, D., 1995, A new species ofUrothoe (Amphipoda, Gammaridea) from the Iberian peninsula, Cahiers de BiologieMarine, 36, 9± 13.

Conradi, M., LoÁ pez-GonzaÁ lez, P. J. and GarciÁ a-GoÁ mez, J. C., 1997, The AmphipodCommunity as a bioindicator in Algeciras Bay (Southern Iberian Peninsula) based ona spatio-temporal distribution, P.S.Z.N.I: Marine Ecology, 18(2), 97 ± 111.

Costa, S., 1960, Premier apercË u sur la reÂ partition des Caprelles dans la reÂ gion de Villefranche-sur-Mer, Recueil des travaux de la Station marine d’Endoume, 33, 99± 101.

Czerniaski, V., 1868, Materialia ad zoographiam Ponticam comparatam, Travaux de laSocieÂ teÂ des naturalistes de St. PeÂ tersbourg, 1, 19± 136.

Fiorencis, A., 1940, I Caprellidi del mare Adriatico presso Rovigno, Thalassia, 4, 34 pp.Fischetti, E., 1937, Cenobiosi della scogliera di S. Giuliano-Boccadasse, con speciale riguardo

agli An® podi, Bollettino dei Musei e laboratori di zoologia e anatomia comparata dellaR. UniversitaÁ di Genova, 17, 17 pp.

Fredj, G., Bellan-Santini, D. and Meinardi, M., 1992, EÂ tat des connaissances sur la faunemarine meÂ diterraneÂ enne, Bulletin de L’Institut occeÂ anographique , Monaco, 9, 133± 145.

Gofas, S., 1998, Marine molluscs with a very restricted range in the Strait of Gibraltar,Diversity and Distributions, 4, 255± 266.

Goodsir, H. D. S., 1842, On a new genus and six new species of Crustacea, Edinburgh NewPhilosophy Journal, 33, 188± 189.

Grube, A. E., 1864, UÈ ber die Crustaceenfauna des Adriatischen und Mittelmeeres, Jahres-Bericht der Schelesischen Gesellschaft fuÈ r vaterlaÈ ndische Cultur, 59± 64.

GueÁ rin, M. F. E., 1836, CrustaceÂ s, Iconographie du ReÁgne Animal de G. Cuvier, 2, 24± 25.Guerra-GarciÁ a, J. M. and Takeuchi, I., 2000. Redescription of Pseudoprotella inermis

Chevreux, 1927, a rare species of caprellidean amphipod (Crustacea) from Ceuta, NorthAfrica, Proceeding of the Biological Society of Washington, 113(4), 980 ± 988.

Haller, G., 1879, VorlaÈ u® ge Notizen uÈ ber die Systematik der im Mittelmeer vorkommendenCaprelliden, Zoologischer Anzeiger, 2(27), 230 ± 233.


Haller, G., 1880, Miscellanea arthropodologica. I. Beschreibung zweier neuen Caprellen,Zeischrift fuÈ r die gesammten Naturwissenschaften, 6, 742± 749.

Harrison, R. J., 1944, Synopsis of the British Fauna. 2. Caprellidea (Amphipoda, Crustacea),The Linnean Society of London, Synopses of the British Fauna, 155, 1± 27.

Heller, C., 1866, BeitraÈ ge zur naÈ heren Kenntniss der Amphipoden des Adriatischen Meeres,Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissen-schaftliche Klasse, 26, 1 ± 62.

Jimeno, A., 1993. ContribucioÂ n al estudio de los anfõ Âpodos de las costas mediterraÂ neas catalanas.Estudio faunõ Âstico, ecoloÂ gico, bioloÂ gico y biogeograÂWco, unpublished phD thesis,Universidad de Barcelona, Spain, 573 pp.

Krapp-Schickel, T., 1993, Suborder Caprellidea, in S. RuŒo (ed.). The Amphipoda of theMediterranean, MeÂmoires de lIÂ nstitute Oceanographique , Monaco, 13 (3), 773± 809.

Krapp-Schickel, T. and Vader, W., 1998, What is, and what is not, Caprella acanthiferaLeach, 1814 (Amphipoda, Caprellidea)? Part 1: the acanthifera-group, Journal ofNatural History, 32, 949± 967.

Kroyer, H., 1843, Beskrivelse af nogle arter og slaegter af Caprellina: med indledendebemaerkninger om Laemodipoda og deres plads i systemet, Naturhistorisk tidsskrift,4, 585.

Kroyer, H., 1844. Karcinologiske bidrag, Naturhistorisk tidsskrift, 2, 290.Larsen, K., 1998, Caprellidea (Crustacea; Amphipoda) from Faroe Islands waters, with a

key to the North-Atlantic species, Faroes Journal of Arts and Sciences, 46, 81 ± 90.Laubitz, D. R., 1972, The Caprellidae (Crustacea, Amphipoda) of Atlantic and Artic Canada,

National Museums of Canada Publications in Biological Oceanography, 4, 1 ± 82.Laubitz, D. R. and Sorbe, J. C., 1996, Deep-water Caprellids (Amphipoda: Caprellidea)

from the Bay of Biscay: A new species and a new locality record, Journal of CrustaceanBiology, 16 (3): 626 ± 632.

Leach, W. E, 1814, Article Crustaceology, in The Edinburgh Encyclopaedia, 7, 429± 437LoÁ pez de la Cuadra, C. M. and GarciÁ a-GoÁ mez, J. C., 1995, Zoogeographical study of the

Cheilostomatida from the Strait of Gibraltar, Biology and Paleobiology of Bryozoans,107± 112.

Lucas, H., 1849, Exploration scientiWque de l’AlgeÂ rie pendant les anneÂ es 1840, 1841, 1842.Sciences physiques, zoologie. Histoire naturelle des animaux articuleÂ s (Paris: Imprimerienationale), pp. 51± 57.

Marques, J. C. and Bellan-Santini, D., 1985, Contribution aÁ l’eÂ tude systeÂ matique et eÂ colo-gique des Amphipodes (Crustacea-Amphipoda) des coÃ tes du Portugal. Premierinventaire des espeÁ ces (Gammariens et Caprelliens), CieÂncia BioloÂ gica, 5, 299± 353.

Mayer, P., 1882, Die Caprelliden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte, Fauna und Flora des Golfes von Neapel, 6, 1± 201.

Mayer, P., 1890, Die Caprelliden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte, Fauna und Flora des Golfes von Neapel, 17, 1± 55.

Mayer, P., 1901, Description d’une nouvelle espeÁ ce de CrustaceÂ Amphipode de la famille desCaprellideÂ s (Caprella erethizon), in H. Gadeau de Kerville, 1900 Recherches sur lesFaunes marine et maritime de la Normandie; 3e voyage, reÂ gion d’Omonville-la Rogue(Manche) et Fosse de la Hague, Juin± Juillet 1889, Extrait du Bulletin de la SocieteÂ Amisdes Sciences Naturelles de Rouen, 239± 245.

Mayer, P., 1903, Die Caprelliden der Siboga-Expedition, Siboga Expeditie, 34, 1± 160.McCain, J. C., 1968, The Caprellidea (Crustacea: Amphipoda) of the Western North Atlantic,

Bulletin of the United States National Museum, 278 (I± IV ), 1 ± 116.McCain, J. C. and Steinberg, J. E., 1970, Amphipoda-I, Caprellidea-I, Crustaceorum

Catalogus , 2, 1± 78.Montagu, G., 1804, Description of several marine animals found on the south coast of

Devonshire, Transactions of the Linnean Society of London, 7, 61 ± 85.Monteroso, B., 1915, Caprellidae del golfo di Catania, Atti della Accadema Gioenia di scienze

naturali in Catania, 8(24), 16 pp.Naranjo, S., Carballo, J. L. and GarciÁ a-GoÁ mez, J. C., 1998, Towards a knowledge of

marine boundaries using ascidians as indicators: characterising transition zones forspecies distribution along Atlantic-Mediterranean shores, Biological Journal of theLinnean Society, 64, 151± 177.


Nicolaidou, A. and Karakiri, M., 1989, The distribution of Amphipoda in a brackish-waterlagoon in Greece, P.S.Z.N.I: Marine Ecology, 10(2), 131 ± 139.

Procaccini, G. and Scipione, M. B., 1992, Observations on the Spatio-Tamporal Distributionof crustacean Amphipods in the Fusaro coastal lagoon (central Tyrrhenian Sea, Italy)and some notes on their presence in Mediterranean lagoons, P.S.Z.N.I: Marine Ecology,13 (3), 203± 224.

SaÁ nchez-Moyana, J. E., Carballo, J. L., and Estacio, F. J., 1995a, Pedoculina garciagomezi(Amphipoda: Caprellidea), new species from Bahõ Â a de Algeciras (Southern Spain),Crustaceana, 68, 418± 427.

SaÁ nchez-Moyano, J. E., JimeÁ nez-MartiÁ n, J. A. and GarciÁ a-GoÁ mez, J. C., 1995b, Caprellasantosrosai n. sp. (Amphipoda: Caprellidea) from the Strait of Gibraltar (SouthernSpain), Ophelia, 43 (3), 197 ± 204.

Say, T., 1818, An account of the Crustacea of the United States, Journal of the Academy ofNatural Sciences of Philadelphia, 1(2), 374 ± 401.

Schellenberg, A., 1928, Report on the Amphipoda, Transactions of the Zoological Societyof London, 22, 633± 692.

Schellenberg, A., 1936, The ® shery grounds near Alexandria. 10. Amphipoda benthonica,FisheriesRresearch directorate, 18, 1± 27.

Schellenberg, A., 1942, Krebstiere oder Crustacea. IV: Flohkrebse oder Amphipoda, DieTierwelt Deutschlands und der angrenzenden Meeresteile, 40, 1± 204.

Slabber, M., 1769, Natuurkundige Verlustigingen, behelzende microscopise Waarneemingen vanin- en uitlandse water- en land-Dieren (Te Haarlem: J. Bosh), 166 pp.

Smaldon, S., 1990, Suborder Caprellidea, in P. J. Hayward and J. S. Ryland (eds) The MarineFauna of the British Isles and North-West Europe. Vol. 1. Introduction and Protozoansto Arthropods (Oxford: Clarendon Press), pp. 482± 488.

Stebbing, T. R. R., 1878, Notes on sessile-eyed crustaceans, with description of a new species,Annals and Magazine of Natural History, Series 5, 1, 31 ± 37.

Stebbing, T. R. R., 1879, Sessile-eyed Crustacea of Devonshire. Supplementary list, Reporton the Transactions of the Devonshire Association, 11, 516± 524.

Stebbing, T. R. R., 1888, Report on the Amphipoda collected b H.M.S. Challenger duringthe years 1873 ± 76, Report on the ScientiWc Results of the Voyage of H.H.S. ChallengerDuring the Years 1873 ± 76, Zoology , 29, 1737 pp.

Stephensen, K., 1929, Amphipoda, in G. Grimper and E. Wagler (eds) Die Tierwelt der Nord-und Ostsee, Leipzig, 188 pp.

Takeuchi, I., 1993, Is the Caprellidea a monophyletic group?, Journal of Natural History,27, 947± 964.

Takeuchi, I., 1995, Suborder Caprellidea, in S. Nishimura (ed.), Guide to Seashore Animalsof Japan with Color Pictures and Keys, Vol. 2 (Osaka: Hoikusha), pp. 193± 206.

Takeuchi, I. and Hirano, R., 1991, Growth and reproduction of Caprella danilevskii(Crustacea: Amphipoda) reared in the laboratory, Marine Biology, 110, 391± 397.

Takeuchi, I. and Hirano, R., 1992, Growth and reproduction of the epifaunal amphipodCaprella okadai Arimoto (Crustacea: Amphipoda: Caprellidea), Journal of ExperimentalMarine Biology and Ecology, 161, 201± 212.

Takeuchi, I. and Hirano, R., 1995, Clinging behaviour of the epifaunal caprellids(Amphipoda) inhabiting the Sargassum zone on the Paci® c Coast of Japan, with itsevolutionary implications, Journal of Crustacean Biology, 15 (3), 481± 492.

Touldmond, A. and Truchot, J. P., 1964, Inventaire de la faune marine de RoscoŒ;Amphipodes, Supplementaux Travaux de la Station Biologique de RoscoŒ, 1± 42.

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