EVOLUTION & SYSTEMATIC BIOLOGY Animal Cells and Systems, 2014 Vol. 18, No. 5, 351–357, http://dx.doi.org/10.1080/19768354.2014.967293 Synalpheus jejuensis n. sp. (Crustacea: Decapoda: Alpheidae) from Korea based on morphological and molecular study a b* Hyeyoung Koo and Won Kim a b Department of Biological Science, College of Natural Science and Engineering, Sangji University, Wonju 220-702, Korea; School of Biological Sciences, Seoul National University, Seoul 151-747, Korea (Received 7 July 2014; received in revised form 1 September 2014; accepted 12 September 2014) We describe Synalpheus jejuensis n. sp. on the basis of 17 specimens collected around Jejudo Island, Korea. The new species is most similar to S. tumidomanus but can be distinguished from that species by shorter scaphocerite, upturned blunt tubercle rather than an acute spine on the superior margin of major chela, and teeth on the ventral margin of the pleura. The DNA barcoding using the mitochondrial cytochrome c oxidase subunit I (COI) gene fragment was employed in these two species. The intraspecific sequence divergence of the COI gene of S. jejuensis n. sp. ranges between 0% and 2.1%, while that of S. tumidomanus ranges between 0.2% and 0.4%. The sequence divergence between two species is 20.1%. DNA barcoding results provide further evidence that these two species are distinct. The discovery of S. jejuensis n. sp. is biogeographically important because the species diversity of Synalpheus is low in temperate region. Keywords: Alpheidae; Synalpheus jejuensis n. sp.; Korea; DNA barcoding Introduction further reports. S. neomeris can be easily distinguished from S. tumidomanus by having longer and stouter lower The decapod genus Synalpheus Bate, 1888 is a member of process of dactyli of third, fourth, and fifth pereopods and snapping shrimp of the family Alpheidae and widely by the presence of movable spines on the inferior margin of distributed in tropical and subtropical waters worldwide. merus of the third pereopod (Koo & Kim 2003). Many species of Synalpheus live in coral reefs and they are In recent inventories of the Korean decapod collec- often associated with sessile invertebrates such as sponges, tions, several specimens belonging to S. tumidomanus corals, bryozoans, and crinoids (Banner & Banner 1975; complex were found. These specimens could be morpho- Chace 1988). Although more than 160 species of the genus logically sorted as two groups. One group was identified have been described worldwide (Hermoso-Salazar & as S. tumidomanus with keys presented by the major Alvarez 2005; Rios & Duffy 2007; Hermoso-Salazar et al. monographs (Miya 1972; Banner & Banner 1975; Chace 2008; De Grave & Fransen 2011; Hultgren et al. 2014), 1988; Hayashi 1996). But the other group showed several clear morphological characters and intra-specific variations morphological characteristics consistently different from for many species have been poorly known. The specimens S. tumidomanus. Individually considered, each of these of Synalpheus are usually small and their appendages characteristics falls in the range of variation of S. are easily detached during collection. Therefore it is very tumidomanus reported from the previous literature difficult to identify the species of Synalpheus morphologi- (Banner & Banner 1975). However, if considered to- cally and the situation is more troublesome in case of young gether, these characteristics appear to differentiate this specimens. In Far East, 30 species from Philippine (Chace group as a new species from S. tumidomanus. The best 1988), 17 from Japan (Hayashi 1996), and only 2 from way to provide clear evidence as a new species is to find Korea have been reported. In Korea, Synalpheus tumido- out whether these different characteristics are intra-spe- manus (Paulson, 1875) was first reported from Hongdo cific or inter-specific variation. Because the authors cannot Island, southern part of Geojedo Island, Kyongsangnam-do examine all other species of Synalpheus in the world and in 1979 without description (Kim et al. 1979). Later in specimens of other worldwide geographic regions of S. 1994, this species was reported from Ulreungdo Island, Kyongsangbuk-do with description and illustrations (Kim tumidomanus, the DNA barcoding was employed as an alternative way to verify specific status of the current new & Moon 1994). Since then, there have been two more species. reports from Bogildo Island, Jeollanam-do (Kim et al. 1996) and Geojedo Island, Kyongsangnam-do (Kim & Kim DNA barcoding is a molecular method to identify a 1998). The second species, Synalpheus neomeris (De Man, species by DNA sequences of short segment of the marker 1897), was reported in 2003 from Jejudo Island, Jeju-do gene. In the species identification with DNA barcodes, it is with description and illustrations (Koo & Kim 2003) and no very important to figure out the range of the intra-specific *Corresponding author. Email: email@example.com © 2014 Korean Society for Integrative Biology 352 H. Koo and W. Kim and inter-specific variation of each species tested. If intra- Bank of Korea (MADBK), Seoul National University, specific variation is much less than inter-specific variation, Korea. reliable species identification can be made and this molecular identification system can be used in the finding Molecular data of cryptic species or to verify taxonomic specific status. In animals, the mitochondrial cytochrome c oxidase I gene Nine specimens of S. jejuensis n. sp. and 3 specimens of (COI) has been widely used in the DNA barcoding (Hebert S. tumidomanus were sampled (Table 1). The genomic DNA from all samples was extracted using the Qiagen et al. 2003; Hebert & Gregory 2005). DNeasy Blood and Tissue kit. The target DNA segment of In the present study, the authors sequenced the mito- the COI gene was amplified with the polymerase chain chondrial cytochrome c oxidase subunit I (COI) genes of reaction (PCR) method. The primers used for the PCR the specimens of S. tumidomanus complex. Then we were LCO1490 (5′-GGT CAA CAA ATC ATA AAG compared the sequences and examined the intra- and ATA TTG G-3′) and HCO2198 (5′-TAA ACT TCA GGG inter-specific variation. As a result, the specimens of two TGA CCA AAA AAT CA-3′) for COI (Folmer et al. groups of S. tumidomanus complex collected from Korea 1994). All reactions carried out for the amplification of the showed enough inter-specific variation to be considered as COI-5′ region were run by the following thermal cycling two different species. Herein we describe Synalpheus program: 5 min at 94°C followed by 35 cycles of 0.5 min jejuensis n. sp. with illustrations. at 94°C, 1 min at 48°C, 1.5 min at 72°C, and a final extension of 72°C for 10 min. The 25 µL PCR reaction mix included 14.7 µL ultrapure water, 5 µL of 5× PCR Materials and methods buffer, 2 µL of each primer (10 µM), 1 µL of dNTP Synalpheus specimens were collected in the vicinity of (10 mM), 0.3 µL of Taq polymerase (5 U), and 2 µL of Jejudo Island, Korea by SCUBA diving. Collecting the DNA template. All the sequences were aligned using locations and details are shown in Figure 1 and Table 1, the MUSCLE software (Edgar 2004). Sequence diver- respectively. The abbreviation “cl” refers to carapace gences among individuals were quantified with the Kimura length, which is measured from the tip of the rostrum 2-parameter (K2P) distance model (Kimura 1980). A to the posterior dorsal margin. The holotype and para- neighbor-joining (NJ) tree of K2P distances was created types are deposited in Marine Arthropod Depository by MEGA version 4.0.2 (Tamura et al. 2007). Figure 1. A map showing the areas from which specimens of the new species were collected. Animal Cells and Systems 353 Table 1. Information on the specimens of S. jejuensis n. sp. and S. tumidomanus. Carapace GenBank length accession Species Sex (cl, mm) Collection locality Scuba Collection date Collector No. S. jejuensis Holotype ♂ 6.0 Munseom Island, Jeju-do, Korea 20 m 11 March 2009 S.H. Kim n. sp. (33.1338 N, 126.3404 E) Paratype ovig. ♀ 5.9 Munseom Island, Jeju-do, Korea 15 m 30 June 1993 W. Kim Paratype ovig. ♀ 5.7 Munseom Island, Jeju-do, Korea 10 m 1 July 1993 W. Kim Paratype ♂ 4.6 Seongsanpo, Jeju-do, Korea 20 m 5 November 2000 S.H. Kim Paratype ovig. ♀ 7.7 Chaguido Island, Jeju-do 20 m 6 November 2000 S.H. Kim Paratype ♂ 4.6 Beomseom Island, Jeju-do, Korea 20 m 21 February 2001 S.H. Kim Paratype ♂ 6.1 Munseom Island, Jeju-do, Korea 20 m 28 October 2005 S.H. Kim JN368144 Paratype ♂ 4.3 Munseom Island, Jeju-do, Korea 20 m 11 March 2009 S.H. Kim JN368137 Paratype ♂ 5.9 Heukgeomdo Island, Jeju-do, Korea 25 m 31 March 2009 S.K. Lee JN368136, JN368142 Paratype ♂ 4.8 Jikgudo Island, Jeju-do, Korea 20 m 31 March 2009 S.K. Lee Paratype ♂ 6.4 Chujado Island, Jeju-do, Korea 25 m 8 September 2009 K.J. Lee and T.S. Park JN368139 (33.5614 N, 126.1869 E) Paratype ♂ 5.6 Chujado Island, Jeju-do, Korea 25 m 8 September 2009 K.J. Lee and T.S. Park JN368140 (33.5614 N, 126.1869 E) Paratype ♂ 5.1 Chujado Island, Jeju-do, Korea 25 m 8 September 2009 K.J. Lee and T.S. Park JN368141 (33.5614 N, 126.1869 E) Paratype juv. 3.2 Chujado Island, Jeju-do, Korea 25 m 8 September 2009 K.J. Lee and T.S. Park JN368143 (33.5614 N, 126.1869 E) Paratype ovig. ♀ 7.0 Chujado Island, Jeju-do, Korea 25 m 8 September 2009 K.J. Lee and T.S. Park JN368138 (33.5614 N, 126.1869 E) Paratype ovig. ♀ 7.1 Chujado Island, Jeju-do, Korea 25 m 8 September 2009 K.J. Lee and T.S. Park (33.5614 N, 126.1869 E) Paratype ovig. ♀ 8.1 Geomundo Island, Jeollanam-do, Korea 20 m 15 October 2001 S.H. Kim JN368135 S. tumidomanus – 5.0 Beomseom Island, Jeju-do, Korea 20 m 21 February 2001 S.H. Kim JN368132 – 6.9 Munseom Island, Jeju-do, Korea 20 m 11 March 2009 S.H. Kim JN368133 – 5.9 Heukgeomdo Island, Jeju-do, Korea 20 m 31 March 2009 S.K. Lee JN368134 juv., juvenile; ovig., ovigerous; –, unable to determine the sex. 354 H. Koo and W. Kim Results Paratypes: All specimens other than holotype of S. jejuensis n. sp. are designated as the paratypes. See Family Alpheidae Rafinesque, 1815 Table 1. Genus Synalpheus Bate, 1888 Synalpheus jejuensis n. sp.(Figure 2) Description Rostrum (Figure 2A, 2B) reaching to middle of first Material examined antennular segment, overreaching slightly lateral ocular teeth, arising from anterior margin of carapace; tip slightly Holotype: male, cl 6.0 mm, Munseom Island, Jeju-do, upturned with several setae. Ocular hoods separated from Korea (33°13′38″N, 126°34′04″E), coll. Sa Heung Kim, 11 March, 2009 [MADBKh3]. rostrum by slightly depressed groove, with teeth slightly Figure 2. S. jejuensis, new species, holotype male, cl 6.0 mm: A, anterior region, dorsal view; B, anterior region, lateral view; C, large (right) ﬁrst pereopod, inner face; D, same, upper face; E, same, outer face; F, small (left) ﬁrst pereopod, inner face; G, same, outer face; H, right second pereopod; I, right third pereopod; J, same, dactylus, magniﬁed; K, abdomen; L, telson and uropods (scale bar, 1 mm). Animal Cells and Systems 355 broader than rostrum and directed inward. Pterygostomial with nine spines throughout length in addition to distal margin produced as a narrowly rounded lobe below pair. Carpus slightly more than half as long as propodus basicerite. with 1 movable spine at distal end of inferior margin. Merus approximately 3.4 times as long as broad, with no Stylocerite narrow, elongated, slightly overreaching movable spine. distal end of first antennular segment, being nearly as long Carpus of fourth pereopod with one movable spine at as sum of length of second and third segments; second distal end of inferior margin. Merus with no movable spine. segment slightly longer than third segment. Scaphocerite with lateral spine falling far short of Carpus and merus of fifth pereopod with no movable spine. distal end of third antennular segment; inner blade well Pleura (Figure 2K) of first to fifth abdominal somites developed, reaching to distal end of second antennular of male with sharp tooth on ventral margin posteriorly; segment; cleft between inner blade and lateral spine deep, pleura of third and fourth bearing additional minute tooth arising from proximal 1/3 of scaphocerite. Basicerite on ventral margin anteriorly. (Figure 2A, 2B) with spine dorsally; lateral spine almost Telson (Figure 2L) with dorsal surface armed typically reaching proximal 1/2 of scaphocerite. Carpocerite over- with two pairs of spines; anterior pair situated at proximal reaching distal end of antennular peduncle by length of 1/3 of telson; posterior pair located at distal 2/5 of telson; third antennular segment. posterior margin with two pairs of strong outer spines; Third maxilliped (Figure 2B) with ultimate segment inner pair more than 2.0 times as long as outer pair. bearing several movable spines at tip, overreaching distal Uropodal exopod with lateral margin slightly convex; end of carpocerite by distal 1/3 of ultimate segment. diaeresis with two immovable spines laterally; one mov- Major chela of first pereopod (Figure 2C, 2D, 2E) able spine between these two immovable spines. approximately 2.5 times as long as broad; palm, approxi- mately 2.0 times as long as fingers, terminating in blunt tubercle on superior distal margin; tip directing upward. Etymology Merus approximately 2.2 times as long as broad. The specific name is from the Jeju-do province, Korea, the Minor chela of first pereopod (Figure 2F, 2G) approxi- type locality of the new species. mately 3.1 times as long as broad; palm 1.8 times as long as fingers. Merus nearly 3.5 times as long as broad. Chela of second pereopod (Figure 2H) with fingers 1.6 Type locality times as long as palm. Carpus approximately 2.0 times as Munseom Island, Seogipo, Jeju-do, Korea. long as chela and composed of five segments; first segment slightly shorter than sum of four distal segments; DNA barcoding analysis second, third, and fourth subequal; fifth almost same length as sum of third and fourth segments. Merus shorter The nucleotide sequences of the COI gene fragment (Gen- than carpus. Ischium shorter than merus. Bank accession numbers, JN368132–JN368144, Table 1) Third, fourth, and fifth pereopods (Figure 2I, 2J) with from both species were compared, and a NJ tree was dactyli biunguiculate; lower process with tip turned down- constructed (Figure 3). The intraspecific sequence divergence ward and base nearly as broad as that of upper process. of the COI gene of S. jejuensis n. sp. ranges between 0% and Propodus of third pereopod (Figure 2I) slightly more 2.1%, while that of S. tumidomanus ranges between 0.2% than 6.0 times as long as dactylus; inferior margin armed and 0.4%. The sequence divergence between two species is Figure 3. An NJ tree of S. jejuensis n. sp. and S. tumidomanus based on COI gene sequences. 356 H. Koo and W. Kim 20.1%, and this value provides further evidence that two in S. tumidomanus, the rostrum is relatively long, exceeds species are distinct. the lateral ocular teeth and arises from anterior slope slightly behind anterior margin of carapace (Kim & Moon 1994, Figure 1A, 1B); (2) the scaphocerite does not Discussion reach the distal margin of the third antennular segment Synalpheus Bate, 1888 is one of the most diverse genera (Figure 2A), while in S. tumidomanus, it overreaches in all of Alpheidae. Since Coutière (1908, 1909)’s division into specimens (Kim & Moon 1994, Figure 1A, 1B); (3) the six species groups of Synalpheus, there have been several shape and direction of the tooth on the superior distal attempts to change and modify the species groups. The margin of the palm of the major chela shows a upturned current S. jejuensis n. sp. and S. tumidomanus show the blunt tubercle (Figure 2A) rather than an acute spine characteristics of the Synalpheus paulsoni group recog- directing forward as seen in S. tumidomanus (Kim & nized by Coutière (1908, 1909). However, the S. paulsoni Moon 1994, Figure 1C, 1D); (4) the lower process of the group was not retained by Banner and Banner (1975) and biunguis on the dactyli of the third to the fifth pereopods the phylogenetic analysis of this group from the American directs more or less downward (Figure 2I, 2J), while it is Pacific showed that it is paraphyletic (Hermoso-Salazar less deflexed in S. tumidomanus (Kim & Moon 1994, et al. 2008). Furthermore, the recent worldwide molecular Figure 1H, 1I); and (5) the teeth are present on the ventral phylogenetic analysis of the genus Synalpheus also margins of the first to fifth pleura (Figure 2K), but on the showed that the S. paulsoni group is not monophyletic first pleuron only in S. tumidomanus (Kim & Moon 1994, (Hultgren et al. 2014). Thus morphological character Figure 1J). In terms of the presence of a tooth on the variations and taxonomic status of species previously pleuron, S. jejuensis n. sp. is similar to the eastern Pacific belonged to the S. paulsoni group should be reexamined. species S. stylopleuron (Salazar & Hendrickx 2006), but S. jejuensis n. sp. is most similar to S. tumidomanus,in differs from S. stylopleuron in that S. jejuensis n. sp. has a terms of having a dorsal spine on the basicerite and a blunt tubercle rather than a spine on the superior distal distally located tubercle or spine on the superior margin margin of the palm of the major chela, and a far shorter on the palm of the major chela. scaphocerite. It has been well known that S. tumidomanus shows We collected all COI sequences of the genus Synal- extensive variations and these variations were well dis- pheus registered in NCBI and a NJ tree using MEGA cussed in the previous literatures (Miya 1972; Banner & version 4.0.2 was constructed from 364 sequences of 112 Banner 1975). Especially Banner and Banner (1975) listed species including current sequence data (tree is not described the incredible morphological diversity of S. shown here). The sequences of S. tumidomanus and S. tumidomanus. Among the characteristics shown in their jejuensis were not mingled with the sequences of other species. The 3 sequences of S. tumidomanus and 10 Table 2, the three characteristics are worthy of noting here: sequences of S. jejuensis comprise each of independent (1) ‘length, orbital teeth to rostrum,’ (2) ‘length, lateral spine of scaphocerite to antennular articles,’ and (3) ‘nature clades. Thus the current two species can be distinguished of protrusion above dactylus, large chela.’ In the first by the other species of Synalpheus of which COI characteristic, the Table 2 shows ‘0.7 length, (from plate)’ sequences are available. in Paulson’s description and figures (Paulson 1875), ‘from The intraspecific sequence divergence of the COI gene 0.5 to 0.8 length’ in 9 specimens from Torres Staits, BAU of S. jejuensis n. sp. ranges between 0.0% and 2.1%, 27, ‘from 0.5 to 0.9 length’ in 27 specimens from around (1.0% average, n = 10) while that of S. tumidomanus Australia. In the second characteristic, it shows ‘end of ranges between 0.2% and 0.4% (0.2% average, n = 3). third’ in Paulson’s description and figures, ‘slightly shorter The sequence divergence between two species is 20.1%. to slightly longer than third’ in 9 specimens from Torres The ratio of interspecific to intraspecific variation (20.1– Staits, BAU 27, and ‘slightly shorter to slightly longer than 100.5×) is much higher than a standard sequence thresh- third’ in 27 specimens from around Australia. In the third old (10×) proposed by Hebert et al. (2004) as a species characteristic, it shows ‘strong, rounded’ in Paulson’s boundary. The genetic distance threshold for cryptic description and figures, ‘from no protrusion to acute tooth’ species of the genus Synalpheus by Hultgren et al. in 9 specimens from Torres Staits, BAU 27, and ‘from no (2014) is 10.2%. Therefore the current 20.1% provides protrusion to acute tooth’ in 27 specimens from around further evidence that two species are distinct. Australia. In the case of S. tumidomanus from Korea, this The species of the genus Synalpheus are found in first characteristic falls in the range of variation, the second tropical and warm temperate waters and usually associated one is consistently ‘longer than third,’ and the third one is with coralline algae, sponges, and coral heads (Chace ‘acute spine.’ However, S. jejuensis n. sp. differs from S. 1988). The present S. jejuensis n. sp. and S. tumidomanus tumidomanus with respect to the following characteristics: in Korea were associated with sponges. The species (1) the rostrum exceeds slightly the lateral ocular teeth and diversity of the genus Synalpheus is very low in temperate arises from anterior margin of carapace (Figure 2A), while waters. 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Animal Cells and Systems
– Taylor & Francis
Published: Sep 3, 2014
Keywords: Alpheidae; Synalpheus jejuensis n. sp.; Korea; DNA barcoding