revision of the genus deutella (crustacea: amphipoda

J N H, 2003, 37, 1059–1084

Revision of the genus Deutella (Crustacea: Amphipoda: Caprellidea)with description of a new species, redescription of Deutella venenosaMayer, 1890 and a key to the species of Deutella

J. M. GUERRA-GARCIA

Laboratorio de Biologıa Marina, Departamento de Fisiologıa y BiologıaAnimal, Facultad de Biologıa, Universidad de Sevilla, Apdo. 1095,E-41080 Sevilla, Spain; e-mail: [email protected]

(Accepted 4 October 2001)

The genus Deutella is reviewed and a key to the species is given. A new speciesof Deutella is described from Isla Margarita, Venezuela. Full description andillustrations are given for the adult male and female. Diagnostic features of thisspecies are: body dorsally smooth, flagellum of antenna 1 with 14 articles inmales and 10 in females, setal formula of the mandibular palp 1-9-1, molar flakepresent, propodus palm of gnathopod 1 with a pair of grasping spines, maleabdominal appendages two-articulate. Deutella venenosa Mayer, 1890 is rede-scribed based on the material newly collected from Coquimbo, Chile. This speciesis recorded for the first time since the original description by Mayer (1890). Thespecies Luconacia vemae McCain and Gray, 1971 is transferred to the genusDeutella.

K: Crustacea, Amphipoda, Caprellidea, Deutella, taxonomy.

IntroductionThe genus Deutella was established by Mayer (1890). So far, six species of this

genus have been described: Deutella aspiducha Gable and Lazo-Wasem, 1987, D.californica Mayer, 1890, D. incerta (Mayer, 1903), D. mayeri Stebbing, 1895, D.schieckei Cavedini, 1981 and D. venenosa Mayer, 1890. In the present paper, a newspecies Deutella margaritae is described and Luconacia vemae McCain and Gray,1971 is transferred to the genus Deutella. Consequently, the genus Deutella ispresently composed of eight species, which are reviewed and illustrated in this paper.

During a short stay at the Universidad Catolica del Norte, Coquimbo, Chile,the author collected abundant material of Deutella venenosa Mayer, 1890, the firstrecord for the species since the original description. The species is fully redescribedand illustrated in the present paper. As a result of consulting material of the genusDeutella from the collections of the National Museum of Natural History,Smithsonian Institution, Washington, DC, an undescribed species of Deutella col-lected from Isla Margarita, Venezuela in 1964 was found. Detailed examination

Journal of Natural HistoryISSN 0022-2933 print/ISSN 1464-5262 online © 2003 Taylor & Francis Ltd

http://www.tandf.co.uk/journalsDOI: 10.1080/00222930110104267

J. M. Guerra-Garcıa1060

revealed that these specimens of Deutella belonged to a new species, described asDeutella margaritae in the present paper.

Figures of all Deutella species and a key for easy identification have also beenincluded based on careful examination of type and non-type material from Museums.The location of the specimens of Deutella examined is indicated by the followingabbreviations: AMNH, American Museum of Natural History, New York, USA;CMNC, Canadian Museum of Nature, Canada; MCSN, Museo Civico di StoriaNaturale di Verona, Italy; MCZ, Museum of Comparative Zoology, HarvardUniversity, MA, USA; MNHN, Museo Nacional de Historia Natural de Santiago,Chile; USNM, National Museum of Natural History, Smithsonian Institution,Washington, DC, USA.

Systematics

Family PARIAMBIDAE Laubitz, 1993

Diagnosis. Antenna 2 flagellum two-articulate. Mandible molar present, palpthree-articulate; setal formula 1-x-1 or one apical setae; incisor five-toothed; leftlacinia mobilis five-toothed; right lacinia serrate or broadly three-toothed. Lowerlip with inner lobes well demarcated. Maxilla 1 with six spiniform setae on outerplate. Maxilliped outer plate larger than inner; inner plate truncate with few apicalsetae. Gnathopod 1 propodus triangular to slender. Gills on pereonite 3 and 4.Pereopod 3 and 4 greatly reduced; pereopod 5 weaker than 6 and 7.

Genus Deutella Mayer, 1890

Diagnosis. Flagellum of antenna 2 with two articles, swimming setae absent.Mandibular palp three-articulate; setal formula for distal article 1-x-1; molar present.Outer lobe of maxilliped longer than inner lobe. Pereopod 5 with six articles. Maleabdomen with pair of appendages and pair of setose lobes.

Type species. Deutella californica Mayer, 1890.

Deutella aspiducha Gable and Lazo-Wasem, 1987(figure 1)

Deutella aspiducha Gable and Lazo-Wasem, 1987: 629–639, figures 1–3.

Material examined. Male holotype, USNM 195178, Little River Cave, Bermuda,8 May 1982; female allotype, USNM 195177, Little River Cave, Bermuda, 8 May1982.

Distribution. Only known from the type locality, Bermuda, USA.Ecology. Among hydroids, shallow subtidal.Remarks. Deutella aspiducha was recently described by Gable and Lazo-Wasem

(1987). The most striking differences from all other Deutella species is the size andthe shape of the last two articles (propodus and dactylus) of pereopod 5.Furthermore, this species lacks a triangular projection on the penultimate article ofthe palp of the maxilliped. Nevertheless, Gable and Lazo-Wasem (1987) classifiedthis species as Deutella on the basis of the following characters: structure andproportions of the antennae, most of its mouthpart morphology, the various pereon-ite projections, the delineation of the palm of gnathopod 2 in males, the stronganterior insertion of gnathopod 2 on pereonite 2, the shape, length and insertion

Revision of the genus Deutella 1061

F. 1. Deutella aspiducha Gable and Lazo-Wasem, 1987. Lateral view and detail ofgnathopod 2. (A, C) Male; (B, D) female. Scale bars: (A, B) 1 mm; (C, D) 0.3 mm.


patterns of pereopods 3 and 4, and the structure of the male abdomen. Even thesmallest individuals of D. aspiducha show short dorsal spines on the cephalon andpereonite 2 (Gable and Lazo-Wasem, 1987). This species is, so far, endemic toBermuda.

Deutella californica Mayer, 1890(figure 2)

Deutella californica Mayer, 1890: 27–28, pl. 1, figures 3, 4, pl. 3, figures 15, 16, pl. 5, figure 18;Dougherty and Steinberg, 1953: 44; 1954: 169, 171, figure 82f; Steinberg and Dougherty,1957: 279–281, figures 15, 21–23, 28; McCain, 1968: 54, figure 52; McCain and Steinberg,1970: 48; Laubitz, 1970: 16–18, figures 3, 26.

Material examined. Seven males, three females, one juvenile, syntypes MCZ 1059,Cape Mendocino, California, 5 July 1860; three males, one female (non-types),CMNC 1989-0944, Tillamook, Oregon, 16 August 1966; one male (non-type),USNM 42667 Point Mugu, California, 5 December 1962.

Distribution. Type locality: Cape Mendocino, California. Other localities:Monterey Bay and Mussel Point, California; San Juan area, Washington; Oregon,Washington and Vancouver Island; Juan de Fuca Strait, B.C., southern PrinceWilliam Sound, Alaska.

Ecology. Collected from Obelia dichotoma (Linnaeus, 1758) on caparace andlegs of Loxorhynchus grandis Stimpson, 1857 (ca 10–15 m depth). Also collectedfrom flat fine sand, sandstone and shale bedrock.

Remarks. The shape of the gnathopod 2 propodus in males is the best character-istic to differentiate easily this species among the remaining species of Deutella.Deutella californica was originally described by Mayer from Cape Mendocino,California and later reported from Monterey Bay by Dougherty and Steinberg(1953). Steinberg and Dougherty (1957) reported it as occurring at ‘probably PortArkansas, Texas’. As has been pointed out by McCain (1968) and Laubitz (1970),the Texas locality record for this species is very doubtful; it is probable that Deutellacalifornica is endemic to the Pacific.

Deutella incerta (Mayer, 1903)(figure 3)

Protellopsis stebbingii Kunkel, 1910: 111–113, figure 43.Deutella incerta; Steinberg and Dougherty, 1957: 281, 285–286; Gable and Lazo-Wasem,

1987: 635–636, figure 4.Luconacia incerta; Mayer, 1903: 49–50, pl. 2, figures 11–14; pl. 6, figures 73–75; pl. 9, figures

21, 40, 57; McCain, 1968: 53–54, 68–72, figures 33–35; McCain and Steinberg, 1970: 53.

Material examined. One male, one female (types), USNM 026001, three females(paratypes), USNM 123527, 213–234 m depth, 10 February 1934; one male, onefemale (non-types), USNM 42667, Gulf of Mexico.

Distribution. Type locality: off Mobile Bay, Alabama, 29°24∞N, 88°04W. Otherlocalities: Bermuda, east coast of North America from Woods Hole, Massachusettsto Straits of Florida; Gulf of Mexico from Cedar Keys, Florida to Yucatan; VirginIslands; Barbuda; Barbados; Isla de Margarita, Aruba.

Ecology. This species has been collected on mangrove roots, Sargassum,Thalassia, sponges, hydroids, alcyonarians, ascidians and it has occasionally beentaken in plankton tows.

Remarks. Deutella incerta is widely distributed in the temperate and tropical


F. 2. Deutella californica Mayer, 1890. Lateral view and detail of gnathopod 2. (A, C)Male; (B, D) female. Scale bars: 1 mm.

areas of the western North Atlantic (McCain, 1968) and seems to be one of themost commonly encountered caprellids in Bermuda (Gable and Lazo-Wasem, 1987).Steinberg and Dougherty (1957) synonymized Luconacia of Mayer (1903) withDeutella, transferring Luconacia incerta Mayer, 1903 to the genus Deutella. McCain


F. 3. Deutella incerta (Mayer, 1903). Lateral view and detail of gnathopod 2. (A, C)Male; (B, D) female. Scale bars: 1 mm.

(1968) re-established Luconacia on the basis of the following differences: apical setaeare present on the terminal article of the maxilliped palp and a projection is presenton the penultimate article in Deutella, while both are lacking in Luconacia incerta;the terminal article of the mandibular palp in Deutella has no knobs and in Luconaciaincerta there is a knob; in Deutella the tip of the male abdominal appendage is


without papillae and fringed or not fringed, and in Luconacia incerta both papillaeand fringes are present; the female abdomen in Deutella is lobed and in Luconaciaincerta it is not; pereopod 5 is inserted posteriorly on pereonite 5 in Deutella, butin Luconacia incerta at mid-length. The discovery of Deutella aspiducha by Gableand Lazo-Wasem (1987) provided these authors with a transitional form that allowedthem to re-unite Deutella and Luconacia. Curiously, Gable and Lazo-Wasem (1987)overlooked the existence of the species Luconacia vemae McCain and Gray, 1971.This species had been described in a revision of the Caprellidea from the Antarcticand Subantarctic by McCain and Gray (1971). However, the differences betweenLuconacia vemae and Deutella incerta further support the combination of the twogenera. The abdominal appendages of L. vemae do not bear as complete a fringe ofsmall teeth around the papillae as in D. incerta. Furthermore, the knobs on theterminal article of the mandibular palp, one of the most important characteristicsused by McCain (1968) to differentiate between Deutella (knobs absent) andLuconacia (knobs present), are not present in L. vemae. Therefore, the speciesLuconacia vemae must be transferred, as Luconacia incerta, to the genus Deutellawith the genus Luconacia becoming a synonym.

Deutella margaritae new species(figures 4–7)

Material examined. Holotype male (USNM 1000206), allotype female (USNM1000207) and paratypes (three males, seven females, USNM 1000208), 11 January1964, Venezuela.

Type locality. Puente de la Restinga, Isla Margarita, Venezuela.Etymology. Deutella margaritae is dedicated to Isla Margarita, the beautiful place

where the specimens were collected.Ecology. The specimens were collected in boulders near Rhizophora.

DescriptionHolotype male

Body length. 7.80 mm.Lateral view (figure 4A). Body dorsally smooth. Anterolateral margin of pereon-

ites 2, 3 and 4 with triangular projection. Pereonites 3 and 4 subequal. Pereonite 5the longest.

Gills (figure 4A). Elongate, length about 3.5 times width.Mouthparts. Upper lip (figure 5F) symmetrically bilobed, smooth. Mandibles

(figure 5A, B) with three-articulate palp; distal article of palp with knobs and a setalformula 1-9-1; mandibular molar process strong, bordered by teeth; left mandiblewith incisor five-toothed, lacinia mobilis four-toothed followed by a row of threeserrate spines; incisor of right mandible divided into five teeth, lacinia mobilis deeplyserrate followed by a row of two serrate spines; molar flake present, serrate distallyand carrying minute setulae. Lower lip (figure 5G) with inner lobes well demarcated;outer lobe without setae. Maxilla 1 (figure 5D) outer lobe carrying five spine-likesetae; distal article of the palp with four distal spine-like setae and one seta medially.Maxilla 2 (figure 5C) outer lobe with three setae distally; inner lobe rectangular,two-thirds of outer lobe in length, provided with four setae. Maxilliped (figure 5E)inner plate small and elongate with one tooth, three simple setae distally and aplumose seta laterally; outer plate about 2.5 times as long as inner plate, with four


F. 4. Deutella margaritae new species. Lateral view. (A) Male; (B) female. Scale bar:1 mm.

setae; penultimate article of palp without distal projection; dactylus wide, with rowsof setulae.

Antennae. Antenna 1 (figure 6A) about half of body length; peduncle not setose;flagellum 14-articulate. Antenna 2 (figure 6B) about two-thirds of antenna 1 length;proximal article of peduncle with an acute projection; flagellum two-articulate;peduncle and flagellum scarcely setose.

Gnathopods. Gnathopod 1 (figure 6C) basis as long as ischium to carpus com-bined; propodus elongate with a pair of grasping spines; grasping margin of propoduspalm and dactylus smooth. Gnathopod 2 (figure 6D) inserted on the anterior halfof pereonite 2; basis as long as pereonite 2; ischium rectangular; merus rounded;carpus short and triangular; propodus somewhat longer than basis; palm providedwith a rectangular projection proximally, provided with a small tooth and smalldistal projection; dactylus thickened medially with three fine setae.

Pereopods. Pereopods 3 and 4 (figure 7A, B) with two articles; distal articlecarrying six setae; pereopods 5–7 missing; insertion of pereopod 5 posteriorly inpereonite 5.

Penes (figure 7C, D). Large, length 2.5 times width.


F. 5. Deutella margaritae new species. Male. (A) Left mandible; (B) right mandible;(C) maxilla 2; (D) maxilla 1; (E) maxilliped; (F) upper lip; (G) lower lip. Scalebars: 0.1 mm.


F. 6. Deutella margaritae new species. (A–D) Male. (A) antenna 1; (B) antenna 2; (C)gnathopod 1; (D) gnathopod 2. (E) Female gnathopod 2. Scale bars: (A, B) 0.5 mm;(C) 0.3 mm; (D, E) 0.5 mm.


F. 7. Deutella margaritae new species. (A–D) Male. (A) Pereopod 3; (B) pereopod 4;(C) abdomen (ventral view); (D) abdomen ( lateral view). (E) Female abdomen(ventral view). Scale bars: (A, B) 0.2 mm; (C–E) 0.1 mm.

Abdomen (figure 7C, D). Abdomen with a pair of two-articulate appendages, apair of lateral lobes and a single dorsal lobe. Distal article of appendages minutelytoothed, carrying small setulae distally.

Allotype femaleBody length 6.20 mm. Flagellum of antenna 1 with 10 articles (figure 4B).

Propodus of gnathopod 2 (figure 6E) lacking the rectangular projection present inmales; dactylus not thickened medially, without setae. First pair of oostegites scarcely


setose; second pair smooth (figure 4B). Abdomen (figure 7E) without appendages;lateral lobes not setose and single dorsal lobe provided with two plumose setae.

RemarksDeutella margaritae nov. sp. posses two characteristics unique in the genus

Deutella: male abdominal appendages two-articulate and a left lacinia mobilis four-toothed. In connection with other morphological characters, D. margaritae resemblesD. mayeri and D. schieckei in the absence of dorsal projections. Nevertheless, themandibular palp is very different; there are knobs in D. margaritae and the setalformula is 1-9-1 while D. mayeri and D. schieckei lack knobs and have only one ortwo setae distally on the terminal article of the mandibular palp.

Deutella mayeri Stebbing, 1895(figure 8)

Deutella mayeri Stebbing, 1895: 400–402, pl. 15a; Mayer, 1903: 44–45; McCain, 1968: 54–57,figures 25–26, 52; McCain and Steinberg, 1970: 48.

Material examined. One male, one female (non-types) USNM 225512, 18 mdepth, Gulf of Mexico, 25°45∞58◊N, 82°01∞08◊W.

Distribution. Type locality: Antigua. Other localities: Bahıa Fosforescente,Puerto Rico.

Ecology. UnknownRemarks. Deutella mayeri was first described by Stebbing (1895). McCain (1968)

redescribed and illustrated the species in detail. The gnathopod 2 of males is verycharacteristic. It has a triangular process proximally, provided with a grasping spine.The striking gnathopod can be used to distinguish this species easily from theremaining species of Deutella.

Deutella schieckei Cavedini, 1981(figure 9)

Deutella schieckei Cavedini, 1981: 515, figure 9; Krapp-Schickel, 1993: 794, 796: figure 542.

Material examined. Holotype male, MCSNV 606-607, 2 June 1974, Sardinia,Capo Caccia, Italy.

Distribution. Only known from the type locality, Sardinia, Capo Caccia, Italy.Ecology. Interstitial ( Krapp-Schickel, 1993).Remarks. The presence of the genus Deutella in Mediterranean waters provided

the Mediterranean fauna with a very interesting biogeographical element because ofthe affinity with species from the Caribbean fauna. McCain’s (1968) suggestionthat the removal of D. mayeri from the genus Deutella would make it an exclusivePacific Ocean genus and Luconacia, then, an exclusive Atlantic Ocean genus, losesfoundation for further consideration with Cavedini’s (1981) discovery of theMediterranean D. schieckei (Gable and Lazo-Wasem, 1987).

Deutella vemae (McCain and Gray, 1971)(figure 10)

Luconacia vemae McCain and Gray, 1971: 123, figure 8.

Material examined. Holotype male, AMNH 13132, 81 m depth, 23 May 1961,41°57∞S, 59°03∞W. Paratypes: two males, one female, AMNH 13133 collected together


F. 8. Deutella mayeri Stebbing, 1895. Lateral view and detail of gnathopod 2. (A, C)Male; (B, D) female. Scale bars: (A, B) 1 mm; (C, D) 0.3 mm.


F. 9. Deutella schieckei Cavedini, 1981. Lateral view and detail of gnathopod 2. (A, C)Male; (B, D) female. Scale bars: (A, B) 1 mm; (C, D) 0.3 mm.

with the holotype; one female, USNM 123749, 92–101 m depth, 5 February 1964,52°56∞S, 75°00∞W.

Distribution. Type locality 41°57∞S, 59°03∞N. Other localities: off coast ofArgentina and Southern Chile.

Ecology. Depth range 24–101 m.Remarks (as for Deutella incerta).

Deutella venenosa Mayer, 1890(figures 11–15)

Deutella venenosa Mayer, 1890: 28, pl. 1, figures 5–9; pl. 3, figures 7–14; pl. 5, figures 19–21;pl. 6, figures 11–24.


F. 10. Deutella vemae (McCain and Gray, 1971). Lateral view and detail of gnathopod 2.(A, C) Male; (B, D) female. Scale bars: 1 mm.


F. 11. Deutella venenosa Mayer, 1890. Lateral view. (A) Male; (B) female. Scale bar:1 mm.


F. 12. Deutella venenosa Mayer, 1890. Male. (A) Right mandible; (B) left mandible;(C) maxilliped; (D) upper lip; (E) lower lip; (F) maxilla 1; (G) maxilla 2. Scalebars: 0.1 mm.


F. 13. Deutella venenosa Mayer, 1890. (A–D) Male. (A) Antenna 1; (B) antenna 2; (C)gnathopod 1; (D) gnathopod 2. (E) Female gnathopod 2. Scale bars: (A, D) 1 mm;(B) 0.3 mm; (C) 0.2 mm; (E) 0.5 mm.


F. 14. Deutella venenosa Mayer, 1890. (A–F) Male. (A) Pereopod 3; (B) pereopod 4; (C)pereopod 5; (D) pereopod 6; (E) pereopod 7; (F) abdomen (ventral view). (G) Femaleabdomen (ventral view). Scale bars: (A, B) 0.2 mm; (C–E) 1 mm; (F, G) 0.1 mm.


F. 15. Deutella venenosa Mayer, 1890. Ontogenetic development. (A) Male lateral anddorsal view; (B) male gnathopod 2. Scale bars: (A) 1 mm; (B) 0.3 mm.

Material examined. Two hundred and four males, 73 premature females, 257mature females, 73 juveniles, 0–5 m depth, November to December 2000, Coquimbo,Chile. Male ‘a’, female ‘b’ and several other specimens have been deposited in the


Museo Nacional de Historia Natural de Santiago de Chile, MNHN AMPHNo. 11312.

Distribution. Only known from the type locality, Coquimbo, Chile.Ecology. Attached to buoys of seaweed cultures, mainly on hydroids, and bryo-

zoans such as Bugula neritina (Linnaeus, 1758). Also under rocks in the intertidalzone clinging to the hydroid Obelia dichotoma (Linnaeus, 1758).

RedescriptionMale ‘a’

Colour of live specimen. Light brown with dark spots.Body length. 5.71 mm.Lateral view (figure 11A). Head provided with a dorsal, anteriorly directed acute

projection. Pereonite 1 with a projection medially. Pereonite 2 with three dorsalprojections, one small anteriorly, one acute at the middle and a large rounded oneposteriorly; a pair of lateral acute projections anteriorly directed. Pereonites 3–7dorsally smooth, except pereonites 3 and 4 with humps. Side plates of pereonites 3,4 and 5 projecting ventrolaterally, with margins distinctly crenulate.

Gills (figure 11A). Oval, length about twice width.Mouthparts. Upper lip (figure 12D) symmetrically bilobed, smooth. Mandibles

(figure 12A, B) with three-articulate palp; distal article of the palp with a setalformula 1-x-1x being=4; penultimate article carrying three setae; mandibular molarprocess strong, bordered by robust teeth; left mandible with incisor and laciniamobilis five-toothed, followed by a row of three serrate spines; right mandible withincisor five-dentate and lacinia mobilis deeply serrate followed by a row of twoserrate spines; molar flake present, rectangular and setose distally. Lower lip(figure 12E) with small inner lobes, although well demarcated; outer lobes slender,provided with short setae apically. Maxilla 1 (figure 12F) outer lobe carrying sixspine-like setae; distal article of the palp with five distal robust setae and one strongseta medially. Maxilla 2 (figure 12G) outer lobe oval, with four setae distally; innerlobe rectangular, two-thirds of outer lobe in length, carrying five setae. Maxilliped(figure 12C) inner plate small and elongate with one tooth and three single setae;outer plate about three times as long as inner plate, with five setae; penultimatearticle of the palp with a distal projection; dactylus with setulae distally and with afine seta subdistally.

Antennae. Antenna 1 (figure 13A) about half of body length; peduncle denselysetose; flagellum 11-articulate. Antenna 2 (figure 13B) as long as the first twoarticles of peduncle of antenna 1; proximal article of peduncle provided with anacute projection; the remaining articles of peduncle profusely setose; flagellumtwo-articulate.

Gnathopods. Gnathopod 1 (figure 13C) basis as long as ischium to carpus com-bined; propodus triangular with a proximal grasping spine; grasping margin ofpropodus palm and dactylus minutely serrate. Gnathopod 2 (figure 13D) insertedon the anterior half of pereonite 2; basis as long as pereonite 2; ischium short andrectangular; merus acute ventrally; carpus short and triangular; propodus abouttwice as long as basis, setose dorsally and ventrally; palm provided with a rectangularprojection proximally provided with a small tooth and a distal knotch; dactyluslarge, curved medially, carrying short setae ventrally on the distal half.

Pereopods. Pereopods 3 and 4 consisting of one article. Pereopod 3 (figure 14A)with nine single long setae apically and one short plumose seta subdistally. Pereopod


4 (figure 14B) with seven single setae and one plumose seta. Pereopods 5–7(figure 14C–E) similar, increasing in length, respectively; palm of propodus with aproximal row of three, four and seven short spines on pereopods 5, 6 and 7,respectively; merus to propodus setose; dactylus smooth, about half as long aspropodus.

Penes (figure 14F). Penes triangular, laterally placed.Abdomen (figure 14F). Abdomen with a pair of appendages, a pair of lateral

lobes and a single dorsal lobe. Appendages serrate distally. Lateral lobes very setose.Dorsal lobe carrying a pair of short plumose setae.

Female ‘b’Body length 4.82 mm. Pereonite 1 smooth. The lateral acute projections on

pereonite 2, present in male, are lacking in female (figure 11B). Flagellum of antenna1 with seven articles (figure 11B). Propodus of gnathopod 2 (figure 13E) shorter andwider than in male, without rectangular projection proximally; dorsal surface scarcelysetose; dactylus shorter than in male and not curved medially. First pair of oostegitesscarcely setose; second pair smooth (figure 11B). Abdomen (figure 14G) withoutappendages; a pair of lateral setose lobes and a single dorsal lobe carrying twosimple setae.

Intraspecific variationThe antennae and body projections are very constant in mature specimens. In

connection with the mouthparts, the setal formula of the apical article of mandiblepalp is 1-4-1 or 1-5-1; the presence of six setae on the outer lobe in maxilla 1 andone tooth on the inner plate of maxilliped is very constant. The structure of themandibles is also constant in all specimens; left mandible with five-toothed incisor,five-toothed lacinia and three spines; right mandible with five-toothed incisor, serratelacinia, two spines and molar flake present.

Ontogenetic developmentThe acute projection on the head is already present in the juvenile stages, although

other dorsal and lateral projections appear through development (figure 15).Although the grasping spine on the palm of gnathopod 2 propodus is present in thefirst stages, the rectangular projection becomes more evident in final phases. Withdevelopment, the ratio length/width of the gnathopod 2 propodus increases, thenumber of dorsal and ventral setae of gnathopod 2 also increases and the proximalprojection on the propodus palm becomes constricted, being more elongate near thecarpus; the dactylus increases in length during development and becomes curvedand setose only in the final stages.

RemarksDeutella venenosa Mayer, 1890 was described by Mayer (1890) based on the

specimens collected from Coquimbo, Chile, during the ‘Siboga Expedition’. A cen-tury later, the presence of Deutella venenosa in Chilean waters is confirmed. Thepresent description is in good agreement with the sparse description and figures ofMayer (1890), except for the structure of pereopods 3 and 4 which are two-articulatein specimens described by Mayer (1890) versus one-articulate in the recently col-lected and herein redescribed material. The feature of gnathopod 2 and the body


ornamentation of this species allow Deutella venenosa to be differentiated from theremaining species of Deutella.

Key to the species of DeutellaThe following key is based on examination of type and non-type male specimens.

The species key provided is meant as an easy field guide to be used without dissection.

1 Body with dorsal projections . . . . . . . . . . . . . . . . 2– Body dorsally smooth . . . . . . . . . . . . . . . . . 6

2 Dorsal projections on head to pereonite 4. Pereopods 3 and 4 generally one-articulate. . . . . . . . . . . . . . . Deutella venenosa (figures 11–15)

– Pereonites 3 and 4 without dorsal projections. Pereopods 3 and 4 two-articulate . . 3

3 Dorsal projections only on head . . . . . . . . Deutella vemae (figure 10)– Dorsal projections on head to pereonite 2 . . . . . . . . . . . . 4

4 Pereopod 5 abnormal, dactylus extremely reduced. Antennae densely setose . . . .. . . . . . . . . . . . . . . . . Deutella aspiducha (figure 1)

– Pereopod 5 not reduced. Antennae not setose . . . . . . . . . . . 5

5 Propodus of gnathopod 2 widened distally, profusely setose. Pereopods 3 and 4, one-sixth as long as gills . . . . . . . . . . Deutella californica (figure 2)

– Propodus of gnathopod 2 not widened distally, scarcely setose. Pereopods 3 and 4,one-third as long as gills . . . . . . . . . . Deutella incerta (figure 3)

6 Propodus of gnathopod 2 with a triangular projection proximally . . . . . . .. . . . . . . . . . . . . . . . . Deutella mayeri (figure 8)

– Propodus of gnathopod 2 with a rectangular projection proximally . . . . . 7

7 Body elongate. Antenna 1 scarcely setose. Pereopods 3 and 4 about half as long as gills. . . . . . . . . . . . . . . Deutella margaritae (figures 4–7)

– Body not elongate. Antenna 1 setose. Pereopod 3 and 4 about one-quarter as longas gills . . . . . . . . . . . . . . . Deutella schieckei (figure 9)

DiscussionThe detailed comparison of morphological characters among the species of

Deutella (table 1) supports the combination of the genus Luconacia and Deutella asthe author accepts throughout the present paper. There is no evident pattern whichwould enable us to recognize species groups within the genus since the similarityamong species depends on the selected character.

The genus Deutella is widely distributed along the latitudinal gradient fromAlaska in the Northern Hemisphere to Subantarctic waters in the SouthernHemisphere (figure 16). Nevertheless, this genus is quite restricted along the longitud-inal gradient. Apart from D. schieckei, which reaches Mediterranean waters, theremaining species are limited to American coasts (Pacific or Atlantic). So far, noDeutella species has been found in the Indian Ocean.

In a revision of Caprellid taxonomy, Laubitz (1993) considered 10 families inthe suborder Caprellidea and she transferred the genus Deutella from the familyProtellidae McCain, 1970 to the new family Pariambidae Laubitz, 1993, mainly onthe basis of lacking a molar flake, a different setal formula than 1-x-y-1 and sixinstead of seven spiniform setae on the outer plate of maxilla 1. As Larsen (1997)pointed out, the classification of Laubitz does not agree with the relationshipssuggested by Takeuchi (1993) and are inconsistent, since the genera AbyssicaprellaMcCain, 1966 and Paradicaprella Hirayama, 1990 are not included in the new family

J.M

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uerra

-Garcıa

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Table 1. Detailed comparison among the species of Deutella based on consulting material from Museums, specimens newly collected and literature.

D. aspiducha D. californica D. incerta D. margaritae D. mayeri D. schieckei D. vemae D. venenosa

Body length (mm)Male 4.8 5.5 9 7.8 4 4.5 13.5 5.7Female 4.2 3.8 8 6.2 3 3.5 12 4.8

Dorsal projections Present Present Present Absent Absent Absent Present PresentAntenna 1 flagellar articles

Male 6–7 12 10–11 14 6–7 6–8 11 11Female 4 12 10 10 6–7 6 9 7

Antenna 2 flagellar articles 2 2 2 2 2 2 2 2Mandibles

Knobs on distal article Present Absent Present Present Absent Absent Absent Absentof palp

Setal formula 1-x-1, x=5 1-x-1, x=3–5 1-x-1, x=3–10 1-x-1, x=9 1 or 2 setae 2 setae 1-x-1, x=11 1-x-1, x=4–5Left and right incisor 6-toothed 5-toothed 5-toothed 5-toothed 5-toothed 5-toothed 5-toothed 5-toothedLeft lacinia mobilis Serrate 5-toothed 5-toothed 4-toothed 5-toothed 5-toothed 5-toothed 5-toothedRigth lacinia mobilis Serrate Serrate Serrate Serrate Serrate Serrate SerrateMolar flake Absent Absent Absent Present Absent Absent Absent Present

Maxilla 1Distal spines of outer 5 6 5–6 5 4–5 5 6 6

lobeDistal spines of palp 3 3 4 4 3–4 4 4 5

Maxilla 2Setae of outer lobe 4 4 5–7 3 5–6 4 5 4Setae of inner lobe 3 5 4–5 4 4–5 4 5 5

MaxillipedInner plate No tooth No tooth No tooth 1 tooth No tooth 1 tooth No tooth 1 tooth

2 setae 3 setae 4 setae 4 setae 3–4 setae 3–4 setae 3 setae 3 setaeOuter plate 4–5 setae 5–6 setae 5–7 setae 4–5 setae 12 setae 7 setae 4–5 setae 5 setae

Distal projection on Absent Present Absent Absent Absent Present Absent Presentpenultimate article ofpalp

Gnathopod 1 grasping 2 1 2 2 1 1 2 1spines

Proximal projection of the Rectangular Rectangular Rectangular Rectangular Triangular Rectangular Rectangular Rectangularpropodus ofgnathopod 2 male

Pereopods 3 and 4 2-articulate 2-articulate 2-articulate 2-articulate 2-articulate 2-articulate 2-articulate 1 or2-articulate

Pereopod 5 Reduced Normal Normal Normal Normal Normal Normal NormalMale abdominal 1-articulate 1-articulate 1-articulate 2-articulate 1-articulate 1-articulate 1-articulate 1-articulate

appendages


F. 16. Biogeographical distribution of the Deutella species.

and also lack a molar flake. Furthermore, the present study has revealed thatDeutella venenosa Mayer, 1890 and the new species Deutella margaritae possess amolar flake. This indicates that the presence/absence of a molar flake should not beconsidered as a good character to differentiate among families. At present, takinginto account that the phylogeny of the Caprellidea is still under debate, it is probablybetter to maintain a smaller number of families. So, Takeuchi (1993) consideredonly five families and placed the genus Deutella in the family Caprellidae White,1847 as defined by Vassilenko (1974). As Takeuchi (1993) pointed out, the followingstudies are needed urgently to address the phylogeny of the Caprellidea: (1) redescrip-tion of each genus and species in order to find further generic characters, (2)discovery of presently unknown genera and species from the tropics to the SouthernHemisphere and the deep sea, (3) molecular analysis based on sequencing mitochon-drial DNA or ribosomal RNA and (4) ecological and behavioural studies.

AcknowledgementsThe author is very grateful to Elizabeth Nelson (National Museum of Natural

History), Elizabeth Borda (The American Museum of Natural History), ArdisB. Johnston (Museum of Comparative Zoology) and Judith C. Price (Canadian Museumof Nature) for the loan of type and non-type material of Deutella species. Special thanksto E. Nelson for her kindness and hospitality during the short stay at the NationalMuseum of Natural History. The author also thanks Martin Thiel for hospitality andfacilities provided during the stay at the Facultad de Ciencias del Mar, UniversidadCatolica del Norte, Coquimbo Chile. The work was partially supported by a grantAP98 28617065 from the Ministry of Education, Culture and Sport from Spain.

Revision of the genus Deutella1084

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revision of the genus deutella (crustacea: amphipoda

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