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ZooKeys 407: 9-28 (20 1 4) 
doi: I0.3897/zookeys.407.7457 
www.zookeys.org 



RESEARCH ARTICLE 



^ZooKeys 

Launched to accelerate biodiversity research 



A new species of shrimp of the genus Anachlorocurtis 
Hayashi, 1 975 from the Red Sea, with range 
extension of A. commensalis Hayashi, 1 975 
(Crustacea, Decapoda, Pandalidae) 

Ivona Horka 1 ' 2 *, Sammy De Grave 3 '*, Zdenek Duris' 5 

I University of Ostrava, Faculty of Science, Department of Biology and Ecology, Chittussiho 10, CZ-71000 
Ostrava, Czech Republic 2 Charles University in Prague, Faculty of Science, Department of Ecology, Vinicnd 
7, CZ- 12844 Prague, Czech Republic 3 Oxford University Museum of Natural History, Parks Road, Oxford, 
0X1 3PW, United Kingdom 

t http://zoobank.org/CCA47494-EA9C-46D9-B579-90772B584F35 
% http:llzoobank.org/2EBlCBFA-2B2B-4740-A177-B9D5290ECE01 
§ http:llzoobank. org/0CF5D3F9-9663-4B76-BF91-713D9BE50BC3 

Corresponding author: Sammy De Grave (sammy.degrave@oum.ox.ac.uk) 



Academic editor: N, Bruce | Received 8 March 2014 | Accepted 19 April 2014 | Published 8 May 2014 

http-.llzoobank. org/91 7BDB2D-AF4F-44BB-9C34-0C09C68DFDE2 

Citation: Horka I, De Grave S, Duris Z (2014) A new species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from 
the Red Sea, with range extension of A commensalis Hayashi, 1975 (Crustacea, Decapoda, Pandalidae). ZooKeys 407: 
9-28. doi: 10.3897/zookeys.407.7457 



Abstract 

A new species of pandalid shrimp Anachlorocurtis occidentalis sp. n., associated with antipatharian corals, 
is described and illustrated from the north-eastern Red Sea. This new species is closely related to Anachlo- 
rocurtis commensalis Hayashi, 1975, the only other species in the genus, and can be distinguished by the 
more slender body and appendages; the carapace with 3 large, and one small, subtriangular lobes in the 
middorsal line; a flattened dorsal outline of the third abdominal segment; the sixth abdominal segment 
twice as long as fifth one; propodi of the ambulatory pereiopods bearing only a single posterior spinule; 
and harbouring 3—5 pairs of dorsolateral spines on the telson. A revised generic diagnosis is provided 
here to accommodate the present new species. The genetic divergence of mitochondrial gene cytochrome 
c oxidase subunit I (COI) between Anachlorocurtis occidentalis sp. n., and A. commensalis is 15.2—15-4%. 
Molecular analysis also confirmed a sister position of the genus Anachlorocurtis to Miropandalus. The 
present records of A. commensalis from Taiwan constitute an extension of the known range of the species. 



Copyright Ivona Horka et at. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), 
which permits unrestricted use. distribution, and reproduction in any medium, provided the original author and source are credited. 



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Ivona Horkd et al. I ZooKeys 407: 9-28 (2014) 



Keywords 

Caridea, Pandalidae, Anacblorocurtis occidentalism new species, Antipathes, Antipatharia, Red Sea 



Introduction 

Shrimps of the family Pandalidae include 23 genera and 190 species (De Grave and 
Fransen 201 1, Komai and Tsuchida 2014), with their greatest diversity occurring in 
boreal to temperate Atlantic and Pacific waters, especially in low-latitudinal areas of the 
Indo-West Pacific, and with relatively few species at similar latitudes in the southern 
hemisphere. In tropical regions, numerous pandalid species are known, but largely 
restricted to deeper, colder water (Bauer 2004). Overall, this is a diverse group of medi- 
um to large -sized, mainly epibenthic shrimps with a typical natant body form possess- 
ing well-developed rostra and long slender or robust legs (Bruce 1983, Bauer 2004). 
With almost no exception, species of Pandalidae are free-living and the association 
with other invertebrates is restricted to three species: Chlorotocella gracilis Balss, 1914, 
Anacblorocurtis commensalis Hayashi, 1975, and Miropandalus hardingi Bruce, 1983. 
Chlorotocella gracilis has been reported in association with a jellyfish and gorgonarian 
corals (Hayashi and Miyake 1968, Li and Davie 2006, Hayashi 2007), although those 
associations are perhaps best regarded as accidental, as numerous reports exist on free- 
living specimens (Bruce 1972). The other two species are obligate symbionts of an- 
tipatharian black corals (Anthozoa, Antipatharia) Antipathes spp., Myriopathes japonica 
(Brook, 1889) and Cirrhipathes spp. (Hayashi 1975, Bruce 1983, 1991). 

The genus Anacblorocurtis Hayashi, 1 975 has, up to now, only included a single 
species, A. commensalis known from Japan (Hayashi 1975, 2007) and recorded herein 
from Taiwan. The new species A. occidentalis sp. n. was found in the Red Sea, thus 
considerably extending the genus' geographical range, equally associated with antipa- 
tharian corals. The external morphology of both Anacblorocurtis and Miropandalus are 
unique, with their body being well adapted to life with their hosts. They are small, 
slender, and compressed animals with a cryptic colour pattern (Kawamoto and Okuno 
2003, Minemizu 1996, 2013), short {Anacblorocurtis), or fully reduced {Miropandalus) 
rostra, specialised mouth-parts, and comparatively large and few eggs (Hayashi 1975, 
Bruce 1983, 1991). 

The following abbreviations are used: COI, mitochondrial gene cytochrome c oxi- 
dase subunit I; fen, field collection numbers for Aqaba (Aq), Eilat (ISR) and Taiwan 
(Tw, TAI); SEM, scanning electron microscope; PoCL, post orbital carapace length; 
spm(s), specimen(s); RL, rostrum length (measured from the posterior orbital margin 
to the distal edge of the most anteriorly advanced tooth of the rostrum); TL, total 
length of the body (measured from tip of the rostrum to the distal end of the telson, 
posterior telson spines not included); MSS, Marine Science Station, Aqaba, Jordan; 
NTOU, National Taiwan Ocean University; OUMNH.ZC, Zoological Collection, 
Oxford University Museum of Natural History, Oxford; RMNH, Naturalis Biodiver- 
sity Center, Leiden; UO, University of Ostrava. 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



11 



Material and methods 

Specimens of Anachlorocurtis occidentalis sp. n. were collected in the Red Sea (Aqaba, 
Jordan, in 2008 and 2009; Eilat, Israel, in 2011), and A. commensalis in the South 
China Sea (southern Taiwan, in 2009, 2011, 2012), respectively, from antipatharian 
corals using hand nets and SCUBA equipment. Samples were preserved in 80% etha- 
nol for morphological studies and in 96-99% ethanol for molecular analysis. 

The terminal segments of the first and second pereiopods of both species examined 
were photographed under SEM. One specimen of each species {A. occidentalis sp. n. 
UO Aq09-30A and A. commensalis UO Twl2-79) was dehydrated by a series of ac- 
etone/ethanol solutions of concentrations 30, 50, 70, 80, 90, 95 and 100% acetone in 
incremental 30 minute steps. Acetone was removed by the C0 2 critical point method 
(POLARON E3000 Critical Point Drying Apparatus) with dried specimens gold- 
coated (Automatic Sputter Coater: JEOL JFC-1300) and examined under a scanning 
electron microscope (SEM -JEOL JSM-6610LV). 

Total genomic DNA was extracted from abdominal muscle tissue or eggs using 
the DNeasy Blood & Tissue isolation Kit (QIAGEN) according to the manufacturer's 
instructions. For amplifying the segment of mitochondrial protein-coding gene COI, 
with a polymerase chain reaction, the universal pair of primers LCO1490/HCO2198 
was used (Folmer et al. 1994). The PCR was conducted in 25-30 ul reaction volume 
containing: 2-3 ul DNA template, 0.3 uM each primer, 0.15 mM dNTP, 0.7 units 
of Taq polymerase, distilled water, 10 x PCR buffer and 2.5 mM of MgCL,. PCR 
cycling profile was as follows: 2.5 min at 94 °C for initial denaturation, followed by 
40 cycles of 30 s at 90 °C, 1 min at 48 °C, 1 min at 72 °C and the final extension step 
at 72 °C for 1 0 min. PCR products were purified using GenElute PCR clean-up kit 
(Sigma). Sequencing reactions were carried out using the ABI3730XL DNA Sequencer 
at Macrogen, Inc. Sequences obtained were deposited in GenBank (Table 1), from 



Table I . Species used in the molecular analysis. Sampling location is given, as is GenBank accession 
numbers (COI), and voucher identification numbers of specimens examined, (ii, v) — see Material; * - 
Sequences obtained from GenBank. 



Taxa 


Sampling location 


GenBank 
accession # 


Voucher ID 


Pandalidae 








Anachlorocurtis occidentalis sp. n. (ii) 


Jordan, Aqaba 


KJ690257 


RMNH.CRUS.D. 56174 


Anachlorocurtis occidentalis sp. n. (v) 


Jordan, Aqaba 


KJ690256 


RMNH.CRUS.D. 56177 


Anachlorocurtis commensalis Hayashi, 1 975 


Taiwan, Nanvan 


KJ690258 


RMNH.CRUS.D. 561 82 


Miropandalus hardingi Bruce, 1983 


Taiwan, Nanvan 


KJ690259 


UOTwll-20A 


Heterocarpus ensifer A. Milne-Edwards, 1881 


Guadeloupe 


*AY612858 


NTOU 


Heterocarpus gibbosus Spence Bate, 1888 


Philippines, Panglao 


*GQ302742 


NTOUM00797 


Pandalus borealis Krayer, 1838 


Canada, Quebec 


*FJ581839 


PB01CN0406 


Pandalus montagui heach, 1814 


Canada, New Brunswick 


*FJ581840 


GSL31-52 


Stenopodidae (outgroup) 








Stenopus hispidus (Olivier, 1811) 


Vietnam, Nhatrang Bay 


KJ690260 


UOV10-17 



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Ivona Horkd etal. I ZooKeys 407: 9-28 (2014) 



which additional sequences were used. Sequences were aligned using MUSCLE (Ed- 
gar 2004). The divergence of the analysed mitochondrial gene between both species 
was detected using the Kimura 2-parameter model. A further seven pandalid species 
(including Miropandalus, a morphologically closely related genus) and one stenopod 
shrimp (outgroup) were included in the phylogenetic analysis. The best-fit nucleotide 
substitution model under Bayesian Information Criterion (GTR+G+I, General Time 
Reversible) was selected. Phylogenetic reconstruction of COI data set was performed 
in a maximum likelihood (ML), bootstrap was done with 1,000 replicates. All analyses 
were conducted using MEGA v5. 2.1 (Tamura et al. 201 1). 



Systematics 

Superfamily Pandaloidea Haworth, 1825 
Family Pandalidae Haworth, 1825 

Genus Anachlorocurtis Hayashi, 1975 

http://species-id.net/wiki/Anachlorocurtis 

Generic diagnosis (modified from Hayashi 1975). Small -sized shrimps. Rostrum 
short, deep, not reaching end of eyes; anterior and dorsal margins with 2-6 small teeth 
in adult females; rostrum styliform, anteriad in adult males. Carapace with 2 large 
compressed dorsal lobes, with small epigastric tooth and one small lobe near poste- 
rior margin; anterior (i.e., postrostral) lobe just behind orbit, similar and subequal to 
rostrum, with 2-4 small teeth anteriorly and small epigastric lobe posteriorly; large 
posterior, anteriorly hooked lobe on distal third of dorsal midline; antennal tooth mar- 
ginal, acute; supraorbital, hepatic and pterygostomial teeth absent. Abdomen smooth, 
without spines or spiniform processes, sixth segment elongate, with posterior lobes 
obtuse. Telson with 3-5 pairs of small dorsal spines on lateral margins; posterior mar- 
gin rounded with 5 pairs of spines. Eyes well developed, cornea with apical tubercle, 
accessory pigment spot lacking, stalk slightly longer than corneal length. Antennular 
peduncle elongate; basal segment with deep medioventral keel; stylocerite short, an- 
teriorly truncate with produced distolateral tooth and medial angle pointed; distal 2 
peduncular segments short; upper and lower flagella uniramous, short. Scaphocer- 
ite well developed, distolateral spine not exceeding lamella. Mandible without palp. 
Maxillula with palp bearing 2 long setae, upper lacinia broader that lower lacinia. 
Maxilla with bi-setose palp, with simple, broad, distal endite and bilobed proximal 
endite, scaphognathite well developed, posterior lobe not particulary elongate. First 
maxilliped with large palp; exopod without flagellum but with elongate setose caridean 
lobe; endites feebly separated; podobranch lacking. Third maxilliped long, slender, 
with elongate lateral lobe; merus, basis and ischium (i.e., antepenultimate segment) 
fused; exopod, epipod and arthrobranch absent. Pleurobranchs present above all perei- 
opods. First pereiopods slender, not chelate, with dactylus reduced; ischium without 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



13 



lamellar expansion. Second pereiopods slender, chelate, equal with small subspatulate 
fingers bearing irregularly denticulate cutting edges; carpus three-segmented. Ambula- 
tory pereiopods with dactylus slender, simple; propodus with 1-2 spinules on ventral 
margin, distoventral spinules lacking or present, small; pereiopod 3-5 meral spinula- 
tion 2-2-(0-2), respectively. Endopod of first pleopod reduced to small setose lobe 
far overreached by functional appendix interna in males, and more reduced, without 
cincinnuli, in females. Uropod with elongate branches. 

Generic distribution. Kii Peninsula to Ruykyus, Japan, at depths of 8-15 m 
(Hayashi 1975, Kawamoto and Okuno 2003); southern Taiwan, depths 10-27 m 
(this report), and northeastern Red Sea, depths 4-55 m (this report). 

Anachlorocurtis occidentalis sp. n. 

http://zoobank.org/AA95EADC-5095-4590-A5D5-16B902C88249 

http://species-id.net/wiki/Anachlorocurtis_occidentalis 

Figs 1-4, 5A-F, 6A-C, 7A-D, 8 

Material examined. Type series, (i) 1 female juvenile (paratype), PoCL 1.6 mm 
(OUMNH.ZC.2014. 01. 016), Marine Science Station area, Aqaba, Jordan, reef wall, 
40 m depth, from Antipathes sp., leg. Z Duris, 17.06.2008, (fcn-Aq08-25B); (ii) 2 
ovigerous females (paratypes), PoCL 2.1 mm (RMNH.CRUS.D.56174, GenBank 
KJ690257), and 2.6 mm (dissected, UO Aq08-34B), Marine Science Station area, 
Aqaba, Jordan, from antipatharian coral in crevice among concrete blocks of pier, 5 
m depth, leg. Z Duris, 16.06.2008 (fen Aq08-34B); (iii) 1 ovig. female (holotype), 
PoCL 3.3 mm (RMNH.CRUS.D.56175), Marine Science Station pier, Aqaba, Jor- 
dan, from antipatharian corals in crevice among concrete blocks of pier, 4 m depth, 
leg. Z Duris, 20.06.2008 (fen Aq08-39); (iv) 1 ovigerous female (paratype), PoCL 
2.4 mm (OUMNH.ZC.2014.01. 017), Marine Science Station area, Aqaba, Jordan, 
reef wall, imm Antipathes sp., 45 m depth, leg. Z Duris & I Horka, 11.06.2009 (fen 
Aq09-8); (v) 1 male (allotype), PoCL 1.9 mm (RMNH.CRUS.D.56176), 1 oviger- 
ous female (paratype), PoCL 2.2 (RMNH.CRUS.D.56177, Genbank KJ690256), 1 
ovigerous female, PoCL 2.5 mm, 5 males PoCL 1.6—1.9 mm, 1 spm PoCL 1.6 mm 
(paratypes) (RMNH.CRUS.D. 56179), Marine Science Station area, Aqaba, Jordan, 
rock on sandy bottom ca 15 m out of reef wall, horn Antipathes sp., 47 m depth, leg. 
Z Duris & I Horka, 21.06.2009 (fen Aq09-28B); (vi) 1 ovigerous female (paratype), 
PoCL 3.1 mm (dissected, dried for SEM; UO Aq09-30A), Marine Science Station area, 
Aqaba, Jordan, reef wall, from Antipathes sp., 55 m depth, leg. Z Duris & I Horka, 
22.06.2009 (fen Aq09-30A); (vii) 1 ovigerous female PoCl 2.5 mm, 2 females 1.9 
mm, 1 male PoCL 1.5 mm (paratypes) (UO Aq09-65), Marine Science Station area, 
Aqaba, Jordan, sandy slope, antipatharian coral on rock, 50-55 m depth, leg. Z Duris 
& I Horka, 01.06.2009 (fen Aq09-65); (viii) 1 ovigerous female (paratype), PoCL 2.3 
mm (RMNH.CRUS.D. 56 180), Marine Science Station area, Aqaba, Jordan, from 
Antipathes sp., 55 m depth, leg. Z Duris & I Horka, 05.06.2009 (fen Aq09-93). 



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Ivona Horkd et al. I ZooKeys 407: 9-28 (2014) 




Figure I. Anachlorocurtis occidentalis sp. n., total aspect. A holotype, ovigerous female (RMNH. 
CRUS.D.56175) B allotype, male ( RMNH. CRUS.D. 56176). Scale bar equals 1 mm. 



Non-type material, (ix) 1 female, PoCL 3.3 mm (OUMNH.ZC.201 1.05.078), 
Eilat, Israel, 29°30'07"N, 34°55'05"E, 40 m depth, from Antipathes sp., leg. S De 
Grave & ML Johnson, 04.04.2011 (fen ISR-092); (x) 3 ovig. females (PoCL 2.6, 
3.0, 3.2 mm) (OUMNH.ZC.201 1.05.079), Eilat, Israel, 29'51"N, 34°55'39"E, 22 
m depth, horn Antipathes sp., leg. S De Grave & ML Johnson, 29°, 05.04.2011 (fen 
ISR-112). 

Diagnosis. Carapace dorsally trilobate in adults. Rostrum short, not reaching end 
of eyes; deep and anteriorly serrated with up to 6 small secondary teeth of which upper 
second or third tooth extending furthest forward; posterior margin smooth and con- 
vex; lower margin unarmed, slightly convex. Posterior dorsal lobe on carapace strong- 
ly hooked in adults. Posterior propodal spinule of ambulatory pereiopods small and 
single. Third abdominal segment distinctly angulated in lateral aspect, with dorsally 
straight outline. Sixth abdominal segment 2.5 times longer that deep. Mesial margin 
of endopod of male first pleopod with 9 mesial setae. Appendix masculina of second 
male pleopod longer than appendix interna. Telson with 3-5 pairs of dorsal spines. 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



15 



Description of female holotype. Carapace (Fig. 1A) smooth, about twice as long 
as height, with 2 large triangular teeth on dorsal midline. Anterior tooth placed just 
behind posterior level of orbit, similar to but slightly smaller than rostrum, anteriorly 
serrated with 4 small teeth; small obtuse epigastric tooth placed posteriorly on base of 
anterior tooth. Large posterior tooth placed on posterior third of carapace, higher than 
anterior tooth and strongly hooked with apex pointed forwards, and with anterior and 
posterior margins convex, smooth, without serration; small acute anteriad directed 
tooth situated posterodorsally near margin of carapace. Dorsal margin of orbit con- 
tinuous with short dorsal midrib extending to middle of rostrum near most anteroven- 
trally tooth of rostrum; lower orbital angle broadly rounded, not produced. Antennal 
tooth well developed, marginal, acute; supraorbital, hepatic and pterygostomial teeth 
absent; pterygostomial angle subquadrate, obtuse. 

Abdomen smooth and compressed, with all pleurae rounded posteriorly. Third 
segment produced posterodorsally, with straight dorsal outline. Sixth segment twice as 
long as fifth segment and more than twice as long as maximal depth; posterolateral and 
posteroventral angles obtusely produced. Abdominal sternites unarmed, sixth sternite 
with small preanal tubercle. 

Telson (Fig. 2B) slender, as long as sixth abdominal segment and 4 times longer 
than maximal width at anterior fourth; with 5 pairs of small (less than 0.05 of tel- 
son length), irregularly placed dorsolateral spinules, first pair placed at 0.35 of telson 
length; the following 4 pairs distributed along distal half of telson. Posterior margin 
broadly rounded, with 5 pairs of spines, lateral shorter than dorsal spines, remaining 
spines longer, with submedian pair longest but not reaching 0. 1 of telson length. 

Eyes (Fig. 5F) long and cylindrical; cornea well pigmented, shorter than stalk, with 
a distinct pointed apical tubercle; without accessory pigment spot. 

Antennular peduncle (Fig. 2A) with basal segment long, more than twice as long 
as distal 2 segments combined, with several long plumose seta on distal dorsal margin; 
mesial margin with deep and thin ventral keel along whole segment. Stylocerite reach- 
ing about middle of basal segment; outer margin nearly straight, ending in acute spine 
far overreaching transverse anterior margin; inner distal angle angulate, pointed. Distal 
2 segments subequal, short and broad. Upper flagellum short, composed of about 9 
segments with 4 basal segments swollen, and with about 6 groups of aesthetascs on 
distal segments; lower flagellum slightly longer than upper flagellum. 

Antenna (Fig. 2C) with basicerite bearing well developed distolateral tooth; car- 
pocerite cylindrical, reaching middle of scale. Scaphocerite distinctly exceeding anten- 
nular peduncle, about 3 times as long as broad; outer margin straight, ending in a stout 
spine, far overreached by angulate distomesial part of lamina. Flagellum slender, long, 
about half as long as body. 

First pereiopod (Fig. 2D-E) slender, not chelate, reaching apex of rostrum; dacty- 
lus fully reduced, possibly indicated only as part of terminal spinulation of propodus; 
propodus about 7.0 times longer than basal depth, slightly tapering to apex; carpus, 
merus and ischium elongate, unarmed, almost uniformly wide, and about 0.8, 1.3 and 
0.8 of propodus length, respectively. 



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Ivona Horkd et al. I ZooKeys 407: 9-28 (2014) 




Figure 2. Anachlorocurtis occidentalis sp. n., holotype, ovigerous female (RMNH.CRUS.D. 56175). 
A anterior carapace, antennae and eyes, dorsal B telson and uropod, dorsal C antenna, ventral D right 
first pereiopod E same, terminal segment F right second pereiopod G same, chela, ventrolateral aspect 
H third pereiopod, lateral J same, dactylus and distal propodus K fourth pereiopod L fifth pereiopod. 



Second pereiopod (Fig. 2F-G) slender, chelate, not reaching distal end of eye; chela 
small, spatulate, with large gap between fingers, dactylus curved, with distally den- 
ticulate lateral cutting edge, fixed finger short, stout, with several irregular distal teeth 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



17 



on cutting margin, with pair of large denticulate spines with hooked apices on apex, 
and with several simple setae subterminally; palm slightly longer than dactylus, swol- 
len proximally; carpus about 3 times as long as chela, three-jointed, with length ratios 
1 .0 : 1 .7 : 1 .4 (distal to proximal), proximal segment obliquely articulating intermediate 
one; merus two-thirds length of carpus; ischium as long as merus. 

Third pereiopod (Fig. 2H, J) more robust than first 2 pereiopods, reaching end 
of basal antennular segment; dactylus slender, 5.0 times longer than basal depth, 
curved, with a distinct unguis; propodus 3.5 times as long as dactylus and about 10 
times longer than basal width, tapering distally, with one distinct spinule in middle 
of ventral margin, without distoventral spines near articulation with dactylus; carpus 
about 0.4 of propodus length and slightly deeper distally than propodus, distodorsal 
end extending over base of propodus as flat expansion; merus about 0.8 of propodus 
length and about 6.0 times longer than uniform width, with 2 distinct ventrolateral 
spines, one subterminal and one at midlength of segment; ischium, basis and coxa 
short, unarmed. 

Fourth and fifth pereiopods (Fig. 2K, L) subequal to third pereiopod, fifth slightly 
longer and more slender than fourth; fourth pereiopod similar to third pereiopod in spi- 
nation of both propodus and merus, with proximal spine placed more forwards (distal 
0.6 of ischium length); fifth pereiopod merus armed with one subterminal spine only. 

Endopod of first pleopod reduced, triangular, appendix interna without cincin- 
nuli, with marginal plumose setae. 

Uropod (Fig. 2B) with branches slightly shorter than telson; outer margin of exo- 
pod almost straight, terminating in short fixed spine and with more mesial movable 
spine about twice overreaching fixed tooth; diaeresis well developed, unarmed mesially 
from lateral movable spine. 

Description of mouthparts (paratype - ovigerous female PoCL 2.6 mm; UO 
Aq08-34B). Mandible (Fig. 3A-C) without palp; molar process with broadly truncate 
distal end densely covered by sharp tubercles and some slightly larger marginal teeth 
and subequal stout setae; incisor process basally broad, tapering distally, with obliquely 
truncate distal end comprised of 4 subquadrate, distally serrate, proximal teeth, and 2 
slender simple teeth, terminal one more produced. 

Maxillula (Fig. 3D) with palp truncate distally and bearing 2 long pappose setae; 
broad distal lacinia armed with numerous strong pappose setae; basal lacinia elongate, 
slender, tapering distally, with about 6 pappose or simple setae terminally. 

Maxilla (Fig. 3E) with short palp about 2.5 times longer than broad basally, with 2 
pappose setae distally; scaphognathite well developed, anterior lobe produced, almost 
twice longer than broad basally, posteriorly rounded, marginal setae densely plumose. 
Distal endite divided into 2 parts with plumose setae marginally; basal endite not di- 
vided and with single apical pappose seta. 

First maxilliped (Fig. 3F) with elongate palp, more than 3.0 times longer than 
wide basally, with 3 simple setae distally; exopod composed of caridean lobe with long 
plumose marginal setae, flagellum absent; endites separated by faint incision, with 
sparsely distributed pappose setae; epipod broadly triangular, faintly bilobed. 



18 



Ivona Horkd et al. I ZooKeys 407: 9-28 (2014) 




Figure 3. Anachlorocurtis occidentalis sp. n., mouthparts (ovigerous female, PoCL 2.6 mm, UO Aq08-34B). 
A mandible, anterior aspect B same, molar and incisor processes, anteroventral C same, apex of molar process, 
anteromesial D maxillula E maxilla F first maxilliped G second maxilliped H third maxilliped. 

Second maxilliped (Fig. 3G) with dactylar segment completely fused to propodus, 
mesial margin with more than 15 stout pappose setae; single pappose seta situated 
distolaterally on merus; ischium broader and longer than carpus; basis and coxa fused, 
with division faintly indicated; exopod absent; epipod large, broadly ovate. 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



19 



Third maxilliped (Fig. 3H) slender and long, failing to reach end of basal antennu- 
lar segment; lateral coxal plate ovate, with small spiniform apex; antepenultimate seg- 
ment (i.e., merus, ischium and basis fused) with subdivisions feebly indicated, about 
12 times longer than uniform width and 1.5 times as long as distal 2 segments com- 
bined; setose concavity present on ventral margin of ischium; penultimate segment 
(i.e., carpus) about 4.0 times longer than wide basally and 0.4 of preceding segment 
length, with several serrulate setae ventrally; ultimate segment about 1 .2 times longer 
than penultimate segment, furnished with about 8 dorsomesial rows of serrulate setae. 

Other specimens. Males generally similar to adult females but distinctly smaller 
and more slender in lateral aspect (see Fig. 1). Allotype male (Figs IB, 5D) with ros- 
trum simple, styliform, with anteriad directed, bifid, dorsal tooth placed just above 
level of posterior orbital margin, and with obtuse epigastric tooth more posteriorly; 
posterior hooked tooth well developed at posterior third of carapace, posterior sub- 
marginal tooth small but distinct. Upper antennular flagellum with basal swollen part 
(corresponding to fused flagella) more elongate than in adult females, with stronger 
developed aesthetascs. Endopod of first pleopod (Fig. 4A, B) reduced to small setose 
lobe with bilateral setae and pair of pappose terminal setae, appendix interna far over- 
reaching reduced endopod but falling short of midlength of exopod, elongate triangu- 
lar, with 9 simple hooked spiniform setae on mesial margin, and with group of about 
8 cincinnuli on oblique apex. Endopod of the second pleopod (Fig. 4C) almost as long 
as exopod, with pair of appendices emerging from proximal third of mesial margin; 
appendix masculina slender, more than 6.0 times longer than basal width, reaching 
midlength of endopod; with 4 stout simple setae on apex. Appendix interna reaching 
0.75 of appendix masculina length, with about 8 terminal cincinnuli (Fig. 4D). 

Variation. The main morphological variation observed within the samples of the 
new species is in the dorsal armament of the carapace and rostrum (Fig. 5A-E). While 
in adult and subadult females the carapace is trilobate, comprised of 2 high, anteriorly 
denticulate, similar "lobes", i.e., the rostrum and the anterior carapacial lobe, and a 
large posterior hooked tooth, in males only the postrostral lobe is developed, being 
small and bidentate, in addition to a short styliform rostrum and the posterior hooked 
tooth. In juveniles, the posterior hooked tooth is absent, and the anterior ornamenta- 
tion generally consists of a short simple rostrum with a single posterior dorsal tooth. 
The rostral formula (number of anterior dentition of the postorbital lobe + same for 
rostrum) is 3-4 + 2-6 for adult females, 2-3 +1-3 for subadult females, 2+1 (i.e., 
simple rostrum) for males, and 1 + 1-3 for juveniles. 

The small epigastric tooth is placed some distance from the posterior end of the 
base of the small anterior tooth on the carapace in juveniles, not on the base of the lobe 
as in adults. The posterior dorsal submarginal tooth, well developed in adult females, 
is small but distinct in males and subadult females, and may present also in juveniles. 

The dorsal telson dentition generally consists of 3 pairs of spinules, with the first 
pair situated anteriorly to the midlength of the telson. Some females possess 4 pairs of 
those spinules, e.g., in a subadult female (UO Aq09-30A), or 5 pairs (ovigerous female 
holotype, RMNH.CRUS.D.56175). 



20 



Ivona Horkd et al. I ZooKeys 407: 9-28 (2014) 




Figure 4. Anachlorocurtis occidentalis sp. n., allotype, male (RMNH.CRUS.D. 56176). A first pleopod 
B same, endopod C second pleopod D same, mesial margin of endopod with appendices interna and 
masculina. 



The specimens are consistent in the number of the meral spines on the 3 rd -5 th 
pereiopods, 2-2-1, respectively. A limited variation is present in the propodal ventral 
spinulation in the ambulatory pereiopods. In the females examined, the spinulation is 
1-1-1, respectively, with the spinules very minute, while in adult males the spinules are 
stronger, numbering 2-2-1/2, respectively. 

Colour in life (Fig. 6A-C). Cryptic, mimicking antipatharian host, ventral part 
of body reddish brown (similar to axial coral branch pattern), dorsal part of body 
transparent with yellowish to light reddish stripes emerging from internal axis of body 
(similar to host polyps), with narrow lighter band across carapace; antennulae light 
brown, scaphocerites with 2 light brown patches (distal and proximal), connected by 
narrow medial stripe; abdomen with irregular narrow light bands across each segment; 
tail fan (Fig. 6C) with 3 wide light brown transverse bands across uropods, with colour 
merged along mesial margin of endopod. Ovigerous females with greyish-brown finely 
marbled pattern on abdominal pleurae covering egg mass. 

Measurements. Holotype ovigerous female: PoCL 3.3 mm, RL 0.8 mm, TL about 
12 mm, eggs (without eye-spots) diameters 0.58 x 0.42 mm. Allotype male: PoCL 
2.0 mm, RL 0.23 mm, TL 8.7 mm. Paratypes: males (5 spms) PoCL 1.6—1.8 mm, 
TL 7.1—8.2 mm; ovigerous females (8 spms) PoCL 2.2—3.1 mm, TL 8.7—11.5 mm; 
subadult females (2 spms) PoCL 1.9 mm; juveniles (2 spms) PoCL 1.5-1.6 mm; eggs 
without eyespots diameters (7 spms) 0.44-0.56 x 0.36-0.40 mm, eggs with eyespots 
(1 spm) 0.62 x 0.48 mm. 

Host and habitat. Specimens were caught on black corals {Antipathes sp.), found 
in deeper waters along the reef wall and on a boat wreck at 22-55 m, but also in dark 
crevices in shallow waters, only 4-5 m deep. 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



21 




Figure 5. Anachlorocurtis occidentalis sp. n., (A— F Aqaba, 2009) and A. commensalis Hayashi, 1975 
(G-P Taiwan, 2009), outline of carapace, lateral aspect (A-E, G-L), third pereiopod (M-N), and eye 
(F, O-P). A holotype, ovigerous female PoCL 3.3 mm (RMNH.CRUS.D. 56175) B ovigerous female 
PoCL 2.5 mm (RMNH.CRUS.D. 56179) C subadult female PoCL 1.9 mm (UO Aq09-65) D allo- 
type, male PoCL 1.9 mm (RMNH.CRUS.D. 56176) E juvenile specimen PoCL 1.6 mm (RMNH. 
CRUS.D.56179) F ovigerous female PoCL 2.5 mm (RMNH.CRUS.D. 56179) G post-ovigerous female 
PoCL 2.6 mm (OUMNH.ZC.20 10-02-0 10 ) H ovigerous female PoCL 2.6 mm J subadult female 
PoCL 1.9 mm K male PoCL 2.2 mm L juvenile specimen PoCL 1.6 mm (OUMNH.ZC.2010.02.061) 
M-O post-ovigerous female PoCL 2.6 mm (OUMNH.ZC.2010-02-010) P male PoCL 2.2 mm. Scale 
bars equal 1 mm. 



22 



Ivona Horkd etal. I ZooKeys 407: 9-28 (2014) 




Figure 6. Colour patterns. Anachlorocurtis occidentalis sp. n. A ovigerous female holotype (RMNH. 
CRUS.D. 56175) on antipatharian host B detail of the eye with produced apical tubercle C colour patterns 
of the tail fan. Anachlorocurtis commensalis Hayashi, 1975 D ovigerous female (UO Twl2-79). 



Associated fauna. The specimens of the present new species were collected from 
their hosts together with specimens of the pontoniine shrimps, Manipontonia psamathe 
(De Man, 1902) and Periclimenes cf lepidus Bruce, 1978, both not previously reported 
from the Red Sea. 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



23 



Etymology. The specific name is from the Latin occidentalis, western, reflexing 
the geographic range of the new species in the westernmost region of the Indo-West 
Pacific area, the Red Sea, as opposed to the East Asian distribution of the type species 
of the genus. 

Distribution. Currently only known from the type locality, Gulf of Aqaba, in the 
north-eastern Red Sea. 

Anachlorocurtis commensalis Hayashi, 1975 

http://species-id.net/wiki/Anachlorocurtis_commensalis 
Figs 5G-P, 6D, 7E-G, 8 

Anachlorocurtis commensalis Hayashi 1975: 172-182, figsl-3.; Hayashi 2007: 147— 
150, figs 538-539.; Kawamoto and Okuno 2003: 64, colour photo.; Minemizu 
2013: 129, colour photo. 

Material examined, (i) 8 spms (1 ovigerous female PoCL 2.6 mm, 1 female subadult 
PoCL 2 mm, 3 males PoCL 1.8-2.1 mm, 3 juveniles PoCL 1.6-1.8 mm) (OUM- 
NH.ZC.2010.02.061), Gudanshr reef, Namwan Bay, Pingtung County, Taiwan, 
21°56.531'N, 120°45.546'E, from bushy black coral, 10 m depth, leg. S De Grave, 
07.12.2009 (fen TAI-244); (ii) 10 spms (4 females ovigerous PoCL 2.5-2.6 mm, 1 
female post ovigerous PoCL 2.6 mm, 1 female PoCL 2.3 mm, 1 female subadult PoCL 
1.9 mm, 3 males PoCL 1.9-2.2 mm) (OUMNH.ZC.20 10-02-0 10), Gudanshr reef, 
Namwan Bay, Pingtung County, Taiwan, 21°56.531'N, 120°45.546'E, from Cirrhi- 
pathes sp., 27 m depth, leg. S De Grave, 12.12.2009 (fen TAI-327); (iii) 1 juvenile 
PoCL 1.8 mm (RMNH.CRUS.D.56181), Gudanshr reef, Namwan Bay, Pingtung 
County, Taiwan, from antipatharian on overhanging rock wall, 10.9 m depth, leg. Z 
Duris, 1 1.1 1.201 1 (fcnTwl 1-11); (iv) 1 juvenile (damaged) PoCL 1.8 mm (OUTwl 1- 
20B), Gudanshr reef, Namwan Bay, Pingtung County, Taiwan, upper side of rocky 
reef, from yellow branching antipatharian, 18.6 m depth, leg. Z Duris, 03.11.2011 
(fen Twll-20B); (v) 1 ovigerous female PoCL 2.3 (RMNH.CRUS.D.56182, Gen- 
bank KJ690258), 1 ovigerous female, PoCL 2.4 mm, 1 male PoCL 2.1 mm (NTOU 
M0 1305), Gudanshr reef, Namwan Bay, Pingtung County, Taiwan, upper side of reef, 
from dense bushy antipatharian, 22.1 m depth, leg. Z Duris , 03.1 1.201 1 (fen Twl 1- 
21); (vi) 1 ovigerous female PoCL 2.5 mm (dissected, dried for SEM; UO TW12-79), 
Gudanshr reef, Namwan Bay, Pingtung County, Taiwan, reef slope, from antipathar- 
ian, 25.3 m depth, leg. Z Duris & I Horka, 12.09.2012 (fen Twl2-79). 

Remarks. The specimens examined agree well with the original description by 
Hayashi (1975). As in A. occidentalis sp. n., the specimens examined of A. commensalis 
vary in the dorsal armament of the carapace and rostrum (Fig. 5G-L). In adult and 
subadult females the carapace with the rostrum is triple lobate, in contrast in males 
only the anterior, bidentate, lobe is developed, in addition to the posterior hooked 
tooth and a short styliform rostrum. In juveniles the posterior hooked is absent and the 



24 



Ivona Horkd etal. I ZooKeys 407: 9-28 (2014) 




Figure 7. Anachlorocurtis occidentalis sp. n., ovigerous female paratype, PoCL 3. 1 mm (UO Aq09-30A). 
A second pereiopod chela B same, tips of fingers C mandible, molar process, antero-mesial aspect (from 
side of incisor process) D eye. Anachlorocurtis commensalis Hayashi, 1975, ovigerous female PoCL 2.5 mm 
(UO Twl2-79) E second pereiopod chela F same, tips of fingers G eye. 



anterior ornamentation generally consists of a short simple rostrum with a single pos- 
terior dorsal tooth. The posterior dorsal submarginal tooth is small in adults and sub- 
adults, and is absent in juveniles. The rostral formula (see above) is 2-4 + 4-5 for adult 
females, 2-3 + 1-2 for subadult females, 2+1 (i.e. simple rostrum) for males, and 1-2 + 
1 for juveniles. Cornea in males is almost globular, with the apical tubercle indistinct, 
in females the tubercle is more produced but not as developed as A. occidentalis sp. n. 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



25 



100 [ Anachlorocurtis occidentalis sp. n. (v) 



59 



Anachlorocurtis occidentalis sp. n. (ii) 



100 



Anachlorocurtis commensalis 



Miropandalus hardingi 



50 



Heterocarpus ensifer 



Heterocarpus gibbosus 



76 



Pandalus borealis 



100 



Pandalus montagui 



Stenopus hispidus 



0.1 



Figure 8. Phylogenetic tree obtained by maximum likelihood analysis (GTR+G+I substitution model) 
of partial sequence of COI (658 bp). 



Colour in life. Generally similar to that of A. occidentalis sp. n., but the light trans- 
verse bands are greyish, more wide and distinctive in A. commensalis (Fig. 6D); also see 
colour photos in Minemizu (1996: 8; 2000: 1 13; 2013: 129), Kato and Okuno (2000: 
58) and Kawamoto and Okuno (2003: 64). 

Distribution. Kii Peninsula (type locality), Izu-Oshima and Hachijyo-jima, Izu Is- 
lands, and Kume-jima, Ryukyus, Japan (Minemizu 1996, 2000, 2013, Kato and Okuno 
2000), Kawamoto and Okuno (2003); South (Namwan) Bay, Taiwan (present report). 



Molecular data 

Molecular analysis (Maximum Likelihood, Fig. 8) supports a sister position of Ana- 
chlorocurtis occidentalis sp. n. (Red Sea) to A. commensalis (Taiwan). Both species form 
a monophyletic clade with Miropandalus hardingi. The maximum Kimura 2-parameter 
distance between A. occidentalis sp. n. and A. commensalis reached 15.4% (for more 
details see Table 2). 



Discussion 

Anachlorocurtis occidentalis sp. n. is morphologically closely related to the only other 
species in the genus, A. commensalis. Both species have a small, slender body and am- 
bulatory legs. The third to fifth pereiopods have the propodi about 10 times longer 
than deep basally A. occidentalis, whilst 8 times longer in A. commensalis, and the dac- 
tyli are 5.5 times longer than deep basally in the new species, and 3 times longer than 



26 Ivona Horkd et al. I ZooKeys 407: 9-28 (2014) 



Table 2. Kimura's 2-parameter distances of COI gene sequences among species studied. 





A. occidentalis sp. n. 


A. occidentalis sp. n. 


A. commensalis 


1 
1 


H. ensifer 


ii 
3 

-0 

•a 
So 

«! 


3 

-a 
*s 


P. montagui 


S. hispidus 


Anachlorocurtis occidentalis sp. n. (RMNH. 
CRUS.D.56177) 




















A. occidentalism, n. (RMNH.CRUS.D. 56174) 


0.002 


















A. commensalis 


0.154 


0.152 
















Miropandalus hardingi 


0.156 


0.154 


0.192 














Heterocarpus ensifer 


0.254 


0.252 


0.267 


0.268 












H. gibbosus 


0.300 


0.297 


0.295 


0.272 


0.211 










Pandalus borealis 


0.278 


0.275 


0.286 


0.293 


0.221 


0.236 








P. montagui 


0.268 


0.266 


0.276 


0.270 


0.220 


0.246 


0.123 






Stenopus hispidus 


0.253 


0.251 


0.237 


0.248 


0.219 


0.226 


0.251 


0.230 





deep in A. commensalis. The new species also differs from A. commensalis by the poste- 
rior dorsal lobe on the carapace being strongly hooked in adults (vs. feebly hooked), 
and more developed posterior submarginal tooth dorsally on the carapace in adults; by 
a more elongate apical tubercle on cornea; by a small and single posterior propodal spi- 
nule on the 3 rd - 5 th pereiopods (vs. stronger, with their number 2-2-2, respectively, in 
A. commensalis); by the distinctly angulated third abdominal segment in lateral aspect, 
with dorsally straight outline (vs. rounded); by the more elongated sixth abdominal 
segment (2.5 times longer that deep in A. occidentalis vs. 2.0 in A. commensalis); by 
the greater number of setae on the mesial margin of the endopod of the first pleopod 
of the male (9 versus 3, respectively); by the length of the appendix masculina of the 
second male pleopod (longer than the appendix interna versus shorter), and possibly 
a higher number of dorsal telson spines (3-5 pairs in A. occidentalis, versus 3 pairs in 
A. commensalis). 

The morphological distinction between the two species is further supported by 
small differences in their colour pattern, but more importantly by their COI diver- 
gence being 15.2—15.4%. 



Acknowledgements 

The authors are grateful to Dr. Ken-Ichi Hayashi (Shimonoseki, Japan) for provid- 
ing unavailable literature and bibliographic data, Dr. Nanette E. Chadwick (Auburn 
University, Alabama, U.S.A.), Dr. Fuad Al-Horani (Marine Science Station in Aqaba, 
Jordan), Dr. Tin-Yam Chan (National Taiwan Ocean University, Keelung), and Dr. 
Chia-Wei Lin (Kenting Marine National Park, Taiwan) for providing facilities and/or 



Anew species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea. 



27 



project possibilities to examine the Red Sea and Taiwanese shrimps, and two referees 
for valuable comments. This study was partially supported by the US-Jordan Project 
'Summer 2009' (NSF-IRES), projects of University of Ostrava (sgs05/PfF/2012 and 
sgs/29PrF/20l4) (IH), the project IET (CZ.l. 05/2.1.00/03.0100) financed by the 
Structural Funds of the European Union and Project LO1208 of the National Fea- 
sibility Programme-I of the Czech Republic, the Czech National Committee for co- 
operation with IOC UNESCO (ZD), and the EU-FP7 ASSEMBLE program (SDG). 



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