Anoplodactylus maritimus, Hodgson, 1914

Lucena, Rudá Amorim & Christoffersen, Martin Lindsey, 2018, Anoplodactylus (Pycnogonida: Phoxichilidiidae) from Brazil, new records and two new species, Turkish Journal of Zoology 42 (4), pp. 372-388 : 378-381

Anoplodactylus maritimus : Stock, 1974: 1069-1074, figure 54; Child, 1992: 52-53, figure 23.

Diagnosis: see Child (1992).

Distribution: Brazil (São Paulo, Rio de Janeiro), Amphi-Atlantic, Belize, Caribbean Sea, Gulf of Mexico to USA (Virginia), Central Atlantic (24°55′N, 44°00′W), Macaronesian Islands: Cape Verde Islands, Madeira Island, Canary Islands, Azores ( Child, 1992; Stock, 1992; Müller, 1993; Turpaeva and Raiskiy, 2014).

Depth: Up to 4379 m deep.

Anoplodactylus massiliformis Stock, 1974

Anoplodactylus massiliformis Stock, 1974: 1063-1066 , figures 48 and 49.

Diagnosis: see Stock (1974).

Distribution: Brazil (off Amapá), French Guyana, Guyana, Marguerita Island, Aruba, Barbados, Caribbean coast of Panama ( Stock, 1974, 1986; Müller, 1993).

Depth: up to 100 m deep.

Anoplodactylus mirim Lucena, de Araújo & Christoffersen, 2015

Anoplodactylus mirim Lucena et al. 2015: 432, figures 6–11.

Diagnosis: see Lucena et al. (2015).

Distribution: Brazil: Paraíba ( Lucena et al., 2015).

Depth: Up to 30 m deep.

Anoplodactylus monotrema Stock, 1979 ( Figures 2a– 2g)

Halosoma robustum : Marcus, 1940: 68-71, figures 8a– 8c ( Dohrn, 1881).

Anoplodactylus robustum : Stock, 1955: 237, figures 12b and 12c.

Anoplodactylus monotrema : Stock, 1979: 15-18, figures 4 and 5; Müller and Krapp, 2009: 105-107, figure 56.

Material examined: Alagoas: ( UFPB.PYC–127) 1 ♂, Galés de Maragogi, intertidal, 14 Jun. 2014, coll. R. A. Lucena, J. Prata and J.P. de Araújo; ( UFPB.PYC–129) 1 ♀, Praia de Peroba , intertidal, 13 Jun. 2014, coll. R. A. Lucena, J. Prata and J.P. de Araújo .

Diagnosis: Trunk robust. Trunk segments 3 and 4 fused. Lateral processes separated by one-fourth of their diameter, with a dorsal seta. Proboscis very robust, cylindrical. Cheliphores moderately robust. Chela with curved fingers having 3 teeth. Oviger with 5 articles. Tube of cement gland oval and long, placed at end of first fourth of femur. Propodus slightly curved, heel with 2 robust spines and 2 smaller subventral spines. Sole with 6 small spines, curved towards distal region, and bordered by 5 short setae on both sides. Main claw robust. Auxiliary claws strongly reduced.

Distribution: Brazil (São Paulo, off Espírito Santo, Alagoas), Colombia; Curaçao, Aruba, Bonaire, Caribbean coast of Panama, Virgin Islands, Jamaica, Bahamas,

Mexico, USA (Florida), Galapagos ( Stock, 1992; Müller, 1993; Müller and Krapp, 2009).

Depth: Up to 41 m deep.

Remarks: A. monotrema belongs to a group of species of Halosoma Cole, 1904 (later synonymized with Anoplodactylus ) that have 5 articles on the oviger instead of 6, as is more common in Anoplodactylus , known as the A. robustus group ( Dohrn, 1881) ( Stock, 1979; Child, 1982a). A. robustus , A. virescens (Hodge, 1864) , and a few other species, together with A. monotrema , are characterized for being small, stout, with contiguous lateral processes, and short articles on appendages ( Child, 1982a).

We may differentiate the three main species of the group by the absence of teeth on the fingers in A. robustus and A. virescens , segmentation completely absent in the trunk of A. robustus (absent only in the segments 3 and 4 in the remaining species), 3–6 pores in the cement gland in A. robustus and A. virescens (only one in A. monotrema ). The abdomen surpasses the lateral process 4 in A. virescens (in A. monotrema it reaches the margin of the process) and coxa 1 has dorso-distal tubercles in A. robustus (smooth in A. monotrema ) ( Stock, 1979; Child, 1979, 1982b; Müller and Krapp, 2009).

In the material examined here, all the characters are equal to the original description ( Figures 2a–2f) but the pore of the cement gland is longer ( Figure 2f) than that described by Stock (1979) and Müller and Krapp (2009). Besides, the auxiliary claws are ornamented by teeth ( Figure 2g), a character not observed in other species of the genus. Both characters differ from the original description by Stock (1979) for specimens representing the first record in Brazil (see Stock, 1992).

The female observed herein has the proboscis somewhat longer and more robust, the abdomen is more erect, and the lateral processes are more widely separated in the female, as also described for the female by Stock (1992). Segmentation is complete in our material, as in the description of A. monotrema . More specimens have to be examined in order to determine if the reduced auxiliary claws observed in the material from Alagoas is a useful character for the characterization of this species. Here we record this species for the first time in the northeastern State of Alagoas.

Anoplodactylus petiolatus (KrØyer, 1844)

For older synonyms and references, see Müller (1993: 239).

Anoplodactylus petiolatus : Melzer et al., 1996: 167-171, figures 1 and 2; Ros-Santaella, 2004: 7, figure 6; Bamber and Costa, 2009: 168, figure 2g; Lehmann et al., 2014: 167, figures 47 and 48.

Material examined: Espírito Santo: ( UFPB.PYC–126) 1 ♂, Ilha do Boi , intertidal, 01 Nov. 2005, coll. K. Paresque.

Diagnosis: Trunk elongate with all segments fused. Lateral processes with rounded tubercles. Scapus of cheliphore slender. Movable finger with one seta on inner and outer margin. Tube of cement gland thin, directed obliquely backwards, beginning in the middle region of femur. Propodus curved. Heel with 2 to 4 spines. Sole almost straight, with 3 or 4 distally curved spines along proximal half, and a cutting lamella on distal half. Main claw slightly curved, thin. Auxiliary claws strongly reduced.

Distribution: Brazil (off Rio Grande do Sul, Santa Catarina, off Paraná, São Paulo, Rio de Janeiro, Espírito Santo), Argentina, Uruguay, French Guiana, Surinam, Guyana, Bonaire, Venezuela, Curaçao, Caribbean coast of Colombia, Mexico, Bahamas, USA (Florida, Alabama, Texas, Georgia), Sargasso Sea, Cape Verde, Morocco, Mediterranean, Spain, Black Sea, Ireland, United Kingdom, Belgium, Holland, Denmark, Faroe Islands, Russia, Terra del Fuego (Beagle Channel), Chile ( Stock, 1986, 1992; Müller, 1993; Bamber and Costa, 2009; Lehmann et al., 2014; Turpaeva and Raiskiy, 2014).

Depth: Up to 4825 m deep.

Remarks: A. petiolatus and A. maritimus are very similar species ( Müller and Krapp, 2009). They may be distinguished, superficially at least, by the more tenuous and slender habitus of the first species, with larger appendages than the second species ( Child, 1992).

The examined specimen is a little more robust than the specimens illustrated by Stock (1974), Child (1992), and Müller and Krapp (2009). It is very similar to the specimen described by Marcus (1940). It has a cutting lamella on the propodus, the fingers of the chelae and the tubercles of the lateral processes are similar to those described by Stock (1974) and Müller and Krapp (2009). Furthermore, it complies with the description by Stock (1974) in the following characters: the auxiliary claws small, but easily discernible, the rectangular palm of the chela, and the third oviger more than 5 times as long as broad.

As noted by Stock (1992) for Brazilian specimens, our specimen has 8 spines in the sole of the propodus, 4 of which are more robust, being located medially on anterior half of sole, and there are median-sized tubercles on the lateral processes that are as broad as long. We were unable to observe vestigial tubercles on coxa 1, as described by Stock (1992).

Anoplodactylus spurius Stock, 1992

Anoplodactylus spurius Stock, 1992: 132-134 , figures 65–75.

Diagnosis: See Stock (1992).

Distribution: Brazil: São Paulo, Rio de Janeiro ( Stock, 1992).

Depth: From 19 to 220 m deep.

Anoplodactylus stictus Marcus, 1940

Anoplodactylus stictus Marcus, 1940: 65-68 , plate VI, figures 7e–7f, plate VII, figures 7a–7d; Sawaya, 1945: 231- 234, figures 1 and 2; Bremec et al., 1986: 36-37, figures 7 and 8; Stock, 1992: 134, figure 62.

Material examined: Rio Grande do Norte: ( UFPB. PYC–112) 1 ♀, Ponta do Mel , Brazil, 23 Jun. 1982, coll. A.I. Kanagawa. Ceará: ( UFPB.PYC–111) 1 ♂ and 1 ♀, Ponta Grossa, 24 Jun. 1982, coll. M.L. Christoffersen and A.I. Kanagawa.

Diagnosis: Trunk elongate with segmentation between segments 3 and 4, or segmentation absent. Lateral processes separated by less than their diameter, with 1 or 2 hairs and a rounded dorsal tubercle. Ocular tubercle elongate. Proboscis of female with a cordiform protuberance. Scapus and palm with few setae that are concentrated at the basis of the movable finger. Third article of oviger with a pseudo-segmental fold. Legs long. Coxa 2 with a small dorso-median tubercle. Femur with a distal tubercle and a long terminal seta. Tube of cement gland short. Tibiae 1 and 2 with a long distal seta. Heel of propodus with 1 or 2 spines. Sole with 6 to 10 spines in proximal region, and a cutting lamina in distal region. Auxiliary claws varying in size within the same individual.

Distribution: Brazil (off Santa Catarina, Paraná, São Paulo, Rio de Janeiro, Espírito Santo, Rio Grande do Norte and Ceará), Argentina ( Marcus, 1940; Bremec et al., 1986; Stock, 1992).

Depth: Up to 250 m deep.

Remarks: As described by Marcus (1940), A. stictus has nondenteate alar processes on the proboscis of the female, a tubercle on the lateral processes of the male, and a cutting lamina on the distal third of the propodal sole, which is different from A. californicus , the closest known species.

We did not observe the presence of a cutting lamina on the sole of the propodus, this being the only important difference detected in relation to the original description. We also observed that individuals of A. californicus tended to be larger than individuals of A. stictus , a character hypothesized by Marcus (1940) as diagnostic to differentiate the two species.

An association of A. stictus with the hydrozoan Tubularia crocea (Agassiz, 1862) was recorded ( Genzano, 2002), probably representing a predator ( Varoli, 1994). However, they were also recorded as parasites of a new hydrozoan species of the genus Podocorina Sars, 1846 , an association established by chance in an incrusting community of experimental plates in Paranaguá, Paraná ( Bettim and Haddad, 2013). According to Marcus (1940), A. stictus is the closest to A. californicus but may be distinguished mainly by the larger size, by the presence of a remnant of the propodus in A. californicus , by the presence of a cutting lamina in the propodal sole in A. stictus , and by a small protuberance on the movable finger of the chela ( Stock, 1992).

This species is known to exist in shallow waters, most species occurring along the upper continental platform in Brazil ( Marcus, 1940; Sawaya, 1945) and northern Argentina ( Stock, 1992).

In the specimens examined by us, the male had a more elongate and conical ocular peduncule than the female. Segmentation between segments 3 and 4 was almost imperceptible and, in some cases, completely absent. On the coxae 1, there were small lateral projections, each with a terminal seta, more conspicuous in males. Some females and juveniles had small tubercles on the lateral processes, more developed in young specimens. According to Marcus (1940), these were only present in males. Vestiges of auxiliary claws were not observed in any of the four examined specimens.

Here we recorded this species for the first time in northeastern Brazil, in the States of Rio Grande do Norte and Ceará.

Anoplodactylus typhlops Sars, 1888

Anoplodactylus typhlops Sars, 1888: 341-342 , plate 2, figures 3a–3c; Sars, 1891: 29-31, plate II, figures 3a–3e; Carpenter, 1905: 175, plate III, figures 12–19; Stephensen, 1933: 44-45, figure 12; Hedgpeth, 1948: 228-229, figures 29a–29c; Stock, 1955: 235-236, figure 12a; Turpaeva, 1973: 181-183, plate I, figures 1–8; Stock, 1991: 207, figure 55.

Anoplodactylus neglectus Hoek, 1898: 293-295 , figures 7–10.

Anoplodactylus pelagicus Flynn, 1928: 25-27 , figures 14a–14b; Barnard, 1954: 128, figures 19a– 19g.

Diagnosis: See Barnard (1954).

Distribution: Brazil (off Pernambuco), Bermuda, northwest coast of Africa, South Africa, Spain, Ireland, Norway, from Prince Edward to Crozet Islands, Tasman Sea, Costa Rica (Cocos Ridge), Gulf of Alaska ( Hedgpeth, 1948; Child, 1992; Müller, 1993; Raiskiy and Turpaeva, 2006).

Depth: From 900 to 3620 m deep.