TWO NEW SPECIES OF DEEP-WATER CAPRELLA … · crustaceana 87 (1) 41-63 two new species of deep-water caprella (peracarida, amphipoda, caprellidae) from the pacific coast of mexico

Crustaceana 87 (1) 41-63

TWO NEW SPECIES OF DEEP-WATER CAPRELLA (PERACARIDA,AMPHIPODA, CAPRELLIDAE) FROM THE PACIFIC COAST OF MEXICO

COLLECTED DURING THE TALUD XIV CRUISE, WITH A CHECKLIST OFSPECIES OF CAPRELLIDAE RECORDED FOR THE EASTERN PACIFIC

BY

MICHEL E. HENDRICKX1,3) and MANUEL AYÓN-PARENTE2)1) Laboratorio de Invertebrados Bentónicos, Unidad Académica Mazatlán,

Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México,P.O. Box 811, Mazatlán, Sinaloa 82000, Mexico

2) Departamento de Ecología, CUCBA-Universidad de Guadalajara, Carretera a Nogales km 15.5,Las Agujas Nextipac, Zapopan, Jalisco 45110, Mexico

ABSTRACT

Two new deep-water species of the genus Caprella are described from the central Gulf ofCalifornia, Pacific coast of Mexico. Caprella calderoni new species is distinguished from the32 previously recorded species of Caprella from the eastern Pacific by its smooth forehead andpereonites, body not particularly slender, gnathopod 2 inserted at about mid-length of the secondpereonite, elongated gills and gnathopod 1, the relative length of antenna 2 articles, length of theflagellum of antenna 1, the lack of an antero-lateral projection on pereonites 2 and 3, and thelength-height proportion of pereonite 1. It is close to C. striata Mayer, 1903, recorded from Alaska.Caprella striata, however, features a different number of articles in the antenna 1 flagellum, twolatero-posterior spines on pereonites 6, a small dorsal tubercle on pereonites 5 and 6, and a poisonspine on the propodus of gnathopod 2 (all lacking in the new species). The buccal appendages alsopresent significant differences. Caprella mercedesae new species, belongs to a group of easternPacific species with a sharp spine on the forehead. It is distinguished from all these species by acombination of characters, including the general shape of the body (not robust), the insertion level ofgnathopod 2, its general shape, and the relative length of its dactylus, the relative length of antennae1 and 2, the shape and relative length of the gills, the proportionally shorter or longer pereonites, andthe presence of dorsal tubercles on at least pereonites 5-7 (absent in the new species).

Key words. — Caprella, new species, deep water, Pacific Mexico

RÉSUMÉ

Deux nouvelles espèces d’eaux profondes du genre Caprella sont décrites du golfe de Californiecentral, sur la côte pacifique du Mexique. Caprella calderoni nouvelle espèce se distingue des 32

3) Corresponding author; e-mail: [email protected]

© Koninklijke Brill NV, Leiden, 2014 DOI:10.1163/15685403-00003277

42 MICHEL E. HENDRICKX & MANUEL AYÓN-PARENTE

espèces de Caprella connues du Pacifique est par la tête et les segments du péréion lisses, le corpsqui n’est pas particulièrement svelte, le gnathopode 2 fixé environ à la moitié du second segment dupéréion, les branchies et le gnathopode 1 allongés, la longueur relative des segments de l’antenne2 et du flagelle de l’antenne 1, l’absence de projections antero-latérales sur les segments 2 et 3du péréion, et la proportion entre la longueur et la hauteur du segment 1 du péréion. Elle estproche de C. striata Mayer, 1903, connue de l’Alaska. Cependant, C. striata possède un nombredifférent d’articles du flagelle de l’antenne 1, deux épines postero-latérales sur le segment 6, unpetit tubercule sur les segments 5-6 du péréion, et une épine venimeuse (“poison spine”) sur lepropodus du gnathopode 2 (tous absents chez la nouvelle espèce). Les pièces buccales présententaussi des différences importantes. Caprella mercedesae nouvelle espèce, appartient à un petit grouped’espèces du Pacifique est qui ont une épine frontale effilée. Elle se distingue de toutes les autresespèces de ce groupe par une combinaison de caractères dont on peut souligner l’aspect général ducorps (pas robuste), le niveau d’insertion du gnathopode 2, sa forme générale et la longueur relativedu dactylus, la longueur relative des antennes 1 et 2, la forme et la longueur relative des branchies,les segments du péréion qui sont soit proportionnellement plus courts ou plus longs, et la présencede tubercules dorsaux sur les segments 5-7 du péréion (absents chez la nouvelle espèce).

Mots clés. — Caprella, nouvelles espèces, eaux profondes, Pacifique mexicain

INTRODUCTION

To date, the amphipods of the family Caprellidae Leach, 1814, of the Pacificcoast of Mexico have not been studied extensively. Records for this region concernonly three species: Caprella californica Stimpson, 1856, C. equilibra Say, 1818and C. scaura Templeton, 1836. In addition, as many as 20 species of CyamidaeRafinesque, 1815 (whale lice) have been reported for Mexican waters, mostlyoff the west coast of the Baja California Peninsula (Brusca & Hendrickx, 2005;García-Madrigal, 2007).

As far as the eastern Pacific is concerned, however, a compilation of old andrecent literature indicates that 61 species of Caprellidea, including 52 speciesof Caprellidae, have been recorded in this region: 49 Caprellinae Leach, 1814(1 species of Abyssicaprella McCain, 1966, 1 Aciconula Mayer, 1903, 32 CaprellaLamarck, 1801, 3 Deutella Mayer, 1890, 2 Mayerella Huntsman, 1915, 3 Meta-caprella Mayer, 1903, 2 Paracaprella Mayer, 1890, 1 Pseudoliropus Laubitz,1970, and 4 Tritella Mayer, 1890), 1 Paracercopinae Vassilenko, 1972 (genus Cer-cops Krøyer, 1843), and 2 Phtisicinae Vassilenko, 1968 (1 Caprellina Thompson,1879, and 1 Perotripus Dougherty & Steinberg, 1953).

As a general rule, shallow water caprellids are much better known than deep-water species, essentially due to the difficulty to sample in great depths. Duringsampling operations in the northern part of the central Gulf of California, manyspecimens of Caprella were collected in two deep-water benthic samples. Thismaterial belongs to two as yet undescribed species, which will be described andfigured herein.

CAPRELLA CALDERONI NOV. AND C. MERCEDESAE NOV. 43

MATERIAL AND METHODS

The material reported in this study was obtained while sampling with the R/V“El Puma”, Universidad Nacional Autónoma de México, in the northern sectionof the central Gulf of California, roughly between 28°10′ and 29°10′N. A total of30 stations were visited, with depths ranging from 148 to 1346 m (see Hendrickx,2012). Specimens of caprellids were collected with a 2.35 m wide by 0.95 m high,standard benthic sledge equipped with an outer collecting net of ca 5.5 cm (2.25inch) stretch mesh and an inner net of ca 2.0 cm (0.75 inch) stretch mesh. Trawlinglasted 30 min at an average speed of 1.75 knots (approx. 3.25 km/h). Samplingdepths were estimated with a digital SIMRAD echo sounder. Epibenthic tempera-ture and oxygen concentration were measured ca 10 m above bottom level with aSeabird CTD-O2 probe. Oxygen concentrations were also double-checked with theWinkler method using water samples collected in closing bottles near the bottom.

Most of the specimens examined, including the holotype, allotype, and para-types, are deposited in the Regional Collection of Marine Invertebrates at theMazatlán Marine Station, UNAM, in Mazatlán, Mexico (EMU), with their re-spective catalogue number. Other paratypes are deposited in the National Crus-tacean Collection of the Instituto de Biología, UNAM, Mexico City (CNCR), andin the crustacean collection of the Los Angeles County Museum of Natural History(LACM).

A compilation of old and recent literature dealing with Caprellidae recorded inthe eastern Pacific was performed and provided a checklist of 52 species for thiszoogeographic region. A result of this compilation is included in this contribution(Appendix), including the major sources of information and the general geographicdistribution of each species.

Abbreviations used are: TL, total body length; ovig., ovigerous; juv., juveniles.The classification used follows Horton & De Broyer (2013). Species descriptionessentially follows Guerra-García et al. (2001), Guerra-García & García-Gomez(2003), and Guerra-García & Takeuchi (2004).

TAXONOMIC ACCOUNT

Caprella calderoni new species(figs. 1-4)

Material examined.— Holotype �, TL 9.7 mm, TALUD XIV, St. 21 (29°00′53′′N 112°51′31′′W),9/April/2012, benthic sledge, 412-415 m depth (EMU-9858). Allotype �, TL 9.5 mm, same station(EMU-9859). Paratypes, all from same station: 1 �, TL 12.9 mm (EMU-9860); 15 �� (TL 2.5-11.4 mm), 5 �� (TL 7.7-9.5 mm), and 1 ovig. � (TL 7.6 mm) (EMU-9861); 14 �� (TL 4.1-8.2 mm),7 �� (TL 7.8-9.0 mm), and 1 ovig. � (TL 8.9 mm) (EMU-9862); 4 �� (TL 7.7-12.6 mm) and 3 ��(TL 8.5-8.7 mm) (LACM-CR20130001); 13 �� (TL 3.4-15.5 mm) and 9 �� (TL 5.7-13.6 mm)


(CNCR-28309). Paratype, TALUD XIV, St. 30, 1 � (TL 7.7 mm) (28°32′57′′N 112°59′26′′W),11/April/2012, benthic sledge, 270-309 m depth (EMU-9865A); same station, 1 �, TL 9.5 mm, and1 ovig. �, TL 7.5 mm (EMU-9865B).

Additional material.— TALUD XIV, St. 21 (29°00′53′′N 112°51′31′′W), 9/April/2012, benthicsledge, 412-415 m depth, 14 �� (TL 5.8-10.8 mm) and 3 �� (TL 4.8-8.1 mm) (EMU-9863); samestation, 25 �� (TL 4.7-19.7 mm), 13 �� (TL 6.9-7.9 mm) and 1 ovig. � (TL 8.5 mm) (EMU-9864);same station, 11 �� (TL 3.9-10.5 mm) and 2 �� (TL 3.9-10.5 mm) (EMU-9992).

Diagnosis.— Head and body smooth, except for a pair of small laterodistalspines on pereonite 5, pereonite 5 the longest. Eyes well developed. Antenna 1about 2/3 body length. Antenna 2 slightly longer than 1/2 of antenna 1. Gnathopod2 inserted posterior to half of pereonite 2 in males, slightly ahead of segment mid-length in females, propodus ovate, twice as long as basis, basis slightly shorterthat 1/2 pereonite length, one grasping spine at about 1/3 length of ventral margin,followed by row of irregularly-set short spines and setae, 1-2 triangular projectionsin distal third of margin; dactylus without setae or serrations. Gills slender, oval-shaped, about 3 (P3) to 4 (P4) times as long as wide. Pereiopods 5-7 long, witha pair of grasping spines at mid-length of propodus palm. Abdominal appendages2-articulated, articles of similar length. Penes moderately large, situated medially.

Description.— Holotype male (fig. 1A). Body length 9.7 mm. Head rounded,without dorsal spine or projection; eyes with distinguishable ommatidia; suturebetween head and pereonite 1 present, incomplete. Pereonites 2-7 smooth, exceptfor a pair of small laterodistal spines on pereonite 5; pereonites 5 the longest, 7 theshortest, 2-4 subequal. Gills on pereonites 3 and 4 slender, about 3 (P3) to 4 (P4)times as long as wide, first pair slightly longer than second.

Upper lip (fig. 2A) symmetrically bilobed, partly setose apically. Mandibles(fig. 2B) without palp, mandibular molar present, left mandible with 5-toothedincisor and lacinia mobilis, followed by 3 plumose setae; right mandible with 5-toothed incisor and lacinia mobilis, followed by 2 plumose setae, both mandibleswithout molar flake. Lower lip (fig. 2A) with inner lobes well demarcated, innerand outer lobes with short setae. Maxilla 1 (fig. 2C) outer lobe with 7 forked spines;distal article of palp with 6 strong spines and 2 rows of setae laterally. Maxilla2 (fig. 2D) inner lobe oval, with about 20 simple setae and a lateral row of 5setulae; outer lobe rectangular, with about 20 simple setae. Maxilliped (fig. 2E)outer plate long, oval, inner margin with a row of 6 short teeth and 10 simplemarginal and submarginal setae; inner plate rectangular, with a row of 10 setae ondistal margin; palp 4-articulated, setose, dactylus with row of setulae on distal halfof upper margin and one, short, subterminal simple setae.

Antenna 1 (fig. 1A, B) about 2/3 of body length; second article of peduncle thelongest, distal article long, about 2/3 length of previous one; flagellum with 14articles.


Fig. 1. Caprella calderoni new species. Male holotype, TL 9.7 mm (EMU-9858) A, lateral view; B,antenna 1; C, antenna 2; D, abdomen, ventral view. Female allotype, TL 9.5 mm (EMU-9859) E,lateral view; F, abdomen, dorsal (up) and ventral views. Scale bars, A, B, E, 2 mm; D, 0.2 mm; C, F,

0.5 mm.


Fig. 2. Caprella calderoni new species. Male holotype, TL 9.7 mm (EMU-9858) A, upper (above)and lower lips; B, left (L) and right (R) mandibles; C, maxilla 1; D, maxilla 2; E, maxilliped. Scale

bars = 0.1 mm.

Antenna 2 (fig. 1A, C) slightly longer than 0.5 of antenna 1, about as long aspeduncle of antenna 1, swimming setae present, distal setae on ultimate article;flagellum with 2 articles.


Fig. 3. Caprella calderoni new species. Male holotype, TL 9.7 mm (EMU-9858) A, male gnathopod1, lateral view; B, male gnathopod 2, lateral view; C, female allotype, TL 9.5 mm (EMU-9859),

gnathopod 2, lateral view. Scale bars = 0.2 mm.

Gnathopod 1 (fig. 3A) basis longer than length of ischium to carpus combined,one distal seta on basis ventral margin; two ventral setae on ischium; merus and


carpus setose; propodus triangular, length about 2 times width, tapering distally,width at base equal to maximum width of carpus, palm with a pair of proximalgrasping spines, with mixed long and short setae along the palm; dactylus slightlycurved, grasping margin serrate, with a row of short submarginal setae and a longersubdistal setae.

Gnathopod 2 (fig. 3B) inserted posterior to half of pereonite 2; basis slightlyshorter than half pereonite 2 length, provided with a triangular projection distally;ischium trapezoidal; merus rectangular, with distal spine; carpus very reduced,triangular; propodus ovate, elongate, twice as long as basis, one grasping spine atabout 1/3 length of ventral margin, followed by row of irregularly-set short spinesand setae, 1-2 triangular projections in distal third of margin; dactylus slightlycurved, without ventral setae or serrations.

Pereopods 5-7 (from male paratype EMU-9865, TL 7.7 mm) (fig. 4) increasingin length, scarcely setose; pereopod 5 long, basis long, rectangular, naked, withstrong distal triangular tooth, ischium small, subquadrate, naked, merus, carpusand propodus with few distal and marginal setae, ventral margin of propoduswith pair of strong grasping spines at about mid-length and short, robust setaein distal half, dactylus naked; pereopod 6 similar to 5, but articles stronger,

Fig. 4. Caprella calderoni new species. Male paratype TL 7.7 mm (EMU-9865) pereopods 5-7 (topto bottom), lateral view. Scale bar = 0.5 mm.


wider; pereopod 7 similar to pereopods 5-6, but distal margin of merus withmore pronounced dorsal, squarish projection, and carpus proportionally shorter,squarish, with stronger distoventral setae.

Abdomen (fig. 1D) with one pair of 2-articulated appendages, articles of similarlength, each with 1-2 distal setae; a pair of lateral lobes and 1 dorsal lobe. Penesmoderately large, situated medially.

Allotype female (fig. 1E, F) (EMU-9859) TL 9.5 mm. Gnathopod 2 (fig. 3C)similar in shape to male, basis 0.6 times as long as pereonite 2, inserted slightlyahead of segment mid-length, spines on margin of palm shorter than in male, morerobust. Oostegites setose; gills slender than in male (fig. 1E). Abdomen withoutappendages.

Etymology.— This species is named after our colleague Luis Eduardo CalderonAguilera, in recognition of his generosity and the numerous occasions in which hekindly provided support to the academic activities of the first author’s laboratoryand to ALCARCINUS (Asociación Latinoamericana de Carcinología).

Habitat and biology.— Caprella calderoni new species, was found on the dorsalportion of the sea-urchin Spatangus californicus H. L. Clark, 1917. Except for onespecimen collected at station 30 (270-309 m depth; epibenthic oxygen and tem-perature 2.48 ml O2/l and 12.7°C, respectively), all the specimens were obtainedat station 21: 412-415 m depth; epibenthic oxygen 1.79 ml/l and temperature 11-21°C, respectively. Maximum size for this species is 19.7 mm TL. Male/female(M : F) proportion was 3.2 : 1. Of the 46 females collected only four (8.7%) wereovigerous and ranged in size from TL 7.6 to 9.5 mm.

Remarks.— The holotype lacks pereopods and these appendages were illus-trated using one of the paratypes.

There are 32 species of Caprella currently recorded from the eastern Pacific(see Appendix), seven with a well-developed frontal spine on the head (see below)which is lacking in C. calderoni new species. Of the remaining 25 species, 10species have the male body with either dorsal tubercles or spines on all or mostpereonites, and sometimes also on the head (i.e., C. acanthifera humboldtiensisMartin, 1977; C. borealis Mayer, 1903; C. ferrea Mayer, 1903; C. kinkaidi Holmes,1904; C. mutica Schurin, 1935; C. natalensis (Mayer, 1903); C. paulina Mayer,1903; C. pilipalma Dougherty & Steinberg, 1953; C. pustulata Laubitz, 1970; andC. scabra Holmes, 1904). Caprella calderoni new species lacks spines or tuberclesaltogether.

Another series of eastern Pacific species (i.e., C. brevirostris Mayer, 1903; C.equilibra Say, 1818; C. irregularis Mayer, 1890; C. laeviuscula Mayer, 1903; C.mendax Mayer, 1903; and C. pilidigitata Laubitz, 1970) has the second gnathopodinserted close to the posterior margin of pereonite 2, while in C. calderoni newspecies it is inserted at about mid-length of this pereonite. Caprella cilliata


Sars, 1883, and C. gracilior Mayer, 1903, are either very or extremely slenderspecies compared to C. calderoni new species. Caprella depranochir Mayer, 1890,features almost rounded gills and short, robust pereonites 2-3 (vs. elongated,oval-shaped gills and slender pereonites in C. calderoni new species). Caprellagreenleyi McCain, 1969 and C. ungulina Mayer, 1903, feature a stout, moremassive body, with first gnathopod almost round in shape (C. greenleyi) or robust,with a very short dactylus (C. unguilinea).

Of the remaining four species, Caprella alaskana Mayer, 1903, shows aconsiderable variation, from the very spiny form to the spineless form (Laubitz,1970). Characters separing this species from C. calderoni new species are the tipof antenna 2 clearly falling short of half the length of antenna 1 second pedunculararticle, gnathopod 2 inserted more posteriorly on pereonite 2, a proportionallyshorter antenna 1 flagellum, and the presence in C. alaskana of a antero-lateralprojection of pereonites 2 and 3. Caprella alaskanensis Holmes, 1904, was poorlydescribed (based on a mutilated, unique specimen) and apparently never foundagain. According to Holmes (1904), C. alaskanensis features a low tubercle onthe forehead, the pereonite 1 is about 3 times as long as deep and gnathopod 2 isinserted almost at the posterior extremity of pereonite 2, all features which separateit from C. calderoni new species. Caprella rudiuscula Laubitz, 1970, features a pairof tubercles on the head, which C. calderoni new species lacks, and short antenna1 vs. long antenna 1 in C. calderoni new species.

Of all species of Caprella recorded from the eastern Pacific (Appendix), theclosest to the newly described species is C. striata Mayer, 1903, a primarily north-ern, deep-water caprellid recorded between 7 and 150 fm (13-278 m) (Laubitz,1970). Both species share a similar shape, size and proportions of antenna 1-2,pereonites 2-4 are almost equal in length in both species, and gnathopods 1 and2 are very similar. There are, however, significant difference between C. striataand C. calderoni new species. When comparing the redescription of C. striata byLaubitz (1970), antenna 1 flagellum possesses at least 21 articles (vs. 14 in the newspecies), there are two latero-posterior spines on pereonites 6 and a small dorsaltubercle on pereionites 5 and 6 (lacking in the new species); mandibles of bothspecies are very similar, but the molar processes are much stronger in C. striata;maxilla 1 is different, with the palp second article ending in numerous setae in C.striata (vs. six strong setae in the new species), and the endopod distal setae arevery robust and with a bifid or trifid tip, vs. slender and with single tip; maxil-liped of both species is also very similar, but the outer plate in C. striata is muchshorter (not overreaching distal margin of the palp first article) and the dactyl ofthe palp of C. striata lacks the dorsal setulae on its distal half. Illustrations of thetype material by Mayer (1903) correspond to specimens with numerous tubercleson the pereonites and female gills proportionally longer than in C. calderoni new


species. Other characters that separate both species are: the palp of maxilla 1 whichis much shorter and proportionally wider in C. striata than in the new species, al-though this might be the result of a different orientation of the appendix when itwas illustrated; the propodus of gnathopod 1 features a small proximal pair of se-tae in C. striata, while these setae are robust in the new species; the propodus ofgnathopod 2 of C. striata features a grasping spine, a poison spine and a subtri-angular distal tooth on the ventral margin, while the poison spine is lacking in thenew species, but this might be related to its development stage.

Caprella mercedesae new species(figs. 5-8)

Material examined.— Holotype �, body length 11.1 mm, GUAYTEC I, St. 11 (27°N 111°50′W),14/February/1987, trawl, 260 m depth (EMU-9866, ex 9196). Allotype �, TL 10.6 mm, TALUD XIV,St. 30 (28°32′57′′N 112°59′26′′W), 11/April/2012, benthic sledge, 270-309 m depth (EMU-9867).Paratypes, all from same station: 1 �, TL 10.2 mm (EMU-9868); 2 ��, TL 7.6 and 19.8 mm(EMU-9869); 1 �, TL 14.4 mm (EMU-9871); 2 ��, TL 6.7-8.5 mm (EMU-9872); 1 �, TL 8.2 mm(LACM-CR20130002); 1 �, TL 8.7 mm (CNCR-28310).

Diagnosis.— Head with sharp anterodorsal spine, pereonites 2-4 smooth, pere-onite 5-7 with spines, pereonites 2-3 the longest. Eyes well developed. Antenna 1almost as long as body length. Gnathopod 2 inserted in middle of pereonite 2 inmales, slightly ahead of segment mid-length in females, propodus ovate, twice aslong as basis, basis slightly longer that 1/2 pereonite length, ventral margin withrow of short, proximal spines, a pair of grasping spine, a row of 7 short, strong se-tae, a strong triangular tooth (poison spine), a deep notch with some short setae, asecond moderately strong, triangular tooth, and a distal row of 5 short, robust setae;dactylus more than 1/2 gnathopod length, finely crenulated dorsally on proximalhalf. Gills slender, subequal, about 3 times as long as wide. Pereopods 5-7 short,articles short, robust, subtriangular, a pair of proximal grasping spines on propoduspalm. Abdominal appendages 2-articulated, articles of similar length. Penes large,subtriangular, situated medially.

Description.— Body (fig. 5A, D) length 11.1 mm. Head rounded, with sharpanterodorsal spine; suture between head and pereonite 1 present, incomplete.Pereonites 2-4 smooth, pereonite 5 with two pairs of distolateral spines, second(posterior) largest, pereonite 6 with one pair of large dorsal spine on posteriormargin and two pairs of smaller, posterolateral spines, pereonite 7 with one large,median spine and a pair of shorter lateral spines on posterior margin, pereonites2-3 subequal, pereonites 4 and 5 about 0.8 length of pereonite 2, pereonite 7 theshortest. Gills on pereonites 3 and 4 slender, subequal in length, about 3 times aslong as wide.

Upper lip (fig. 6A) symmetrically bilobed, partly setose apically. Mandibles(fig. 6B) without palp, mandibular molar present, left mandible with five-toothed


Fig. 5. Caprella mercedesae new species. Male holotype, TL 11.1 mm (EMU-9866) A, lateral view;B, antenna 1; C, antenna 2; D, pereonites 6-7, dorsal view; E, abdomen, ventral view. Femaleallotype, TL 10.6 mm (EMU-9867) F, lateral view; G, pereonite 7, dorsal view; H, same, ventral

view. Scale bars: A, B, D, F, 1 mm; C, G, H, 0.5 mm; E, 0.1 mm.


Fig. 6. Caprella mercedesae new species. Male holotype, TL 11.1 mm (EMU-9866) A, upper (above)and lower lips; B, left (L) and right (R) mandibles; C, maxilla 1; D, maxilla 2; E, maxilliped. Scale

bar = 0.2 mm.


incisor and lacinia mobilis, followed by 3 plumose setae; right mandible with five-toothed incisor and 3-toothed lacinia mobilis, followed by 2 plumose setae, molarflake absent. Lower lip (fig. 6A) with inner lobes well demarcated, inner and outerlobes with short setae. Maxilla 1 (fig. 6C) outer lobe with 5 forked spines; distalarticle of palp with 5 robust spines and a series of setae laterally, setae increasing innumber from mid-length to distal part. Maxilla 2 (fig. 6D) inner lobe oval, with 21simple setae and a lateral row of 7 tiny setulae; outer lobe rectangular, with >20simple setae. Maxilliped (fig. 6E) outer plate long, subrectangular, inner marginwith a row of 6 spines and a submarginal row of long, simple setae; inner platesquarish, distal margin with a row of 12 robust setae; palp 4-articulated, setose,dactylus naked except for a short distal setae.

Antenna 1 (fig. 5A, B) almost as long as body length; second article of pedunclethe longest, distal article long, about 2/3 length of previous one; flagellum with 24articles.

Antenna 2 (fig. 5A, C) slightly longer than 1/2 of antenna 1, about as long aspeduncle of antenna 1, swimming setae present, distal setae on ultimate article;flagellum with 2 articles.

Gnathopod 1 (fig. 7A) basis about as long as ischium to carpus length combined,a few short setae on basis ventral margin and a cluster of 3 longer distal setae;5 ventral setae on ischium; merus and carpus setose; carpus with a heel-shapedventral projection, propodus triangular, length about 1.7 times width, taperingdistally, width at base equal to maximum width of carpus, palm with a pair ofproximal grasping spines, with mixed long and short setae along the palm; dactylusslightly curved, grasping margin serrate on distal 2/3, tip with 4 large teeth, a rowof short submarginal setae, and a pair of longer subdistal setae.

Gnathopod 2 (fig. 7B) inserted in middle of pereonite 2; basis slightly longerthan half pereonite 2 length, provided with a sharp triangular projection distally,upper margin serrate; ischium trapezoidal, merus squarish with distal spine, carpusvery reduced, triangular, ventral margin serrate; propodus ovate, elongate, twice aslong as basis, row of short spines on ventral margin proximal third, followed bypair of grasping spine, a row of 7 short, strong setae, a strong triangular tooth(poison spine), a deep notch with some short setae, a second moderately strong,triangular tooth, and a distal row of 5 short, robust setae, dorsal margin with shortspines, more densely set proximally; dactylus slightly curved, finely crenulateddorsally on about proximal half, with a short row of short distal setae.

Pereopods 5-7 (from male paratype EMU-9868, TL 10.2 mm) (fig. 8) increasingin length, scarcely setose, articles short, robust; pereopod 5 short, basis with strongtriangular dorsal projection with serrate margin, distal angle acute, ischium short,subquadrate, merus short, with strong subtriangular dorsal projection, distal angleforming a lobe with an acute terminal spine and a few long setae, carpus short, with


Fig. 7. Caprella mercedesae new species. Male holotype, TL 11.1 mm (EMU-9866) A, malegnathopod 1, lateral view; B, male gnathopod 2, lateral view. Female allotype, TL 10.6 mm (EMU-

9867) C, gnathopod 2, lateral view. Scale bar = 0.5 mm.


Fig. 8. Caprella mercedesae new species. Male paratype TL 10.2 mm (EMU-9868) pereopods 5-7(top to bottom), lateral view. Scale bar = 1 mm.


strong suboval, dorsodistal projection and a few long, terminal setae, propodusoval, dorsal margin with long setae, ventral margin with basal protuberance withtwo grasping spines, followed by row of short setae, dactylus naked; pereopods 6and 7 similar to pereopod 5, but setae on ventral margin of propodus more robust.

Abdomen (male) (fig. 5E) with one pair of 2-articulated appendages, articles ofsimilar length; a pair of lateral lobes and 1 dorsal lobe. Penes large, subtriangular,situated medially.

Allotype female (EMU-9867). Body (fig. 5F, G, H) length 10.6 mm. Gnathopod2 (fig. 7C) similar to male, basis dorsal margin serrate, propodus slender, basis 0.5times as long as pereonite 2, inserted slightly ahead of segment mid-length, spineson margin of palm less robust than in male. Oostegites setose; gills slender than inmale. Abdomen without appendages.

Etymology.— This species is named after M.E.H.’s wife, Mercedes CorderoRuiz, for her constant support during his career and the numerous and significantcontributions she made to the Laboratorio de Invertebrados Bentónicos, ICML,UNAM, in the last 18 years.

Habitat and biology.— The holotype of Caprella mercedesae new species wasfound in a jar together with a specimen of Hesperocidaris perplexa (H. L. Clark,1907), but there is no evidence that it lived associated with this sea urchin. Onthe contrary, many specimens of the material collected during the TALUD XIVsurvey were found grasping small colonies of unidentified Hydrozoa. The depthrange for C. mercedesae new species is set at 260-309 m. Environmental conditionsat bottom level of the TALUD material were as follows: 2.48 ml O2/l, 12.7°C.Maximum size for this species is 19.8 mm TL. Of the 16 specimens collectedonly three were females (M : F = 5.3 : 1) and of these two were ovigerous (TL,7.5-10.6 mm).

Remarks.— Along the eastern Pacific coast, there are seven species of Caprellafeaturing a well-developed frontal spine on the head, similar to the one found inCaprella mercedesae new species: Caprella angusta Mayer, 1903, C. californicaStimpson, 1856, C. incisa Mayer, 1903, C. penantis Leach, 1814, C. scauraTempleton, 1836, C. uniforma La Follete, 1915, and C. verrucosa Boeck, 1871.Of these seven species, Caprella angusta presents a much shorter dactylus ofgnathopod 2 (much less than 0.5 its length vs. well over 0.5 the gnathopodlength in C. mercedesae new species) and it is apically truncated vs. pointedin C. mercedesae new species; antenna 1 is also proportionally much shorterin C. augusta than in C. mercedesae new species. In C. californica, the bodyand gnathopod 2 are very slender compared to C. mercedesae new species, andpereonite 5 in the former features a sharp spine in dorsal posterior third, which isabsent in the latter; gills are also strickingly longer in the male of C. californica.Caprella incisa has a proportionally much shorter first antenna, the second antenna


is clearly much longer than the first, and this species features paired or unpaireddorsal protuberances on all pereonites vs. none in C. mercedesae new species.Caprella penantis is a robust species, with pereonites 2-5 short and narrow (lessthan twice as long as wide in lateral view), while C. mercedesae new species ismore slender, with pereonites 2-5 twice as long as wide; in addition, antenna 1is proportionally shorter and gills are rounded in C. penantis vs. oval and narrowin C. mercedesae new species. Caprella scaura is a very slender species, withpereonite 1 almost 3 times as long as the head and gnathopod 2 is inserted closeto posterior margin of pereonite 2, while in C. mercedesae new species the headis only slightly shorter than pereonite 1 and the first pair of gnathopod is insertedat about mid-length of pereonite 2; the shape of gnathopod 2 is also strickinglydifferent. Caprella uniforma possesses pereonites 3-4 that are more robust andproportionally shorter than in C. mercedesae new species, and it features a seriesof strong dorsal tubercles in pereonites 5-7, a character absent in C. mercedesaenew species. Finally, C. verrucosa pereonites 1-7 have paired or unpaired strongtubercles (“blunt spines”) dorsally, a character absent in C. mercedesae newspecies.

ACKNOWLEDGEMENTS

The authors thank Bruno David, Université de Lyon, for the identificationof Spatangus californicus and the Instituto Tecnólogico de Monterrey, CampusGuaymas, for donation of the material collected during the GUAYTEC I cruise.The TALUD XIV cruise aboard the R/V “El Puma” was supported by CTIC,UNAM. We also thank Mercedes Cordero Ruiz for preparing a Caprellidea database on which the Appendix is based, and for the final edition of the manuscript.We are grateful to two anonymous reviewers for their suggestions. One of us(M.E.H.) also warmly thanks Dr. J. Carel von Vaupel Klein, editor of Crustaceana,for making arrangements for the publication of this contribution, as well as forhis patience, assistance, and friendship during my 34 years of publishing in thejournal.

REFERENCES

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AUSTIN, W. C., 1985. Amphipoda. In: An annotated checklist of marine invertebrates in the coldtemperate northeast Pacific: 589-623. (Khoyatan Marine Laboratory, Cowichan Bay, BC).


BRUSCA, R. C. & M. E. HENDRICKX, 2005. Cap. 12. Crustacea 4. Lophogastrida, Mysida,Amphipoda, Tanaidacea & Cumacea. In: M. E. HENDRICKX, R. C. BRUSCA & L. T.FINDLEY (eds.), A distributional checklist of the macrofauna of the Gulf of California, Mexico.Part I. Invertebrates. [Listado y Distribución de la Macrofauna del Golfo de California, México,Parte I. Invertebrados]: 139-154. (Arizona-Sonora Desert Museum, Tucson, AZ).

CHESS, J. R., 1989. Aciconula acanthosoma, new species, a caprellid amphipod from SouthernCalifornia, with notes on its ecology. Journ. Crust. Biol., 9: 662-665.

GARCÍA-MADRIGAL, M. S., 2007. Annotated checklist of the amphipods (Peracarida: Amphipoda)from the tropical eastern Pacific. In: M. E. HENDRICKX (ed.), Contributions to the study ofeastern Pacific crustaceans [Contribuciones al Estudio de los Crustáceos del Pacífico Este], 4(2): 63-195. (Instituto de Ciencias del Mar y Limnología, UNAM, Mexico).

GUERRA-GARCÍA, J. M. & J. C. GARCÍA-GOMEZ, 2003. A new species of Caprella (Amphipoda,Caprellidae) from deep sea waters. Crustaceana, 76: 581-590.

GUERRA-GARCÍA, J. M., J. E. SÁNCHEZ-MOYANO & J. C. GARCÍA-GÓMEZ, 2001. Two newhairy species of Caprella (Amphipoda) from the Strait of Gibraltar, with a redescription ofCaprella grandimana. Journ. Crust. Biol., 21: 1014-1030.

GUERRA-GARCÍA, J. M. & I. TAKEUCHI, 2004. The Caprellidae (Crustacea: Amphipoda) fromTasmania. Journ. Nat. Hist., 38: 967-1044.

GUERRA-GARCÍA, J. M. & M. THIEL, 2001. The caprellid fauna (Crustacea: Amphipoda: Caprel-lidea) from Coquimbo, northern-central Chile with a taxonomic key for species identification.Rev. Chilena Hist. Nat., 74: 873-883.

HENDRICKX, M. E., 2012. Pandalid shrimp (Crustacea: Decapoda: Caridea: Pandalidae) collectedduring the TALUD XIV cruise in the Gulf of California, Mexico, and rediscovery of Plesionikacarinirostris Hendrickx, 1989. Cah. Biol. Mar., 53: 495-504.

HOLMES, S. J., 1904. Amphipod crustaceans of the expedition. Harriman Alaska Expedition,Alaska, 10 (Crustaceans): 233-246.

HORTON, T. & C. DE BROYER, 2013. Caprelloidea. In: T. HORTON, J. LOWRY & C. DE

BROYER (eds.), World Amphipoda database. Accessed through: World Register of Ma-rine Species, available online at http://www.marinespecies.org/aphia.php?p=taxdetails\&id=196121 on 2013-08-09.

LAUBITZ, D. R., 1970. Studies on the Caprellidae (Crustacea, Amphipoda) of the American NorthPacific. Natl. Mus. Can. Publ. Biol. Oceanogr., 1: 1-89.

MAYER, P., 1890. Die Caprelliden des Golfes von Neapel und der angrenzenden Meeresabschnitte.Nachtrag zur Monographie derselben. Fauna und Flora des Golfes von Neapel, 17: 1-157.

— —, 1903. Die Caprelliden der Siboga-Expedition. Siboga Exped. Monogr., 34: 1-160.MCCAIN, J. C. & W. S. GRAY, 1971. Antartic and Subantartic Caprellidea (Crustacea: Am-

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(eds.), A taxonomic list of common marine invertebrate species of southern California.Technical Report Allan Hancock Foundation, 3: 196-223. (University of Southern California,Los Angeles, CA).

First received 28 August 2013.Final version accepted 29 December 2013.


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ain

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einb

erg,

1970

;Aus

tin,1

985

Bri

tish

Col

umbi

a,C

anad

a.T

ierr

ade

Fueg

o,C

hile

.Sou

thA

tlant

icC

apre

lla

unif

orm

aL

aFo

llete

,191

5M

cCai

n&

Stei

nber

g,19

70;A

ustin

,198

5L

agun

aB

each

,sou

ther

nC

alif

orni

a,U

.S.A

.C

apre

lla

verr

ucos

aB

oeck

,187

1M

cCai

n&

Stei

nber

g,19

70;L

aubi

tz,1

970;

Wic

kste

n,19

80;A

ustin

,198

5;G

uerr

a-G

arcí

a&

Thi

el,2

001

Japa

n.B

ritis

hC

olum

bia,

Can

ada,

toso

uthe

rnC

alif

orni

a,U

.S.A

.Coq

uim

bo,C

hile

Deu

tell

aca

lifo

rnic

aM

ayer

,189

0M

cCai

n&

Stei

nber

g,19

70;L

aubi

tz,1

970;

Wic

kste

n,19

80So

uthe

rnPr

ince

Will

iam

Soun

d,A

K,t

oso

uthe

rnC

alif

orni

a.[?

]Po

rtA

rans

as,T

exas

,nor

ther

nG

ulf

ofC

alif

orni

a,U

.S.A

.D

eute

lla

vem

ae(M

cCai

n&

Gra

y,19

71)

McC

ain

&G

ray,

1971

Off

coas

tsof

Arg

entin

aan

dSo

uthe

rnC

hile

Deu

tell

ave

neno

saM

ayer

,189

0M

cCai

n&

Stei

nber

g,19

70;G

uerr

a-G

arcí

a&

Thi

el,2

001

Coq

uim

bo,C

hile

May

erel

laba

nksi

aL

aubi

tz,1

970

Lau

bitz

,197

0;W

icks

ten,

1980

;Aus

tin,1

985

Chi

chag

ofIs

land

,AK

,to

Sout

hern

Cal

ifor

nia,

U.S

.A.


(AP

PE

ND

IXC

ON

TIN

UE

D)

Spec

ies

Ref

eren

ces

Dis

trib

utio

nM

ayer

ella

mag

ella

nica

(McC

ain

&G

ray,

1971

)M

cCai

n&

Gra

y,19

71H

uasc

o,C

hile

Met

acap

rell

aan

omal

a(M

ayer

,190

3)M

cCai

n&

Stei

nber

g,19

70;A

ustin

,198

5;L

aubi

tz,1

970

Japa

nA

rchi

pela

goan

dof

fSu

mdu

mG

laci

er,A

K,

toC

alif

orni

a,U

.S.A

.M

etac

apre

lla

ferr

ea(M

ayer

,190

3)M

cCai

n&

Stei

nber

g,19

70C

alif

orni

a,U

.S.A

.M

etac

apre

lla

kenn

erly

i(St

imps

on,1

864)

McC

ain

&St

einb

erg,

1970

;Aus

tin,1

985;

Lau

bitz

,197

0A

lask

ato

sout

hern

Cal

ifor

nia,

U.S

.A.

Para

capr

ella

barn

ardi

McC

ain,

1967

McC

ain

&St

einb

erg,

1970

Cul

ebra

Isla

nd,P

anam

aPa

raca

prel

lapu

sill

aM

ayer

,189

0M

ayer

,189

0;G

uerr

a-G

arcí

a&

Thi

el,2

001

Chi

na.F

lori

daan

dno

rthe

rnG

ulf

ofM

exic

o,U

.S.A

.Car

ibbe

anIs

land

s.B

razi

l.T

ropi

calW

est

Afr

ica

and

Sout

hA

fric

a.Ta

nzan

ia.S

uez

Can

al.

Coq

uim

bo,C

hile

.Haw

aii,

U.S

.A.

Pse

udol

irop

usva

nus

Lau

bitz

,197

0L

aubi

tz,1

970;

Aus

tin,1

985

Que

enC

harl

otte

Isla

nd,B

C,C

anad

aTr

itel

lala

evis

May

er,1

903

McC

ain

&St

einb

erg,

1970

;Lau

bitz

,197

0;W

icks

ten,

1980

;Aus

tin,1

985

Van

couv

eran

dQ

ueen

Cha

rlot

teIs

land

,BC

,C

anad

a,to

sout

hern

Cal

ifor

nia,

U.S

.A.

Trit

ella

orna

taM

ayer

,190

3M

cCai

n&

Stei

nber

g,19

70Po

poff

Stra

its,S

hum

agin

Isla

nds,

AK

,U.S

.A.

Trit

ella

pili

man

aM

ayer

,189

0M

cCai

n&

Stei

nber

g,19

70;L

aubi

tz,1

970;

Wic

kste

n,19

80;A

ustin

,198

5Ju

ande

Fuca

Stra

it,A

K,t

oSo

uthe

rnC

alif

orni

a,U

.S.A

.Tr

itel

late

nuis

sim

aD

ough

erty

&St

einb

erg,

1953

McC

ain

&St

einb

erg,

1970

;Wic

kste

n,19

80;

Aus

tin,1

985

Sout

hern

Cal

ifor

nia,

U.S

.A.

Para

cerc

opin

aeC

erco

psco

mpa

ctus

Lau

bitz

,197

0L

aubi

tz,1

970

Ore

gon,

U.S

.A.

Phtis

icin

aeC

apre

llin

alo

ngic

olli

s(N

icol

et,1

849)

McC

ain

&St

einb

erg,

1970

;Gue

rra-

Gar

cía

&T

hiel

,200

1So

uth

Afr

ica.

New

Zea

land

.Coq

uim

bo,C

hile

Pero

trip

usbr

evis

(La

Folle

te,1

915)

McC

ain

&St

einb

erg,

1970

;Lau

bitz

,197

0V

anco

uver

Isla

nd,B

C,C

anad

a,to

Lag

una

Bea

ch,

sout

hern

Cal

ifor

nia,

U.S

.A.

Documents

TWO NEW SPECIES OF DEEP-WATER CAPRELLA … · crustaceana 87 (1) 41-63 two new species of deep-water caprella (peracarida, amphipoda, caprellidae) from the pacific coast of mexico