Saltipedis (Spinosaltipedis) puertoricensis, Morales-Núñez, Andrés G., Heard, Richard W. & Alfaro, Mónica, 2010

Morales-Núñez, Andrés G., Heard, Richard W. & Alfaro, Mónica, 2010, Saltipedis (Spinosaltipedis) puertoricensis, a new subgenus and species of apseudomorphan (Crustacea: Tanaidacea: Parapseudidae) from coastal waters off Culebra Island, Puerto Rico, with keys and taxonomic observations, Zootaxa 2578, pp. 25-46 : 28-44

Saltipedis (Spinosaltipedis) puertoricensis n. sp.

Figs. 2–6 , 7 G, 8A–C, 9G; 10I

Material examined. Holotype, terminal adult male, length 5.1 mm, ( USMN 1145550), 18˚16’21.81’’N, 65˚19’09.01’’W, 3.2 km Southwest of Culebra Island, Puerto Rico, depth 28 m, collected between October 2002 and October 2003. Paratypes (same collection data as for holotype): three males (1 terminal, 2 subterminal), three females ( USMN 1145551); one male ( GCRL 2981), two females ( GCRL 2982, 2983). Additional specimens from the type locality are in the collection of authors.

Etymology. The species is named for the Territory of Puerto Rico, where the material used in this study was collected.

Type locality. 18˚16’21.81’’N, 65˚19’09.01’’W, 3.2 km Southwest of Culebra Island, off eastern Puerto Rico, depth 28 m, soft substrata (sand).

Distribution. Presently known only from the type locality.

Description. Adult female (with oostegites).

Body ( Fig. 2 A): Length about 4.3–5.2 mm, about 5.7–6.0 times as long as wide. Pereonites 1–5 with oostegites, greatly reduced, vestigial on pereonite 1. Hyposphenia lacking on pereonites 1–5, greatly reduced and inconspicuous, on pereonite 6. Pleonites with hyposphenia.

Cephalothorax ( Figs. 2 A–B): Carapace smooth, longer than wide, subquadrate in dorsal aspect. Rostrum with broad base becoming abruptly constricted anteriorly forming small acute tip ( Fig. 2 B). Ocular lobes with compound eyes having visual elements present. Second thoracomere (thoracic segment) with strongly developed hyposphenia between bases of chelipeds and directed anteriorly over the base of the maxilliped. Epistomal spine undeveloped, appearing absent.

Pereonites ( Fig. 2 A): Lacking spination, subquadrate in dorsal aspect, generally increasing slightly in length posteriorly; first two pereonites subequal in length; third pereonite longer than first and second; fourth widest, longer than third; fifth pereonites greater than length of any of the first four pereonites; sixth becoming narrower posteriorly, about equal in length with fifth.

Pleon ( Figs. 2 A, C): Width distinctly less than pereon; compressed, combined length of pleonites 1–5 about equal to that of last pereonite; pleonites decreasing slightly in width posteriorly; each pleonite about twice as wide as long; pleonite 1 with a dorsolateral row of setae; pleonites 1–3 each with ventrolateral margin tapering and forming acute posterior process; pleonites 4–5 with acute posteroventral margin; each pleonite with single hyposphenia, decreasing in size posteriorly ( Fig. 2 C).

Pleotelson ( Figs. 2 A, D): Subquadrate, about 1.2 times as long as wide, length slightly more than combined lengths of pleonites 3–5; sparsely setose with lateral margins each having two or three simple setae; apex of posterior margin with pair of small setulose setae between two longer simple setae ( Fig. 2 D).

Antennule ( Figs. 2 E–F): Peduncle of four articles: Article 1 about 2.5 times as long as wide, with eight broom setae along inner and distal margin, and numerous simple setae on both margins; article 2 about half length of first, with three sensory (two distal and one on mid-inner margin); article 3 about half length of second, with one brush-setae on distal inner margin and one near mid outer margin; article 4 shorter than third. Accessory flagellum with nine articles. Main flagellum with 13 articles, distal margins of articles 8, 10 and 12 each with single aesthetasc ( Fig. 2 F).

Antenna ( Fig. 2 G): Basal article with two setae and finely serrate inner margin. Article 2 with four short setae; article 3 with three setae (two short and one long); article 4 lacking setae; article 5 with one setae; article 6 longest (about 1.5 times as long as the article 3) with long and short seta at middle of inner and outer margins. Squama at junction of articles 2 and 3, well-developed, longer than peduncle article 2, bearing 18 long simple marginal setae. Flagellum with eight articles bearing long simple setae.

Mouthparts: Labrum ( Fig. 3 A): As illustrated.

Mandibles ( Figs. 3 B–E): Left mandible, incisor process with tip having four teeth, lacinia mobilis ( Fig. 3 D) with five teeth (distal three largest), setiferous lobe with four bifurcate setae, two distal-most short and flattened ( Fig. 3 C). Right mandible, similar to left but lacking lacinia mobilis; setiferous lobe with five setae ( Fig. 3 B); Molar process relatively slender, with grinding surface having fine denticulation ( Fig. 3 E). Palp with three articles; first article shortest with 3–5 simple setae, with serrate margin; second article longest, about twice that of first, with 6 simple setae on distal margin; third article longer than the first and slightly more than half second with one proximal spiniform seta, one lateral simple setae, and with eight short and one long simple setae distally ( Fig. 3 E). Margin between base of palp and molar process with five or six distinctive, robust, buttressed, simple setae ( Fig. 3 E).

Maxillule ( Fig. 3 F): Inner endite with five setulate setae, both margins finely setose; outer endite with eleven spiniform setae (at least one being serrate) and two subdistal setulate setae, margin finely setose; palp biarticulate with 4–5 distal or subdistal whip-like “cleaning” setae.

Maxilla ( Figs. 3 G–H): Unexceptional, outer lobe of fixed endite with 3–4 spiniform distally trifurcate setae ( Fig. 3 H), and several pectinate and setulate setae; inner lobe of fixed endite with row of about 30 basely swollen setae; outer lobe of movable endite with six long simple setae and three serrated setae.

Labium ( Fig. 3 I): Outer margin serrate, inner margin setulose. Palp, length about 1.5 times width, distal margin with three spiniform curved setae.

Maxilliped ( Figs. 4 A–D): Basis, longer than wide, usually with three to five small spines on distolateral margin ( Fig. 4 B). Endite with 9-10 short, basally swollen, setulose setae on inner anterior margin and four coupling hooks on the inner posterior margin ( Figs. 4 A, C); subdistal edge with stout seta with microspinules covering distal half; inner distal margin with “chisel-like, apically bidentate setae and multiform setae; large, curved, simple seta medially ( Figs. 4 C–D). Palp: Article 1 having long seta (about half length of article 2) on inner margin, distolateral margin attenuated with cluster of three simple setae on apex ( Fig. 4 B); article 2 with inner margin having two rows of setae (including four long simple setae, distolateral margin with cluster of five long simple setae and relatively short lateral seta ( Fig. 4 A); article 3 bilobed ( Fig. 4 B), wider than long, with simple attenuated and serrate setae on inner distal margin; lateral margin asetose ( Fig. 4 A); article 4 with two simple setae laterally, and six pectinate setae medially ( Fig. 4 A).

Epignath ( Fig. 4 E): sub-oval, cup-shape with minute, stout seta on inner subdistal margin.

Cheliped ( Figs. 5 A–B): Slender, smaller and much less robust than that of adult male. Basis inflated with three spiniform setae on posterior margin (proximal smallest), and two long simple setae near posterodistal margin; four or five small simple setae along anterodistal margin. Merus about half length of carpus with cluster of four simple subdistal setae lateral face; posterodistal margin with 9–10 simple setae of varying lengths. Carpus long, narrow, inserted in anteroproximal surface of merus; small distal seta near anterior margin; posterior margin with 9–10 marginal and submarginal simple setae. Propodus about half length of carpus, with cluster of four simple setae on lateral face at junction of fixed and movable finger and on distofrontal margin adjacent to base of movable finger, inner face as illustrated ( Fig. 5 B); fixed finger with rows and clusters of simple setae on posterolateral and inner margin near tip, grasping edge with row of 9–12 articulated “blade” setae having finely pectinate or serrate upper margins, unguis with bidentate tip for insertion of tip of movable finger; dactylus (movable finger) subequal in length to palm of propodus, with row of 6–7 pectinate blade setae along grasping edge, unguis well developed. Exopod having three articles, second article with small seta on anterior margin, third article with four plumose setae ( Fig. 4 A).

Pereopod 1 ( Figs. 5 F–G): Coxal process present, reduced to small, rounded lobe bearing three simple setae, not visible from dorsal aspect ( Fig. 5 G). Basis with length slightly more than twice width; anterior margin (i.e. dorsal, flexor) with proximal half having four or five small, simple setae; posterior margin with proximal 2/3 armed with four spiniform setae (distal-most smallest); posterodistal margin with stout spiniform seta and narrow simple seta. Ischium short, with three simple setae (one long, reaching over half length of carpus) on posterodistal margin. Merus slightly longer than wide; anterodistal margin having two simple setae (longest distal margin of carpus) and two acutely tipped spiniform setae (larger twice length of smaller); posterior margin with about 9–10 simple setae of varying lengths and stout, conical, spiniform seta distally. Carpus as long as wide with anterior margin having stout spiniform seta and two simple setae and submarginal oblique row of six simple setae; posterior margin with three stout conical spiniform setae (becoming smaller proximally) and seven simple setae. Propodus slightly longer than wide, anterior margin with distal half bearing three long simple setae, and two spiniform setae (largest adjacent to and over ½ length of dactylus); posterior margin with four stout conical spiniform setae (becoming longer distally) Dactylus about as long as propodus with minute setae on anterior margin proximal to unguis, four small teeth on posterior (flexor) margin; unguis simple, less than half total length of dactylus. Exopod with three articles; article 1 and 2 lacking setae, article 3 with six plumose setae ( Fig. 5 G).

Pereopod 2 ( Figs. 6 A–B): Basis about 3.5 times as long as wide; anterior margin asetose; posterior margin with small spiniform seta on proximal third and five simple setae (one short and four long) on distal margin. Ischium with length about twice width with posterior margin having five long distal setae. Merus slightly shorter than carpus; anterodistal margin with long narrow curved spiniform seta; posterior margin having two narrow spiniform setae and nine setae of varying length. Carpus slightly longer than propodus; anterodistal margin with three spiniform setae of differing lengths; posterior margin having three spiniform and eight simple setae, inner face with two oblique rows of long setae ( Fig. 6 B). Propodus subequal in length and narrower than carpus; anterodistal margin with two spiniform and two long simple setae. posterior margin with four subequal spiniform and seven simple setae. Dactylus subequal in length with propodus; anterior margin with small seta; posterior margin with seta at or near junction with unguis; unguis simple, about half total length of dactylus.

Pereopod 3 ( Fig. 6 C): Basis with length about 3 times width, posterior distal margin with three long setae. Ischium slightly wider than long; anterior margin with small seta; posterior margin having seven (one distal) setae. Merus slightly longer than wide; anterodistal margin with small spiniform seta; posterior margin with distal half bearing three spiniform and five simple setae. Carpus longer than propodus; anterodistal margin with spiniform seta; posterior margin with three spiniform and six simple setae. Propodus slightly shorter and narrower than carpus; anterior margin with broom seta medially and one spiniform and two simple setae distally; posterior and anterodistal margin with four spiniform (distal most immediately adjacent to dactylus) and six subequal simple setae. Dactylus about equal in length to propodus; anterior margin with small seta; posterior with minute seta at or near junction with unguis; unguis simple, slightly less than half total length of dactylus.

Pereopod 4 ( Figs. 6 D–E): Basis swollen, about twice as long as wide, with proximal spine and distal seta on posterior margin. Ischium slightly longer than wide, with seven simple setae on posterodistal margin. Merus slightly shorter than propodus, with three spiniform setae and five simple setae on posterior margin; anterior margin with distal spiniform seta. Carpus about twice as long as wide, with six spiniform and two simple setae on posterior margin; frontal margin with two spiniform setae proximally. Propodus shorter and narrower than carpus, with five acute spiniform setae and four simple setae on posterior margin; frontal margin with one broom setae and two long, acute spiniform setae proximally, and four (one small and three long) spiniform setae distally. Dactylus as in pereopods 2–3, but appearing slightly shorter.

Pereopod 5 ( Fig. 6 F): Similar to pereopod 4, with slight differences in setation (e.g. presence of two broom seta and minute seta on anteroproximal margin of basis and broom seta on mid anterior margin of propodus).

Pereopod 6 ( Figs. 6 G–I): Basis swollen, about twice as long as wide; anterior margin with 22–24 plumose setae and three small broom setae inserted between third and fourth proximal plumose setae). Ischium longer than wide, posterior margin with four simple setae (one proximal and three distal). Merus slightly shorter than carpus; frontal margin with spiniform seta on medially and plumose seta distally; posterior margin having three spiniform and four simple setae. Carpus longer than propodus; anterior margin with three or four plumose setae proximal two thirds and spiniform seta distally; posterior margin with about 10 spiniform and four simple setae. Propodus shorter than carpus; anterior margin with broom seta on distal half, ( Figs. 6 G–H); posterior margin with eight spiniform setae (longest adjacent to dactylus) and one simple seta; distal margin anterior to dactylus armed with three elongate spiniform setae becoming shorter posteriorly (anterior most about as long as dactylus), distolateral margin with dense row of short fringing setulate setae ( Figs. 6 H–I). Dactylus with minute simple seta on both anterior and posterior margins; unguis simple, slightly less than half total length of dactylus.

Pleopods ( Figs. 6 J–K): Five well-developed, biramous pairs decreasing in size posteriorly. Exopods and endopods uniarticulate. First pleopod with basal article having 5-6 plumose setae on each margin; exopod and endopod each with 18-20 long, plumose, marginal setae ( Fig. 6 J). Fifth pleopod well-developed, but smaller than first pleopod; basal article distinctly shorter than that of pleopod 1, with 1 to 2 plumose setae on each margin; exopod shorter than endopod; exopod with about 12 and endopod with about 16 long plumose marginal setae, respectively ( Fig. 6 K).

Uropod ( Figs. 2 A, D): Basal article twice as long as wide, three or four simple, setae on distolateral margin; three simple setae on distal inner margin, three small setae along inner margin ( Fig. 2 D). Exopod with 10–12 articles and/or pseudoarticles, distal-most article having four apical simple setae. Endopod with division of articles unclear, apparently having from 50-60 articles and/or pseudoarticles, distal-most article with three (two short and one 1ong) simple apical setae.

Subadult female. Body: Usually smaller than adult female. Oostegites buds undeveloped. Thoracic hyposphenia present on pereonites 1–2, greatly reduced or appearing absent on pereonite 6.

Terminal (ultimate) adult male. Body: Similar to female, except slightly narrower and with hyposphenia present on pereonites 1 and 2 ( Fig. 2 C). Pereonite 6 having male genital cone well-developed, conspicuous, with posteriorly directed, acute tip.

Antennule ( Fig. 2 E): Similar to that of female, but with main flagellum usually having 14 articles, distal margins of articles 4, 6, 8, 10 and 12 each with pair of aesthetasc.

Cheliped ( Figs. 5 C–D): Robust, distinctly more massive than in female. Basis with anterior region excavate (for insertion of anterior or extender part of carpus) forming lateral and inner margins; inner medial margin with blunt spine or tubercle; posterior margin having two stout spiniform setae, and three (one short and two long) distal setae. Merus with posterior margin having cluster of 9–12 simple setae; lateral face with pair of simple setae. Carpus deeper than long, posterior margin with 5–6 setae (one proximal and two distal) and three ventral setae (one proximal and two distal), small seta subproximally on lateral face. Propodus stout with fixed finger having well-developed, weakly bidentate tooth; edge between tooth and unguis with row of five to six articulated "blade" setae with finely pectinate or serrate upper margins (as on female chela), and six to seven simple setae; unguis small, brownish ( Fig. 5 D). Dactylus with large, blunt, proximal tooth; six spines on anterior margin, and two simple setae lateral on distal margin; unguis well developed, brownish. Exopod with three articles: article 1 with two small spiniform setae; article 2 with small simple seta near anteroproximal margin; article 3 with four plumose setae.

Remarks. Saltipedis ( Spino s altipedis) puertoricensis n. sp., is distinguished from five of the six species currently attributed to the genus by its mandible having robust simple setae along the margin between the base of the palp and middle region at the base of molar process. The presence or absence of such setae is undetermined for S. tetracanthus , since its mouth parts (e.g., mandibles, maxilla 1 and 2 and maxilliped are presently undescribed). The females of S. puertoricensis , however, can be distinguished from those of S. tetracanthus by (1) the first pereopod having the inner frontal margin of basis with three (instead of one) blunt tubercles, and the posterior margin of the propodus armed with four (instead of five) stout spiniform setae and (2) the presence of a few simple (rather than plumose) setae on the frontal margin of the basis. The apparent absence of an epistomal spine on S. puertoricensis and S. achondroplasia distinguishes them from the other members of Saltipedis . Table 2 and the Keys A–C may be used to further distinguish S. puertoricensis from the other six currently recognized members of the genus.

Species Carpus Propodus Basis

Anterior margin Posterior margin Podictenius (“ Apseudes ”) espinosus (Moore 1901) , which was described from St. Thomas in the Virgin Islands ( Richardson 1905), was incompletely described and inadequately illustrated. Its description is based on a single female holotype collected at depth of about 53 m (“20 fathoms”) about 60 km east of the type locality for S. puertoricensis . Based on Moore’s incomplete illustrations, P. espinosus , at least in some aspects, superficially resembles S. puertoricensis (e.g., shape of rostrum and body form). The lack of dorsal setation on the first abdominal segment and the stronger developed row of spiniform setae on the frontal margin of the first pereopodal basis (Moore 1901, Plate 7, figs. 1 and 4) appears to distinguish P. e s p i n o s u s from S. puertoricensis . Notwithstanding P. espinosus needs to be carefully reexamined and redescribed to determine if it is correctly referable to Podictenius and to remove any reservations about its status in relation to S. puertoricensis .

Key A. Separation females for the currently recognized nominal species of the genus Saltipedis Gu ţ u, 1995

1 Pereopod 1 with posterior margins of carpus and propodus with four and six stout spiniform setae, respectively ( Fig. 7 B) ..................................................................................................................... S. robustispinosus Guţu, 1996 [ Brazil]

- Pereopod 1 with posterior margins of carpus and propodus with three or fewer and five or fewer spiniform setae, respectively ................................................................................................................................................................... 2

2 First pereopod with carpus having distoanterior margin armed with two stout spiniform setae ( Fig. 7 D).................... ................................................................................. S. navassensis Hansknecht, Heard, and Martin, 2001 [Caribbean]

- First pereopod with carpus having distoanterior margin armed with one stout spiniform seta................................... 3

3 First pereopod with carpus having two stout spiniform setae on posterior margin ..................................................... 4

- First pereopod with carpus having three stout spiniform setae on posterior margin .................................................... 6

4 First pereopod with propodus having posterior margin armed with five stout spiniform setae ( Fig. 7 C) ..................... ................................................................................................................................ S. bacescui Guţu, 1998 [ Tanzania]

- First pereopod with propodus having posterior margin armed with four stout spiniform setae................................... 5

5 Cheliped of female with anterior l margin of basis having blunt spine or tubercle; fixed and moveable fingers each with small tooth ( Fig. 9 D) ........................................................................................ S. paulensis (Brum, 1971) [ Brazil]

- Cheliped of female with anterior margin of basis lacking blunt spine or tubercle; fixed and movable fingers lacking tooth ( Fig. 9 A) .................................................. S. achondroplasia Bamber, Bird, and Angsupanich, 2003 [ Thailand]

6 Mandible with 5–6 stout accessory simple setae between the base of palp and base of molar process ( Fig. 3 E, 10H). First pereopod with posterior margin of propodus having four short, stout spiniform setae ( Fig. 5 F, 7G). Epistomal spine undeveloped ............................................................................................. S. puertoricensis , n. sp. [ Puerto Rico]

- Mandible with lacking accessory buttressed naked setae between the base of palp and base of molar process. First pereopod with posterior margin of propodus armed with at least 5 short, stout spiniform setae ( Fig. 7 F). Epistomal spine present, distinct ............................................................. S. tetracanthus Guţu & Angsupanich, 2004 [ Thailand]

1. Differences in the setation of the first pereopod. The known species of Saltipedis display considerable interspecific variation in the numbers and shapes (i.e. short, long, stout, conical, acute) of the spiniform setae on the first pereopods (see Fig. 7 ). This degree of interspecific variation in these setal characters appears to be diverse as those exhibited by several other genera within the Tribe Parapseudini . The first pereopod of some species, especially that of the S. paulensis ( Fig. 7 A), have relatively long acute spiniform setae on the propodus, carpus, and merus, while others such as western Atlantic species, S. robustispinosus and S. puertoricensis , have stout, short, and blunt spiniform setae ( Figs. 7 B, and 7G, respectively).

Like the chelipeds and first pereopods, the second through sixth pereopods exhibit an array of diverse setal types and patterns making each distinctive enough for identification of the species to which it belongs ( Fig. 7 ). We have presented examples of keys for the separation of the species of Saltipedis using characters displayed by the chelipeds or first pereopods since they are usually retained on specimens after the sieving and storing of samples; whereas, pereopods 2–6 are often lost during these processes.

2. Male cheliped forms exhibited by the genus Saltipedis sensu lato. Saltipedis (Spinosaltipedis) puertoricensis , n. sp., represents the fifth nominal species of the genus for which the male is known. The male chelipeds of these five species (i.e. S. achondroplasia , S. bacescui , S. navassensis , S. tetracanthus , and S. puertoricensis ) are somewhat similar, but can be distinguished by differences in the setal patterns on the basis and the dentition of the fixed and movable fingers (see Key B). Of these, three species, S. puertoricensis , S. achondroplasia , and S. tetracanthus have a tooth present on the fixed finger ( Figs. 8 A, 8D and 8J, respectively). Based on the setation of the posterior margin of the basis, the male cheliped of S. puertoricensis appears most similar, at least superficially, to that of S. achondroplasia ( Figs. 8 A, D). In contrast the presence on the basis of four tubercles on the frontal margin and four distinct spiniform setae on the posterior margin distinguishes S. tetracanthus from S. puertoricensis the other species of this genus, as well as, the other parapseudids of the Tribe Parapseudini for which males are known ( Figs. 8 J, A).

The males of Saltipedis bacescui and S. navassensis lack a tooth on the fixed finger, and have a similar setal pattern on the posterior margin of the basis, but they differ in overall shape and the in the presence [i.e. S. navassensis ] or absence [i.e. S. bacescui ] of tubercles on the frontal and distolateral margins ( Figs. 8 H, F). Based on its overall less compact shape, especially the relatively elongate carpus ( Fig. 8 F), the male cheliped illustrated by Guţu (2006) for S. bacescui is reminiscent of the penultimate male cheliped of S. puertoricensis ( Fig. 8 C). This indicates the possibility that Guţu's figure of S. bacescui may not represent the terminal male form. If so, a comparison of chelipeds of S. bacescui and S. navassensis would be much less meaningful. Notwithstanding this, the distinctive distolateral lobe on the basis and apparent lack of an unguis on fixed finger of S. navassensis appear to be unique ( Fig. 8 H), at least in comparison to the known males of Saltipedis and other related parapseudid genera.

Key B. Separation of known males for the genus Saltipedis Gu ţ u, 1995 based on the cheliped.

1 Fixed finger lacking a tooth. Posterior margin of basis having two to four distinct spiniform setae, the proximal most being larger than the distal-most .................................................................................................................................. 2

- Fixed finger with tooth. Posterior margin of basis having two spiniform setae, proximal most very small or minute ...................................................................................................................................................................................... 3

2 Fixed finger apparently lacking unguis. Basis with anterior inner margin with subacute tubercle; disto-lateral margin with distinct lobe or tubercle. Exopod having article 2 with two small simple setae on proximal margin ( Fig. 8 H) .... ....................................................................................... Saltipedis navassensis Hansknecht, Heard, and Martin, 2002

- Fixed finger with distinct unguis. Basis with anterior inner margin entire [lacking tubercle]; distolateral margin not lobate. Exopod with article 2 lacking setae ( Fig. 8 F) ...................... Saltipedis bacescui Guţu, 1998 (see Guţu, 2006)

3 Basis with posterior margin having four distinct spiniform setae ( Fig. 8 J).................................................................... .................................................................................................. Saltipedis tetracanthus Guţu and Angsupanich, 2004

- Basis with posterior margin having two distinct, subequal spiniform setae................................................................ 4

4 Dactylus with basal tooth acute. Exopod with article 2 having two small simple setae ( Fig. 8 D)................................. ................................................................................ Saltipedis achondroplasia Bamber, Bird, and Angsupanich, 2003

- Dactylus with basal tooth subacute. Exopod with article 2 having one small simple seta ( Fig. 8 A) ............................ .................................................................................................................................... Saltipedis puertoricensis , n. sp.

3. Female chelipeds of Saltipedis sensu lato. The female chelipeds of Saltipedis represent a range of characters as diverse as those found within the other members of the Tribe Parapseudini (e.g. Parapseudes ; Podictenius ). Some of the characteristics for the cheliped in conjunction with others (i.e. setation of pereopod 1, basal article of uropod, and pleotelson) may indicate that members of the genus represent polyphyletic assemblage. The following key distinguishes the seven nominal species of Saltipedis based on characters exhibited by the female cheliped.

The chelipeds for the females of S. bacescui , S. puertoricensis , and to a lesser degree of S. robustispinosus appear to share a number of features. In contrast those of S. achondroplasia , S. navassensis , S. paulensis , and S. tetracanthus each distinctively different. Saltipedis paulensis , the generotype, is distinguished by having (1) a tooth on the fixed and movable finger (male character), (2) a tubercle on the frontal margin of the basis, and (3) an exopod with only three plumose setae on the distal-most article ( Fig. 9 D). The dentition of the cheliped for S. paulensis immediately distinguishes it from the other species of the genus; as well as, the other genera and species within the Tribe Parapseudini . The swollen basis and setation (e.g. reduced spiniform setae on posterior margin of basis) of S. navassensis appears unique among Saltipedis species, but is reminiscent of that on the female for Longipedis fragilis Larsen and Shimomura, 2006 . The basis of S. achondroplasia has a distinctive setation pattern (i.e. frontal margin with two long distally attenuated setae and posterior margin with long simple seta medially and the distal-most spiniform seta being the smallest, instead of largest) shows no resemblance to that of male ( Figs. 8 D–E, and 9A). In contrast, though the spiniform setae are less robust, the setation of the posterior margin of basis for the female of S. tetracanthus is generally similar to that of the male ( Figs. 8 J–K, and 9F).

Key C. Separation of the nominal species of Saltipedis Gu ţ u, 1995 based on the female cheliped.

1 Cutting edges of fixed and movable fingers with small, but distinct tooth. Anterior margin of basis with subacute tubercle. Exopod with three plumose setae on distal article ( Fig. 9 D) ................................ S. paulensis (Brum, 1971)

- Cutting edges of fixed and movable fingers lacking tooth. Anterior margin of basis lacking tubercle. Exopod with 4– 6 plumose setae on distal article .................................................................................................................................. 2

2 Posterior margin of basis with three distinct, elongate spiniform setae proximally, one short stout spiniform seta distally, and cluster of 3 subdistal simple setae. Exopod with five plumose setae on distal article ( Fig. 9 F) .................... ................................................................................................................ S. tetracanthus Guţu and Angsupanich, 2004

- Posterior margin of basis without this setal arrangement. Exopod with four or six plumose setae on distal article .. 3

3 Basis with frontal margin armed with two long attenuated setae; posterior margin with one long simple seta proximal to spiniform setae, distal-most spiniform seta distinctly smaller than more proximal one. Exopod with six plumose setae on distal article ( Fig.9 A) .................................... S. achondroplasia Bamber, Bird, and Angsupanich, 2003

- Basis with frontal margin lacking long attenuated setae; posterior margin without long simple seta proximal to spiniform setae. Exopod with four plumose setae on distal article .................................................................................... 4

4 Basis with posterior margin having one spiniform seta. Exopod with two small setae on outer margin of article 2... 5

- Basis with posterior margin having two or three spiniform (some may be minute). Exopod with one or without a small seta on outer margin of article 2 .......................................................................................................................... 6

5 Basis swollen, with posterior margin having stout spiniform seta and one simple distal seta. Posterior margin of carpus with 1–3 medial setae ( Fig. 9 C) .......................................... S. navassensis Hansknecht , Heard, and Martin, 2002

- Basis not swollen with posterior margin having small spiniform seta and two simple, distal setae. Posterior margin of carpus with medial cluster of 6–7 setae ( Fig. 9 E) .................................................... S. robustispinosus Guţu, 1996

6 Basis having posterior margin with two spiniform setae, the proximal most one minute; subdistal pair of setae relatively short, less than 1/3 length of merus posterior margin. Exopod without seta on outer margin of article 2 ( Fig. 9 B) ............................................................................................................................................. S. bacescui Guţu, 1998

- Basis having posterior margin with three spiniform setae, proximal most two minute; posterior margin with subdistal pair of relatively long simple setae, about ½ length posterior margin of merus. Exopod with one small seta on outer margin of article 2 ( Fig. 5 F; 9G)...................................................................................... S. puertoricensis n. sp.

4. Mandibular accessory setae. The presence of "accessory setae" on the mandibular trunk between the palp and molar process in Saltipedis puertoricensis is not unique, since setae have been reported from this part for some members of the apseudid Pectinapseudes Bǎcescu and Williams, 1988 (see Bǎcescu and Williams 1988, Guţu 2002), and the parapseudid Platylicoa Guţu, 2006 (subfamily Parapseudinae ) (see Guţu 2006, 2008).

Another parapseudid belonging to the monotypic genus, Biropalostoma Guţu and Angsupanich, 2004 (Tribe Pakistanapseudini Guţu, 2008 ), has a distinctive spinose "bludgeon -like" process adjacent to palp ( Fig. 10 H); but this apparently unique structure may be independently derived. The 5–6 simple "accessory setae" on S. (S.) puertoricensis are robust and buttressed, appearing distinctly different and longer than the thornshaped types occurring on P. carolinensis and P. m e s t e r i Guţu, 2002 ( Figs. 10 A–B) and the row of small narrow simple setae occurring on P. magnus Guţu, 2002 ( Fig. 10 C).

The possibility of the accessory mandibular setae on S. puertoricensis being homologous with those found in the parapseudid genus Platylicoa (Tribe Pakistanapseudini ), is more problematical. The presence of this character in Platylicoa is represented by a compact cluster of five small, stout spiniform setae in P. s e t o s a Guţu, 2006 ( Fig. 10 F) and a single spiniform seta in P. p e c t i n i s ( Bamber, 1998) ( Fig. 10 G). In S. puertoricensis the accessory setae, which are distinctly larger and buttressed, occur in two groups, three or four larger setae near the base of the palp and two smaller setae adjacent to the molar process ( Figs. 3 E, 10I). For comparison, illustrations depicting the accessory setae and/or tubercles occurring on the mandibles of the apseudids [ Pectinapsuedes (Pectinapseudes) carolinensis , P. (P.) mesteri , P. (P.) magnus , and P. (Impectinatus) sicilianus ], the parapseudids [( Platylicoa setosa , Biropalostoma spiniferum and S. (S.) puertoricensis ] and the kalliapseudid [ Paraleiopus macrochelis ] are presented in Figure 10 .

The distinctive buttressed, accessory setae on the mandible of S. puertoricensis may, or may not, have the systematic significance purported for the apseudid subgenus Pectinapseudes and the parapseudid Platylicoa . Unlike the other members of its genus Pectinapseudes , P. (Impectinatus) sicilianus (Bǎcescu, 1980), lacks accessory setae; however, the presence of small tubercles or spinules on outer convex margin of the incisor process (between palp and setiferous row) ( Fig. 10 D) is one of the diagnostic characters for the subgenus ( Guţu 2006). Similar tubercles are reported for other apseudomorphan taxa, including the kalliapseudid Paraleiopus macrochelis Brum, 1978 (see Guţu 1996) ( Fig. 10 E), and to a lesser extent on Biropalostoma spiniferum Guţu and Angsupanish, 2004 ( Fig. 10 H). There are also small tubercles with associated small, short, bristle setae in the same location on the mandible of S. puertoricensis ( Fig. 10 I). We believe that the presence of small tubercles or spinules between the palp and setiferous row of some species within the families Apseudidae , Kalliapseudidae , and Parapseudidae ( Fig. 10 ) were independently derived. Further, their presence or absence appears not to be related to the presence of the “accessory setae.”

Figures

FIGURE 2. Saltipedis (Spinosaltipedis) puertoricensis n. subgen., n. sp., Adult female A, B, D, F, Adult male C, E, G. A, body, dorsal view; B, enlargement of rostrum; C, lateral aspect of adult male body showing hyposphaenia; D, enlargement of pleotelson; E, antennule, male; F, inner flagellum, female; G, antenna, male. Scale line: A = 1.0 mm; C = 0.3 mm; E – G = 0.5 mm.

FIGURE 3. Saltipedis (Spinosaltipedis) puertoricensis n. subgen., n. sp., Adult Female. A, labrum; B, right mandible; C, left mandible; D, enlargement of lacinia mobilis and tip of incisor process; E, mandible, excluding molar process, showing accessory setae; F, maxillule; G, maxilla; H, enlargement of trifurcate setae; I, labium. Scale line: A and F = 0.1 mm; B, C and E, = 0.5 mm; G and I = 0.2 mm.

FIGURE 4. Saltipedis (Spinosaltipedis) puertoricensis n. subgen., n. sp., Adult Female. Maxilliped A – D: A, Maxilliped showing setation; B, inner face with inner plate, most setae excluded; C, inner plate; D, enlargement of spiniform setae from different aspect; E, epignath. Scale line: A = 0.2 mm; F = 0.1 mm.

FIGURE 5. Saltipedis (Spinosaltipedis) puertoricensis n. subgen., n. sp., Chela, adult female: A – B: A, lateral view; B, inner aspect of half chela. Male: C – E: C, adult chela, inner distal aspect; D, adult cheliped lateral view; E, sub-adult cheliped, lateral view. F, pereopod 1, lateral view; G, exopod and reduced coxal process, ventral aspect. Scale line: A, C, E, and G = 0.2 mm; F = 0.5 mm.

FIGURE 6. Saltipedis (Spinosaltipedis) puertoricensis n. subgen., n. sp., Pereopods. Adult female: A, right second, lateral, view; B, right second, inner view of merus, carpus, propodus, and dactylus; C, right third; D, right fourth, lateral view; E, right fourth, inner view of merus and carpus; F, right fifth, lateral view; G, right sixth, lateral view; H, right sixth, inner view of dactylus and distal part of propodus; I, setulate seta. Pleopods: J, first; K, fifth. Scale line: A, C, D, F and G = 0.5 mm; J and K = 0.2 mm.

FIGURE 7. Known first pereopods for species of the genus Saltipedis. Except for exopods on B and E, all setae and exopods are illustrated. A, Saltipedis paulensis; B, S. robustispinosus; C, S. bacescui; D, S. navassensis; E, S. achondroplasia; F, S. tetracanthus; G, S. puertoricensis. [Figures modified from: Guţu 1996 (A – B); Guţu 1998 (C); Hansknecht et al. 2001 (D); Bamber et al. 2003 (E); and Guţu and Angsupanich 2004 (F)].

FIGURE 8. Known male chelae for the genus Saltipedis. Except for spiniform setae, all other setae are not shown. A – C, Saltipedis puertoricensis: A, adult male; B, adult female, posterior margin of basis; C, penultimate male; D – E, Saltipedis achondroplasia: D, adult male; E, female, posterior margin of basis; F – G, S. bacescui: F. penultimate? male; G, adult female, posterior margin of basis; H – I, Saltipedis navassensis; H, adult male; I, female, posterior margin of basis; J – K Saltipedis tetracanthus: J, adult male; K, female, posterior margin of basis. [Figures modified from: Bamber et al. 2003 (D – E); Guţu 1998 (G); Guţu 2006 (F); Hansknecht et al. 2001 (H – I); and Guţu and Angsupanich 2004 (J – K)].

FIGURE 9. Known female chelipeds for Saltipedis. A, S. achondroplasia; B, S. bacescui; C, S. navassensis; D, S. paulensis; E, S. robustispinosus; F, S. tetracanthus; G, S. puertoricensis. [Figures modified from Bamber et al. 2003 (A); Guţu, 1998 (B); Hansknecht et al. 2001 (C) Guţu 1996 (D – F); and Guţu and Angsupanich 2004 (G)].

FIGURE 10. Examples of apseudomorphan mandibles having accessory simple or spiniform setae between base of molar process and palp and examples of species with small tubercles on the convex margin of incisor process between palp and setiferous row. A, Pectinapseudes carolinensis; B, P. m e s te r i; C, P. magnus; D, P. sicilianus; E, Paraleiopus macrochelis; F, Platylicoa setosa; G, P. pectinis; H, Biropalostoma spiniferum; I, Saltipedis (Spinosaltipedis) puertoricensis. [Figures modified from: Bǎcescu and Williams 1988 (A); Guţu 2002 (B – C); Guţu 2006 (D); Guţu 1996 (E); Guţu 2006 (F); Bamber, 1998 (G), and Guţu and Angsupanich 2004 (H)].

Tables

TABLE 2. Some distinguishing characters for the species of Saltipedis based on the first pereopod (for carpus and propodus the number of spiniform setae on the anterior / posterior margins are indicated).

achondroplasia bacescui	1/2* 1/2	2/4 2/5	Few simple setae Few simple setae	3 small simple setae 2 small stout setae
navassensis	2/3	2/5**	4-5 plumose setae	No medial stout setae
paulensis puertoricensis	1/2 1/3	2/4 2/4	No evident setae Few simple setae	4 small stout setae 3-4 spiniform setae
robustispinosus	1/4	1/6	Few simple setae	Few minute simple setae
tetracanthus	1/3	2/5	Few plumose setae	Few small simple setae
* Anterior (Frontal)	margin/Posterior	margin; **A	minute, proximal, submarginal	spiniform setae not included.