Phoxichilidiidae Sars, 1891

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 : 373-378

Family Phoxichilidiidae Sars, 1891

Diagnosis: see Bamber (2007).

Key for the genera of the Phoxichilidiidae

1. Ovigers present in male and female ......... Phoxiphilyra − Ovigers present only in male ...................................... 2

2. Without dorsal segmentation, segmentation only visible ventrally. Cheliphores smaller than proboscis ........... ............................................................................... Pycnosomia

− Segmentation present dorsally, or completely absent. Cheliphores larger than proboscis ....................................... 3

3. Auxiliary claws long, located distally to main claw ...... .......................................................................... Phoxichilidium

− Auxiliary claws reduced or absent, located on dorsal basis of main claw ........................................... Anoplodactylus

Genus Anoplodactylus Wilson, 1878

Type species: Anoplodactylus lentus Wilson, 1878 , by original designation.

Diagnosis: see Child (1998a).

Remarks: Anoplodactylus is the largest and the most abundant genus of Phoxichilidiidae . It is among the three most numerous genera of Pycnogonida, with 134 species. It is considered a shallow-water genus, having a pantropical distribution. Bamber (1998) proposed that Anoplodactylus has species that disperse passively in the plankton, riding on medusae, thus explaining their wide global distribution ( Bamber and Costa, 2009).

Wilson (1878), in the original description, stressed that the main diagnostic characters of Anoplodactylus are the ovigers, with 6 articles, and the absence of auxiliary claws. However, for Marcus (1940) the only distinctive character of the group is the elongation of the neck over the proboscis. This applies particularly after the synonymization of Halosoma Cole, 1904 , a genus with specimens having 5 articles in the oviger, the discovery of a new species, and the transfer of species of Phoxichilidium Milne Edwards, 1840 to Anoplodactylus . The latter species are distinguished only by the position of the auxiliary claws (reduced and basal in relation to the main claw in Anoplodactylus , long and terminal in Phoxichilidium ) ( Child, 1995).

Despite being considered a stable, morphologically homogeneous ( Arango and Maxmen, 2006), and monophyletically robust group ( Arango and Wheeler, 2007), as suggested by the absence of palps and ovigers in the females, and by a stable number of segments in the ovigers of males ( Hedgpeth, 1948; Child, 1998a; Arango and Maxmen, 2006; Lucena et al., 2015), the species of Anoplodactylus display large variations in the structure of the cement glands, in body shape and segmentation, and in the size and shape of the proboscis ( Arango, 2002).

According to Hedgpeth (1948), the species of Anoplodactylus form a variable group whose identification is uncertain. That is because their distinction is based on a set of one or two distinct characters, or by a unique combination of distinct characters, particularly of the male ( Arango and Krapp, 2007). This situation indicates the necessity for a revision of the group, as is pointed out by Williams (1941), Hedgpeth (1948), Arango and Maxmen (2006), and Müller and Krapp (2009).

Given the large morphological variations and high number of described species, taxonomic groups were proposed on the basis of species distributions and morphological characters ( Arango and Krapp, 2007). A series of species-complexes were recognized: A. pygmaeus ( Stock, 1974) , A. insignis ( Stock, 1974) , A. robustus ( Stock, 1979) , A. gestiens ( Nakamura and Child, 1991) , A. californicus-digitatus ( Arango and Maxmen, 2006) , A. tenuicorpus ( Arango and Krapp, 2007) , and A. angulatus (Krapp et al., 2008; Lucena et al., 2015).

We point out the great difficulty in defining species in the genus ( Arango and Krapp, 2007), resulting from incomplete diagnoses, absence of illustrations ( Sawaya 1949), and similarity among species due to lack of variations in some characters ( Child, 1998a). Furthermore, the males have the main diagnostic characters: the ovigers and the cement gland. Females are impossible to identify when not accompanied by males ( Nakamura and Child, 1991; Child, 1998a). As a consequence, a large number of species remain to be named ( Williams, 1941; Hedgpeth, 1948).

To date, this is the only genus of Phoxichilidiidae recorded from Brazil, being one of the most abundant genera in shallow waters in all regions of Brazil.

Key to the Brazilian species of Anoplodactylus (adult specimens)

1. Oviger with 5 articles .................................................. 2

− Oviger with 6 articles ................................................... 3

2. Trunk partially segmented, segmentation absent between segments 3 and 4. Chelae well developed. Fingers with teeth ......................................................... A. monotrema

− Trunk without segmentation. Chelae with movable finger well developed, immovable finger reduced. Fingers without teeth .......................................................... A. marcusi

3. Segments 1 and 2 of trunk with a bifid tubercle near margin of segment ................................................ A. evelinae

− Segments of trunk without ornamentation ............... 4

4. Proboscis styliform, with terminal region directed upwards. Tube of cement gland with transverse rings .......... ............................................................................ A. batangensis

− Proboscis not styliform. Tube of cement gland without transverse rings ....................................................... 5

5. Animal with very small trunk, less than 0.6 mm ..... 6

− Animal with trunk not very small, larger than 0.6 mm .......................................................................................... 8

6. Lateral processes contiguous. Movable finger of chela reduced ................................................................. A. marcusi

− Lateral processes not contiguous. Movable finger of chela equal to fixed finger ..................................................... 7

7. Trunk segmentation present between segments 1–2 and 2–3 ............................................................... A. brasiliensis

− Trunk segmentation completely absent ........ A. mirim

8. Ocular tubercle low or absent, eyes absent or very small and unpigmented ........................................ A. typhlops

− Ocular tubercle regular, eyes present and pigmented .. ................................................................................................... 9

9. Lateral processes smooth or ornamented only with setae ...................................................................................... 10

− Lateral processes ornamented with tubercle; with or without setae ........................................................................ 14

10. Tube of cement gland long and narrow, extended forwards. Lateral processes with small dorsal spine and two smaller lateral spines ...................................... A. aragaoi

− Tube of cement gland short ....................................... 11

11. Trunk without segmentation ................ A. maritimus

− Trunk with segmentation complete or absent between segments 3 and 4 .................................................................. 12

12. Trunk very long. Lateral processes separated by twice their diameter. Legs with long dorsal spores on femur and tibia 1 .......................................... A. massiliformis

− Trunk short. Lateral processes separated at most by their diameter. Legs without dorsal spores ....................... 13

13. Coxa 2 of all legs with long spurs on ventral region in males. Lateral processes with three long dorsal setae ....... .................................................................................. A. eroticus

− Coxa 2 of legs 3 and 4 with spurs not very long, remaining legs with very short spur. Lateral processes with many small setae .................................................... A. ricardoi

14. Lateral processes with large, triangular or elongate tubercles ............................................................................... 15

− Lateral processes with small, rounded tubercles ....... 17

15. Third article of oviger with a small hook directed backwards, near base ..................................... A. ganchiformis

− Third article of oviger straight, without a hook ...... 16

16. Chelifore with anterior tubercle. Opening of cement gland elongate ........................................................ A. insignis

− Chelifore without tubercle, but with many setae. Tube of cement gland robust, conical large, directed obliquely to distal region ............................................................. A. spurius

17. Trunk not segmented .............................. A. petiolatus

− Trunk partially or completely segmented ................ 18

18. Ocular tubercle conical. With a small rounded tubercle on lateral processes. A small constriction present near base of third article of oviger ........................... A. stictus

− Ocular tubercle rounded. With small rounded tubercles dorsally on lateral processes. No constriction at the base of third article of oviger ..................... A. californicus

Anoplodactylus aragaoi Sawaya, 1949

Anoplodactylus aragaoi : Sawaya, 1949: 63–67, figures 1–9; Stock, 1992: 130, figures 63–64.

Diagnosis: Segments 3 and 4 of trunk fused. Dorso-distal margin of lateral processes 1–3 with a small median seta and a smaller seta on each side. Palps reduced to a small cylindrical tubercle. Length of legs circa 3 times the length of the trunk. Coxae 1 and 3 with 2 lateral setae. Femur with a conical tubercle on distal margin, and a long apical seta. Tube of cement gland thin and elongate, tilting forwards. Propodus with a spine on heel, and 2 smaller spines on sole. Sole with 9 quadrangular spines, and a small terminal cutting lamina. Auxiliary claws absent (modified from Sawaya, 1949).

Distribution: Brazil: Espírito Santo ( Stock, 1992).

Depth: 35–38 m.

Anoplodactylus batangensis (Helfer, 1938)

For older synonyms and references, see Müller (1993: 220).

Anoplodactylus batangensis : Müller and Krapp, 2009: 88-90, figure 47; Lucena et al. 2015: 430-432, figures 1–5.

Material examined: Alagoas: ( UFPB.PYC–173) 2 ♂♂ and 1 f ♀, Pontal do Peba , intertial, 15 Jun. 2014, coll. R .A. Lucena, J. Prata and J.P. de Araújo . Paraíba: ( UFPB. PYC–150) 1 ♀, Coqueirinho, intertidal, 17 Apr. 2014, coll. R .A. Lucena; J. Prata and J.P. Araújo . Rio Grande do Norte: ( UFPB.PYC–144) 1 ♀, Tourinhos, 16 Jul. 2014, coll. R .A. Lucena and L.M. Falkenberg . Ceará: ( UFPB.PYC–135) 3 ♂♂ , Praia da Redonda, 09 Aug. 2014, coll. R .A. Lucena and J. Prata .

Diagnosis: Animal small. Segmentation incomplete, absent between segments 3 and 4. Chelae small with short fingers. Proboscis cone-shaped and curved upwards. Lateral processes with a rounded dorsal tubercle. Distal region of the 2 anterior lateral processes more expanded than the 2 posterior lateral processes. Legs short. Tube of cement gland narrow, formed by small transversal rings. Auxiliary claws very small or absent.

Distribution: Circumtropical. Brazil (Bahia, Alagoas, Pernambuco, Paraíba, Rio Grande do Norte, Ceará), Curaçao, Venezuela, Caribbean coast of Colombia, Atlantic and Pacific coasts of Panama, Caribbean coast of Costa Rica, Belize, Gulf coast of Mexico, Barbados, Martinique, Guadalupe, Antigua, Dominican Republic, Jamaica, Ilhas Cayman, Bahamas, USA (Flórida), Bermuda, Cameroon, Madagascar, Tanzania, Australia, New Caledonia, Papua New Guinea, the Philippines, Sulu Sea, Tonga ( Müller, 1993; Arango, 2003; Müller and Krapp, 2009).

Depth: Up to 40 m deep.

Remarks: Easily recognized by the shape of the proboscis (styliform and directed upwards) and by the structure of the cement gland (formed by transversal rings) ( Child, 1996; Müller and Krapp, 2009). The first record for Brazil was given by Lucena et al. (2015), for the States of Pernambuco and Paraíba. Subsequently it was recorded for the coast of Bahia ( Lucena and Christoffersen, 2017), which is the most southern occurrence in the southern Atlantic. Here we provide the first references for the States of Alagoas, Rio Grande do Norte, and Ceará.

Anoplodactylus brasiliensis Hedgpeth, 1948

Anoplodactylus pygmaeus Marcus, 1940: 63–65 , Pl. VI figure 6a–d.

Diagnosis: Trunk small, completely fused. Lateral processes separated by a distance smaller than their diameter. Ocular tubercle wide and rounded. Scape very short and wide, less than twice as long as chela. Chelae with strong spines. Legs with many setae. Coxa 2 with a dorsal tubercle. Femur and tibias with a long distal seta. Tube of cement gland long and narrow. Propodus curved, heel with 2 spines. Sole with 2–6 median spines and a cutting lamina occupying one-third to half of the sole. Main claw little more than half length of propodus. Auxiliary hooks very small or absent (modified from Marcus, 1940).

Distribution: Brazil (São Paulo), USA ( Georgia) ( Marcus, 1940; du Bois-Reymond Marcus and Marcus, 1967).

Depth: Infralittoral.

Anoplodactylus californicus Hall, 1912

For older synonyms and references, see Müller (1993: 222).

Anoplodactylus californicus : Arango and Maxmen, 2006: 60, 62 [key], figure 3f; Melzer et al. 2006: 238–240, figures 2b, 2f, 2g; Müller and Krapp, 2009: 90-93, figure 48; Weis and Melzer, 2012: 200-201, figures 2b, 10f, 10g, 11a– 11g.

Material examined: Bahia: ( UFPB.PYC–182) 1 ♀ and 4 juveniles , Praia das Conchas, 16 May 2015, coll. R .A. Lucena, L.M. Falkenberg and J. Prata . Paraíba: ( UFPB. PYC–099) 2 ♂♂ and 7 juveniles , Cabo Branco , intertidal in Sargassum vulgare , 01 Feb. 2014, coll. E. Canuto and K.S. Pacheco; ( UFPB.PYC–100) 1 ♂♂ and 1 juvenile , Cabo Branco , intertidal in Gracilaria ferox , 31 Jan. 2014, coll. E. Canuto and K.S. Pacheco; ( UFPB.PYC–149) 1 ♂, 1 ♀ and 1 juvenile , Cabo Branco, intertidal in Gracilaria sp. , 31 Jan. 2014, coll. E. Canuto and K.S. Pacheco; ( UFPB.PYC–151) 3 ♀♀ and 3 juveniles, Coqueirinho , intertidal, 17 Apr. 2014, coll. R .A. Lucena; J. Prata and J.P. Araújo; ( UFPB. PYC–168) 2 ♀♀, Seixas , Paraíba, 09 Nov. 2014, coll. T . Dias . Rio Grande do Norte: ( UFPB.PYC–219) 2 ♂♂, 2 ♀♀ and 5 juveniles, Tourinhos , intertidal, 6 Jul. 2014, coll. R .A. Lucena and L.M. Flakenberg . Ceará: ( UFPB.PYC–218) 24 ♂♂, 15 ♀♀ and 23 juveniles, Jericoacoara , intertidal, 11 Aug. 2014, coll. R .A. Lucena and J. Prata; ( UFPB.PYC–220) 1 ♂, 4 ♀♀ and 3 juveniles, Flecheiras , intertidal, 13 Aug. 2014, coll. R .A. Lucena and J. Prata; ( UFPB.PYC–244) 1 ♂, Jericoacoara , intertidal in hydrozoans, 11 Aug. 2014, coll. R .A. Lucena and J. Prata . Piauí: ( UFPB.PYC–221) 2 ♂♂, 1 ♀ and 11 juveniles , Praia do Coqueiro , intertidal, 19 Mar 2015, coll. R .A. Lucena, J. Prata and J.P. de Araújo .

Diagnosis: Body elongate, trunk segments 3 and 4 fused. Two small setae inserted laterally near the bases of the cheliphores. Chela with a row of small setae in the direction of palm-finger, and 3 or 4 setae on ventral side of palm. Movable finger with 5 median dorsal setae. Legs very long. Femur with a small dorso-distal tubercle. Spur on legs 3 and 4 long in both sexes. Tube of the cement gland short and oval. Propodus with 2 spines on heel. Female with a pair of wing-like tubercles on proboscis. Palp vestigial.

Distribution: Pantropical, pantemperate. Strait of Magellan, Brazil (Santa Catarina, São Paulo, Bahia, Pernambuco, Paraíba, Rio Grande do Norte, Ceará, Piauí), Colombia, Curaçao, Bonaire, USA (Florida, Hawaii), Egypt, Turkey, Israel, Italy, Chile ( Marcus, 1940; Sawaya, 1949; Stock, 1974; Child, 2004; Müller and Krapp, 2009).

Depth: Up to less than 100 m deep.

Remarks: Anoplodactylus californicus may be identified by the robust appearance, the strigilum on the oviger, with many setae, cement gland with a short tube pointing distally, long spur on coxa of leg 4 of male, the peculiar alate process on the proximo-ventral region of the proboscis of the female ( Child, 1992; Bamber and Takahashi, 2005) and the large, abnormal sprout representing the palp ( Child, 1995).

This species was referred to erroneously under several names because the type-species was poorly described and insufficiently illustrated by Hall (1912). The material corresponding to this species was considered lost until it was rediscovered and reexamined by Child (1987). Apparently, this is one of the few species of Pycnogonida that have a high thermal tolerance. They occur in Subantarctic habitats, temperate localities in the South America, tropical latitudes in the eastern Pacific, and temperate latitudes of the northern hemisphere ( Child, 1995).

We detected a certain plasticity in certain characters that differ from the original description and from those presented by Müller and Krapp (2009). Some specimens had the ocular tubercle conical, while others had a rounded ocular tubercle. Complete segmentation was observed in some specimens, and a small rounded tubercle occurred above the insertion of the proboscis in some specimens. Some individuals had the abdomen slightly curved backwards, and some presented a small elevation, less than a tubercle, on the lateral processes ornamented by setae. Some individuals had a tubercle only on the lateral process 4, a femur with one elongate dorso-terminal tubercle and one apical seta. Furthermore, there were differences among the cement glands in specimens from different localities along Brazil, some being longer, others being more robust. Here, we recorded this species for the first time for Bahia, Paraíba, Ceará, and Piauí States .

Anoplodactylus eroticus Stock, 1968

Anoplodactylus eroticus Stock, 1968: 49 , figure 18; Arango and Maxmen, 2006: 55, figures 2a–2e, 3a, 3b, 4, 5; Lucena et al. 2015: 435, figures 12–28.

Material examined: Rio Grande do Norte: ( UFPB. PYC–216) 1 ♂, 4 ♀♀ and 1 juvenile, Tourinhos, intertidal, 16 Jul. 2014, coll. R .A. Lucena and L.M. Falkenberg . Piauí: ( UFPB.PYC–120) 2 ♂♂, 2 ♀♀ and 1 juvenile , Praia do Coqueiro , intertidal, 19 Mar. 2015, coll. R .A. Lucena, J. Prata and J.P. de Araújo .

Diagnosis: Trunk elongate. Segmentation complete. Lateral processes ornamented with three long setae. Legs long with many setae. Coxa 2 with long spurs on all legs. Tube of cement gland short and erect distally. Propodus with 2 spines on heel, and a small cutting lamina on distal region. Auxiliary claws reduced.

Distribution: Brazil (Alagoas, Paraíba, Rio Grande do Norte, Ceará, Piauí, and Maranhão), India, USA (Hawaii) ( Stock, 1968; Arango and Maxmen, 2006; Lucena et al., 2015).

Depth: Not cited.

Remarks: This species was recorded for the first time in Brazil by Lucena et al. (2015), for the States of Alagoas, Paraíba, Rio Grande do Norte, Ceará, and Maranhão. This is the second record for Brazil, and the first from the State of Piauí.

Anoplodactylus evelinae Marcus, 1940

Anoplodactylus (Labidodactylus) evelinae : Stock, 1954.

Anoplodactylus evelinae Marcus, 1940: 55 , figure 4; Hedgpeth, 1948: 232, figure 31; Fage, 1949: 27-28, figure 3; Müller and Krapp, 2009: 93-94, figure 49.

Material examined: Bahia: ( UFPB.PYC–170) 1 ♂ and 1 juvenile , Praia de Apuã, intertidal, 17 Apr. 2015, coll. R.A . Lucena, L.M. Falkenberg and J. Prata . Alagoas: ( UFPB.PYC–128) 2 ♂ and 1 ♀ , Praia de Peroba, intertidal, 13 Jun. 2014, coll. R.A . Lucena, J. Prata and J.P. de Araújo .

Diagnosis: Body small. Trunk segments 3 and 4 fused. Cuticulum with small tubercles. Segments 1 and 2 with a bifurcate median-dorsal tubercle, near posterior border of segment. Proboscis robust, of similar size to segment 1, and with small ventral tubercles near median line. Abdomen small, with a rounded tubercle at base. Fingers with small teeth. Legs short and robust. All articles of legs with small median and dorsal tubercles. Cement gland opening onto femur by a median-dorsal pore. Propodus with 1 spine on heel. Main claw two-thirds length of propodus. Auxiliary claws strongly reduced.

Distribution: Brazil (São Paulo, Rio de Janeiro, Bahia and Alagoas), Colombia, Atlantic and Pacific Panama, Anguilla, Jamaica, Belize, Mexico, USA (Florida), Congo ( Marcus, 1940; Sawaya, 1949; Müller, 1993; Müller and Krapp, 2009).

Depth: Up to 30 m deep.

Remarks: The species is clearly distinguished from the remaining species of the genus for having some rare characters, otherwise commonly found in the genus Pycnogonum (Ström, 1762) : The short dorso-median tubercles on the trunk very rare or unique to this genus, abdomen inflated and extending horizontally from trunk, very short legs, with tibia as long as wide ( Stock, 1979; Child, 2004), and propodus with very pronounced heel having a robust spine ( Child, 1982b).

For Müller and Krapp (2009), A. evelinae is related to A. arescus du Bois-Reymond Marcus, 1959 and A. tarsalis Stock, 1968 , for having a compact structure with similar tarsus and propodus. They may be distinguished by the cement gland, which in A. evelinae does not have a tubular opening.

Here we record this species for the first time in the Brazilian State of Alagoas.

Anoplodactylus insignis ( Hoek, 1881)

Phoxichilidium insigne Hoek, 1881: 82-84 , plate XIV, figures 5–7.

Anoplodactylus insignis bermudensis Cole, 1904: 325- 327 , plate XX, figures 1–3, plate XXII, figures 21–29.

Anoplodactylus insignis calcaratus : Stock, 1986: 437- 438, figures 15a–15b.

Anoplodactylus insignis : Hedgpeth, 1948: 226-228, figures 28d– 28g; Stock, 1974: 1056-1058, figure 44; Child, 1992: 46-49, figure 21.

Material examined: Maranhão: ( UFPB.PYC–110) 1 ♀, Cedral, 19 Jul. 1982, coll. M.L. Christoffersen and I.A . Kanagawa.

Diagnosis: Trunk long, tenuous. Trunk segments 3 and 4 fused or partially fused. Lateral processes short, with setose dorso-distal tubercles that diminish in size in the posterior pairs. Ocular tubercle conical, directed obliquely forwards. Proboscis of male with a pair of medio-ventral alar protuberances. Scapus of chelifore with anterior setose tubercles. Chela setose without teeth on fingers. Legs with long ventro-distal tubercles on coxa 2, dorso-distal tubercles on femur and tibia 1. Cement gland opening into long groove. Propodus with distal tubercles, a pronounced heel with 3 spines, without a lamina. Auxiliary claws absent.

Distribution: Brazil (off Espírito Santo, Bahia, Maranhão, Pará, off Amapá), French Guyana, Suriname, Guyana, Trinidad and Tobago, Venezuela, Caribbean coast of Colombia, Curaçao, Guadeloupe, Gulf of Mexico, Bermuda, USA (Florida, Georgia, North Carolina) ( Stock, 1986, 1992; Child, 1992; Müller, 1993; Arango, 2000).

Depth: Up to 90 m deep.

Remarks: This species has tubercles on the cheliphores and long legs, which makes it easy to distinguish from the remaining species of the genus and from the species considered the closest to it, A. lentus Wilson, 1878 ( Child, 1992; Child, 1998b).

Stock (1986) proposed the variety calcaratus based on the difference in the size and number of the tubercles on the lateral processes and legs; the larger tubercles occur in calcaratus. This difference may represent an intraspecific or interspecific variation ( Arango, 2000). However, further analyses are necessary to determine the validity of this character and of this variation. In the specimens examined herein, the tubercles are short, differing from those described by Stock (1986) and Arango (2000), both from specimens coming from northern South America.

There are two recognized varieties of A. insignis , Anoplodactylus insignis var. bermudensis Cole, 1904 and Anoplodactylus insignis var. calcaratus Stock, 1986 , both having the same geographical and bathymetric distribution.

The variety proposed by Cole does not have lateroventral tubercles on the femur, nor segmentation between segments 3 and 4, but has the cement gland in the shape of a sieve ( Hedgpeth, 1948). The variety by Stock has a small anterior tubercle and a long posterior tubercle on the dorsal surface of coxa 1, a long tubercle in the distal region of the femur, and two small tubercles on the anterolateral and postero-lateral surfaces of the femur, some long tubercles on the proximal region of tibia 1, and a small tubercle on the dorsal margin of tibia 2 ( Stock, 1986).

The specimen examined by us is more similar to the material examined by Arango (2000), which differs from the variety by Stock for not possessing a dorsal tubercle on the lateral processes. However, the ocular tubercle is longer than that described by Arango, and a latero-distal tubercle is present on the femur and two dorsal proximal tubercles occur on tibia 1, as indicated by Stock, except that they are extremely reduced, being almost imperceptible on some legs. On tibia 2, the present specimen has two small tubercles with an apical seta at the distal apex, in a latero-distal position. The remaining characters agree with the description by Hoek (1881).

We agree with Arango (2000) and Stock (1986) in that a larger number of animals from different locations are necessary for deciding upon the intra- or interspecific value of these coxal morphological variations. Here we recorded this species for the first time in the State of Maranhão.

Anoplodactylus marcusi ( Mello-Leitão, 1949)

Halosoma marcusi Mello-Leitão, 1949: 168-173 , figures 1–4.

Anoplodactylus robustus : Zilberberg, 1963: 25-27, figures 5–7 ( Dohrn, 1881).

Diagnosis: Cuticle with many hypodermic glands. Trunk small and robust. Lateral processes contiguous. Proboscis slightly shorter than half length of trunk. Abdomen strongly reduced. Cheliphore and chela with few setae. Fixed finger almost half as long as movable finger. Oviger with 5 articles. Legs robust with few setae. Propodus only slightly smaller than femur, with 2 spines on heel. Sole with small spines along its length. Auxiliary claw strongly reduced (modified from Mello-Leitão, 1949).

Distribution: Brazil (Rio de Janeiro, Ilha de Trindade) ( Mello-Leitão, 1949; Zilberberg, 1963).

Depth: Intertidal to 40 m deep.

Anoplodactylus maritimus Hodgson, 1914