Alpheus intrinsecus Spence Bate, 1888

Cunha, Andressa Maria, Terossi, Mariana, Mantelatto, Fernando L. & Almeida, Alexandre O., 2017, Morphological and molecular analyses support the amphi-Atlantic distribution and taxonomic status of the snapping shrimp Alpheus intrinsecus Spence Bate, 1888 (Crustacea: Decapoda: Alpheidae), Zootaxa 4303 (4), pp. 573-589 : 577-580

Alpheus intrinsecus Spence Bate, 1888

( Figs. 1–2 )

Alpheus intrinsecus Spence Bate 1888, p. 557 , pl. 100, fig. 1; Osorio 1892: p. 201, 313; 1898, p. 186, 194; Coutière 1899, p. 44, 94–97, 147, 236, 237, 352, fig. 64; Rathbun 1900, p. 313; Moreira 1901, p. 10; 1906, p.131; Balss 1916, p. 20; 1925, p. 292, fig. 75; Monod 1927, p. 594; Holthuis 1951, p. 87, 88; 1952, p. 44; 1959, p.103; Sourie 1954, p. 112, 253; Buchanan 1958, p. 19–21; Longhurst 1958, p. 91; Crosnier & Forest 1964, p. 358; 1965, p. 607; 1966, p. 286, fig. 26a–e; Fausto- Filho 1970, p. 56; 1974, p. 13; 1978, p. 66; 1980, p. 113; Chace 1972, p. 68; Coelho & Ramos 1972, p. 148; Christoffersen 1979, p. 316, figs. 9, 10; 1980, p. 71, figs. 16, 17; 1982, p. 105, 106; 1998, p. 358; Coelho et al. 1980, p. 63; 2006, p. 51; Ramos-Porto 1980, p. 299; Rodriguez 1980, p. 147, fig. 594; Sampaio & Fausto-Filho 1984, p. 13; Martinez-Iglesias et al. 1997, p. 425; Coelho-Santos & Coelho 1998, p. 75, figs. 14, 15; Costa et al. 2000, p. 779; 2016, p. 616; Prieto et al. 2000, p. 236; Anker 2001, p. 202; Almeida et al. 2006, p. 6, 9, 19, 24; 2007a, p. 13; 2007b, p. 9, 18; 2012, p. 11; 2013b, p. 1399, 1401; Felder et al. 2009, p. 1057; De Grave & Fransen 2011, p. 384; Boos et al. 2012, p. 1024; Santos et al. 2012, p. 149, 151, fig. 3f; Soledade & Almeida 2013, p. 101, 107, 113, 114, fig. 5e; Furlan et al. 2013, p. 87, 89; Pantaleão et al. 2016, p. 386; Santos et al. 2016, p. 5; Mantelatto et al. 2016, p. 313, 315; Anker et al. 2016, p. 6.

Alpheus intrinseccus Luederwaldt 1919, p. 430 ; 1929, p. 52 (erroneous spelling).

Crangon intrinsecus Schmitt 1926, p. 23 .

Type material. Not examined (deposited in the Natural History Museum, London, England [ NHMUK 1888.22 ]) .

Material examined. WESTERN ATLANTIC— Mexico. 1 m, Morro de la Mancha, Mpio of Actopan, Veracruz , col. R. Álvarez, 16.v.1996, CNCR 20277 (specimen used in genetic analysis) ; Venezuela. 1 m, Sucre, Rio de Oro , 16–20 m, Mercator XIV expedition, 30.xii.1937, RMNH . Crus.D. 8588; Suriname. 1 m, Suriname River , 7 m, 3.v.1957, RMNH . Crus.D. 11462 (specimen used in genetic analysis); Brazil. 1 ovf, 1 m, Pernambuco, Goiana, Ponta de Pedras Beach , on the sponge Desmapsamma anchorata , col. U. Pinheiro, 16.v.2014, MOUFPE 15626 ; 1 f, Bahia, Camamu , 06.viii.2005, UESC 705; 1 ovf, Bahia, Ilhéus, São Domingos, col. R . O. Vasques, 02.v.2003, UESC 188; 1 ovf, Bahia, Ilhéus, Ilhéus coast, 21.i.2004, UESC 292; 2 ovf, 1 f, Bahia, Ilhéus , Cachoeira River , 18.ix.2004, UESC 584; 1 m, 1 ovf, Bahia, Ilhéus , Cachoeira River , 13.i.2005, UESC 619; 2 m, 1 ovf, Bahia, Ilhéus , Malhado Beach , col. F.J. Guimarães, 20.ix.2011, UESC 1571 (one specimen used in genetic analysis); 2 m, 2 ovf, Bahia, Prado , Corumbau , 27.iv.2006, UESC 1572 (one specimen used in genetic analysis); 1 f, Rio de Janeiro, col. A.O. Almeida, 05.xii.2007, MOUFPE 15610 (specimen used in genetic analysis) ; 1 ovf, 1 f, 2 m, São Paulo, Santos , cols. A.L. Castilho et al., 24.x.2011, CCDB 1591 ; 1 m, 1 ovf, São Paulo, Ubatuba, Ubatuba Bay , cols. F.L. Mantelatto et al., 03.v.2007, CCDB 1951 ; 2 m, 3 ovf, São Paulo, Ubatuba, Ubatuba Bay , col. F.L. Mantelatto, 18.xiii.2007, CCDB 2219 ; 2 ovf, São Paulo, Ubatuba, Ubatuba Bay , cols. F.L. Mantelatto et al., 02.vi.2008, CCDB 2357 ; 2 ovf, 1 f, São Paulo, Ubatuba, Ubatuba Bay , col. F.L. Mantelatto, 13.iii.2005, CCDB 3410 ; 1 ovf, 1 f, São Paulo, Ubatuba, Ubatuba Bay , 7 m, col. F.L. Mantelatto, 27.v.2009, CCDB 3496 ; 3 m, 1 ovf, São Paulo, Ubatuba, Ubatuba Bay , 04.v.2009, CCDB 3497 ; 3 m, 5 f, São Paulo, Ubatuba, Ubatuba Bay , col. F.L. Mantelatto, 12.v.2010, CCDB 3499 (one specimen used in genetic analysis) ; 1 f, São Paulo, Ubatuba, Ubatuba Bay , 25 m, col. F.L. Mantelatto, viii.2002, CCDB 3500 ; 2 m, São Paulo, Ubatuba, Ubatuba Bay , col. F.L. Mantelatto, 20.viii.2002, CCDB 3502 ; 1 m, 1 f, São Paulo, Ubatuba, Ubatuba Bay , col. F.L. Mantelatto, 23.iv.2012, CCDB 3957 ; 1 f, São Paulo, Ubatuba, Grande Beach , col. F.L. Mantelatto, 01.v.2002, CCDB 1126 ; 2 ovf, São Paulo, Ubatuba, Anchieta Island , cols. F.L. Mantelatto et al., 01.viii.2002, CCDB 1416 .; 1 m, 2 f, São Paulo, Ubatuba, Anchieta Island , cols. F.L. Mantelatto & R . Biagi , iii.2001, CCDB 3490 ; 1 f, São Paulo, Ubatuba, Anchieta Island , cols. F.L. Mantelatto & R . Biagi , xi.2002, CCDB 4096 ; 1 f, São Paulo, Ubatuba, Ubatuba Bay , Cedro Beach, cols. R . C. Costa et al., 07.viii.2011, CCDB 425 ; 7 m, 6 ovf, São Paulo, Ubatuba, Ubatuba Bay, Cedro Beach , col. F.L. Mantelatto, 04.iv.2011, CCDB 3414 ; 2 m, 1 f, São Paulo, Ubatuba, Ubatuba Bay , Cedro Beach, cols. A.L. Castilho & R . C. Costa , 06.vii.2011, CCDB 3441 ; 3 m, 11 f, São Paulo, Ubatuba, Ubatuba Bay, Cedro Beach , cols. A.L. Castilho et al., 07.vii.2011, CCDB 3817 ; 3 m, 18 f, São Paulo, Ubatuba , offshore, col. D. Rosa, 05.ix.2011, CCDB 3618 ; 1 m, São Paulo, Caraguatatuba, Caraguatatuba Bay , cols. A. Meireles & R . Biagi 01.ix.2002, CCDB 3504 ; 1 ovf, Santa Catarina, Penha, Itapocoroy , 17 m, 25.v.2014, MOUFPE 15611 (specimen used in genetic analysis) ; EASTERN ATLANTIC— Sierra Leone. 2 ovf, in front of the Sierra Leone River , 19.iii.1953, RMNH . Crus.D. 9313; 1 ovf, Freetown, Freetown Harbour Station , 28.iii.1953, RMNH . Crus.D. 9314; Ghana. 1 ovf, 1 f, 1 m, Gold Coast, 14.iii.1951, RMNH . Crus.D. 10209; Benin. 1 ovf, Cotonou, 11.iv.1964, RMNH . Crus.D. 19933; Cameroon. 1 ovf, 2 f, Kribi, 09.viii.1964, RMNH . Crus.D. 21736 (one specimen used in genetic analysis); Democratic Republic of Congo. 1 ovf, 2 f, 2 m, Banana, 1968, RMNH . Crus.D. 24888; Angola. 1 ovf, Mercator Expedition XIV, 11.i.1930, RMNH .Crus.D.8587.

Previous records. Gulf of Mexico ( Felder et al. 2009).— Republic of Trinidad and Tobago ( Chace 1972).— Venezuela ( Rodriguez 1980; Prieto et al. 2000).— Suriname ( Holthuis 1959; Christoffersen 1979).— Brazil: Piauí ( Fausto-Filho 1980; Coelho et al. 2006), Ceará ( Fausto-Filho 1970; 1978; 1980; Christoffersen 1979; Sampaio & Fausto-Filho 1984; Coelho et al. 2006), Rio Grande do Norte ( Fausto-Filho 1980; Coelho et al. 1986), Paraíba ( Fausto-Filho 1980), Pernambuco ( Coelho & Ramos 1972; Fausto-Filho 1980; Ramos-Porto 1980; Coelho-Santos & Coelho 1998), Alagoas ( Fausto-Filho 1980; Coelho et al. 2006; Santos et al. 2016), Sergipe ( Santos et al. 2016), Bahia ( Spence Bate 1888; Almeida et al. 2006; 2007a; 2007b; 2012; 2013b; Coelho et al. 2006; Santos et al. 2012), Espírito Santo ( Christoffersen 1979), Rio de Janeiro ( Christoffersen 1979; 1982; Costa et al. 2016; Pantaleão et al. 2016), São Paulo ( Moreira 1906; Luederwaldt 1919; 1929; Christoffersen 1979; 1982; Costa et al. 2000; Furlan et al. 2013; Mantelatto et al. 2016; Pantaleão et al. 2016), and Santa Catarina ( Christoffersen 1979; 1982; Boos et al. 2012).— Senegal ( Sourie 1954).— Republic of Guinea ( Holthuis 1951; 1952; Crosnier & Forest 1965; 1966).— Sierra Leone ( Longhurst 1958).— Ghana ( Buchanan 1958).— Togo ( Crosnier & Forest 1966).— Benin ( Crosnier & Forest 1966).— Cameroon ( Balss 1925).— Democratic Republic of São Tomé and Príncipe ( Osorio 1892; Balss 1916; Crosnier & Forest 1965; 1966).— Gabon ( Crosnier & Forest 1966).— Congo ( Crosnier & Forest 1964; 1966).— Democratic Republic of Congo ( Schmitt 1926).

Colour in life. A photograph of the colour pattern of a live specimen from Bahia, Brazil, was provided by Santos et al. (2012).

Distribution. Western Atlantic—Gulf of Mexico, Suriname, Venezuela, Trinidad and Tobago, and Brazil ( Piauí to Santa Catarina). Eastern Atlantic—western Sahara to Angola ( Crosnier & Forest 1966; Christoffersen 1979; 1998; Coelho et al. 2006; Felder et al. 2009; Soledade & Almeida 2013; this study) ( Fig. 3 ).

Type locality. Off Salvador , Bahia, Brazil .

Ecology. Mud, clay, sand, sand with gorgonians, Halodule meadows, coarse sand with shells, in calcareous algae, coral gravel and crevices; pieces of decaying wood; depth range: 0 to 40 m. ( Christoffersen 1979; Almeida et al. 2006; Santos et al. 2012, this study).

Measurements. Largest specimen analyzed: male, CL: 13.48 mm (UESC 1573); smallest specimen analyzed: female, CL: 2.13 mm (RMNH.Crus.D.21736).

Redescription (based on specimens from Ilhéus, Bahia). Carapace smooth, without teeth or tubercles on midline; rostrum dorsally flat, inverted Y-shaped, reaching distal limit of first segment of antennular peduncle; adrostral furrows deep and broad, clearly delineated; ocular hoods dorsally inflated, with strong acute tooth arising from dorsomesial slope ( Fig. 1A ); basal segment of antennula with small triangular tooth on ventromesial surface, shark fin-shaped ( Fig. 1C ). Pterigostomial angle rounded. Cardiac notch well developed.

Abdominal somites smooth, ventral and posterior margins of pleurae rounded; telson broad; dorsal surface convex, without median groove, with two pairs of large spiniform setae, inserted approximately at 1/3 and 2/3 of telson length; posterior margin convex, with row of small spiniform setae; posterolateral angles each with two pairs of spiniform setae, lateral setae much shorter than mesial; anal tubercles well-developed; uropods with bifid protopods, each lobe ending in sharp tooth; exopod and endopod subequal in length; distolateral setae of exopod unpigmented; endopod with series of spines irregularly distributed along the posterior margin ( Fig. 1J ).

Eyes not visible in dorsal and lateral view; cornea rounded, well-developed; antennular peduncle with stylocerite distally acute, not reaching distal margin of first segment of antennular peduncle; antenna basicerite broad, with acute distolateral tooth directed forward; scaphocerite with lateral margin slightly concave; distolateral tooth strong, reaching far beyond narrow blade; carpocerite moderately enlarged, not exceeding distal portion of third segment of antennular peduncle ( Fig. 1B ).

Mouthparts typical for Alpheus ( Figs. 1 D–I); mandible with palp; incisive process robust, with eight or nine teeth distally rounded; molar process robust ( Fig. 1D ); first and second maxilliped as illustrated, not speciesspecific; third maxilliped robust, longer than antennular peduncle and carpocerite when extended; pleurobranchia absent at base; coxa with lateral plate, slightly truncated distally; antepenultimate segment of endopod narrow, not flat, with rough ventral margin and few setae; penultimate segment approximately half length of last segment; lateral margin slightly rough; last segment tapering distally, with lateral margin slightly rough and very long setae; exopod of third maxilliped not reaching distal end of endopodal penultimate segment, with long, flexible setae on posterior margin (Fig. 11).

Major cheliped merus ventrally flattened, with half-length merus of minor cheliped; ventromesial and ventrolateral margins serrated; carpus cup-shaped, with long distodorsal setae; palm laterally compressed in dorsal view; surface smooth, covered with setae; linea impressa well-marked; dorsal and ventral notches present; ventral notch with three small tubercles; presence of strong distodorsal and distoventral acute tooth on mesial and lateral region of the palm; dorsal shoulder not protruding, rounded, sloping gently into notch; ventral shoulder not protruding, rounded; fingers shorter than half length of palm; pollex tip curved upwards, ventral margin rounded, not notched on cutting edge anterior to fossa, plunger weakly developed ( Figs. 2A, B ).

Minor cheliped merus with approximately twice the length of the major cheliped merus; ventromesial and ventrolateral margins serrated; carpus cup-shaped, slightly longer than carpus of major cheliped; palm laterally compressed, with smooth surface, covered with setae; presence of strong distodorsal and distoventral acute tooth on mesial and lateral region of palm; linea impressa well-marked; fingers slender and straight, with balaenicepstype setae in males; pollex with sharp cutting edge; dactylus with serrated margin ( Figs. 2C, D ).

Second pereiopod with ischium and merus subequal in length; first segment of carpus the longest; carpal proportion of segments (proximal to distal) 3: 2: 1: 1: 2; chela simple; fingers subequal to palm, with few tufts of setae on distal portion ( Fig. 2E ). Third and fourth pereiopods with spiniform setae on ventrolateral surface of ischium; merus armed with distal tooth on ventral margin; carpus unarmed, with half length of merus; propodus with seven spiniform setae on ventral margin plus pair of distoventral spiniform setae near articulation with dactylus; dactylus about 1/3 length of propodus, simple, gradually curved towards sharp tip ( Figs. 2F, G ). Fifth pereiopod with ischium and merus unarmed; carpus armed with distoventral tooth, about half length of merus; propodus slightly longer than carpus with six spiniform setae on ventral margin plus pair of distoventral setae near articulation with dactylus; dactylus simple, about 1/3 length of propodus, slightly curved, distally sharp ( Fig. 2H ).

Variation. There was no consistent morphological variation in the characters analyzed for all western and eastern Atlantic specimens. The morphological analysis showed neither significant variation between sexes nor during ontogeny.

Figures

FIGURE 1. Alpheus intrinsecus Spence Bate, 1888, male (A–C, G–J) and ovigerous female (D–F) Malhado Beach, Ilhéus, Bahia, Brazil (UESC 1571): A, frontal carapace and cephalic appendages, dorsal view; B, frontal carapace and cephalic appendages, lateral view; C, tooth on mesioventral carina of first segment of antennular peduncle, lateral view; D, left mandible, mesial view; E, first maxilla, lateral view; F, second maxilla, lateral view; G, first maxilliped, lateral view; H, second maxilliped, lateral view; I, third maxilliped, lateral view; J, telson and uropods, dorsal view. Scale bars = 1 mm.

FIGURE 2. Alpheus intrinsecus Spence Bate, 1888, male, Malhado Beach, Ilhéus, Bahia, Brazil (UESC 1571): A, major cheliped, mesial view; B, major cheliped, lateral view; C, minor cheliped, mesial view; D, minor cheliped, lateral view; E, second pereiopod, lateral view; F, third pereiopod, lateral view; G, fourth pereiopod, lateral view; H, fifth pereiopod, lateral view. Scale bars = 1 mm.

FIGURE 3. Presently known distribution of the snapping shrimp Alpheus intrinsecus Spence Bate, 1888 based on material examined and literature records.