A NEW SPECIES OF EMERITA (DECAPODA, ANOMURA, HIPPIDAE) FROM TAIWAN, WITH A KEY TO SPECIES OF THE...

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Crustaceana 88 (3) 247-258

A NEW SPECIES OF EMERITA (DECAPODA, ANOMURA, HIPPIDAE)FROM TAIWAN, WITH A KEY TO SPECIES OF THE GENUS

BY

PAN-WEN HSUEH1)

Department of Life Sciences, National Chung Hsing University, Taichung, Taiwan

ABSTRACT

A new species, Emerita taiwanensis sp. nov., is described from Taiwan and represents the firstrecord of the genus from East Asia. The species differs from its congeners by having (1) a longand sharply triangular median frontal lobe, which is approximately equal in length to the laterallobes, (2) the highest ratio of the maximum length of the merus of the 3rd maxilliped to that samesegment’s maximum width, (3) a narrow and elongate P1 dactylus, combined with a ratio of the P1dactylus length to its maximum width of approx. 2.78, (4) 1-3 spines present on the inner margin ofthe P1 dactylus, and (5) a peculiar colour pattern on the dorsum of the carapace. A comparison of themorphological characters of the present species with those of its congeners with a similar, elongateP1 dactylus is presented in the form of a table.

RÉSUMÉ

Une nouvelle espèce, Emerita taiwanensis sp. nov., est décrite de Taiwan, elle représente lepremier signalement de ce genre en Asie de l’Est. L’espèce diffère de ses congénères en ayant (1)un lobe frontal median long, en triangle aigu, qui est approximativement égal en longueur aux lobeslatéraux, (2) le rapport le plus élevé de la longueur maximale du mérus du 3ème maxillipède à lalargeur maximale de ce segment, (3) un dactyle de P1 étroit et allongé, combiné à un rapport dela longueur de P1 dactyle à sa largeur maximale d’environ 2,78, (4) 1-3 épines présentes au bordinterne du dactyle de P1, et (5) un patron de couleur particulier sur la partie dorsale de la carapace.Une comparaison des caractères morphologiques de la présente espèce avec ceux de ses congénèresayant un dactyle de P1 similaire allongé, est présentée sous forme de tableau.

INTRODUCTION

The family Hippidae Latreille, 1825 currently contains three genera, EmeritaScopoli, 1777, Hippa Fabricius, 1787 and Mastigochirus Miers, 1878 (Boyko &McLaughlin, 2010). A recently published monograph on crab-like anomurans from

1) e-mail: pwhsueh@dragon.nchu.edu.tw

© Koninklijke Brill NV, Leiden, 2015 DOI 10.1163/15685403-00003413

248 PAN-WEN HSUEH

Taiwan and nearby islands recorded six species from two genera: Hippa adactylaFabricius, 1787, H. admirabilis (Thallwitz, 1892), H. marmorata (Hombron &Jacquinot, 1846), H. ovalis (A. Milne-Edwards, 1862), H. truncatifrons (Miers,1878) and Mastigochirus gracilis (Stimpson, 1858) (Osawa et al., 2010). Membersof the third hippid genus, Emerita, which are mostly known from the coasts ofthe Americas and in the Caribbean, have not been recorded from East Asianwaters, although some species have been reported from the Indo-West Pacific(Niazi & Haque, 1974; Efford, 1976; Boyko & Harvey, 1999). Recently, specimensof a mole crab were collected from the northeastern and southwest coasts ofTaiwan. These specimens have a narrowly ovoid carapace with strongly convexdorsal surface, slender and elongate eyestalks, large and long second antennae,and lamellate dactyli of the first pereopods (P1), characters, which agree withthe diagnosis of Emerita given by Barnard (1950) and Boyko & Harvey (1999).Therefore, all three genera of Hippidae are present in Taiwanese waters (Osawa etal., 2010; present study).

In 1777, Scopoli erected the monotypic genus Emerita (Boyko & McLaughlin,2010), which is now represented by 10 species (Boyko & McLaughlin, 2010).Rathbun (1900) and Heegaard & Holthuis (1960) moved Hippa analoga Stimpson,1857, and Hippa talpoida Say, 1817, to Emerita. Schmitt (1935, 1937) describedfive new species from the Americas and the Caribbean, while Sankolli (1965) andNiazi & Haque (1974) both reported one new species each from the Indian Ocean.The present study adds one species to the genus, the first from East Asia.

MATERIAL AND METHODS

Thirty-six specimens were collected using shovels and 2-mm-mesh sieves on30 March, 1 and 7 September 2013 from the surf zone on sandy beaches of thenortheastern and southwestern coasts of Taiwan. Environmental parameters, suchas surface seawater salinity and sand grain size of the bottom substrate, were alsomeasured and collected from both sampling locations in early September. Sand wasanalysed in the laboratory for grain size and composition. The abbreviations usedare: CL, carapace length (mm); P1-P3, pereopods 1-3; NMNS, National Museumof Natural Science (Taichung, Taiwan); NCHU, National Chung Hsing University(Taichung, Taiwan).

TAXONOMY

Family HIPPIDAE Latreille, 1825Genus Emerita Scopoli, 1777

Emerita taiwanensis sp. nov. (figs. 1-3)

Material examined.— Holotype, female (13.1 mm CL) (NMNS7194-01), the river mouth of Zu-An Creek (24°50′30′′N 121°49′40′′E), Touchien, I-Lan County, Taiwan, coll. P.-W. Hsueh & H.-T.

EMERITA TAIWANENSIS NOV. 249

Hong, 7 September 2013. Paratypes: 3 females (5.2 mm, 10.1 mm, 11.6 mm CL) (NMNS7194-02through NMNS7194-04), 30 females (5.4-9.7 mm CL) (NCHU), same data as holotype; 1 female(5.9 mm CL), same locality as holotype, coll. P.-W. Hsueh & H.-T. Hong, 1 September 2013; 1female (10.3 mm CL) (NMNS7194-05), Ah-Kon-Dein Creek (22°48′10′′N 122°12′49′′E), Mi-Tou,Kaohsiung, Taiwan, coll. H.-T. Hong, 30 March 2013.

Diagnosis.— Adult female with long, distally acute, sharply triangular medianfrontal lobe, approximately equal in length to lateral lobes; merus of 3rd maxillipedmaximum length to maximum width ratio about 1.81; P1 dactylus narrow, 2.78times as long as greatest width, lower margin with 1-3 spines, no spine on uppermargin.

Description.— Body subcylindrical; anterior carapace with 3 lobes, separatedon either side by a U-shaped sinus, median lobe sharply triangular, distally acute,lateral lobes subacute, all 3 lobes approximately equal in length, curved slightlydownward distally (figs. 1A, 2A). Dorsum of carapace distinctly convex, coveredwith numerous straight or sinuous, uneven transverse ridges (except near posteriorand posterolateral margins), each transverse ridge formed by denticles, cervicaland branchial grooves clearly visible (figs. 1A, 2A). Post-frontal furrow absent,post-gastric furrow clearly present (figs. 1B, 2B). Setose pits present on marginsor submargins of carapace (with exception of margins of frontal lobes and U-shaped sinuses); each setose pit accompanied by a minute ridge, giving serratedappearance (figs. 1B, 2B). Posterolateral carapace margins rounded (figs. 1B, 2B).

Eyes with slender, elongated peduncles, length (including cornea) exceeding tipof longest spine on second antennal segment (figs. 1A, 2A). Antennular flagellumwith 19 articles (figs. 1B, 2A). Second antennal segment with 3 large spines distally(fig. 2C), all with corneous tips, median spine longest; oblique longitudinal ridgepresent on base of second antennal segment, extending from base of outer spine toposterior of median spine, with 12 setae (fig. 2C).

Merus of third maxilliped subrectangular, approximately 1.8 times as long asgreatest width, setose pits present on all margins except on articulations, each se-tose pit accompanied by minute ridge (fig. 3A); anterolateral angle of outer marginrectangular, not produced, anterointernal lobe high, triangular, distally rounded;surface of propodus with long oblique ridge (fig. 3A). Inner surface of merus withoblique shaft, extended from middle of anterior margin to posterolateral corner,groove present on lower half of shaft (fig. 3B).

P1 dactylus elongated, 2.78 times as long as greatest width, distally subacutewith one spine, ovate at base, with corneous tip; lower margin with 1 (left P1dactylus) to 3 (right P1 dactylus) spines, no spine on upper margin, a row of 21minute spines along anterior lower margin (fig. 3C-D); oblique ridge present neardactylus base (fig. 3C). P2-3 dactylus falcate, basally broad, distally subacute, thin(fig. 1B).

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Fig. 1. Fresh specimen of Emerita taiwanensis sp. nov. Holotype, female (13.1 mm CL)(NMNS7194-01): A, dorsal view; B, lateral view; C, ventral view. This figure is published in colourin the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.

com/content/journals/15685403.

EMERITA TAIWANENSIS NOV. 251

Fig. 2. Emerita taiwanensis sp. nov. Holotype: female (13.1 mm CL) (NMNS7194-01): A, dorsalview; B, lateral view showing carapace and first to fourth abdominal somites; C, second segment of

right antenna. All setae omitted from figure. Scale bars = 1.0 mm.

First abdominal somite small, fused to carapace; second abdominal somitewidth approximately 2/3 carapace width, not overlapping with carapace, dorsumwith 2 long oblique ridges (fig. 1B). Third to sixth abdominal somites narrowingprogressively, each with 1 long oblique ridge. Telson sharply triangular, resemblingspearhead; lateral margins slightly convex, distally rounded (fig. 1C).

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Fig. 3. Emerita taiwanensis sp. nov. Holotype, female (13.1 mm CL) (NMNS7194-01): A, outersurface of third right maxilliped; B, inner surface of third right maxilliped; C, distal portion of right

pereopod I; D, distal right pereopod I. Setae in A, B omitted from figure. Scale bars = 1.0 mm.

Etymology.— The name is derived from the name of the island where thespecimens were collected.

Type locality.— Sandy beach adjacent to the river mouth of Zu-An Creek, I-LanCounty, northeastern Taiwan.

EMERITA TAIWANENSIS NOV. 253

Habitat.— The species inhabits sandy beach coasts of Taiwan with a brackishwater environment and medium + coarse sand dominated bottom substrates (grainsize 0.25-0.5 mm and 0.5-1.0 mm).

Distribution.— Only known from two distantly separated locations in Taiwan.Colour.— In life, body and dorsum of crab dark blue at anterior areas of

carapace. Dark blue fades and becomes brown to greyish brown at areas posteriorto cervical groove. A white longitudinal median line runs along carapace; whitelongitudinal line more distinct on smaller size mole crabs. Symmetrical anduneven sized brownish-white patches (triangular, polygonal or ovate in shape)noticeable on lateral gastric, cardiac and posterior branchial regions. SymmetricalC-shaped marks, formed by evenly-spaced small and bold white-brownish shortlines visible, apparently extension of branchial groove when observed with thenaked eye; another symmetrical C-shaped mark, formed by evenly-spaced, boldwhite lines of various lengths, present lateral to previous ones. Dorsum of carapaceat areas posterior to branchial groove toward posterior margin covered withnumerous, evenly distributed, minute white specks (fig. 1A). Dorsum of abdominalsomites greyish-blue with mixing of grey and brown; brownish white symmetricalpolygonal patches and minute white speckles visible near posterior margin andlateral areas of second to fifth abdominal somites (fig. 1B). Ventral body andappendages, as well as telson, are generally whitish (fig. 1C). Blue colour ondorsum of the carapace and abdomen fades and replaced by brown when animalsare kept in −20°C for two weeks or preserved in 70% ethanol; but above-mentioned colour patterns on dorsum and abdominal somites remain visible afterpreservation.

Remarks.— A single specimen was first collected from southwestern Taiwanon 30 March 2013 by H.-T. Hong, and although the same site was re-visited fourtimes in July, Hong and the author failed to collect any additional specimens.On 1 September 2013, Hong collected one more specimen from northeasternTaiwan. We revisited the site the following weekend and collected an additional34 individuals of various sizes during the ebb tide. This suggests the presence of ahealthy population of the present species at that location.

Most Emerita species are endemic to continental landmasses, but this is nottrue of the new species from East Asia. The author considered the possibility thatthe species is not native and may have reached Taiwan by anthropogenic means(for example, larvae transported by ballast water of cargo ships). It is somewhatsurprising that the new species has been missed until now. This is especiallyconsidering the prolonged larval stages and the high dispersal ability of Emeritaspecies (see Johnson & Lewis, 1942; Rees, 1959; Knight, 1967; Efford, 1970,1976; Tam et al., 1996; Sorte et al., 2001; Haye et al., 2002; Siddiqui & Ghory,2006). However, the morphological differences between the present species andknown congeners suggest otherwise. With regards to the elongate P1 dactylus, the

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present species falls into the same species-group as E. benedicti Schmitt, 1935, E.portoricensis Schmitt, 1935, E. holthuisi Sankolli, 1965 and E. karachiensis Niazi& Haque, 1974 (see Appendix for the Key to the species). However, E. benedictiand E. portoricensis are known to be endemic to warm regions of the westernAtlantic (Efford, 1976), and E. benedicti can be distinguished morphologicallyfrom the present species by having (1) a blunt-tipped, equilateral triangular medianfrontal lobe which is shorter than lateral lobes, (2) a higher P1 dactylus length/max.width ratio, and (3) a second abdominal somite width not exceeding 2/3 of carapacewidth, whereas E. portoricensis can be distinguished from the present speciesby having (1) a blunt-tipped, equilateral triangular median frontal lobe which isshorter than lateral lobes, and (2) a lower P1 dactylus length/max. width ratio(table I). On the same token, E. holthuisi and E. karachiensis are known endemicto the Indo-West Pacific (Niazi & Haque, 1974; Efford, 1976). Emerita holthuisidiffers from the present species by having (1) a blunt-tipped, equilateral triangularmedian frontal lobe which is shorter than lateral lobes, (2) equilateral triangularlateral lobes, (3) a merus of 3rd maxilliped max. length/max. width ratio about1.43, (4) a P1 dactylus length/max. width ratio about 2.00, and (5) three spines onthe P1 dactylus lower margin (table I). Emerita karachiensis can be distinguishedfrom the present species by having (1) a blunt-tipped, equilateral triangular medianfrontal lobe which is shorter than lateral lobes, (2) a merus of 3rd maxilliped max.length/max. width ratio about 1.67, (3) a P1 dactylus length/max. width ratio about2.06-2.09, and (4) 5-8 spines (excluding distal spine) on the P1 dactylus lowermargin (table I).

No male individuals were collected in the present study. A possible explanationfor this phenomenon could be that males are very small and was easily overlookedduring sampling. Efford (1967) suggested that neoteny in males is a characteristicof Emerita. He reported that five out of seven species studied (E. analoga(Stimpson, 1857), E. emeritus (Linnaeus, 1767), E. portoricensis, E. rathbunaeSchmitt, 1935 and E. talpoida (Say, 1817)) had neotenous males. Neotenous malesof E. rathbunae can be as small as 2.3 mm, whereas females can reach 33 to44 mm CL (Efford, 1967). Subramoniam (1981) and Subramoniam & Gunamalai(2003) supported the idea of widespread of neoteny in Emerita species. If malesof the present species are also neotenic, the collecting method used in the presentstudy (with the larger mesh size of the sieves) may have failed to obtain the smallmale individuals. The largest female specimen collected during the present studymeasured only 13.1 mm CL, and male individuals would probably be small enoughto go through the 2-mm-mesh sieve used. Efford (1967) reported that neotenousmales often attached to various body parts of mature female E. rathbunae. In orderto find male individuals of the present species, collection and examination of aneven larger number of mature females will be required.

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The dispersion of the present species in Taiwan may be tied with environmentalconditions, which consists of a brackish water influence with medium + coarsesand dominated bottom substrates. Surface seawater salinity at the collectingsites was between 21 and 26� (northeastern Taiwan) or 30� (southwesternTaiwan). Sand grain size compositions of both collecting sites were all with a highpercentage (90.7% and 76.6%) of medium + coarse sand.

ACKNOWLEDGEMENTS

I thank Drs. P. K. L. Ng, C. B. Boyko and two anonymous reviewers for theirvaluable comments on the manuscript. Dr. C. B. Boyko also provided the referenceon E. karachiensis. Mr. H.-T. Hong helped in the field work, and Y.-W. Tzengparticipated in preparation of photographs and line drawings of the new species.

APPENDIX

KEY TO THE SPECIES OF Emerita

Key to species of Emerita (modified from Schmitt, 1935, 1937; Niazi & Haque, 1974):

1. Dactylus of P1 broadly ovate, less than twice as long as greatest width . . . . . . . . . . . . . . . . . . 2– Dactylus of P1 narrow, basally ovate, distally subacute, more than twice as long as greatest

width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

2. Dactylus of P1 broadly ovate, distally subacute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3– Dactylus of P1 broadly ovate, distally rounded, obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Spines on dactylus of P1 occupy nearly entire distal third of lower margin. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. emeritus (Linnaeus, 1767)

– Spines on dactylus of P1 occupy half to less than two-thirds of distal part of lower margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. austroafricana Schmitt, 1937

4. Transverse ridges cover most of dorsum of carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5– Transverse ridges not covering most of dorsum of carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Transverse ridges more or less continuous, densely packed . . . . E. rathbunae Schmitt, 1935– Transverse ridges distinctly interrupted, not densely packed . . E. brasiliensis Schmitt, 1935

6. Center portion of carapace posterior to cervical groove smooth, shining . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. talpoida (Say, 1817)

– Center portion of carapace posterior to cervical groove not smooth or shining . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. analoga (Stimpson, 1857)

7. Median frontal lobe lower than lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8– Median frontal lobe not lower than lateral lobes . . . . . . . . . . . . . . . . . . E. taiwanensis sp. nov.

8. Width of second abdominal somite less than 2/3 of carapace width . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. benedicti Schmitt, 1935

– Width of second abdominal somite not less than 2/3 of carapace width . . . . . . . . . . . . . . . . . . 9

9. Lateral frontal lobes elongated, tapered, distally subacute . . . E. portoricensis Schmitt, 1935– Lateral frontal lobes not elongated, tapered, distally blunt . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

EMERITA TAIWANENSIS NOV. 257

10. Merus of third maxilliped approximately barrel-shaped, short; distal inner margin rounded,not produced; usually with 3 spines (excluding distal spine) on lower margin of P1 dactylus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. holthuisi Sankolli, 1965

– Merus of third maxilliped approximately cone-shaped, elongate; distal inner margin pointed,produced; usually with 5 or more spines (excluding distal spine) on lower margin of P1dactylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. karachiensis Niazi & Haque, 1974

REFERENCES

BARNARD, K. H., 1950. Descriptive catalogue of South African decapod Crustacea (crabs andshrimps). Ann. South African Mus., 38: 1-837.

BOYKO, C. B. & A. W. HARVEY, 1999. Crustacea Decapoda: Albuneidae and Hippidae ofthe tropical Indo-West Pacific region. In: A. CROSNIER (ed.), Résultats des CampagnesMUSORSTOM, 20. Mém. Mus. Natl. Hist. Nat., 180: 379-406.

BOYKO, C. B. & P. A. MCLAUGHLIN, 2010. Annotated checklist of anomuran decapod crustaceansof the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of theGalatheoidea) Part IV — Hippoidea. Raffles Bull. Zool., Suppl. No. 23: 139-151.

EFFORD, I. E., 1967. Neoteny in sand crabs of the genus Emerita (Anomura, Hippidae). Crus-taceana, 13: 81-93.

— —, 1970. Recruitment to sedimentary marine populations as exemplified by the sand crab,Emerita analoga (Decapoda, Hippidae). Crustaceana, 18: 293-308.

— —, 1976. Distribution of the sand crabs in the genus Emerita (Decapoda, Hippidae). Crustaceana,30: 169-183.

HAYE, P. A., Y. K. TAM & I. KORNFIELD, 2002. Molecular phylogenetics of mole crabs (Hippidae:Emerita). J. Crust. Biol., 22: 903-915.

HEEGAARD, P. E. & L. B. HOLTHUIS, 1960. Proposed use of the plenary powers to validatethe generic name Idotea J. C. Fabricius, 1798 (class Crustacea, order Isopoda) and mattersconnected therewith. Z.N. (S.) 412. Bull. Zool. Nomencl., 17: 178-184.

JOHNSON, M. W. & W. M. LEWIS, 1942. Pelagic larval stages of the sand crabs Emerita analoga(Stimpson), Blepharipoda occidentalis Randall, and Lepidopa myops Stimpson. Biol. Bull.,83: 67-87.

KNIGHT, M. D., 1967. The larval development of the sand crab Emerita rathbunae Schmitt(Decapoda, Hippidae). Pac. Sci., 21: 58-76.

NIAZI, M. S. & M. M. HAQUE, 1974. On a new species of mole-crab (Emerita karachiensis sp.nov.) with a key to common Indo-Pacific species. Rec. Zool. Sur. Pakistan, 5: 1-6.

OSAWA, M., C. B. BOYKO & T.-Y. CHAN, 2010. Part I. Hippoidea (Mole crabs). In: T.-Y.CHAN (ed.), Crustacean fauna of Taiwan: crab-like anomurans (Hippoidea, Lithodoidea andPorcellanidae): 1-41. (National Taiwan Ocean University, Keelung).

RATHBUN, M. J., 1900. The decapod crustaceans of West Africa. Proc. U. S. Natn. Mus., 22: 271-316.

REES, G. H., 1959. Larval development of the sand crab Emerita talpoida (Say) in the laboratory.Biol. Bull., 117: 356-370.

SANKOLLI, K. N., 1965. On a new species of Emerita (Decapoda, Anomura) from India, with anote on Emerita emeritus. Crustaceana, 8: 48-51.

SCHMITT, W. L., 1935. Crustacea Macrura and Anomura of Porto Rico and the Virgin Islands.Scientific Survey of Porto Rico and the Virgin Islands, New York Acad. Sci., 15: 125-227.

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First received 27 October 2013.Final version accepted 13 February 2015.