ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2014 Magnolia Press

Zootaxa 3815 (2): 215–232

www.mapress.com/zootaxa/Article

http://dx.doi.org/10.11646/zootaxa.3815.2.3

http://zoobank.org/urn:lsid:zoobank.org:pub:2A5EC5EA-EBC9-4E79-B8F0-1D17F1E97ECD

Taxonomic notes on some Brazilian species of Synalpheus Spence Bate, 1888,

with new records and description of a new species (Decapoda, Alpheidae)

ARTHUR ANKER1, 3 & PAULO P.G. PACHELLE2

1Tropical Marine Science Institute, National University of Singapore, 18 Kent Ridge Road, Singapore 119227, Republic of Singapore2Laboratório de Invertebrados Marinhos do Ceará, Departamento de Biologia, Universidade Federal do Ceará, Fortaleza, Ceará,

Brazil3Corresponding author. E-mail: arthuranker7@yahoo.com

Abstract

The present report deals with several western Atlantic species of the snapping shrimp genus Synalpheus Spence Bate, 1888

occurring in Brazilian territorial waters. Two species, Synalpheus cf. ruetzleri Macdonald & Duffy, 2006 and Synalpheus

yano (Ríos & Duffy, 2007), are recorded from the southwestern Atlantic Ocean and Brazil (Ceará) for the first time; S.

yano is also newly reported from the eastern Gulf of Mexico and USA (Florida). Synalpheus androsi Coutière, 1909, a

distinctive species previously known in Brazil only from a single unconfirmed record, is confirmed to be present in Bra-

zilian waters based on material from Bahia and Espírito Santo. Synalpheus tenuispina Coutière, 1909, a poorly known

species from southern Brazil, is redescribed. Synalpheus ul (Ríos & Duffy, 2007), previously known from Bahia and

Alagoas, is reported from Pernambuco based on older “Calypso” material misidentified as Synalpheus longicarpus (Her-

rick, 1891). A new species from the Synalpheus paraneptunus Coutière, 1909 complex is described as Synalpheus maxil-

lispinus sp. nov. based on three specimens dredged from moderate depths (15–55 m) off southern Bahia and Espírito

Santo, eastern Brazil. Finally, an incomplete specimen that appears to belong to a species presently unknown in Brazil is

preliminarily reported as Synalpheus sp. gambarelloides group. Illustrations are provided for all reported taxa except for

S. ul.

Key words: Atlantic Ocean, Brazil, Decapoda, snapping shrimp, Synalpheus, sponge, new record, new species, redescrip-

tion

Introduction

The speciose and taxonomically challenging snapping shrimp genus Synalpheus Spence Bate, 1888 is represented

in the Atlantic Ocean by over 60 species (De Grave & Fransen 2011; Anker et al. 2012). Most of them occur in the

tropical regions of the northwestern Atlantic, mainly in the Caribbean Sea, where most of the recent taxonomic

research on the genus has been done (e.g., Ríos & Duffy 2007; Anker & Tóth 2008; Macdonald et al. 2009;

Hultgren et al. 2010, 2011). In contrast, the synalpheid fauna of the southwestern Atlantic has not studied

exhaustively since Christoffersen (1979) and remains comparatively less known. The only noteworthy recent

studies dealing with the Brazilian material of Synalpheus are Coelho et al. (2006), Bezerra & Coelho (2006) and

Coelho Filho (2006), all simply listing several species, some of them as new records for Brazil; Almeida et al.

(2007, 2012) with an illustrated record of Synalpheus ul (Ríos & Duffy, 2007); and Anker et al. (2012), with

additional records (some new) for Brazil and illustrations of several species based on material collected in Atol das

Rocas. Anker et al. (2012) also commented on some morphologically variable taxa requiring taxonomic revisions,

including Synalpheus minus (Say, 1818), Synalpheus brevicarpus (Herrick, 1891), and Synalpheus apioceros

Coutière, 1909, all three of which occur in Brazil. These and other examples (Hultgren & Duffy 2012) show that

the taxonomy of the American species of Synalpheus is far from being settled.

In September–November 2013, the authors were able to study the unsorted caridean material deposited in the

crustacean collection of Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP). This

material contained numerous samples of Synalpheus, often in mixed lots together with other alpheid shrimps.

Accepted by J. W. Goy: 28 Apr. 2014; published: 13 Jun. 2014 215

Additional specimens examined for this study came from a small collection made by marine zoology students of

the Universidade Federal do Ceará (UFC), material from sponges reported by Bezerra & Coelho (2006) deposited

partly in UFC, samples from a short survey of São Sebastião harbour in 2009 (all later deposited in MZUSP), as

well as the crustacean collections of the Museu de Oceanografia Petronio Alves Coelho, Universidade Federal de

Pernambuco, Recife, Brazil (MOUFPE), and Florida Museum of Natural History, University of Florida,

Gainesville, U.S.A. (FLMNH).

In the present study, two species of Synalpheus are reported for the first time from the southwestern Atlantic

and Brazil (one of them also from USA); one species with a single uncertain Brazilian record is confirmed from

Brazil based on new material; one poorly known species is redescribed; one species is reported as new state record,

based on previously misidentified material; and one species from the Synalpheus paraneptunus Coutière, 1909

species complex (Anker & Tóth 2008) is described as new. Most species reported in this study are abundantly

illustrated. Carapace length (cl, in mm) was measured along the mid-dorsal line from the tip of the rostrum to the

posterior margin of the carapace. The type material of the new species is deposited in the carcinological collection

of MZUSP.

Taxonomy

Alpheidae Rafinesque, 1815

Synalpheus Spence Bate, 1888

Synalpheus androsi Coutière, 1909

(Fig. 1)

Synalpheus androsi Coutière 1909: 82, fig. 50; Chace 1972: 86; Dardeau 1984: 18; Coelho et al. 2006: 52; Macdonald et al.

2006: 169; Macdonald et al. 2009: 7, fig. 2, pl. 1C; Hultgren & Duffy 2011: 357, fig. 2 (in cladogram); Almeida et al.

2012: 28; Anker et al. 2012: 81 (table), 88 (key).

Zuzalpheus androsi—Ríos & Duffy 2007: 9, pl. 1.

Material examined. 1 male (cl 3.4 mm), Brazil, off Espírito Santo, REVIZEE—Comissão Central: 2, R/V Astro

Garoupa, station 21C, 20º42’21’’S 35º27’25’’W, depth: 55 m, 08.XI.1997 [MZUSP 31138]; 1 male (cl 3.0 mm),

same collection data as for previous specimen [MZUSP 31139]; 1 male (cl 3.0 mm, major P1 missing), 1 male (cl

2.6 mm, most pereiopods detached), 1 female (cl 2.6 mm, most pereiopods detached), same collection data as for

previous specimens [MZUSP 31140]; 1 specimen (not sexed), Brazil, Bahia, off Salvador, Saldanha Leste I

Expedition, station AS 1981, 13°40’S 38°48’W, depth: 49 m, sand and gravel, leg. N. Oc. Almirante Saldanha, det.

E. Silva, 23.IX.1968 [MOUFPE 9212] (material examined by J. Fidelis).

Description. See Coutière (1909) for original description and illustrations and Ríos & Duffy (2007, under

Zuzalpheus androsi) for diagnosis; comparative illustrations of the Brazilian material are provided in Fig. 1.

Distribution. Western Atlantic: Bahamas (Andros Island), Belize (Carrie Bow Cay), Jamaica, Barbados

(Coutière 1909; Dardeau 1984; Ríos & Duffy 2007; Macdonald et al. 2009; Hultgren et al. 2011), Brazil (off Bahia

and Espírito Santo: Vitória-Trindade seamounts) (Coelho et al. 2006; present study).

Ecology. In Belize, Synalpheus androsi inhabits large demosponges, Pseudoceratina crassa (Hyatt) and

Hyattella intestinalis (Lamarck), at a depth range of 14–15 m (Ríos & Duffy 2007). The present material was

dredged from much deeper water (49–55 m) and the collection records give no indication on association with

sponges.

Remarks. The present material represents the first confirmed and illustrated record of Synalpheus androsi in

Brazilian waters, closely matching the description and illustrations in Coutière (1909). Almeida et al. (2012)

pointed out that the first record of S. androsi from Brazil by Coelho et al. (2006) required confirmation. This record

was merely a listing of S. androsi in a species table, without a specific locality other than Bahia, and without

illustrations. The corresponding material was deposited in the collections of Museu de Oceanografia Petronio

Alves Coelho in Recife (MOUFPE) under the accession number DOCEAN-9212. At our request, Jesser Fidelis

examined Coelho’s material, a single dry specimen that had to be rehydrated, and confirmed it as S. androsi (see

material examined for collection details).

ANKER & PACHELLE216 · Zootaxa 3815 (2) © 2014 Magnolia Press

FIGURE 1. Synalpheus androsi Coutière, 1909, male (cl 3.4 mm) from off Espírito Santo, Brazil [MZUSP 31138]: A—frontal

region, dorsal; B—same, lateral; C—abdomen, lateral; D—telson, dorsal; E—third maxilliped, lateral; F—major (right)

cheliped, lateral; G—same, chela, mesial; H—minor (left) cheliped, lateral; I—same, carpus and chela, mesial; J—second

pereiopod, lateral; K—third pereiopod, lateral; L—same, distal merus, carpus and proximal propodus, ventrolateral; M—same,

distal propodus and dactylus, lateral; N—uropod, dorsal.

Zootaxa 3815 (2) © 2014 Magnolia Press · 217BRAZILIAN SPECIES OF SYNALPHEUS

Synalpheus androsi is unique within the Synalpheus gambarelloides (Nardo, 1847) species group in having a

deep excavation on the ventral surface of the third and fourth pereiopods, forming very pronounced lateral flanges.

On the third pereiopod, this excavation extends from the distal half of the merus to the proximal half of the

propodus, gradually flattening towards the dactylus, and is the deepest on the merus and the opposed surface of the

carpus (Fig. 1K, L). Thus the third pereiopods can be folded with the lateral and mesial flanges of the carpus fitting

into and mesial to, respectively, the excavation of the merus (Fig. 1K). This probably helps the shrimp to squat

more effectively and gain additional space in the most narrow sponge canals.

Noteworthy and in direct relationship to its morphological distinctiveness, S. androsi appears to have no close

relatives in the western Atlantic Ocean, forming a moderately supported clade with the eastern Atlantic S.

gambarelloides (Hultgren & Duffy 2011).

Synalpheus maxillispinus sp. nov.

(Figs. 2, 3)

Material examined. Holotype: female (cl 3.0 mm), Brazil, Abrolhos Bank off southern Bahia and northern

Espírito Santo, REVIZEE—Comissão Central: 2, R/V Astro Garoupa, station 17C, 18°34’00”S 38°04’00”W,

depth: 55 m, 22.XI.1997 [MZUSP 30093]. Paratypes: 1 male (cl 2.6 mm), same collection data as for holotype

[MZUSP 30094]; 1 male (cl 3.5 mm), Brazil, Espírito Santo, off Rio Doce, TAAF MD-55/Brasil 1987, station 51/

DC–87, 19°39’S 39°42’W, depth: 15 m, 29.V.1987 [MZUSP 30095].

Description. Rostrum slightly lower than orbital hoods, triangular, distinctly longer than wide at base,

subequal or slightly longer than orbital teeth, subacute distally; orbital teeth triangular, subacute, longer than wide

at base; frontal margin between orbital teeth and rostrum deeply concave, forming with rostrum V- to U-shaped

notches, adjacent rostro-orbital area slightly depressed (Fig. 2A, B, L).

Pleuron of first abdominal somite rounded posteroventrally, without hook; pleura of third to fifth somites also

rounded, fifth slightly more angular (Fig. 2C). Telson moderately broad, about 1.5 times as long as anterior width,

tapering distally; dorsal surface with broad, longitudinal, median depression and two pairs of strong spiniform

setae inserted at about 0.4 and 0.7 length of telson, respectively; posterior margin rounded, less than half-length of

anterior width, with two stout spiniform setae, mesial much longer than lateral (Fig. 2D).

Antennular peduncle with second article 1.3–1.6 times as long as wide; stylocerite falling short of distal margin

of first article, with blunt or subacute tip (Fig. 2A, L). Antennal basicerite with blunt, non-projecting distodorsal

tooth, and strong distolateral tooth reaching at least to middle of second article of antennular peduncle;

scaphocerite with strong broad distolateral tooth reaching beyond end of antennular peduncle; blade always

present, slender, not reaching mid-length of scaphocerite, slightly overreaching distolateral tooth of basicerite,

fringed with short setae (Fig. 2A, B, L).

Mouthparts typical for genus. Third maxilliped with ultimate article bearing one dorsal row of six short

spiniform setae, one ventral row of three longer spiniform setae and crown of seven or so stouter spiniform setae on

apex (Fig. 2E, F).

Major cheliped with palm 2.5–2.8 as long as dactylus; distodorsal tooth on palm not greatly protruding, ending

blunt distally, without acute point; dactylus reaching distinctly beyond pollex (Fig. 3A–E). Minor cheliped with

palm about 1.3 times as long as fingers, latter with cutting surfaces excavated and bidentate tips; dactylus with very

few tufts of setae not organised in rows (Fig. 3F, G).

Second pereiopod with five-articulated carpus, first carpal article about four times length of second, about as

long as sum of second to fifth articles (Fig. 2G). Third pereiopod with merus about three times as long as wide;

propodus with six spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin;

dactylus biunguiculate, moderately stout, secondary unguis subparallel to and shorter than terminal unguis (Fig.

2H, I). Fifth pereiopod with propodus armed with five spiniform setae on ventral margin; grooming brush well

developed (Fig. 2J).

Uropodal exopod with distolateral margin bearing two strong teeth (including lateral tooth of diaeresis) and

one or two spiniform seta(e) (Fig. 2K, M).

Colour in life unknown.

Type locality. Brazil: Abrolhos Bank off southern Bahia / northern Espírito Santo (holotype and first paratype)

and off Rio Doce, Espírito Santo (second paratype).

ANKER & PACHELLE218 · Zootaxa 3815 (2) © 2014 Magnolia Press

FIGURE 2. Synalpheus maxillispinus sp. nov., A–K, holotype, female (cl 3.0 mm) from Abrolhos Bank, Brazil [MZUSP

30093]; L, M, paratype, male (cl 3.5 mm) from Rio Doce, Espírito Santo, Brazil [MZUSP 30095]: A—frontal region, dorsal;

B—same, lateral; C—abdomen, lateral; D—telson, dorsal; E—third maxilliped, lateral; F—same, carpus and ultimate article,

mesial; G—second pereiopod, lateral; H—third pereiopod, lateral; I—same, distal propodus and dactylus, lateral; J—fifth

pereiopod, lateral; K—uropod, dorsal; L—frontal region, dorsal; M—uropod, dorsal.

Zootaxa 3815 (2) © 2014 Magnolia Press · 219BRAZILIAN SPECIES OF SYNALPHEUS

FIGURE 3. Synalpheus maxillispinus sp. nov., A, B, holotype, female (cl 3.0 mm) from Abrolhos Bank, Brazil [MZUSP

30093]; C–G, paratype, male (cl 3.5 mm) from Rio Doce, Espírito Santo, Brazil [MZUSP 30095]: A—major (right) cheliped,

lateral; B—same, chela, mesial; C—major (left) chela, lateral; D—same, mesial; E—same, distal palm and fingers, lateral;

F—minor (right) cheliped, lateral; G—same, carpus and chela, mesial.

Distribution. Southwestern Atlantic: presently known only from two localities off eastern Brazil (southern

Bahia and Espírito Santo).

Etymology. The specific name refers to the presence of the characteristic spiniform setae on the ultimate

article of the third maxilliped; used as an adjective.

Ecology. The type specimens were dredged from moderate depths of 15–50 m, without records of their sponge

hosts.

Remarks. Synalpheus maxillispinus sp. nov. plainly belongs to the Synalpheus paraneptunus species complex

recently revised by Anker & Tóth (2008) and with one additional species subsequently added by Hultgren et al.

(2011). The S. paraneptunus species complex was originally defined by three morphological characters: (1) the

relatively long stylocerite, almost reaching or overreaching the distal margin of the first article of the antennular

peduncle; (2) the minor chela fingers broadened and spoon-shaped, i.e. with hollowed cutting surfaces; and (3) the

minor chela dactylus with dorsal and lateral setae not organised in rows forming a dense brush, usually forming a

single row of scarce setae. Anker & Tóth (2008) recognised additional five species in the complex: Synalpheus

duffyi Anker & Tóth, 2008; Synalpheus riosi Anker & Tóth, 2008; Synalpheus brevidactylus Anker & Tóth, 2008;

ANKER & PACHELLE220 · Zootaxa 3815 (2) © 2014 Magnolia Press

Synalpheus bocas Anker & Tóth, 2008; and Synalpheus belizensis Anker & Tóth, 2008, all from the Caribbean Sea

(Panama, Belize, Dominica). Hultgren et al. (2011) added a seventh species, Synalpheus microneptunus Hultgren,

Macdonald & Duffy, 2011, from Barbados. The S. paraneptunus complex was shown to form a well-supported

clade within the S. gambarelloides group in the combined molecular-morphological analysis of Hultgren & Duffy

(2011). However, and interestingly, this clade also contained a morphologically more distinctive Synalpheus

kensleyi (Ríos & Duffy, 2007), which unlike other species of the S. paraneptunus complex is characterised by a

rather well-developed brush of gambarelloid setae and a strong acute tooth projecting from the dorsal margin of the

antennal basicerite.

Synalpheus maxillispinus sp. nov. is morphologically closest to S. bocas and S. belizensis, sharing with them

the presence of two teeth on the distolateral margin of the uropodal exopod, adjacent to spiniform seta. The new

species can be separated from S. bocas and S. belizensis by the shorter stylocerite, not reaching (vs. overreaching)

distal margin of the first article of the antennular peduncle; the major chela with a short, bluntly ending (vs. more

projecting and sharply pointed) distodorsal tooth; specifically from S. bocas by the scaphocerite having (vs. not

having) a blade1; and the shorter dorsal spiniform setae on the telson; and specifically from S. belizensis by the

more slender third pereiopod, with merus about three (vs. 2.5) times as long as wide (cf. Anker & Tóth 2008).

The presence of two teeth on the distolateral margin of the uropodal exopod separates Synalpheus

maxillispinus sp. nov. from S. paraneptunus, S. duffyi, S. riosi, S. brevidactylus, S. microneptunus and S. kensleyi,

which all have three teeth. In addition, S. maxillispinus sp. nov. can be separated from S. microneptunus by the

presence of five (vs. four) articles in the carpus of the second pereiopod; from S. kensleyi by the minor chela

without setae organised in rows (vs. with distinct gambarelloid setae) and the antennal basicerite without (vs. with)

acutely projecting dorsal tooth; from S. duffyi and S. brevidactylus by the moderately slender, distally blunter (vs.

more slender and acute) orbital teeth and the major chela palm with a bluntly ending (vs. sharply pointed)

distodorsal tooth; from S. riosi by the major chela palm with a dorsally non-swollen, bluntly ending (vs. dorsally

swollen and with a small sharp point) distodorsal tooth and the minor chela without setae grouped in rows (vs. with

setae organised in a single dorsal row) (cf. Ríos & Duffy 2007; Anker & Tóth 2008; Hultgren et al. 2011).

The presence of three spiniform setae on the ventral surface of the ultimate article of the third maxilliped

appears to be another important and possibly diagnostic character of Synalpheus maxillispinus sp. nov. Anker &

Tóth (2008) did not illustrate these setae in S. paraneptunus, S. duffyi, S. riosi and S. belizensis, although some

stout setae in the same position can be seen in their figure 4C of S. duffyi. In addition, Hultgren et al. (2011)

illustrated slender spiniform setae (clearly thicker than adjacent serrulate setae) on the third maxilliped of S.

microneptunus. Therefore, these spiniform setae may also be present, although less developed, in other species of

the S. paraneptunus complex.

All species of the Synalpheus paraneptunus complex are associated with sponges (see Table 1 in Anker et al.

2012 for sponge records) and at least three of them (S. duffyi, S. riosi and S. microneptunus) live in social colonies

(Anker & Tóth 2008; Hultgren et al. 2011; Anker et al. 2012). The sponge hosts of S. paraneptunus, S. riosi and S.

maxillispinus sp. nov. remain unknown; all sponges listed as hosts for S. paraneptunus in Table 1 of Anker et al.

(2012) are in fact sponge records for S. paraneptunus sensu lato and not for S. paraneptunus sensu stricto as

redefined by Anker & Tóth (2008). Whether S. maxillispinus sp. nov. is a social or pair-living species presently

remains unknown.

Synalpheus cf. ruetzleri Macdonald & Duffy, 2006

(Fig. 4)

Synalpheus ruetzleri Macdonald & Duffy 2006: 8, figs. 8–13; Anker et al. 2012: 85 (table), 88 (key).

Zuzalpheus ruetzleri—Ríos & Duffy 2007: 61, pl. 4.

1. �Anker & Tóth (2008, p. 21) obviously made a type-error in the diagnosis of Synalpheus belizensis by stating that the

antennal scaphocerite has no blade. Their illustration of the frontal region of the holotype of S. belizensis (fig. 12A)

clearly shows both scaphocerites having a narrow blade and the presence / absence of a scaphocerite blade is also

used to distinguish S. belizensis from S. bocas under remarks (p. 24) and in the key (p. 27). However, an aberrant

specimen of S. belizensis in Anker & Tóth's fig. 13H has no blade on the left scaphocerite and a reduced blade on the

right scaphocerite.

Zootaxa 3815 (2) © 2014 Magnolia Press · 221BRAZILIAN SPECIES OF SYNALPHEUS

FIGURE 4. Synalpheus cf. ruetzleri Macdonald & Duffy, 2006, male (cl 4.1 mm) from off Fortaleza, Brazil [MZUSP 31501]:

A—frontal region, dorsal; B—abdomen, lateral; C—telson, dorsal; D—third maxilliped, lateral; E—major (right) cheliped,

lateral; F—same, chela, mesial; G—minor (left) cheliped, lateral; H—same, carpus and chela, mesial; I—second pereiopod,

lateral; J—third pereiopod, lateral; K—same, distal propodus and dactylus, lateral; L—left uropod, dorsal; M—right uropod,

distolateral margin of exopod, dorsal.

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Material examined. 1 male (cl 4.1 mm), Brazil, Ceará, off Fortaleza, Parque Estadual Marinho da Pedra da Risca

do Meio, station Dive II, in Agelas dispar, leg. L.E.A. Bezerra, 14.VII.2004 [MZUSP 31501].

Description. See Macdonald & Duffy (2006) for description and illustrations and Ríos & Duffy (2007, under

Zuzalpheus ruetzleri) for diagnosis and colour photograph; comparative illustrations of the Brazilian specimen are

provided in Fig. 4.

Distribution. Western Atlantic: Caribbean Sea (Panama: San Blas Islands, Belize: Carrie Bow Cay); possibly

Brazil (Ceará) (Macdonald & Duffy 2006; Ríos & Duffy 2007; present study).

Ecology. In Belize, Synalpheus ruetzleri was found exclusively in the demosponge Hymeniacidon cf. caerulea

Pulitzer-Finali, in very shallow water, 1–2 m (Ríos & Duffy 2007). In contrast, the Ceará material was found inside

a different sponge, Agelas dispar Duchassaing & Michelotti, collected by scuba diving in less than 20 m, whereas

in Panama, S. ruetzleri appears to associate with various sponges, including H. cf. caerulea (Macdonald & Duffy

2006).

Remarks. The present material, if confirmed to be conspecific with the Caribbean Synalpheus ruetzleri, may

represent the first record of this species in the southwestern Atlantic and Brazilian waters. Like most other species

of the taxonomically difficult Synalpheus brooksi Coutière, 1909 species complex, S. ruetzleri appears to be

morphologically highly variable, as exemplified by differences in the shape of the orbital teeth (Macdonald &

Duffy 2006: figs. 8A, 13A) and distodorsal protuberance of the major chela (idem., figs. 10A, D, G, 13C). The

presence of a small scaphocerite blade separates this species from the otherwise almost identical Synalpheus

bousfieldi Chace, 1972 and Synalpheus idios (Ríos & Duffy, 2007). It also differs from the supposedly variable S.

brooksi, for instance, by the shape and direction of the distodorsal protuberance of the major chela (more pointed

and directed somewhat upward in S. brooksi) in addition to the absence of a scaphocerite blade (Macdonald &

Duffy 2006).

The single male specimen from Ceará matches only partly the description and illustrations provided by

Macdonald & Duffy (2006). The shape of the frontal margin of the carapace is rather similar as are most features of

the antennular and antennal peduncles (cf. Fig. 4A and Macdonald & Duffy 2006: fig. 8A, D). The telson of the

Ceará male is rather different from those of the male holotype and female allotype from Belize, especially in the

narrower posterior half (cf. Fig. 4C and Macdonald & Duffy 2006: fig. 8B, G). However, in the number (4 + 5) and

position of dorsal spiniform setae, the telson of the Ceará specimen is clearly aberrant (Fig. 4C). Another important

difference with the type material is the shape of the distodorsal protuberance of the major chela, which is relatively

short and straight in the Brazilian specimen compared to the dorsally slightly swollen and ending in a small,

downward directed point in both the holotype and allotype (cf. Fig. 4E, F and Macdonald & Duffy 2006: fig. 10A,

D, F, G). However, in this last feature, the Ceará specimen approaches the non-type male of S. ruetzleri from

Panama (cf. Fig. 4E, F and Macdonald & Duffy 2006: fig. 13C). Noteworthy, the armature of the distolateral

margin of the uropodal exopod is variable (one or two teeth) in the Ceará specimen (Fig. 4L, M).

The paucity of the Brazilian material (presently one single male specimen) does not allow to evaluate the

importance of these morphological differences. Sequencing of the barcoding gene segment (COI) of the Ceará

specimen is highly desirable to test the correctness of the present identification.

Synalpheus tenuispina Coutière, 1909

(Figs. 5, 6)

Synalpheus latastei tenuispina Coutière 1909: 26, fig. 8; Chace 1972: 93; Coelho & Ramos 1972: 150; Christoffersen 1980:

171; Christoffersen 1998: 362.

(?) Synalpheus latastei tenuispina—Abele 1976: 273; Martínez-Iglesias et al. 1993: 12; Martínez-Iglesias et al. 1996: 35.

Synalpheus tenuispina—Crosnier & Forest 1965: 364; Ríos & Duffy 2007: 78 (key); Anker et al. 2012: 9, 86 (table), 90 (key).

Material examined. 1 male (cl 7.1 mm), Brazil, São Paulo, São Sebastião, Pier da Petrobras, 12.XII.2009

[MZUSP 31503]; 1 male (cl 6.9 mm, major P1 missing), 1 heavily damaged specimen (only major P1, P2-P5 and

sternal portion of cephalothorax), Brazil, Rio de Janeiro, Niteroi, Baia de Guanabara, Praia de Boa Viagem, scuba

diving, fauna associated with Polychaeta, coll. J.B. Mendonça Jr., 04.X.2006 [MZUSP 22133].

Description. Rostrum with base clearly delimited from anteromesial margin of orbital hoods, much longer

than wide at base, arrow-shaped in dorsal view, longer than orbital teeth, acute distally, tip overreaching mid-length

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of first article of antennular peduncle; orbital teeth large, triangular, acute distally, about as long as wide at base;

anteromesial margin between orbital teeth and rostrum abruptly sloping ventrally, forming deep V-shaped notch in

dorsal view; rostro-orbital process well developed (Fig. 5A, B).

Pleuron of first abdominal somite pointed posteroventrally, forming feeble hook; pleura of third to fifth

somites also rounded, fifth slightly more angular (Fig. 5C). Telson broad, about 1.3 times as long as anterior width,

slightly tapering distally; dorsal surface with shallow, longitudinal, median depression and two pairs of stout

spiniform setae inserted at about 0.45 and 0.75 length of telson, respectively; posterior margin broadly rounded,

about 0.6 length of anterior width, with two stout spiniform setae, mesial much longer than lateral (Fig. 5D).

Antennular peduncle with second article 1.5 times as long as wide; stylocerite very strong, reaching far beyond

distal margin of first article, with acute tip (Fig. 5A). Antennal basicerite with acutely projecting distodorsal tooth

and much stronger distolateral tooth, latter reaching well into first third of second article of antennular peduncle

and almost equalling tip of stylocerite; scaphocerite with very strong distolateral tooth reaching beyond end of

antennular peduncle; blade well-developed, moderately broad, reaching to about 0.7 length of scaphocerite, fringed

with long setae; carpocerite reaching slightly beyond end of antennular peduncle but not beyond distolateral tooth

of scaphocerite (Fig. 5A, B).

Mouthparts typical for genus. Third maxilliped with very short penultimate article; ultimate article with crown

of about six stout spiniform setae on apex (Fig. 5E, F).

Major cheliped with merus about 2.3 times as long as wide; palm ovoid, 2.3–2.4 as long as dactylus;

distodorsal margin of palm slightly protruding, forming low blunt protuberance, without acute tooth; dactylus

subequal to pollex (Fig. 6A, B). Minor cheliped with merus about 2.3 times as long as greatest width, with convex

dorsal and ventral margins, distodorsal margin projecting as sharp tooth; carpus cup-shaped; palm about 1.2 times

as long as fingers; dactylus and pollex with sharp cutting edges, laterodorsal surface of pollex slightly depressed

(Fig. 6C, D).

Second pereiopod with five-articulated carpus, first carpal article about four times length of second, equal to

sum of second to fifth articles (Fig. 5G). Third pereiopod with merus almost four times as long as wide; carpus with

stout spiniform seta distoventrally; propodus with eight spiniform setae on ventral margin and one pair of

spiniform setae on distoventral margin; dactylus biunguiculate, moderately stout, secondary unguis subparallel to

and shorter than terminal unguis (Fig. 5H, I). Fifth pereiopod with propodus armed with four spiniform setae on

ventral margin and one pair of spiniform setae on distoventral margin, and with well-developed grooming brush

laterally, consisting of at least six rows of serrulate setae (Fig. 5J, K).

Uropod with protopod ending in large subacute tooth laterally; distolateral margin of exopod with two strong

teeth (including lateral tooth of diaeresis) and one stout spiniform seta (Fig. 5L).

Colour in life unknown.

Distribution. Southwestern Atlantic: Brazil (Rio de Janeiro, São Paulo, Santa Catarina). Records of S.

tenuispina from the Caribbean Sea (Abele 1976; Martínez-Iglesias et al. 1993, 1996) most likely refer to a species

currently known as Synalpheus cf. africanus Crosnier & Forest, 1965, and not to S. tenuispina (see below).

Ecology. Synalpheus tenuispina appears to be a non-symbiotic species, dwelling in pier growth or in crevices

of polychaete reefs.

Remarks. Coutière's (1909) original description of Synalpheus latastei tenuispina was based on a single large

female (tl 30 mm) from Desterro, now Florianópolis, Santa Catarina, Brazil. The description is actually a short

paragraph with comparisons of proportions and lengths between the new form and the supposedly closely related,

eastern Pacific Synalpheus latastei Coutière, 1909 (now junior synonym of Synalpheus spinifrons (H. Milne

Edwards, 1837)) and Synalpheus lockingtoni Coutière, 1909, accompanied by a few figures. The holotype of S.

latastei tenuispina was deposited in the Muséum National d'Histoire Naturelle in Paris, France (MNHN), where,

however, it was missing already in the 1960s (Crosnier & Forest 1965) and was also not found during a more recent

search in 2007 (A. Anker, pers. obs.). Based on the morphological differences between S. latastei tenuispina and S.

spinifrons in Coutière's (1909) account, Crosnier & Forest (1965) elevated the Brazilian form to full species, as S.

tenuispina.

The most complete specimen of the present material is an adult male (cl 7.1 mm, tl ~25 mm) from São

Sebastião (Figs. 5, 6). This male agrees well with figures of the missing female holotype in Coutière (1909), except

for the slightly broader rostrum and stouter merus of the third pereiopod (cf. Fig. 5A, H and Coutière 1909: fig. 8a,

m). Obviously more fresh material is needed to understand the range of morphological variation of S. tenuispina

and also to document the colour pattern of the species.

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FIGURE 5. Synalpheus tenuispina Coutière, 1909, male (cl 7.1 mm) from São Sebastião, Brazil [MZUSP 31503]: A—frontal

region, dorsal; B—same, lateral; C—abdomen, lateral; D—telson, dorsal; E—third maxilliped, lateral; F—same, tip of ultimate

article, mesial; G—second pereiopod, lateral; H—third pereiopod, lateral; I—same, distal propodus and dactylus, lateral;

J—fifth pereiopod, lateral; K—same, propodus and dactylus; L—uropod, dorsal.

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FIGURE 6. Synalpheus tenuispina Coutière, 1909, male (cl 7.1 mm) from São Sebastião, Brazil [MZUSP 31503]: A—major

(left) cheliped, lateral; B—same, chela, mesial; C—minor (right) cheliped, lateral; D—same, carpus and chela, mesial.

The Caribbean records of S. latastei tenuispina (Abele 1976; Martínez-Iglesias et al. 1993, 1996, all as S.

latastei tenuispina) are difficult to assess without examining the material. They most probably refer to a

problematic amphi-Atlantic taxon (possibly more than one species) currently known as Synalpheus cf. africanus

(cf. Anker et al. 2012: fig. 3) and not to S. tenuispina. The differences between S. cf. africanus and S. tenuispina

are rather slight. Anker et al. (2012) noted that the specimens tentatively identified as S. cf. africanus from Atol das

Rocas, Aruba and Panama were closer to S. africanus than to S. tenuispina in the length of the scaphocerite and

shape of the rostrum, and were also much smaller than S. tenuispina (maximum tl ~15 mm compared to 30 mm for

the female holotype of S. tenuispina). As Anker et al. (2012) pointed out, the entire S. africanus—S. tenuispina

complex needs a molecular analysis, but for the time being it is more sensible to treat the larger southern Brazilian

form as S. tenuspina.

Synalpheus ul (Ríos & Duffy, 2007)

Zuzalpheus ul Ríos & Duffy 2007: 63, figs. 27–30, pl. 5.

Synalpheus ul—Macdonald et al. 2009: 50; Hultgren et al. 2010: 234, 251, fig. 9A–F, pl. 6A–D; Almeida et al. 2012: 19, figs.

6–8; Anker et al. 2012: 72, figs. 47–49.

Synalpheus cf. pandionis—Almeida et al. 2007: 15, figs. 3, 4 (not S. pandionis Coutière 1909, see Almeida et al. 2012).

Synalpheus yano—Macdonald et al. 2009: 51, pl. 6A, B (not S. yano (Ríos & Duffy, 2007), see Hultgren & Duffy 2010: p. 3).

Material examined. 3 males (cl 6.4, 6.1, 3.6 mm), Brazil, Pernambuco, SE of Recife, R.V. “Calypso” station 25,

08º22’S 34º44’W, depth 38–52 m, sand and mud, 21.XI.1961 [MZUSP 4552].

Description. See Ríos & Duffy (2007) for detailed description and illustrations; see also Almeida et al. (2012)

for illustrations of the material from Bahia and Anker et al. (2012) for additional colour photographs.

Distribution. Western Atlantic: Caribbean Sea: Belize, Panama, Jamaica, Dominican Republic, St. Martin;

Brazil: Pernambuco, Alagoas, Bahia (Ríos & Duffy 2007; Macdonald et al. 2009; Hultgren et al. 2010; Almeida et

al. 2012; Anker et al. 2012; present study).

Ecology. Synalpheus ul is an obligate sponge symbiont (see Anker et al. 2012 for all sponge records); the

“Calypso” material was dredged from 38–52 m, without notes on association with sponges.

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Remarks. The present specimens from Pernambuco, collected by the French research vessel “Calypso” in

1961, were misidentified as Synalpheus longicarpus (Herrick, 1891) and included by Christoffersen (1979) as part

of his extensive material of S. longicarpus. It is also likely that several other records of S. longicarpus in

Christoffersen (1979) and perhaps in other studies are based on misidentified material. Therefore, the entire

Brazilian material of S. longicarpus, including all the remaining material in Christoffersen (1979), must be re-

examined. Synalpheus ul was previously reported from Brazil, from Camamu Bay, Bahia (Almeida et al. 2007,

2012) and Maceió, Alagoas (Anker et al. 2012); this is the first record of this species from Pernambuco.

Synalpheus yano (Ríos & Duffy, 2007)

(Figs. 7, 8)

Zuzalpheus yano Ríos & Duffy 2007: 69, figs 31–34, pl. 5.

Synalpheus yano—Hernáez et al. 2010: 692, fig. 2; Hultgren et al. 2010: 234; Anker et al. 2012: 76, fig. 50.

Not Synalpheus yano—Macdonald et al. 2009: 51, pl. 6A, B (= S. ul, see Hultgren & Duffy 2010: p. 3).

Material examined. 2 ov. females (cl 4.3 mm, 3.5 mm), Brazil, Ceará, ~15 km N-NE of Fortaleza, scuba diving, in

coral rubble, coll. C. Cerqueira & M. Soares, 21.IV.2013 [MZUSP 31142]; 1 male (cl 3.9 mm), same collection

data as for previous specimens [MZUSP 31141]; 1 ov. female (cl ~4.5 mm), USA, Florida, N-NW of St.

Petersburg, south of Big Bend area, station FI02012-01, 28.55667 N -84.2745 W, dredge, hard bottom, rocks and

coral rubble, depth: 26–30 m, coll. J. Slapcinsky, 24.V.2012 [FLMNH UF 31631].

Description. See Ríos & Duffy (2007) for description and illustrations and Anker et al. (2012) for colour

photographs; comparative illustrations of the Brazilian material and an additional colour photograph of a specimen

from western Florida are provided in Figs. 7, 8.

Distribution. Western Atlantic: Gulf of Mexico (Mexico: Alacranes Reef, USA: western Florida); Caribbean

Sea (Belize, Panama, Jamaica); Brazil (Ceará) (Ríos & Duffy 2007; Anker et al. 2012; present study).

Ecology. Synalpheus yano is an obligate sponge symbiont (see Anker et al. 2012 for all sponge records); all

specimens from Ceará and the single specimen from Florida were extracted from coral rubble that likely contained

cryptic host sponges.

Remarks. The present material represents the first record of Synalpheus yano in Brazil and the southwestern

Atlantic (MZUSP material), as well as USA and the eastern Gulf of Mexico (FLMNH UF material). This species is

closely related to Synalpheus pandionis Coutière, 1909 and Synalpheus ul, which both also occur in Brazil

(Almeida et al. 2012; Anker et al. 2012; see also above), but can be separated from them by the absence of a

scaphocerite blade, and from S. pandionis also by the absence of a rounded protuberance on the mesial face of the

major chela.

Synalpheus sp. gambarelloides group

(Fig. 9)

Material examined. 1 ov. female (cl 5.0 mm, missing posterior part of abdomen and tail fan), Brazil, Ceará, off

Fortaleza, Parque Estadual Marinho da Pedra da Risca do Meio, station Dive II, in Agelas dispar, leg. L.E.A.

Bezerra, 14.VII.2004 [MZUSP 31502].

Remarks. Unfortunately, this adult female from Ceará is damaged, missing the two posterior somites of the

abdomen, telson and uropods. The short stylocerite (Fig. 9A) and the minor chela with well-developed

gambarelloid setae (Fig. 9E) indicate clearly that this specimen belongs to the Synalpheus gambarelloides species

group. However, the combination of the frontal region of the carapace with a short rostral carina extending

posteriorly to about mid-length of corneas and flanked by shallow rostro-orbital depressions (Fig. 9A); the antennal

scaphocerite without a blade and equalling the distolateral tooth of the basicerite (Fig. 9A); the major chela with

distodorsal protuberance poorly developed and bluntly rounded (Fig. 9C, D); and the five-articulated carpus of the

second pereiopod (Fig. 9F) do not allow to assign it unambiguously to any of the species reported so far from

Brazil or the western Atlantic. With a single damaged specimen on hand it is impossible to say whether it belongs

to an undescribed or a morphologically variable species.

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FIGURE 7. Synalpheus yano (Ríos & Duffy, 2007), male (cl 3.9 mm) from off Fortaleza, Brazil [MZUSP 31141]: A—frontal

region, dorsal; B—abdomen, lateral; C—telson, dorsal; D—major (left) cheliped, lateral; E—same, chela, mesial; F—minor

(right) cheliped, lateral; G—same, carpus and chela, mesial; H—second pereiopod, lateral; I—third pereiopod, lateral;

J—same, distal propodus and dactylus, lateral; K—uropod, dorsal.

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FIGURE 8. Synalpheus yano (Ríos & Duffy, 2007): A—ovigerous female (cl 4.3 mm) from Ceará, Brazil [MZUSP 31142],

lateral view; B—ovigerous female (cl ~4.5 mm) from St. Petersburg area, western Florida, USA [FLMNH UF 31631]:

dorsolateral view. Photographs courtesy of F. Monteiro (A) and J. Slapcinsky (B).

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FIGURE 9. Synalpheus sp. gambarelloides group, ovigerous female (cl 5.0 mm) from off Fortaleza, Brazil [MZUSP 31502]:

A—frontal region, dorsal; B—anterior margin of carapace, lateral; C—major (left) chela, mesial; D—same, distal palm and

fingers chela, lateral; E—minor (right) cheliped, lateral; F—second pereiopod, lateral; G—third pereiopod, lateral; H—same,

distal propodus and dactylus, lateral.

Acknowledgements

We are indebted to Marcos Tavares (MZUSP), who invited the senior author to study the MZUSP caridean

collection under the “Coleções e Acervos Científicos” programme of MZUSP, and who also coordinated the

REVIZEE and TAAF campaigns during which some of the above reported material was collected. Mauro Cardoso

Júnior and Joana d’Arc de Jesus Pinto (MZUSP) helped finding caridean specimens in the MZUSP collection and

retrieving collection data. Jesser Fidelis (MOUFPE) confirmed a previous record of S. androsi from Bahia by

locating and examining material reported by Coelho et al. (2006). Colour photographs were taken by Felipe

Monteiro (UFC) and John Slapcinsky (FLMNH). Amanda Bemis (FLMNH) provided collection data for the

specimen of S. yano collected in Florida. Matthew R. McClure (Lamar State College, Orange, USA) kindly

reviewed the originally submitted manuscript.

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