Crustacea: Peracarida: Tanaidacea: Leptocheliidae

Zootaxa 3664 (2): 259–282 www.mapress.com / zootaxa / Copyright © 2013 Magnolia Press

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3664.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:0B8DBC17-E307-4397-A3A2-301021D1AE6D

A new species of Pseudoleptochelia Lang, 1973 (Crustacea: Peracarida: Tanaidacea: Leptocheliidae) from the Northwest Atlantic with observations on the status of the genus ANDRÉS G. MORALES-NÚÑEZ1,4, RICHARD W. HEARD2 & MÓNICA ALFARO3 1

Department of Natural Sciences, University of Maryland Eastern Shore, Princess Anne, MD 21853, USA. E-mail: [email protected] 2 Department of Coastal Sciences, University of Southern Mississippi, Gulf Coast Research Laboratory Campus, Ocean Springs, MS 39564, USA. E-mail: [email protected] 3 Department of Biology University of Puerto Rico Mayagüez Campus, PR 00680, Puerto Rico. E-mail: [email protected] 4 Corresponding author

Abstract Pseudoleptochelia juliae n. sp. is described from Puerto Rican waters in depths from 1 to 28 m. The new species and P. fairgo from western Australia are the only members of the genus as defined in this revision that have a distinct process on the posterior margin of the cheliped carpus. The male of P. juliae is distinguished from that of P. fairgo by this process being sub-acute and having a distinctly narrower propodus. Lang’s generic diagnosis for the female of Pseudoleptochelia is incorrect, which has led to confusion. Based in examination of Mediterranean specimens attributed to the type species P. anomala (Sars), Pseudoleptochelia is rediagnosed to contain seven species; P. anomala; P. fairgo; P. juliae; P. magna; P. mortenseni; P. occiporta; and P. straddi., The genus is reviewed and discussed and the taxonomic status for the remaining nine species previously assigned to the genus is re-evaluated. Pseudoleptochelia shares similarities with the more recently described Pacific genera Konarus and Parakonarus and to a lesser extent to the highly derived genus Makraleptochelia. Key words: Tanaidacea, Leptocheliidae, new generic diagnosis, Pseudoleptochelia, P. anomala, P. juliae n. sp., Caribbean, Puerto Rico

Introduction There have been earlier reports of an apparently undescribed species of Pseudoleptochelia Lang, 1973 from the southern waters of the Northwest Atlantic (Suárez-Morales et al. 2004; Heard et al. 2004; García-Madrigal et al. 2005; Heard et al. 2009) and during a study of the tanaidacean fauna of Puerto Rico by the first author, additional specimens of this species were collected. Based on this material, the description of this new species is presented here. Additionally, access to specimens of the type species, P. anomala (Sars, 1882), has allowed us to discuss its taxonomic status in relation to other nominal species presently assigned to Pseudoleptochelia. This publication represents the fourth in a series (Morales-Núñez et al. 2010; Drumm & Morales-Núñez 2010; and Heard & Morales-Núñez 2011) on the tanaidacean fauna of Puerto Rico. Including these published records, eight nominal species have been previously reported from Puerto Rican and closely adjacent waters (see MoralesNúñez 2011). The only previous records for tanaidomorphans from the coastal waters of Puerto Rico were for the leptocheliids Leptochelia forresti (Stebbing, 1896) and L. dubia (Krøyer, 1842), as reported by Stoner (1986) from shallow-water grass bed habitats near Mayagüez. The present study deals with a review of the leptocheliid genus Pseudoleptochelia with comments on its taxonomic status and the description of a new species from Puerto Rico coastal waters.

Accepted by R. Bamber: 4 Apr. 2013; published: 24 May 2013

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Materials and methods Specimens of an undescribed species of Pseudoleptochelia were collected with a benthic grab during 2008 in the vicinity of North San Cristobal keys, southwest of La Parguera, Puerto Rico (Fig. 1). Additional specimens were diver-collected using a PVC core sampler (8.8 cm in diameter) from a depth of 28 m in the vicinity of Culebra Island off the eastern coast of Puerto Rico (Fig. 1). These collections were obtained every two months between October 2002 and October 2003 and took place on soft bottom substrata in a 120 x 100 m (12,000 square m) area. Sample processing has been previously described by Morales-Núñez and Kornicker (2007). Specimens of P. anomala from a sub-littoral benthic habitat off the Balearic Islands, Spain (Mediterranean) were supplied by Victoria Fernandez-Gonzales. Images were taken with a GEOL 5410LV scanning electronic microscope, the specimens having been prepared using a critical point dryer and counting gold technique. Pictures were taken at different levels with a Canon 7D camera, using a Visionary Digital Laboratory System and stacking with Helicon Focus software. Type material has been deposited at the National Museum of Natural History, Smithsonian Institution, Washington DC, (USNM), and Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi MS, (GCRL). All measurements are in millimetres (mm). Total body length (TL) is measured from the tip of the rostrum to the end or tip of the telson. Terminology generally follows that of Larsen (2003), with following exceptions. The term ‘seta’ is here applied to those processes, thin, compound, or robust, that articulate with tegument of the body or its appendages or that are separated from them by a distinct suture; whereas, a ‘spine’, ‘spinose process’, ‘tooth’, or ‘tubercle’ is considered a process that is fully fused with the tegument of the body or its appendages.

FIGURE 1. Location of North San Cristobal Key at southwest of La Parguera Puerto Rico and Culebra Island, indicating the sampling station where Pseudoleptochelia juliae n. sp., was collected.

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MORALES-NÚÑEZ ET AL.

Systematics

Suborder Tanaidomorpha Sieg, 1980 Superfamily Paratanaoidea Lang, 1949 Family Leptocheliidae Lang, 1973 Genus Pseudoleptochelia Lang, 1973 Revised diagnosis. Male: Functional mouth parts lost (atrophied mass). Maxilliped greatly reduced, vestigial, basis represented by small cluster of simple setae, palps greatly reduced, partially fused. Antennule with peduncular article-3 reduced, forming dorsal saddle-like protrusion over flagellum article-1; flagellum with 8–14 aesthetascbearing articles, and minute terminal article with simple setae. Cheliped appearing subchelate, fixed finger reduced to small distal process; carpus with or without inferior process. Uropod with endopod having four to five (usually four) articles or incipient articles; exopod unarticulated. Female: Antennule compressed, stout, broad at base; composed of three apparent articles, plus minute terminal article. Antenna with article-2 having simple dorsodistal setae (lacking buttressed spiniform setae), article-3 anterodorsal distal margin with simple seta only. Maxillule spines slender. Maxilliped bases unfused distally with sub-distal margin having 3 to 5 simple setae reaching past article-1 of palp. Cheliped with coxal sclerite isolated, not reaching posterior of cephalothorax; disto-lateral margin of carpus extending as cuff to overlap propodus. Pereopod-1 unguis as long as or longer than dactylus. Pereopods 4 to 6 with basis swollen; merus distal margin with pair of basally-swollen, distally attenuated setae (appearing setulate); carpus with stout, sometimes weakly bidentate and/or setulate setae. Uropod with endopod having four to five (usually four) articles or incipient articles; exopod uniarticulate. Type species: Pseudoleptochelia anomala (Sars, 1882). Other species: Pseudoleptochelia fairgo Bamber, 2005; P. juliae, n. sp.; P. magna (Smith, 1906); P. mortenseni Lang, 1973; P. occiporta Błaźewicz-Paszkowycz & Bamber (2012); P. straddi Bamber, 2008. Remarks. The original description and illustrations for Heterotanais anomalus are based solely on two male specimens (Sars 1882, 1886) collected from off Messina, in the Mediterranean. Lang (1973) extracted this species from Heterotanais Sars 1882 to become the type species of the genus Pseudoleptochelia Lang, 1973. In addition to the type species, P. anomala, Lang transferred seven additional species, Heterotanais antarcticus Lang, 1953; H. magnus Smith, 1906; H. provincialis Dollfus, 1898; H. inermis Dollfus, 1898; H. mercantilis Smith, 1906; H. mergellinae Smith, 1906; and Leptochelia filum (Stimpson, 1853) to Pseudoleptochelia (see Table 1). In the same publication he also described a ninth species, P. mortenseni from the southern Caribbean Sea (Tobago). Since then, six additional species have been added to the genus (see Table 1): P. anorexia Bird & Bamber, 2000; P. bulbus Bamber, 2006; P. fairgo; P. longidactylus (Băcescu, 1977), P. straddi and P. occiporta Błaźewicz-Paszkowycz & Bamber, 2012. The 15 nominal species presently attributed to Pseudoleptochelia appear to comprise a heterogeneous grouping, which is in need of revision (Błaźewicz-Paszkowycz & Bamber 2012). When Lang (1973) designated Heterotanais anomalus as the type species for Pseudoleptochelia, he did not have type or topotypical material available, thus as part of his generic diagnosis he incorrectly included features of what he believed to be the female of the non-type species P. mortenseni, described from Tobago in the same publication and as part of the generic diagnosis. It is our opinion that this part of Lang’s diagnosis has no taxonomic standing since it is not based, at least in part, on the female of the type species, P. anomala. Further, based on our review of the literature and the examination of other leptocheliid genera and species, we believe that the female diagnosed and ascribed to P. mortenseni belongs to a different genus. Ideally the original generic diagnosis for the genus should have been based on the original description of H. anomalus Sars, 1882 and males and females attributable to the species from the Mediterranean and supplemented with observations on the male holotype of P. mortenseni. Notwithstanding the lack of a female, the text and illustrations provided by Sars (1882 and 1886, respectively) for the male type are adequate to demonstrate its uniqueness and specific validity. Later, based on material from the Mediterranean Sea collected near Naples, Smith (1906) presented an illustrated supplemental description of H. anomalus and more recently Riggio (1996) reported this species from Italian waters

NEW PSEUDOLEPTOCHELIA FROM THE NORTHWEST ATLANTIC

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as P. anomala. In spite of these efforts, no female has been well-enough described to be definitely confirmed as that of P. anomala. The female illustrated and attributed to P. anomala sensu Riggio (1996) appears to represent a species of Leptochelia sensu lato. TABLE 1. Alphabetical listing of the 15 previously recognized species for the genus Pseudoleptochelia sensu lato, including information on distribution and depth range (based on Bird & Bamber, 2000; Anderson 2012). Species

Geographical area

Depth range (m)

anomala (Sars, 1882) [type species]

Mediterranean (Messina); Indian Ocean (Zanzibar)

?–37

anorexia Bird & Bamber, 2000

Indo-Pacific (South China Sea)

10–30

antarcticus (Lang, 1953)

SW Antarctic (Falkland Island)

0–358

bulbus Bamber, 2006

New Caledonia (Loyalty Islands)

12–20

fairgo Bamber, 2005

Australia (Esperance, Queensland)

23–39

filum (Stimpson, 1853)

NE Pacific (NZ), US Pacific coast

0.5–138

inermis (Dollfus, 1898)

NE Atlantic (Dakar)

7

longidactylus (Băcescu, 1977)

Mediterranean (Libya)

70

magna (Smith, 1906)

Mediterranean (Naples)

Littoral

mercantilis (Smith, 1906)


Littoral

mergellinae (Smith, 1906)


Littoral

mortenseni Lang, 1973

Caribbean (Tobago)

Littoral

occiporta Błaźewicz-Paszkowycz & Bamber, 2012

Australia (Western Port)

3−51

provincialis (Dollfus, 1898)

Mediterranean (St. Tropez)

50

straddi Bamber, 2008

Australia (Queensland)

10

In summary, we consider the genus Pseudoleptochelia to be valid because Lang designated Heterotanais anomalus as the type for the genus. At the time of its generic designation, the unique characters exhibited by the male of H. anomalus fit the criteria of a valid genus. Recently, males and females attributable to type species, P. anomala from the Mediterranean were independently made available to us and to Roger Bamber, who is preparing a detailed description of both sexes (R. Bamber, pers. comm.). Our and Roger Bamber’s specimens were both collected from the Spanish Balearic Islands, about 1000 km west of its type locality near Messina, Italy. Based on our examination of this Mediterranean material, we believe that the female attributed to P. mortenseni appears to represent a distinctly different genus. Lang’s inappropriate diagnosis has led to confusion in separating females belonging to species within the genera Pseudoleptochelia, Pseudonototanais Lang, 1973 sensu lato, and Leptochelia sensu lato, especially when they were not accompanied by males. In effect, this facilitated the transformation of the genus into a “catch all” for some species that could not be placed reliably in other leptocheliid genera (e.g. Pseudoleptochelia filum sensu Sieg (1976); Pseudoleptochelia anomala sensu Riggio (1996); and Pseudoleptochelia anorexia). Compared to the revised generic diagnosis (see above), the female of P. mortenseni sensu Lang (1973) is characterized by (1) having the first antennular article-1 elongate, (2) the antenna having buttressed thorn setae and a distal dorsal spiniform process, respectively, on articles 2 and 3, (3) only two pairs of long simple setae on the sub-distal margin of the maxillipedal basis, (4) the cheliped lacking a cuff on the disto-lateral face of the propodus, and (5) the merus of pereopods 4–6 apparently lacking a pair of proximally swollen and distally attenuated setae. We believe that the female attributed to P. mortenseni by Lang (1973) may represent an unnamed species, possibly belonging to a putative new Leptochelia sensu lato or a Pseudonototanais-like genus. In contrast, the antennule in Lang’s figure 16g, which he attributes to an immature female stage of P. mortenseni, appears to represent that of the true female for this species and is similar to that of P. anomala material examined by us. Notwithstanding, Lang’s description and illustrations for the male of P. mortenseni support its congeneric status with the type species, P. anomala.



The females for P. anomala (examined during the present study), P. fairgo, and the new species described herein as well as those of the similar genera Konarus Bamber, 2006 and Parakonarus Bird, 2011, and to a lesser extent the apparently more highly derived genus Makraleptochelia Araújo-Silva & Larsen, 2012, share a number of similar characters. These include: Antennule composed of three apparent articles, plus vestige of minute terminal article; article-1 broad in dorsal aspect with length slightly greater than breadth. Antenna with article-2 having simple dorsodistal seta (lacking buttressed spiniform seta), article-3 anteriodorsal distal margin with simple seta only (no distal spinose process present). Maxilliped bases with sub-distal margin bearing 3 to 5 simple setae reaching past article-1 of palp. Cheliped with coxal sclerite isolated, not reaching posterior of cephalothorax; distolateral margin of carpus extending as cuff to overlap propodus. Pereopod-1 unguis as long as or longer than dactylus. We consider the carpal process on the males of P. juliae, P. fairgo, and P. occiporta to be a specific or subgeneric rather than generic character and believe that these two species are congeneric with P. anomala, P. magna, P. mortenseni, and P. straddi. The possibility remains, however, that the presence or absence of a prominent carpal process, in combination with other less pronounced morphological features (e.g. setation of pereopods 4–6) may have systematic importance and, if so, more than one genus may be represented (G. Bird, pers. comm. Feb 2011). We concur with this possibility, but for the present include P. juliae, P. fairgo, and P. occiporta within Pseudoleptochelia. The taxonomic status the male and female of P. occiporta, recently described from Bass Strait, Australia in not treated here, but will be clarified in a forthcoming revision of the group by R. Bamber (pers. comm. 2013).

Generic status of those species not retained within Pseudoleptochelia sensu stricto Pseudoleptochelia anorexia Bird & Bamber, 2000 is known only from a single female specimen collected from Hong Kong. Because it is characterized by the first antennular peduncle being relatively narrow, the apparent loss of functional eyes, and a uropod with only three endopodal articles, it is not referable to Pseudoleptochelia as defined here. Based on body form and the setation of the maxilliped, it has some similarities to Mesotanais Dollfus, 1897; however, like Leptochelia and unlike Mesotanais, the third antennal article of P. anorexia has a distinct “Leptochelia-like” disto-dorsal spinose process. Pending examination of additional material, especially the male, we are unable to reliably place this species in any of the currently recognized leptocheliid genera. Therefore we tentatively place it within Leptochelia sensu lato as L. anorexia (Bird & Bamber, 2000), n. comb. Based on Lang (1957), Bamber & Bird (1997:133) suggested that Pseudoleptochelia filum (Stimpson, 1853) sensu Lang (1973), a species originally described from boreal waters off eastern North America, might be referable to Pseudonototanais sensu lato. We agree that “Tanais filum” sensu Stimpson (1853) is not attributable to the Pseudoleptochelia as defined here. It does show some similarities to Leptochelia (e.g. antennules and uropods), the genus to which Lang (1957) tentatively assigned it, but then he later transferred it to Pseudoleptochelia (see Lang 1973). The female as described and illustrated by Lang (1957), like those attributed to Pseudoleptochelia (as defined here), has only simple narrow seta present on antennal article-2, but has an atypically narrow cheliped with long fingers, and the maxillipedal basis is described as having “two to four long ‘ciliated’ setae on the mid-distal margin” (Lang 1957, p. 8). This, in our opinion, precludes this species being placed in either Pseudoleptochelia or Leptochelia as now constituted. Due to this combination of morphological characters, we are unable to place “Tanais filum” sensu Lang (1957) and Wallace (1919) within any of the existing leptocheliid genera as presently defined. The reported ciliated condition of the long setae on the basis of the maxilliped is unknown for other leptocheliids. Lang’s and Wallace’s material should be re-examined to verify that the ciliated condition of the long maxillipedal setae is real and not due to fixation artefacts (e.g. chemical precipitates). The records for “Pseudoleptochelia filum” from the tropical western Atlantic, and boreal and tropical eastern Pacific (see Sieg 1983) are questionable and probably refer to more than one taxon. With the exception of Sieg (1976), the descriptions and illustrations for the specimens attributed to this species from these regions are too vague or incomplete to make a proper specific determination. Based on material from Peru, however, Sieg (1976) presented a detailed supplemental description of a female and primary male attributed to “Pseudoleptochelia filum” sensu Lang (1973). We believe that Sieg described and illustrated what appears to be the female and primary male of an undescribed species of Leptochelia sensu lato, similar to some of the nominal species having males with



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typical chelae (e.g. Leptochelia lusei Bamber & Bird, 1997; L. tanykeraia Bamber, 2009) and by having females with three or fewer pairs of long simple setae on the basis of the maxilliped and narrow, attenuated bodies. The female attributed to Pseudoleptochelia sensu Lang (1973) and that of Pseudonototanais ebriosus Bamber & Bird, 1997 also share these characters and appear somewhat similar, at least superficially. The illustrations and description given for the male ascribed to Pseudoleptochelia (= Heterotanais) antarcticus Lang, 1953, unlike that of P. anomala, are more reminiscent of the primary males of some species of Leptochelia (e.g. the L. dubia/savignyi complex), which have a relatively small and similar male chela lacking a wide gape and with the uropod endopod having only four articles. The male differs from those Pseudoleptochelia as recognized here, by not having a third scale-like peduncle article, but instead appears to have at least one article of the flagellum fused with last peduncular article. Based on Lang’s description and illustrations for the female, the maxillipedal bases each have two pairs of long simple setae. His illustration (Lang 1953: fig. 1D) indicates a fairly well-developed antenna with disto-dorsal setae on articles -2 and -3, but they do not appear to be as well-developed and buttressed as the thorn seta and distal setae or narrow spine present on typical members of the genus Leptochelia; a more detailed comparison is needed. Pending further study and for pragmatic reasons, we tentatively refer Pseudoleptochelia antarcticus to the genus Leptochelia sensu lato to become Leptochelia antarcticus (Lang, 1953), n. comb. It should be noted that Lang’s drawing of the uropod of the “4th”manca stage shows article-1 on the endopod being much longer and actually similar to that of the female illustrated by Shiino (1978) for Pseudonototanais werthi (Vanhöffen, 1914), however, this could be a developmental condition in Lang’s material. In our opinion, the four poorly known species, P. inermis, P. mercantilis, P. mergellinae, and P. provincialis are not referable to Pseudoleptochelia. Of these P. inermis, P. mercantilis, and P. mergellinae have typical primary male chelae and may represent those of Leptochelia sensu lato and they are tentatively referred to this genus. If the illustration and meagre description by Dollfus for Pseudoleptochelia provincialis are correct, then the male cheliped only appears at most to have a vestige of a fixed finger (Fig. 8G); we consider it as a species inquirenda. Table 2 summarizes our tentative perception of the taxonomic status of the species previously assigned to the genus Pseudoleptochelia. The taxonomic status of the two remaining nominal species of “Pseudoleptochelia,” P. bulbus Bamber, 2006 and L. longidactylus (Băcescu, 1977) is currently being re-examined and revised by Bamber (in prep.). Pseudoleptochelia longidactylus, a Mediterranean species, was originally placed in the genus Heterotanais but was removed by Sieg (1983). As exemplified by our remarks above, the large and heterogeneous genus Leptochelia is in urgent need of major systematic revision, especially in light of the many new species described during the past 15 years from the southern Pacific and Indian Oceans. The breadth and plethora of characters currently exhibited by these species of Leptochelia has facilitated it being “a catch-all” or “dumping ground” for leptocheliids of uncertain generic status. These include those treated in this study (see Table 2). Besides Leptochelia sensu stricto (i.e. L. minuta/forresti/ longimana complex) and those within the Leptochelia dubia/savignyi complex, as it now stands several additional genera maybe represented within the Leptochelia sensu lato. Such a formidable and extensive revision will be needed before a better understanding of the taxonomy and systematics of the family Leptocheliidae can be achieved.

Status of the genera Konarus Bamber, 2006 and Parakonarus Bird, 2011 Based on female and neuter forms, Bamber (2006) created the new monotypic genus Konarus designating K. cheirus Bamber, 2006 from the Loyalty Islands and New Caledonia as the type species. Bird & Larsen (2009), in their phylogenetic analysis for some of the genera within the superfamily Paratanaoidea, treated Pseudoleptochelia and Konarus as if they were congeneric. Later, Bamber & Chatterjee (2010) reviewed the status of Konarus and transferred Heterotanais crassicornis Stebbing, 1905 to the genus. As previously mentioned, the females of Konarus share several morphological features with those Pseudoleptochelia, but they also differ in several distinctive ways. As defined here, Pseudoleptochelia is distinguished from Konarus by having a uropod with fewer endopodal articles and a single exopodal article. Based on material attributable to the type species of Pseudoleptochelia, P. anomala, R. Bamber (pers. comm. 2012), is preparing a comparison of the males and females attributable to these three apparently closely-related genera.



TABLE 2. Summary of possible taxonomic affinities or designations for the nine nominal species not considered members the genus Pseudoleptochelia in this study. Original generic designation

Postulated taxonomic status or possible generic affinities (this study)

Known sex(es)

Pseudoleptochelia anorexia Bird & Bamber, 2000

Leptochelia sensu lato

Female

Heterotanais antarcticus (Lang, 1953)


Both

Pseudoleptochelia bulbus Bamber, 2006

Being revised*

Both

Tanais filum (Stimpson, 1853) sensu Wallace, sensu Lang

?Undesignated leptocheliid genus

Both

Pseudoleptochelia filum sensu Sieg (1976)

Leptochelia (undescribed)

Both

Heterotanais inermis (Dollfus, 1898)


Male

Heterotanais longidactylus (Băcescu, 1977)

Being revised*

Both

Heterotanais mercantilis (Smith, 1906)


Male

Heterotanais mergellinae (Smith, 1906)


Male

Heterotanais provincialis (Dollfus, 1898)

Pseudoleptochelia?

Male

*Bamber (in prep.)

Bird (2011) erected the genus Parakonarus to receive the new species P. kopure Bird, 2011, which occurs in New Zealand waters. More recently Edgar (2012) described a second species, P. robertsoni, from Australian waters. As described and illustrated by Bird (2011) and Edgar (2012) for P. kopure and P. robertsoni, respectively, the males and females of both these species resemble those of Pseudoleptochelia fairgo; however, the carpal process on their cheliped is much less pronounced. The females of Parakonarus also are similar to those of Konarus (see Bamber 2006; Bamber & Chatterjee 2010). These two genera are also quite similar to and share a number of characters with those of Pseudoleptochelia as defined here; these include a stout dorsally wide antennule and a transverse suture on the carapace. In a revision of genus Konarus Bamber (in press) has identified the male for this genus. Also, as in the Parakonarus kopure, the females of P. anomala (examined during the present study), P. fairgo, and the new species described herein have pereopods 4 to 6 with the merus bearing a pair of small basally swollen-distally, attenuated setae (i.e. “pedestalbased inferior spines” of Bird 2011). These setae appear to be unknown, undescribed, or possibly overlooked in the females for Parakonarus robertsoni (illustration indistinct), Konarus crassicornis (unknown), Pseudoleptochelia mortenseni (undescribed), P. straddi (female unknown), and P. magna (unknown). Based on the original illustration for K. cheirus, these setae appear more strongly developed and not as finely attenuated as those of Pseudoleptochelia anomala. The homologous spiniform setae occurring on the female of the highly derived genus Makraleptochelia are not greatly attenuated distally. The females of P. kopure and P. robertsoni differ from those of Pseudoleptochelia by the relative lengths of the pereonites and by having more articles in the uropodal endopod. As previously mentioned, the male chela of both species of Parakonarus has a low rounded carpal process (Fig. 8L), which is distinctly less pronounced than those in P. fairgo (Fig. 8D) and the new Puerto Rican species (Fig. 8B) described herein; this process is not recognizable on the carpus of P. anomala and P mortenseni (Figs. 8K; 8A, respectively). The setation on the propodus of pereopods 4–6 for the female for Parakonarus kopure appears to be less similar to P. fairgo than to the new Puerto Rican species. Again, the status of the members of the Pseudoleptochelia complex will be better understood when detailed descriptions for the male and female of P. anomala, which are currently being prepared by Roger Bamber, become available.

Pseudoleptochelia juliae n. sp. Figs. 2–7, 8B, 9–12 Synonyms. Pseudoleptochelia sp. A: Heard et al. (2004), p 112, fig. 151; Suárez-Morales, et al. (2004), pp. 72–76, figs 36–39; and García-Madrigal et al. (2005), p 1173.



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Material examined. Pseudoleptochelia juliae Holotype. Adult male, length 2.2 mm, (USMN 1204767), 17° 56’47.41’’N, 67° 04’56’’W, North San Cristóbal Southwest of La Parguera, Puerto Rico, depth 12.2 m, collected with benthic grab in June 18 of 2008. Paratypes (same collection data as for holotype): two males (USMN 1204768), seven females (USMN 1204769); two males and two females (GCRL 6510). Additional specimens from the type locality are in the collection of authors. Pseudoleptochelia anomala three males, twelve females, Balearic Islands, SpainǤ Diagnosis. Female: Antenna with article-3 having anteriodorsal margin with simple seta only. Maxilliped having inner margin of each endite with two rounded denticles. Carapace with transverse suture indicating second thoracomere. Chela inner comb with two to three stout setulate surrounded by minute setae. Pereopods 4 to 6 posterodistal margin with pair of short, stout pectinate, spiniform seta. Male: Antennule with flagellum article-1 appearing composite with 5–6 bundles of aesthetascs, 10–12 partially fused or unfused articles each with distoventral tuff of aesthetascs, distal article minute with four long simple setae. Antenna with dorsal margins of articles 2 to 4 each with long, anteriorly curved, simple seta; ventral margins of articles 2 to 3 lacking simple setae; distoventral margin of article-4 with long simple seta. Cheliped subchelate, appearing carpochelate, carpus with well-developed sub-acute inferior process; propodus length more than twice width, fixed finger reduced to short distal process with strong spine on inner margin. Uropod in both sexes with endopod usually having four articles or incipient articles. Etymology. This species is named in honour of Julia Stella Núñez, the mother of the senior author. Type locality. Off North San Cristóbal southwest of La Parguera, Puerto Rico, depth 12.2 m, soft substrata (sand). Distribution. Puerto Rico; possibly southern Florida, eastern Gulf of Mexico, and Mexican Caribbean. Description. Adult Female. Body (Figs. 2A–B; 11): Length 2.6 to 3.2 mm, about 7.5 times width. Carapace (Figs. 2A–B; 9A–B): About 18 % of total body length, slightly longer than pereonites 1–2 combined; distinct transverse dorsal suture on posterior third with finer line extending anterolaterally to each ventral edge (apparently indicating demarcation of second thoracomere); mid-dorsal divided-plate structure between dorsal suture and dorsal margin of first pereonite. Eye lobes sub-acute with fine suture separating them from carapace (Figs. 2B−9B). Eyes dark pigmented having 8–10 ommatidia. Four to six simple setae laterally behind eye lobes (Figs. 2A–B; 9A–B). Pereonites (Fig. 2A): About 60 % of total body length; pereonites 1–3 and pereonite-6 wider than long, pereonite 4−5 longer than wide. Pleonites (Fig. 2A): About 16 to 18 % of total body length, pleonites 1–5 sub-equal in length, much wider than long; pleonite-1 with pair of small simple, sub-marginal setae dorsally; lateral margins of pleonites 2–5 each with mid-marginal seta (Fig. 2A). Pleotelson (Fig. 2A): About 5% of total body length, sub-equal in length to pleonites 4–5 combined, posterior margin with two pairs of medial terminal simple setae, innermost pair relatively long reaching nearly past basal article of uropod. Antennule (Figs. 3A; 9A–C, 9E): Relatively short and broad from dorsal and ventral aspect, about one seventh of total body length, with three articles plus terminal article. Article-1 length more than twice that of articles 2–3 combined, dorsal and ventral seta on mid-margin, four sensory setae on mid-ventral margin, simple seta and two sensory setae on sub-distal ventral margin. Article-2 slightly longer than wide, with one dorsal and one ventral simple setae, and sensory seta on dorsodistal margin. Article-3 length about 2.3 times width, with simple seta on dorsodistal margin. Terminal article minute, with five simple setae, one broom seta, and aesthetasc. Antenna (Figs. 3B; 9F): Article-1 reduced, asetose; article-2 slightly longer than wide, with two small simple setae on each distal margin; article-3 quadrate, with simple seta on distodorsal margin. Article-4 with length slightly less than combined lengths of article 2–3, about three times width, with two short and two long simple setae on sub-distal margin. Article-5 with length slightly more than half that of article 4, about 2.5 times width, two long simple setae on sub-distal margin. Article-6 minute, with two short and two long, simple setae. Mouthparts: Labrum (Fig. 3C): As illustrated. Mandible (Figs. 3D–E): Left: lacinia mobilis, broad, sub-triangular with three to four shallow, sub-distal denticles (Fig. 3D), incisor with one small proximal and two weak shallow distal lobes. Right: incisor weakly bidentate distally, with six denticles on upper margin, (Fig. 3E). Molar process stout, with grinding surface having fine denticulation.



FIGURE 2. Pseudoleptochelia juliae n. sp., paratype female: A, dorsal view; B, lateral view. Scale line: A–B = 1.0 mm.



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FIGURE 3. Pseudoleptochelia juliae n. sp., paratype female: A, antennule, lateral view; B, antenna, lateral view; C, labrum; D, left mandible; E, right mandible; F, labium; G, maxillule; H, maxilla. Scale line: A–D and F–I = 0.1 mm.

Labium (Fig. 3F): Bilobed, lobes equal and wide, finely setose distally. Maxillule (Fig. 3G): Endite having 9–10 relatively slender distal spines, sub-distal and outer margin setose. Palp slender with two long terminal setae of unequal length. Maxilla (Fig. 3H): Sub-quadrate.



Maxilliped (Figs. 4A–B): Bases distally unfused, distal margin usually with four, occasionally five, pairs of long simple setae, extending to palp article-2. Endites not fused, each having finely setose lateral margins with relatively long inwardly curved lateral seta, distomedial margin with two stout tooth-like spines, and inner distal margin with stout distally rounded denticle with minutely papillate (under high magnification) surface (Fig. 4B), inner medial margin with two pairs of small narrow coupling hooks. Palp: Article-1 asetose. Article-2 with short simple seta on outer margin and cluster of three simple setae on sub-distal inner margin. Article-3 with five simple setae (proximal three strongly developed) on inner margin. Article-4 with seven distal simple setae, and simple seta sub-distally.

FIGURE 4. Pseudoleptochelia juliae n. sp., paratype female: A, maxilliped, inner face endite showing coupling hooks; B, enlargement of distal part of endites showing “globular denticles”; C, epignath; D, left cheliped, lateral view; E, right cheliped, inner view. Scale line: A, C–E = 0.1 mm.



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FIGURE 5. Pseudoleptochelia juliae n. sp., paratype female: A–F, pereopods 1 to 6, respectively; G, enlargement of styliform setae; H, enlargement of tubercle-like stout carpal spine; I, enlargement of broad simple, stout carpal spine,; J, enlargement of stout pectinate, spiniform propodal seta; K, pleopod; L, uropod. Scale line: A–F and K–L = 0.2 mm.



Epignath (4C): Elongate, strap-like, with margins finely setose. Cheliped (Figs. 4D−E; 9B–D): Coxal sclerite sub-triangular, dorsally inserted with respect to basis, having two simple setae (Figs. 4D; 9D). Basis elongate, length about 2.5 times width; simple seta on sub-distal lateral margin. Merus triangular, having posterior margin with two closely-approximated rows (one with five and other with three) of long simple setae. Carpus length about 1.8 times width, with anterior margin having six small spiniform setae and simple setae distally; three long simple setae on mid-posterior margin; distal margin extending to form cuff partially covering propodus. Propodus length about twice width, simple seta on anterodistal margin just below dactylus, fixed finger with anterior margin having three simple setae on outer incisive margin and two on midposterior margin; Inner aspect (Figs. 4E; 10A): with oblique dense row of minute setae and two to three small, stout setulate setae just proximal to articulation with dactylus. Dactylus (movable finger) longer than fixed finger, inner face with setulate seta proximally. Pereopod-1 (Fig. 5A): Distinctly longer than other pereopods. Coxa with simple seta on anterodistal margin. Basis slender, length about 3.3 times width, and length about equal to that of ischium, merus and carpus combined; anteroproximal margin with simple seta. Ischium reduced, with simple seta. Merus appearing asetose, length about half that of basis. Carpus length about equal to that of merus, with five distal and sub-distal setae. Propodus length about 3.6 times width, slightly longer than carpus; four sub-distal simple setae, three (one minute) on or near anterior margin, and one on posterior margin. Dactylus narrow, styliform, naked; together with unguis longer than propodus. Unguis longer than the dactylus Pereopod-2 (Figs. 5B; 2B): Coxa with simple seta on anterodistal margin (Fig. 2B). Basis length about equal to that of ischium, merus, and propodus combined; proximal third of anterior margin with simple seta and two sensory setae. Ischium with two setae (one minute), on anterior margin. Merus appearing asetose, length about one third that of basis. Carpus length about equal to that of merus; antero- and posterodistal margins each with simple seta. Propodus length slightly more than 1.3 times that of carpus with two relatively-long simple setae (longest about equal in length to that body of propodus), posterodistal margin with minute spiniform seta. Dactylus and unguis length together slightly more than half that of propodus; dactylus with simple accessory seta (not shown) originating just proximal to junction of dactylus and unguis. Unguis with pore at tip for extrusion of tube-building material. Pereopod-3 (Fig. 5C): Similar to pereopod-2, but smaller. Basis slightly less than 40% total length of pereopod, antero-proximal margin with simple seta and sensory seta. Reduced ischium with simple seta on anterior margin. Merus length slightly more one third that of basis; minute spiniform seta on posterodistal margin. Carpus, propodus, and dactylus as in pereopod-2. Pereopod-4 (Figs. 5D; G–J; 10B): Basis, length about 1.4 times that of ischium, merus, and carpus combined; cluster of small sensory setae on mid-anterior margin and two sensory setae on posterior margin. Ischium about three times as wide as long, with mid-posterior simple setae. Merus, length about 1.4 times width, with distal posterior margin bearing two setae attenuated distally into styliform tips (Fig. 5G). Carpus, length about 1.3 times width, articulated obliquely with merus, anterodistal margin with small simple seta; posterodistal margin with three stout (as wide as long or wider), tooth-like dissimilar setae (Figs. 5H–I). Propodus, narrower and slightly longer than carpus; anterodistal margin with three subequal finely pectinate setae (longest about equal in length to dactylus); posterodistal margin (immediately adjacent to articulation with dactylus) with pair of short, stout pectinate, spiniform seta (Figs. 5J; 10B). Dactylus articulated posterodistally with propodus; small simple seta just proximal to unguis; unguis minute, with dactylus forming claw-like tip. Pereopod-5 (Figs. 5E; 10C): Similar to pereopod-4, except basis appearing asetose. Ischium with two simple setae on posterior margin. Pereopod-6 (Figs. 5F, 10D): Similar to pereopods 4–5, except ischium with single simple seta; propodus with anterodistal margin having two sub-equal long finely pectinate and three curved, “comb-like” setulate setae. Pleopods (Fig. 5K): Five similar, well-developed, biramous pairs. Basal article broad, with inner plumose seta. Rami with lengths slightly more than twice widths, both with proximally inflated plumose seta on distal inner margin adjacent to articulation with basis. Endopod with inner and distal margins bearing 10 long plumose setae, distal-most seta modified (greatly attenuated, filament-like distally); with strongly developed plumose seta on midlateral margin. Exopod with inner and distal margins bearing 14 long plumose setae, lateral margin asetose. Uropod (Fig. 5L): Basal article about as long as wide, asetose. Exopod uniarticulate, extending past midpoint of article-2 of endopod, with simple seta on outer mid-margin, two distal simple setae (outer seta longest, about



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equal combined lengths of basis and exopod; inner seta about one third length of outer). Endopod usually with four or five incipient articles; article-1 almost half length of exopod, with inner distal simple setae. Article-2 length about twice width, inner distal margin with sensory seta. Article-3 nearly as long as exopod with simple seta and sensory setae near inner distal margin. Article-4 almost same length as article-3 with simple seta on mid-margin with sensory seta and five (three long, two small) simple setae and on inner distal margin.

FIGURE 6. Pseudoleptochelia juliae n. sp., holotype male: A, lateral view; B, enlargement of antennule tip; C, enlargement of antennule peduncle-flagellum junction; D, antenna; E, maxilliped. Male (form A): F, male, lateral view. Scale line: A and F = 1.0 mm; B–D = 0.2 mm; E = 0.1 mm.



FIGURE 7. Pseudoleptochelia juliae n. sp., holotype male: A, right cheliped, inner view; B, enlargement of margin near at articulation of dactylus; D–G, pereopods 1–2, 4, and 6, respectively; H, enlargement of spiniform carpal seta; I, uropod. Male (form A): C, right cheliped, inner view. Scale line: A and C = 0.1 mm; D–I = 0.2 mm.



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Male. Body (Figs. 6A, 12A): Natatory, smaller than female. Length about 2.0 to 2.3 mm. Cephalothorax (Fig. 6A): Length of carapace nearly equal to that of pereonites 1 to 3 combined, rostrum weak; ocular lobes bearing large eyes with visual elements present, eyes about 5 times larger than in female, about 25 % of total carapace length. Pereonites (Fig. 6A): About 50% of total body length. Pereonites 1 to 4 becoming progressively longer; pereonite-5 same length that as pereonite-4; pereonite-6 same length as pereonite-3; pereonites 1 to 3 wider than long; pereonites 4 to 5 same length, and quadrate. Pleonites (Fig. 6A): About 20% of total body length, combined length about equal to that of combined length of pereonites 5–6; all pleonites sub-rectangular, wider than long; pleonite-1 with one small simple seta on midlateral margin; pleonites 2–5 with two small simple setae on mid-lateral margin. Pleotelson (Fig. 6A): About 5% of total body length with rounded margin, length slightly less than combined length of pleonites 4–5; posterior margin with two pairs of simple setae, inner-most pair relatively long extending well-past uropodal basal article. Antennule (Figs. 6A–C): Peduncle with three articles. Article-1, length about 1.7 times width with disto-dorsal simple setae. Article-2, length about 1.6 times width, with disto-ventral rounded apophysis. Article-3 saddle-shape forming dorsal shelf over first apparent flagellar article, with small simple seta dorsally and laterally (Fig. 6C). Flagellum having ventral margin with 15–16 aesthetasc bundles and 10–13 articles. Article-1 with four to five bundles or short rows of aesthetascs. Article-2 often partially fused with composite article 1. Articles from article2 to penultimate article (10 or 11) each with disto-ventral bundle of aesthetascs. Terminal article minute with simple setae (Fig. 6B). Antenna (Fig. 6D): With six articles. Article-1 compressed, asetose. Article-2 wider than long, with sensory seta ventrally on sub-distal margin; long simple seta dorsally on distal margin. Article-3, smaller than article-2, wider than long, ventral margin with sensory seta sub-distally; dorsal margin with long simple seta distally. Article4 length about 3.7 times width, longer than combined length of articles 2 to 3, dorsal margin with long simple setae medially; ventral margin with sensory seta medially and long sub-distal seta. Article-5 length about 6.5 times width, slightly longer than article-4, with four simple setae distally. Article-6 minute, with three long terminal setae. Mouthparts: Unrecognizable, atrophied mass. Maxilliped (Fig. 6E): Atrophied, area of basis with four simple setae; palps greatly reduced with two recognizable, partially fused articles. Cheliped: Inner aspect (Fig. 7A–B): Subchelate and appearing carpochelate. Basis length about twice width. Merus triangular with posterior margin having three distal and one proximal simple setae. Carpus with anterior margin having six small spines and simple seta distally; posterior margin with disto-ventral apophysis bearing three simple setae. Propodus length about 2.6 times width; fixed finger reduced to small pointed apophysis with small tooth on inner proximal margin, with five inner (2 anterior and 3 posterior) simple setae; inner palm of propodus with comb row of 13 setae (11 short and two longer setae near base of dactylus). Dactylus slender, equal in length to propodus, with one simple seta near to anteroproximal margin; posterior margin with row of fine setae along cutting edge. Pereopod-1 (Fig. 7D): Similar to that of female, except for minor differences in setation and proportionately longer carpus and propodus. Dactylus, long falciform; including unguis, slightly shorter than propodus; unguis slightly longer than dactylus. Pereopod-2 (Fig. 7E): Appearing more elongate than in female, not noticeably modified for tube construction. Coxa with simple seta on anterodistal margin. Basis similar to that of female with length nearly 3.0 times width, simple seta on anterior proximal margin. Ischium reduced with two minute setae on posterior margin. Merus length about one third that of basis; small, stout, spiniform seta on posterodistal margin. Carpus length about 1.8 times that of merus; anterodistal margin with well-developed, narrow, acute spiniform seta; posterodistal margin with two spiniform setae. Propodus length slightly more than 1.2 times that of carpus, anterior margin with sub-distal seta; posterodistal margin with pair of narrow spiniform setae. Dactylus and unguis together weakly curved posteriorly, length slightly more than 40% that of propodus; unguis, about one third length of dactylus. Unguis appearing to lack distal opening. Pereopod-3 (not figured): Similar to pereopod-2.



FIGURE 8. Examples of the male chelae of the Pseudoleptochelia complex, indicating those with a prominent carpal process, underdeveloped carpal process, or lacking carpal process. A, Pseudoleptochelia mortenseni; B, P. juliae; C, P. juliae (male form A); D, P. fairgo; E, P. straddi; F, P. bulbus; G, P. provincialis; H, P. algiricus; I, P. magna; J, P. anomala; K, P. anomala; L, Parakonarus kopure. [Figures not authored from: Lang 1973 (A); this study (B−C); Bamber 2005 (D); Bamber 2008 (E); Bamber 2006 (F); Dollfus 1898 (G–H); Smith 1906 (I); this study (J−K); and Bird 2011 (L)].



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FIGURE 9. Pseudoleptochelia juliae n. sp., female (SEM images): A, carapace showing transverse suture of specimen no.1, dorsal view; B, carapace showing transverse suture of specimen no. 2, lateral view; C, enlargement of thorax/carapace region of specimen no. 3, ventral view; D, enlargement of coxal sclerite of specimen no. 2, lateral view; E, enlargement on antennule and antenna of specimen no. 3, ventral view; F, enlargement tip of antenna of specimen no. 4, lateral view.

Pereopod-4 (Figs. 7F and 7H): Basis, length about 2.8 times width. Ischium reduced, appearing asetose. Merus, length about 2.4 times width, with distal posterior margin bearing two well-developed, narrow, spiniform setae (not proximally inflated as in female). Carpus length slightly longer than merus, anterodistal and sub-distal margins with small sub-distal simple seta and three spiniform setae (Fig. 7H). Propodus length about two thirds that of basis; distal margin with three setae and sub-distal spiniform seta. Dactylus and unguis, narrow, length about 70% that of propodus, weakly curved posteriorly with minute setae on posterior concave margin; unguis about on third length of dactylus. Pereopod-5 (not figured): Similar to pereopod-4.



Pereopod-6 (Fig. 7G): Similar to pereopods 4–5, except for basis slightly longer; propodus shorter with superior distal margin with five, two long finely pectinate and three curved setulate “comb-like” setae. Pleopod (not figured): Five pairs, similar to, but more strongly developed than those of female with longer plumose setae. Uropod (Fig. 7I): Similar to the female, but exopod being a little more setose. Remarks. As redefined here, we currently recognize seven nominal species of Pseudoleptochelia: P. anomala, P. fairgo, P. magna, P. mortenseni, P. straddi, P. juliae n. sp., and the recently described Pseudoleptochelia occiporta. As mentioned previously, P. occiporta which has affinities with P. fairgo, be will treated by R. Bamber in the near future, and it has not been included in the key presented here. An apparently undescribed species of Pseudoleptochelia from the northern Pacific coast of Costa Rica was reported by Heard et al. (2009); but no males are known for this species. The males and females of “Pseudoleptochelia sp. A” (sensu Heard et al. 2004; SuárezMorales et al. 2004; and García-Madrigal et al. 2005) are very similar to those of P. juliae. Although there are some slight morphological variation exhibited by these more northern and western populations, we tentatively consider them as conspecific with the specimens of the new Puerto Rican species. The nominal species of Pseudoleptochelia, excluding P. occiporta, can be separated in the following key.

FIGURE 10. Pseudoleptochelia juliae n. sp., female (SEM images): A, right chela of specimen no. 5, inner view; B, enlargement of dactylus/propodus of pereopod-4 of specimen no. 2; C, enlargement of dactylus/propodus of pereopod-5 of specimen no. 2; D, enlargement of dactylus/propodus of pereopod-6 of specimen no. 2.

Key to the recognized males of Pseudoleptochelia based on the cheliped. 1. 2. -

Cheliped with ventral carpal apophysis or process (appearing carpochelate). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Cheliped lacking ventral carpal apophysis or process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Cheliped with propodus having mid-ventral rounded apophysis just distal to setal comb row (Fig. 8E) … P. straddi [Australia] Cheliped with propodus lacking mid-ventral rounded apophysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3



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3. 4. 5. -

Cheliped having propodus with 17 setae in comb row; fixed finger without distinct sub-distal tooth (Fig. 8I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. magna [Mediterranean] Cheliped having propodus with 14 or less setae in comb row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Cheliped having propodus with 11 anterior setae in oblique, sub-distal, comb row; fixed-finger with distinct sub-distal tooth or tubercle (Fig. 8A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mortenseni [Southern Caribbean: Tobago] Cheliped having propodus with about 14 setae in sub-linear comb row (Fig. 8J); fixed finger without distinct sub-distal tooth (Fig. 8K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. anomala [Mediterranean] Cheliped having the ventral carpal processes and the ventral propodal processes blunt and rounded (Fig. 8D); propodus relatively robust, length less than twice width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fairgo [Australia] Cheliped having the ventral carpal processes and ventral propodal processes sub-acute to acute (Fig. 8B). Propodus relatively narrow, length distinctly more than twice width . . . . . . . . . . . . . . . . P. juliae [Caribbean, Gulf of Mexico, Florida East coast].

FIGURE 11. Pseudoleptochelia juliae n. sp., female (digital image): dorsal view of habitus, specimen no. 7.



FIGURE 12. Pseudoleptochelia juliae n. sp., male and male form A (digital images): A, male, lateral view of habitus, specimen no. 8; B, male form A, lateral view of habitus, specimen no. 9.

The male of P. juliae is distinguished from that of P. mortenseni, the only other nominal member of the genus known from the NW Atlantic, by the latter having chela with a only a shallow, raised bump on the carpus, with three associated simple setae (Fig. 7A, 8A, respectively). The chelae of the two species also differ in the number and form of the comb setae on the inner sub-distal face of the propodus; there are 13 setae becoming gradually longer posteriorly (ventrally) on P. juliae (Fig. 7A), whereas on P. mortenseni the setal row is irregular with 11 setae (Fig. 8A); the first seta is longer than all but the last two, which are subequal and distinctly longer than all the rest (Fig. 8A). Further, the male antennule of P. juliae has ten aesthetasc-bearing articles whereas that of P. mortenseni has seven. The stout female antennule attributed to small females/ neuters of P. mortenseni by Lang (1973: fig. 16g) is quite similar to those of P. juliae (see Remarks for the genus). Until their mouthparts can be carefully compared, we are unable to reliably separate the females of these two Caribbean species. Another form of a Pseudoleptochelia male (form A) co-occurs in Puerto Rican waters with the males and females attributed to P. juliae. Though similar (see Figs. 6F; 12B) in many aspects to the male holotype of P. juliae, there are some distinct differences. “Male form A” is briefly diagnosed as follows:



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Male form A: Antennule (Fig. 6F) with flagellum having nine aesthetasc-bearing articles. Cheliped; lateral aspect (Fig. 6F) carpus with anterior margin having six small spiniform setae and simple setae distally; with distinct narrow process on posterior margin; inner aspect (Fig. 7C) with propodus having sub-distal, oblique combrow of 13 to 14 stout setae gradually increasing in length anteriorly [eleven short and two long setae in juliae]; fixed finger of propodus with proximal tooth adjacent to articulation with dactylus. Dactylus weakly falcate, subequal in length to propodus with proximal simple seta on upper inner margin, three minute setae on inner concave margin. There is a possibility that P. juliae and the ‘Pseudoleptochelia male Form A’ may represent two sympatric, cryptic species for which their respective females appear to be indistinguishable. This may also be the case for P. fairgo and P. straddi (see Bamber 2008). With respect to non-Caribbean species, the male and female of P. juliae appear to be most similar to those of the Australian P. fairgo (see Bamber 2005, 2008). This species has recently been transferred to the genus Parakonarus by Edgar (2012); however, for the reasons given previously under the Remarks for the genus we retain it within Pseudoleptochelia. The males for both P. juliae and P. fairgo have a prominent carpal process making them appear carpochelate (Figs. 8B and 8D). In the male of P. anomalus, a distinctive carpal process is not developed, although possibly represented by a raised ridge bearing a few simple setae (Fig. 8K). In the subchelate male of Parakonarus sensu Bird 2011 (Fig. 8L), there is a much less prominent blunt process with simple setae. The male of Pseudoleptochelia juliae can be distinguished from that of P. fairgo by the cheliped having the propodus distinctly narrower (length more than twice width) and the carpus having a sub-acute carpal process or apophysis. The females are separated by P. juliae having (1) more “spatulate spines” on the distal medial inner margins of the maxillipedal endites, (2) a greater number of setae on the carpus of the cheliped (eight versus five), (3) apparently fewer articles or incipient articles in the uropodal endopod (four versus five), and (4) pereopods 4–6 posterodistal margin with pair of short, stout pectinate, spiniform seta. The females of P. juliae and P. fairgo, as well as those of the genera Konarus and Parakonarus are characterized by having (1) the maxillipedal basis with four or five sub-distal pairs of long simple setae (see Fig. 4A); and (2) the merus of pereopods 4–6 having a pair of distinctive proximally swollen and distally attenuated (styliform) setae (appearing setulate) and a carpus with stout spiniform setae (as wide or wider than long) on the distal margin (Figs. 5G; 5H−I, respectively). The antennae of P. anomala, P. juliae, P. fairgo and members of genera Konarus and Parakonarus also lack distinctive buttressed thorn setae on the distal margins of their second and third antennal articles. This may, at least superficially, link these two genera, and possibly the other members of “anomala group,” to the subfamily Heterotanaidinae sensu Larsen & Wilson, 2002. In a more recent cladistical treatment of the superfamily Paratanaoidea, Bird & Larsen (2009) the two leptocheliid subfamilies were not resolved separately but a more refined approach may do this. Based on our observations on the male of P. juliae and the descriptions and illustrations of Bird (2011) for the males of Parakonarus kopure and P. robertsoni, we believe that the first “apparent” article of the antennular flagellum represents a composite of three to six precursor proximal articles. The first “apparent” article in the antennular flagellum of P. juliae, P. mortenseni, and P. kopure, have five to six, four, and three clusters of aesthetascs, respectively. This condition has not been reported for the males of P. fairgo and P. straddi nor in the earlier descriptions or illustrations for the males of P. anomala (see Sars 1886, Smith 1906, Riggio 1996) and P. magna (see Smith 1906); however, based on examination of the males we attribute to P. anomala, three clusters of aesthetascs are present on the first apparent flagellum article. The significance of this apparent fusion and compression of the first few proximal flagella articles for males within the Pseudoleptochelia complex remains to be determined. Habitat: Pseudoleptochelia juliae occurred on soft bottom sites off La Parguera and Culebra Island in depths ranging from 1 to 28 m. Other tanaidaceans co-occurring with it included: Mesokalliapseudes quadriflagellata Drumm & Morales-Núñez, 2012, Leptochelia spp., Pseudotanais sp., and Swireapseudes sp.

Acknowledgments This research represents in part the results of a dissertation submitted to the Department of Marine Sciences, University of Puerto Rico Mayagüez-Campus (UPRM) by the senior author in partial fulfilment for the degree of



Doctor of Philosophy. We thank participants in this study for their help in the collection of samples used in this research. AGM-N thanks Puerto Rico Sea Grant College Program (PRSGCP), Faculty of Arts and Sciences, and Research and Development Center at University of Puerto Rico Mayagüez-Campus (UPRM) for providing financial support to work at the Gulf Coast Research Laboratory (GCRL) of the University of Southern Mississippi. We extend out thanks to Sara E. Lecroy and Jerry A. McLelland for their comments and suggestions on early version of this paper. We wish to express our gratitude to Graham Bird and Roger Bamber for their helpful comments and constructive suggestions; we take responsibility for any remaining errors what we were unable to address. We gratefully acknowledge Victoria Fernandez-Gonzales for kindly supplying us with specimens of P. anomala from the Mediterranean (Balearic Islands, Spain). We also wish to acknowledge Michael Nemeth Feliciano (UPRM) for preparing Figure 1 and José Almodár for kindly prepare the scanning electronic and digital images, Microscopy Center (UPRM).

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