The Apparent Rediscovery Of Nematopagurus Scutellichelis Alcock, 1905 With Notes On Other Regional Species Of The Genus

13 Pages·2002·5.2 MB·English

by  P A Mclaughlin

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
115(4):741-753. 2002.
The apparent rediscovery of Nematopagurus scutellichelis
Alcock, 1905 (Decapoda: Anomura: Paguroidea: Paguridae) in the
South China Sea, with notes on other regional species of the genus
Patsy A. McLaughlin and Yong-Liang Wang
(PMcL) Shannon Point Marine Center, Western Washington University, 1900 Shannon Point
Road, Anacortes, Washington 98221-9081B, U.SA.;
(Y-LW) Institute of Oceanology, Chinese Academy of Sciences, 7 Nanhai Road, Qingdao
266071, People's Republic of China
—
Abstract. The accuracy of records of species of the hermit crab genus Ne-
matopagurus occurring in the South China Sea has been evaluated using col-
lections from the Hainan Island region. Information gleaned from this material
strongly suggests that previous reports have been inaccurate. None of the re-
ported four species, A^. gardineri Alcock, N. indicus Alcock, N. muricatus
(Henderson), and N. squamichelis Alcock have been recognized among the
extensive collections of the Institute of Oceanology, Chinese Academy of Sci-
ences, Qingdao, People's Republic of China. Nevertheless, a total of seven
species of Nematopagurus have been identified, including what appears to be
N. scutellichelis Alcock. Ifcorrect, this would be the first record ofthis species
since its original description. The other six are also known species; however,
most represent new distributional records.
Nematopagurus A. Milne-Edwards & son, 1896) were added to the Chinese fau-
Bouvier, 1892 is one of several genera of nal fist by Wang (1983). Rahayu (1996)
the hermit crab family Paguridae character- also recorded TV. muricatus from Singapore
ized by a sexual tube produced from, or de- and Malaysia. While absent from the South
veloped in conjunction with, the male right China Sea, A^. vallatus (Melin, 1939) was
gonopore that is usually elongate and often recorded from the East China Sea by Wang
filiform. Typically, a similar, but very much (1992). All five species were included by
shorter tube is produced from the left gon- Wang (1994) in his list of "Marine Species
opore. Females of this genus also develop and Their Distributions in China's Seas".
paired and modified first abdominal pleo- Unfortunately none of these species identi-
pods. At present, 24 species are assigned to fications were correct.
the genus (McLaughlin 1998), all but one Regrettably, Wang's (1994) specimens
of which occur in the IndoPacific region. In have not been available for reexamination.
a recent summary of the hermit crab fauna However, extensive surveys in the Hainan
ofthe South China Sea, Rahayu (2000) list- Island region of the South China Sea were
ed four species of Nematopagurus. For the conducted by Chinese research vessels
most part, Rahayu's report was a compila- (RA'^) during the National Comprehensive
tion from the existing literature. The occur- Oceanographic Survey (NCOS) of 1958-
rences of A^. indicus Alcock, 1905b and A^. 1960, with supplemental sampling in the
squamichelis Alcock, 1905b, in the South East China Sea in 1976, by the Institute of
China Sea had been reported by Wang & Oceanology, Chinese Academy of Sciences
Huang (1979). Nematopagurus gardineri (lOCAS). These surveys resulted in the col-
Alcock, 1905a and A^. muricatus (Hender- lection ofa wealth ofhermit crabs thathave
742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
allowed us to evaluate the species of Ne- (Stimpson, 1858) = A^. vallatus sensu Wang
matopagurus occurring in Chinese Seas. (1992, 1994); A^. alcocki McLaughUn, 1997
We also have reexamined the specimens = ?A^. indicus sensu Wang & Huang (1979),
from Singapore referred by Rahayu (1996) Wang (1983, 1992, 1994), Huang (1989);
to A^. muricatus and deposited in the Raffles N. meiringae McLaughlin, 1998 = ?A^. gar-
Museum ofBiodiversity Research, National dineri sensu Wang (1983, 1992, 1994),
University of Singapore (ZRC); the speci- Huang (1989); and A^. diadema Lewinsohn,
men identified by Balss (1912) as N. indi- 1969, heretofore reported only from the
cus from the Nicobar Islands in the collec- Red Sea and eastern coast of South Africa.
tions of the Naturhistorisches Forschung- Although all are known species, only A^. tri-
sinstitut Museum fiir Naturkunde zu Berlin, carinatus has been reported previously
Berlin, Germany (ZMB); and specimens in from the South China Sea. A complete re-
the collections of the Institute of Oceanol- description is presented herein for A^. scu-
ogy, Chinese Academy of Sciences (lO- tellichelis, whereas the other species in the
CAS), identified by resident scientists as N. Institute of Oceanology collections are dis-
muricatus and A^. squamichelis. cussed only as they pertain to previous mis-
A species not recognized by previous identifications. However, pertinent, albeit
Chinese carcinologists, but present in the not complete, synonymies are included. De-
Institute ofOceanology collections, is atax- scriptions and/or diagnoses ofthese species
on identified by the present authors as Ne- have been presented by McLaughlin (1997,
matopagurus scutellichelis Alcock, 1905b. 1998, 2002) and Komai & McLaughlin
This species was described by Alcock (2001).
(1905b) from a single male specimen col- In the materials examined, one measure-
lected in the Indian Ocean between the ment, shield length (si), as measured from
Maldives and Cape Comorin at a depth of the midpoint of the rostral lobe to the mid-
1507 m. Any subsequent reports ofthis spe- point of the posterior margin of the shield
cies have merely been literature citations of provides an indication of specimen size.
the original record (e.g., Gordan 1956). The The abbreviation Stn. refers to ship stations,
species collected during the Hainan Survey while ovig. refers to the ovigerous condi-
is from much shallower depths. Nonethe- tion of some females. Terminology used in
less, it agrees, with the exception of seta- the description ofTV. scutellichelisgenerally
tion, entirely with Alcock's description and follows that of McLaughlin (1998). The ra-
illustration. The policy of the Indian Mu- tio ofcorneal diameter to peduncularlength
seum, Calcutta, to not lend materials, has was obtained by comparing maximum cor-
precluded personal examination of the ho- neal width to maximum pedunculai" length
lotype; however, in view of the setal vari- measured on the lateral surface of the pe-
ation common to species of this genus duncle from the distal corneal margin to the
(McLaughlin, pers. obs.), we believe that proximal margin of the distal segment of
the specimens redescribed herein as N. scu- the peduncle. Inteipretation of the ocular
tellichelis are true representatives of Al- acicle follows that of Forest et al. (2000),
cock's taxon. not that of Boyko & Harvey (1999). Bor-
In addition to A^. scutellichelis, the Insti- rowed specimens have been returned to the
tute of Oceanology collections contain the lending institutions. The majority of speci-
following species of Nematopagurus: N. mens are deposited in the collections of the
australis (Henderson, 1888) = A^. muricatus Institute of Oceanology, Chinese Academy
sensu Wang (1983) and Rahayu (1996); N. of Sciences, Qingdao, People's Republic of
lepidochirus (Doflein, 1902) = N. squami- China; however, where adequate numbers
chelis sensu Wang & Huang (1979), Huang of specimens have allowed, representatives
(1989), and Wang (1994); N. tricarimitus have also been placed in the collections of
VOLUME 115, NUMBER 4 743
the National Museum of Natural History, sparse tuft of stiff setae at corneal inden-
Smithsonian Institution, Washington, D.C. tation, dorsal and/or dorsomesial surface(s)
(USNM). with sparse tuft(s) of setae; corneal diame-
ter 0.55-0.70 of peduncular length. Projec-
Nematopagurus scutellichelis tion of ocular acicles small, subrectangular;
Alcock, 1905 terminating subacutely, with deeply con-
Fig. 1 cave dorsal surface and well developed sub-
marginal spine.
Nematopagurus scutellichelis Alcock,
Antennular peduncles exceeding distal
1905b:112, pi. 12, fig. 3.
margins of corneas by 0.30-0.50 length of
—
Material examined. S.E. off Hainan Is- ultimate segments. Ultimate segment with
land, South China Sea: RA^2, NCOS Stn sparse tuft of setae at dorsolateral distal
6091, 20°30'N, 112°30'E, 78 m, 7 May margin, few additional sparse tufts on dor-
1959, 1 c? si = 4.1 mm, lOCAS K57B; sal or dorsolateral surfaces; penultimate
RA^S, NCOS Stn 6052, 21°30'N, 114°00'E, segment with few scattered setae. Basal
48 m, 17 Mar 1959, 1 ovig. 9 si = 3.6 mm, segment with acute spine on distal margin
lOCAS S15BS; R/V21, NCOS Stn 6107, of statocyst lobe.
20°00'N, 112°00:E, 92 m, 4 Apr 1960, 1 6 Antennal peduncles overreaching distal
si = 2.7 mm, lOCAS K147B; R/VlOl, margins of corneas by 0.10-0.20 length of
NCOS Stn 6161, 18°15'N, 110°30'E, 106 fifth segments. Fifth and fourth segments
m, 7 May 1959, 1 c? si = 4.2 mm, IOCAS each with few to several tufts of simple se-
N197B; R/V171 NCOS Stn. 6107, tae. Third segment unarmed, but with tuft
20°00'N, 112°00'E, 96.5 m, 20 Apr 1959, of long, stiff setae at ventrodistal angle.
1 2 si = 3.2 mm, lOCAS L57B; RA^171, Second segment with dorsolateral distal an-
NCOS Stn 6166, 16°30'N, 110°30'E, 72 m, gle prominently produced, terminating in
20 Apr 1959, 3 c? si = 2.0-3.0 mm, 2 9 acute spine, lateral and mesial margins with
si = 3.2, 3.6 mm, 1 ovig. 9 si = 3.0 mm, few stiff setae; dorsomesial distal angle
lOCAS L568-62; R/V220, NCOS Stn. with small spine. First segment unarmed.
6017, 22°00'N, 116°30'E, 88 m, 23 Apr Antennal acicle moderately long, reaching
1959, 1 (? si = 2.2 mm, lOCAS S229B; beyond proximal half of ultimate pedun-
R/V220, NCOS Stn 6037, 2r30'N, cular segment; arcuate, terminating in acute
1 15°00'E, 90.4 m, 4 Jul 1959, 1 cJ si = 4.8 spine; mesial margin with tufts oflong stiff
mm, USNM 1003650; R/V220, NCOS Stn. setae. Antennal flagella long, overreaching
6054, 21°00'N, 114°00'E, 80 m, 10 Dec tip ofright cheliped; most articles each with
1959, 2 9 si = 2.0, 2.4 mm, lOCAS 1 or 2 extremely short setae or bristles, at
S141B. — least proximally.
Redescription. Shield (Fig. la) broader Chelipeds subequal; right (Fig. lb)
than long; anterolateral margins sloping; slightly, if at all, longer, but stouter than
posterior margin roundly truncate, usually left, both moderately elongate. Dactyl of
with small median concavity; dorsal surface right cheliped appreciably shorter to nearly
with few sparse tufts of short setae laterally equaling length of palm; cutting edge with
and anteriorly. Rostrum very broadly several, sometimes fused, calcareous teeth
rounded, sometimes produced to level of and short row of corneous teeth in distal
lateral projections. Lateral projections mod- 0.20-0.25, terminating in small corneous
erately well developed; each with mesially claw and slightly overlapped by fixed fin-
or laterally directed, marginal or submar- ger; dorsal surface with longitudinal row of
ginal spinule. scutes, extending onto rounded dorsomesial
Ocular peduncles 0.80-1.00 length of margin, each with marginal row of short
shield; dorsal surfaces each with median stiff setae, few tufts of short setae near tip
744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. \. Nematopaguriis scutellichelis Alcock, 1905, cJ si = 4.8 mm, R/V220, NCOS Stn. 6037. USNM
1003650: a, shield and cephalic appendages; b, carpus and chela ofright cheliped (dorsal view); c, right second
pereopod (lateral view); d, leftthird dactyl (mesial view); e, sternite andcoxae offifthpereopods (ventralview);
f, telson. Scales equal 1.0 mm.
and adjacent to cutting edge; mesial face each with marginal row of short stiff setae;
ventrally and ventral surface with tufts of dorsal surface offixed finger with moderate
long setae. Palm 0.80 to nearly as long as to short transverse rows ofscutes extending
carpus; dorsomesial margin not delimited; nearly to tip and provided with marginal
dorsal surface with 6-8 rows of partially to short stiff setae, few sparse tufts of setae
nearly complete transverse scutes continued near tip; cutting edge with row of calcare-
onto lateral, mesial, and ventral surfaces. ous teeth, terminating in tiny corneous
VOLUME 115, NUMBER 4 745
claw; scutes of mesial and ventral surfaces dorsolateral margin not delimited; dorsal
ofpalm accompanied by longer setae, often surface with several complete or more fre-
particularly dense on mesial face of palm. quently interrupted, marginally setiferous
Carpus slightly longer than merus; dorso- scutes extending onto lateral and mesial
distal margin with row of uniformly short faces; ventral surface with tufts of setae, 1
stiff setae; dorsal surface with complete or small spine at ventrolateral distal angle.
incomplete transverse scutes extending onto Merus subtriangular; dorsal surface with
lateral and mesial faces and provided mar- transverse ridges and stiff setae, distal mar-
ginally with short stiff setae, most scutes gin with row of moderately long stiff setae;
interrupted at dorsomesial margin by mod- lateral and mesial faces with tufts of stiff
erately small to prominent spine; dorsolat- setae; ventral surface with few ridges and
eral margin not delimited; ventral surface tufts of setae; ventromesial margin with 1
with transverse rows of long setae. Merus acute spine usually and 2 or 3 low protu-
subtriangular; dorsal margin, mesial and berances; ventrolateral margin with 1 or 2
lateral faces all with transverse ridges and spines distally. Ischium with acute or blunt
long stiff setae; ventrolateral margin with 2 spine at ventrolateral angle.
spines in distal 0.50; ventromesial margin Ambulatory legs (Fig. Ic, d) overreach-
with 1 spine distally, ventral surface with ing tip of right cheliped by at least 0.50
few low ridges and tufts of setae. Ischium length of dactyl. Dactyls approximately
with small acute or blunt spine at ventro- twice length of propodi; in lateral view
lateral angle. straight or slightly (second) to distinctly
Left cheliped usually reaching nearly to (third) curved ventrally; in dorsal view,
tip of dactyl of right; moderately slender. slightly twisted; each terminating in mod-
Dactyl usually slightly longer than palm; erate to strong corneous claw; dorsal sur-
cutting edge with row of small corneous faces each with row ofmoderately long stiff
teeth, terminating in small corneous claw setae; mesial faces each with row of cor-
and slightly overlapped by fixed finger; dor- neous spinules dorsally and row of 9-15
somesial margin rounded, dorsal surface corneous spines at ventromesial margin (not
with few sparse tufts ofshort setae adjacent visible in lateral view). Propodi approxi-
to cutting edge, 1 longitudinal row of mar- mately 0.35 longer than carpi; dorsal sur-
ginally setiferous scutes, each extending faces presenting "polished" appearance,
onto mesial face; ventral surface with few each circumscribed by 6 or 7 weakly pili-
tufts of setae. Palm approximately 0.75 ferous scutes. Carpi 0.45-0.55 length of
length of carpus; dorsomesial margin meri; dorsal surface of second pereopods
rounded; dorsal surface, like that of right, each with row of spines accompanied by
with 6-8 transverse rows ofcomplete or in- tufts of short stiff setae; third pereopods
complete scutes, each extending onto lateral each with similar tufts of stiff setae as well
and mesial faces and provided with margin- as dorsodistal spine and at least 1 additional
al row of short stiff setae, most proximal spine in proximal half; lateral faces each
scute sometimes with small subacute spine with 1 or 2 short, oblique rows of stiff se-
at dorsomesial margin; dorsal surface of tae, ventral and mesial surfaces glabrous or
fixed finger with several short transverse with few tiny tufts ofvery short setae. Meri
scutes, each with marginal fringe of short laterally compressed; dorsal surfaces each
stiff setae; cutting edge with row of small with row of transverse ridges and stiff se-
calcareous teeth interspersed with corneous tae; lateral faces each with 1 or 2 sparse
teeth. Carpus approximately 1.10-1.30 lon- longitudinal rows ofshort setae; mesial fac-
ger than merus; dorsodistal margin with es glabrous or with very few sparse tufts of
uniform row of short stiff setae, dorsome- short setae; ventrolateral margins of second
sial margin with row of prominent spines. pereopods each with small, acute spine at
746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
distal angle and 3-5 additional spines mar- cock simply did not notice these spines,
ginally, accompanied by sparse tufts of se- which are present in all species of the ge-
tae; third only with tufts of setae. Ischia nus.
each with few setae dorsally and ventrally. Differences in the depths at which the
Anterior lobe of sternite of third pereopods South China Sea specimens and Alcock's
m
roundly subquadrate, with long setae on an- (1905b) were collected, 48-106 and
terior margin. 1507 m respectively, are appreciable. How-
Males with well developed, elongate, ter- ever, depth collection extremes also are not-
minally spiraled sexual tube on coxa of ed for N. longicornis A. Milne-Edwards &
&
right fifth pereopod, left with sexual tube Bouvier, 1892. A. Milne-Edwards Bou-
developed as short to moderate tube, vary- vier (1892) cited the depth distribution for
ing from approximately equal to nearly A^. longicornis as 75—800 m; however, sta-
twice coxal length (Fig. le). Telson (Fig. tion data for "Talisman" sample no. 28,
If) with posterior lobes asymmetrical, one of the stations at which N. longicornis
roundly or subacutely triangular; separated reportedly was collected, listed a depth of
by moderate to deep median cleft; terminal 2600 m (A. Milne-Edwards & Bouvier
margins oblique, each with 3-6 acute 1900). Considering the morphological
spines; lateral margins oblique, each with agreement between Alcock's (1905b) de-
row of sm—aller acute or subacute spines. scription and our South China Sea speci-
Color. —Not known. mens, we feel confident that the latter truly
Habitat. Sand and mud substrates in represent Alcock's taxon, the broad geo-
South China Sea. graphic separation not withstanding. Ne-
—
Distribution. Eastern Indian Ocean, matopagurus diadema, heretofore known
1507 m; Sou—th China Sea, 48-106 m. only from the Red Sea and eastern South
Remarks. As indicated previously, the Africa now also has been found in the
several specimens collected in the South South China Sea.
China Sea agree entirely with Alcock's
(1905b) description ofN. scutellichelis, ex- Nematopagurus alcocki McLaughlin, 1997
cept for setation. The scutes on the propodi INematopagurus indicus—Wang & Huang,
of the ambulatory legs of Alcock's speci-
men reportedly were "nude and polished"; 1979:63, fig. 3.—Wang, 1983:54 (Ust);
1992:61 (list); 1994:571 (fist).—Huang,
the dactyls were "perfectly nude". The
1989:253, fig. 214 (not Nematopagurus
scutes on these appendages in our speci-
indicus Alcock, 1905b).
mens all are provided with short, stiffsetae;
Nematopagurus alcocki McLaughlin, 1997:
each dactyl has a row of long stiff setae on
517, fig. 23a-i.
the dorsal margin. The chelae and other ap-
pendages also are provided with additional Material examined.—R/V21, NCOS Stn.
setae apparently absent from Alcock's spec- 6093, 19°30'N, 112°30'E, 219 m, 17 Feb
imen. However, setal variation in Nemato- 1959, 3 cJ si = 1.8-3.3 mm, 2 ? si = 2.7,
pagurus species is subject to great variation 2.8 mm, lOCAS 6-21; IWlOl, NCOS Stn.
(McLaughlin, pers. obs.). Alcock (1905b) 6123, 18°30'N, lir30'E, 220 m, 8 Feb
also indicated and illustrated ambulatory 1960, 1 c? s—i = 2.3 mm, lOCAS N115B.
dactyls that lacked any spinulation on the Remarks. One of the two lots of N. al-
ventral margins. Although a ventral row of cocki previously had been identified as N.
short corneous spines is present on each muricatus by resident scientists at the Insti-
dactyl in all of our specimens, the twisting tute of Oceanology, although all of the oth-
ofthe dactyls is such that these spines occur er samples similarly identified as Hender-
on the ventromesial surface and are not vis- son's (1896) taxon aie referable to N. aus-
ible in lateral view. It is probable that Al- tralis. Tiirkay (1986) pointed out the very
VOLUME 115, NUMBER 4 747
distinctive armature of the chelae of TV. Nematopagurus australis
muricatus when he described Nematopa- (Henderson, 1888)
gurus lewinsohni Tiirkay, the species Lew-
Catapagurus australis Henderson, 1888:76,
insohn (1969) had misidentified as A', mur-
icatus. Nonetheless, species in the South Nepmia.t8o,pfaigg.ur1u.s gardineri—Haig & Ball,
China Sea with spinulose palms similarly
1988:185 (not Nematopagurus gardineri
continued to be identified as A^. muricatus
Alcock, 1905a).
(e.g., Wang 1994; Rahayu 1996, 2000). Ne- Nematopagurus muricatus—Wang, 1983:
matopagurus australis, another regional
54 (list).—Rahayu, 1996:350 [not TV^-
species, and N. alcocki are morphologically
matopagurus muricatus (Henderson,
quite similar, although the dorsal surfaces
of the chelae of N. australis are usually 1896)]. —
Nematopagurus australis McLaughlin,
somewhat more spinose than those of A^.
1997:505.
alcocki. However, spination, like setation, is —
subject to considerable variation. The most Material examined. RA^K, NCOS Stn.
reliable character for separating the two 6106, 20°30'N, 112°00'E, 71 m, 3 Jul 1959,
taxa is the dorsomesial margin ofthe dactyl 1 9 si = 1.4 mm, lOCAS K50B; RA^S,
of each cheliped. This margin bears a row NCOS Stn. 6030, 21°30'N, 115°30'E, 115
of spines in A^. australis, but is unarmed in m, 14 Jul 1959, 1 d si = 1.8 mm, 1 ovig.
N. alcocki. 9 si = 2.9 mm, IOCAS SIII19B; R/VS,
Another of Alcock's (1905b) species that NCOS Stn. 6038, 21°00'N, 115°00'E, 100
is poorly known is A^. indicus. Until ques- m, 4 Apr 1959, 1 c? si = 2.1 mm, lOCAS
tionably identified from Indonesia by SIII18B; RA^S, NCOS Stn. 6038, 21°00'N,
=
McLaughlin (1997), A^. indicus had only 115°00'E, 103 m, 14 Jul 1959, 1 c? si
been reported correctly from the Indian 2.1 mm, 1 9 si = 2.2 mm, lOCAS S67B-9,
Ocean. Having reexamined Balss's (1912) 10; R/VS, NCOS Stn. 6048, 21°00'N,
ovigerous female specimen of A^. indicus 114°30'E, 85 m, 10 Jul 1959, 1 cj si = 2.0
from the Nicobar Islands (ZMB 16475) we mm, 1 9 si = 1.8 mm, IOCAS SIII8B;
have concluded that TV. alcocki also bears RA^S Stn, 6093, 19°30'N, 112°30'E, 219 m,
considerable similarity to TV. indicus. No 6 Apr 1960, 1 c? si = 2.7 mm, 1 ovig. 9
specimens referable to Alcock's (1905b) si = 3.3 mm, lOCAS K151B-15; RA^21,
taxon were found in the collections of the NCOS Stn. 6093, 19°30'N, 112°30'E, 219
Institute of Oceanology, thus it is probable m, 17 Feb 1959, 2 cJ si = 3.0, 3.1 mm,
that Wang & Huang's (1979), Wang's lOCAS 6-21; RA^lOl, NCOS Stn. 6141,
(1983, 1992, 1994), and Huang's (1989) re- 19°15'N, 111°00'E, 80 m, 10 Feb 1960, 1
ports ofTV. indicus should be referred to TV. c? si = 1.8 mm, lOCAS N120R-32;
alcocki. The reports by Miyake (1961) and R/VlOl, NCOS Stn. 6175, 17°45'N,
Miyake et al. (1962) ofTV. indicus were cor- 110°00'E, 105 m, 10 Mar 1960, 1 9 si =
rected by Miyake (1978) to TV. vallatus. 2.4 mm, lOCAS N114B-54; R/VlOl,
However, McLaughlin & Sandberg (1995) NCOS Stn. 6181, 18°10'N, 109°45'E, 65 m,
showed that Miyake's (1978, 1982) and Mi- 12 May 1960, 1 9 si = 2.4 mm, IOCAS
yake & Imafuku's (1980) specimens actu- N200B-63; R/VlOl, NCOS Stn. 6186,
ally were conspecific with the species 18°00'N, 109°30'E, 74 m, 11 Mar 1959, 1
McLaughlin (1992) attributed to TV. tricar- 9 si = 2.7 mm, lOCAS N148B-43;
inatus (as Ceratopagurus). Therefore, while R/VlOl, NCOS Stn. 6210, 18°15'N,
TV. vallatus does occur in the Japanese Bo- 108°24'E, 38 m, 7 Mar 1959, 2 9 si = 2.0,
nin Islands, the species occurring in the 2.6 mm, lOCAS N158B-6; RA^131, NCOS
South China Sea and southern Japan is TV. Stn. 6141, 19°15'N, 111°00'E, 79 m, 13 Jul
tricarinatus. 1959, 1 (? si = 3.8 mm, lOCAS N120B-
748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
32; R/V131, NCOS Stn. 6215, 20°30'N, and Fiji suggest a potentially broad distri-
=
108°30'E, 50 m, 19 Apr, 1959, 1 c? si bution for this species, the only other ten-
3.3 mm, lOCAS N428-22; RA^lVl, NCOS tative report is that of McLaughlin (2002)
Stn. 6141, 19°15'N, 1 11°00'E, 78 m, 22 Jan for a specimen collected in the Andaman
1959, 1 ovig. 9 si = 2.4 mm, lOCAS 15- Sea. Nematopagurus australis has proven
36; R/V171, NCOS Stn. 6144, 18°30'N, to be the most abundant species of the ge-
=
lirOO'E, 140 m, 27 Jan 1959, 1 c? si nus in the Institute of Oceanology collec-
3.0 mm, 1 9 si = 3.2 mm, USNM tions. As indicated above, it most frequent-
1003651; R/V171, NCOS Stn. 6166, ly was misidentified as A'', muricatus by res-
16°30'N, 110°30'E, 72 m, 20 Apr 1959, 1 ident scientists. Henderson's (1896) A^. mur-
6 si = 3.2 mm, lOCAS L568-62; RA^220, icatus was also initially described in
NCOS Stn. 6030, 2r30'N, 115°30'E, 115 Catapagurus, but was transferred to Ne-
m, 13 Apr 1959, 1 cJ si = 2.0 mm, 1 9 si matopagurus by Alcock (1905a). Although
= 2.0 mm, IOCAS 219B; R/V220, NCOS A^. muricatus was accurately diagnosed by
Stn. 6052, 21°30'N, 114°00'E, 48 m, 17 Alcock (1905a, 1905b), it has repeatedly
Mar 1952, 1 specimen, poor condition, lO- been misinterpreted (e.g., Laurie 1926,
CAS 15-49; R/V220, NCOS Stn. 6054, Thompson 1943, Lewinsohn 1969, Miyake
21°00'N, 114°00'E, 80 m, 10 Dec 1959, 1 1978, Wang 1983; Rahayu 1996, 2000).
9 si = 2.1 mm, lOCAS, S199B-26; Nematopagurus muricatus is distinguished
R/V220, NCOS Stn. 6054, 21°00'N, from all other species in the genus by the
114°00'E, 79 m, 11 Jan 1960, 1 d si = 2.6 very stout, blunt, tubular spines on the dor-
mm, lOCAS S242B; RA^228, NCOS Stn. sal surfaces of the chelae (cf. Tiirkay 1986,
6227, 17°30'N, 108°30'E, 97 m, 7 Nov fig. 9). No specimens actually identifiable
1959, 1 9 si = 2.4 mm, lOCAS Q75B-59; as A^. muricatus have been found in the
R/V307, NCOS Stn. 6168, 18°30'N, South China Sea.
110°15'E, 46 m, 4 Apr 1959, 1 ovig. 9 si Nematopagurus australis is most closely
= 2.9 mm, lOCAS Q25-10; R/V307, allied to A^. meiringae, a species described
NCOS Stn. 6173, 18°15'N, 110°00'E. 85 m, from the eastern coast of South Africa
29 Jan 1959, 1 c? si = 3.5 mm, lOCAS 13- (McLaughlin 1998), but during the present
32; RA^307, NCOS Stn. 6242, 18°30'N, study found in both the South and East Chi-
108°00'E, 72 m, 17 Sep 1959, 1 ovig. 9 si na Seas. No single diagnostic characterwill
= 2.3 mm, lOCAS Q18B-8; R/V307, readily separate the two taxa and intraspe-
NCOS Stn. 6243, 18°00'N, 108°00'E, 86 m, cific variations sometimes make differenti-
16 Apr 1959, 1 9 si = 2.7 mm, IOCAS ation difficult. Nematopagurus australis
Q56B-7; R/V394, NCOS Stn. 6154, most commonly has only a very short mat
19°15'N, 110°45'E, 38 m, 10 Jul 1959, 1 6 ofsetae on the dorsal surfaces ofthe chelas,
si = 3.0 mm, lOCAS R26B; RA^394, although longer setae frequently are present
NCOS Stn. 6219, 19°30'N, 108°30'E, 52 m, marginally; the dactyl of the right chela
25 Jan 1959, 1 ovig. 9 si = 3.9 mm, lO- lacks much surface setae. In contrast, a usu-
CAS 30-8; RA^394, NCOS Stn. 6226, ally dense, moderate to long covering of
17°45'N, 108°30'E, 79 m, 28 Jan 1959, 1 simple and plumose setae is present on the
c? si = 4.7 —mm, lOCAS 16-8. dorsal surfaces of the chelas of N. meirin-
Remarks. Henderson's (1888) N. oiis- gae, together with long plumose setae mai--
tralis initially was described as Catapagu- ginally; the dactyl of the right chela is sim-
rus australis, but was transferred to Ne- ilarly provided with a moderate covering of
matopagurus by McLaughlin (1997) after setae. The corneal diameter of N. australis
an examination of the two syntypes and, at varies from 0.33 to 0.48 of the peduncular
that time, the only known specimens. Al- length in the present material, whereas this
though the type localities ofthe Arafura Sea diameter in N. meiringae varies from 0.46
VOLUME 115. NUMBER 4 749
to 0.72 ofthe peduncular length. Thus, gen- 1978:129 (key).—Wang & Huang, 1979:
erally speaking, the ocular peduncles are 63.—Miyake & Imafuku, 1980:63.—
slightly longer and the corneas slightly less Wang, 1983:55 (list); 1992:61 (list);
dilated in A^. australis. The ambulatory dac- 1994:571 (Hst).——Baba, 1986:201, 303,
tyls, particularly the third left, is typically unnumbered fig. Huang, 1989:254, fig.
longer, more slender, and more strongly 215.—Komai, 1999:28.—Rahayu, 2000:
arched in N. australis than in N. meiringae. 394 (not Nematopagurus squamichelis
However, unless both species are available Alcock, 1905b). —
for comparison, the best means of separat- Nematopagurus lepidochirus Komai &
ing these two taxa is by the density of the McLaughlin, 2001:83, figs. 1-4.
—
setation of the chelas. Material examined. R/V2, NCOS Stn.
6094, 19°00'N, 112°30'E, 290 m, 6 Apr
Nematopagurus diadema Lewinsohn, 1969 1959, 1 (? si = 3.9 mm, 1 2 si = 6.0 mm,
lOCAS K150B; RA^21, NCOS Stn. 6094,
Nematopagurus diadema Lewinsohn, 1969:
74, fig. 13.—McLaughlin, 1998:325, 199°0si0'N=, 41.142°3m0m',E, l2O70CAmS, 77-F1e5b; 1R9A5^92,1,1
fig. 3. NCOS Stn. 6094, 19°00'N, 112°30'E, 270
—
Material examined. R/V171, NCOS m, 19 Apr 1959, 1 c? si = 5.5 mm, IOCAS
Stn. 6108, 19°30'N, 112°00'E, 120 m, 26 K32B-75; R/V171, NCOS Stn. 6109,
Apr 1959, 1 ovig. 9 si = 2.0 mm, lOCAS 19°112'N, 112°00'E, 194 m, 27 Jan 1959,
L58B; RA^171, NCOS Stn. 6120, 20°00'N, 1 9 si = 3.7 mm, lOCAS L59B; R/V1082,
111°30'E, 77 m, 26 Jan 1959, 1 c? si = 2.1 NCOS Stn. 6080, 19°30'N, 113°00'E, 210
mm, IOCAS—47B. m, 10 Feb 1960, 2 S s\ = 4.3, 5.1 mm, 2
Remarks. Nematopagurus diadema was 9 si = 3.0, —4.7 mm, lOCAS K135B.
described from seven males and two ovig- Remarks. Like others of Alcock's
erous females collected at a depth between (1905a, 1905b) species oi Nematopagurus,
62 and 80 m in the Red Sea (Lewinsohn N. squamichelis has been misinterpreted by
1969). One ovigerous female was subse- several authors; however, not all authors
quently collected in the Mozambique Chan- misinterpreted this taxon in the same way.
nel off the coast of eastern South Africa at The species identified in Japanese and Chi-
a depth between 120 and 150 m (Mc- nese waters as A^. squamichelis (e.g., Mi-
Laughlin 1998). The two specimens col- yake 1978, Wang & Huang 1979, Miyake
lected in the South China Sea are well with- & Imafuku 1980, Miyake 1982, Wang
in the depth range recorded for the species. 1983, Baba 1986, Huang 1989, Wang 1994,
Nspeemcaiteospianghuarvuisngdithaedecmhaeliispesdsucahrmaeudniwqiuteh Kshoomwaniby19K9o9,maiRa&haMycuLa2u0g0h0h)nh(a2s001b)eetno
ball-like tubercles and platelets (Lewinsohn actually be N. lepidochirus. Although de-
1969: fig. 13; McLaughlin 1998: fig. 3C) scribed initially in the genus Pagurus (as
that there can be no doubt as to its identity, Eupagurus), an examination of the type
despite the very major extension of its material proved conclusively that Doflein's
known range. (1902) species belonged in Nematopagurus.
It does not appear to be an abundant species
in the South China Sea.
Nematopagurus lepidochirus
(Doflein, 1902)
Nematopagurus meiringae
Eupagurus lepidochirus Doflein, 1902:623. McLaughlin, 1998
— —
Pagurus lepidochirus Miyake, 1978:81 Nematopagurus gardineri Kensley, 1969:
(key). — 163, fig. 6e—h (not Nematopagurus gar-
Nematopagurus squamichelis Miyake, dineri Alcock, 1905a).
750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
—
INematopagurus gardineri Wang, 1983: dactyls of A^. meiringae are provided with
54 (list); 1992:61 (list); 1994:571 (list) long stiff setae. It is most probable that
(not Nematopagurus gardineri Alcock, specimens reported by Wang (1983, 1992,
1905a). 1994) and Huang (1989) as N. gardineri are
Nematopagurus meiringae McLaughlin, actually A^. meiringae.
1998:332, fig. 6. As previously discussed, A^. australis and
— A^. meiringae are morphologically very sim-
Material examined. RA^2, NCOS Stn. ilar species. While their distributions over-
6108, 19°30'N, 112°00'E, 124 m, 4 Apr lap in the northern part of the South China
1960, 1 9 si = 2.6 mm, lOCAS K148B; Sea, only N. meiringae has been found in
R/V171, NCOS Stn. 6108, 19°30'N, the East China Sea. In contrast, onlyN. aus-
112°00'E, 120 m, 20 Apr 1959, 1 ovig. $ tralis is present in the southern part of the
si = 2.6 mm, lOCAS L58B; RA^lTl, South China Sea. Nematopagurus meirin-
NCOS Stn. 6120, 20°00'N, 1 1 1°30'E, 77 m gae also generally appears to inhabit areas
26 Jan 1959, 1 c? si = 2.9 mm, IOCAS of greater depth.
K148B; R/VV, Stn. lV-5, 28°30'N,
125°30'E, 125 m, 4 Jul 1976, 1 c? si = 2.7 Nematopagurus tricarinatus
mm, lOCAS V495B-60; RA^V, Stn. V-3, (Stimpson, 1858)
27°30'N, 124°00'E, 105 m, 4 Jul 1976, 1 6
si = 2.1 mm, 2 ovig. ? si = 2.1, 2.6 mm, Eupagurus tricarinatus Stimpson, 1858:
lOCAS V519B-135; R/VV, Stn. V-6, 251.—Ortmann, 1892:309.—
27°30'N, 125°30'E, 115 m, 4 Jul 1976, 1 S Ceratopagurus tricarinatus McLaughlin,
si = 3.0 mm, lOCAS V499B-15; RA^V, 1992:795, figs. 1, 2. —
Nematopagurus indicus Miyake, 1961:
Stn. V-6, 27°30'N, 125°30'E, 112 m, 5 Jul
1976, 2 S s\ = 2.0, 2.6 mm, 2 ovig. 5 si 12.—Miyake et al., 1962:126 (fist) (not
= 2.3, 3.5 mm, lOCAS V496B-16; R/VV, Nematopagurus indicus—Alcock, 1905b).
Stn. V-7, 27°30'N, 125°00'E, 131 m, 5 Jul Nematopagurus vallatus Miyake, 1978:
4.—
m19m7,6, 11 dovisgi.=?2.1simm=, 23.39 msim,= 2l.O0C-2A.S3 M13i1y,akfieg.&52;Ima1f9u8k2u:,1211,98p0i:.634,1,pif.ig.2, fig.
V498B-29; R/VV, Stn. VI-3, 26°30'N, 8.—Wang, 1992:61 (list); 1994:571 (list)
123°00'E, 98 m, 4 Jul 1976, 2 ovig. 9 si = [not Nematopagurus vallatus (Melin,
2.4, 3.0 mm,—lOCAS V494B-16. Nem19a3t9o)p].agurus tricarinatus—McLaughlin
Remarks. Like A^. muricatus, Alcock's &
Sandberg, 1995:579.
(1905a) N. gardineri has been frequently
—
misinterpreted (e.g., Kensley 1969, Miyake Material examined. R/VS, NCOS Stn.
&
1978, Haig Ball 1988). Having examined 6012, 22°30'N, 117°00'E, 48 m, 20 Jul
the holotype of Alcock's (1905a) taxon, we 1959, 2 (? si = 1.7, 2.1 mm, 1 ? si = 2.0
can confidently say that it is not included mm, 2 ovig. ? si = 2.0, 2.1 mm, lOCAS
in the collections of the Institute of Ocean- SIII33B; R/VS, NCOS Stn. 6021, 22°15'N,
ology. Nematopagurus gardineri is most 116°00'E, 51 m, 9 Jul 1959, 1 specimen,
readily identified by its moderately long oc- poor condition, IOCAS S1II23B; R/VS,
ular peduncles that lack any noticeable cor- NCOS Stn. 6059, 2r30'N, 1 13°45'E, 45 m,
neal dilation. McLaughlin (1998) compared 9 Dec 1959, 1 2 si = 1.7 mm, lOCAS
N. gardineri with N. meiringae, and was S138B; R/V2 NCOS Stn. 6063, 21°30'N,
able to readily separate the two taxa, not 113°30'E, 41 m, 24 Apr 1959, 8 d si =
only by the lack of corneal dilation in A^. 1.7-3.8 mm, 3 5 si = 1.8-2.4, 5 ovig. ?
gardineri, but also by the row of corneous si = 2.4-2.9 mm, lOCAS K42B; R/V121,
spines on the dorsal surface of each am- NCOS Stn. 6140, 19°30'N, 1 1 1°00'E. 39 m,
bulatory dactyl. The dorsal surfaces of the 29 Oct 1959, 1 ovig. 9 si = 3.8 mm, lO-