Table Of ContentPROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
114(1):109-119. 200L
Description of two new species of Polyonyx Stimpson, 1858 from the
Indo-West Pacific, with a key to the species of the Polyonyx sinensis
group (Crustacea: Decapoda: Porcellanidae)
Bemd Werding
Institut fiir Allgemeine und Spezielle Zoologie, Stephanstrasse 24, D-35390 Giessen, Germany
—
Abstract. Two new species of the genus Polyonyx Stimpson, 1858, P. tu-
learis and P. thai are described. The new species belong to the P. sinensis
group, which currently contains 11 species from the Indo-West Pacific, three
from the eastern Atlantic, and three from American waters. A key for all spe-
cies in this group is included.
The Polyonyx sinensis species group was P. sinensis group by Johnson (1958), P. si-
defined by Johnson (1958: 97) for taxa nensis Stimpson, 1858, P. utinomii Miyake,
characterized by him as follows: "Lateral 1943, P. pedalis Nobili, 1905, and P. trans-
margins of carapace without spines. Cara- versus (Haswell, 1882), are currently con-
pace broader than long, transversely ovate, sidered as valid. In addition, P. loimicola
or rectangular with rounded comers. Front SankoUi, 1965, P. maccullochi Haig, 1965,
rather narrow, trilobate with the median P. haigae McNeil, 1968, P. vermicola Ng
lobe broad and rounded andprojecting little & Sasekumar, 1993, and P. bella Hsueh &
beyond the lateral lobes, which are often Huang, 1998, have been described since
scarcely developed so that the front is al- then and adescribed to the P. sinensis group
most straight. Chelipeds with the anterior by the original authors. P. bella is consid-
margins of the carpus and merus unarmed; ered a junior synonym of P. sinensis (see
a more or less marked development ofhairs Discussion). The position of P. plumatus
on their outer, and often also on their inner Yang & Xu, 1994 was not discussed at all
surfaces. Legs hairy dorsally, and often the in the original description, it seems to be
carapace also more or less hairy. Dactyl of identical with P. haigae and is considered
the walking legs with the dorsal claw much a junior synonym of that species. P. com-
smaller than the ventral claw, and bearing etes Walker, 1887 which was included in
two or three accessory spinules." Most of the P. sinensis group by Johnson (1958)
the species are definitely known to live as was placed in a new genus by Ng & Na-
commensals of tube dwelling polychaetes. kasone (1993) but is considered a member
The species of the group are widespread in of the P. sinensis group in this paper since
the Indo-West Pacific where it is represent- it fits in all respects with Johnson's defini-
ed now by 11 species. Three additional spe- tion. The three Polyonyx species from west-
cies are known from the tropical easternAt- em Africa (Chace 1959) correspond with
lantic and three from American waters. the basic morphological features of the P.
When Johnson (1958) revised the Indo- sinensis group and have to be included
West Pacific species ofthe genus Polyonyx, here. The same is true for three species
the genus comprised 14 species, some of from the Americas which were recognized
which were assigned to other genera by lat- by Haig (1960) as clearly belonging to
er workers (Ng & Sasekumar 1993). Ofthe Johnson's P. sinensis group. The type spe-
Indo-West Pacific species attributed to the cies of the genus, P. gibbesi, is the only
110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
3mm
Fig. I. Polyonyx tulearis, new species. Male holotype, Tulear, Madagascar, USNM 296465: dorsal view of
body and chelipeds.
species from the western Atlantic and a basal part of antennae; surface smooth an-
common commensal in the tubes ofthe an- teriorly, slightly rugose with oblique striae
nelid Chaetopterus variopedatus (Renier). near postero-lateral margins; devoid of se-
Study of collections in the National Mu- tae except with scatteredplumose setaenear
seum of Natural History, Smithsonian In- lateral margins and with forward-directed
stitution, Washington D.C. (USNM), re- setae fringing frontal margin. Front broad,
vealed the presence oftwo undescribed spe- slightly trilobed in dorsal view, distinctly
cies of the P. sinensis group from the Indo trilobate in frontal view (Fig. 2A), median
West Pacific. The measurements givenrefer lobe bluntly rounded, extending somewhat
to maximum carapace length (CL) and car- beyond lateral lobes. Orbits well defined;
apace width (CW). eyes large, partly visible from above.
Basal antennal segment with transverse
Family Porcellanidae crest; ventral face partly bent upward, vis-
ible from above. Segments of peduncle
Polyonyx tulearis, new species
smooth.
Figs. 1, 2
— Third thoracic stemite slender, without
Material. Holotype: male, CL 4.1 mm, median lobe. Ischium of third maxilliped
CW 5.2 mm, Madagascar, Tulear, shore, RV (Fig. 2D) about as long as broad, with scat-
Anton Bruun, hand-collected, 11 Aug 1964, tered long setae on outer face.
USNM
296465. Paratype: ovigerous fe- Merus ofchelipeds (Fig. 1) with well de-
male, CL 5.4 rtmi, CW 7.3 mm, same data veloped, anteriorly crenulate rounded lobe,
USNM
as holotype, — 296466. laterally setose, with transverse, setose ru-
Description. Carapace (Fig. 1) sub- gae on upper surface; outer distal angle of
ovate, about 1.3 times as wide as long, smaller cheliped with several conical tuber-
broadest at epibranchial level; moderately cles. Carpus about 1.6 times as long as
convex longitudinally. Cervical and gastric wide, surface smooth except for some low
grooves distinct, epigastric elevations transverse striae near outer border; inner
prominent; anterolateral margin depressed shelf evenly convex throughout length,
at beginning of cervical groove to receive finely denticulate on proximal half of mar-
VOLUME 114, NUMBER 1 111
3mm -• A, B, C, D, E, H
^ G
F,
1mm
Fig. 2. Polyonyxtulearis, new species. Maleholotype,Tulear,Madagascar,USNM296465: A. front,anterior
view; B. left (greater) chela, dorsal view; C. right (smaller) chela, ventral view; D. third left maxilliped, outer
view; E. first left ambulatory leg, ventral view; F. propodus and dactylus ofsecondright ambulatory leg, dorsal
view; G. first left ambulatory leg, propodus and dactylus, dorsal view; H. telson, dorsal view.
gin, fringed with widely-set plumose setae. gaping near tip, movable finger with den-
Chelae (Figs. 1, 2B, C) swollen, smooth, ticulate ridge throughout length starting
except for faint longitudinal ridge starting from outer articulation. Cutting edge ofpol-
from dorsal articulation of movable finger; lex with strong tubercle-like tooth proxi-
external margin with tuberculate ridge vis- mally and smaller tooth distally; dactylus
ible only in lateral view, fringed with row with 2 large tooth, first near base and sec-
ofsimple setae. Fingers slightly curvedout- ond median; fingers of smaller cheliped
ward. Fingers of mayor cheliped somewhat closing completely, movable finger with ac-
112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
centuated longitudinal ridge formed by row ous female, CL 3.4 mm, CW 4, 9 mm, same
USNM
oftubercle-like teeth accompaniedby dense data as holotyp—e, 296464.
fringe of setae; cutting edge of both fingers Description. Carapace (Fig. 3) subrect-
denticulate. angular, 1.4-1.5 times wider than long,
Ambulatory legs (Figs. 2E-G) decreas- broadest at epibranchial level; moderately
ing in size from first to third pair, moder- convex from front to back; anterolateral
ately setose. Merus and carpus unarmed; margin somewhat depressed to receive bas-
propodus about 3 times as long as wide, al portions of antennae; carapace regions
armed on lower margin with 5-7 stout, faintly indicated, surface punctate, without
movable spinules including pair at distal setae except on posterolateral margins and
end, proximal spines sometimes paired. lateral walls. Front broad, sinuously trilo-
Dactylus hooked, with bifid tip and 2 strong bate in dorsal and frontal view (Fig. 4A),
conical spinules on lowermargin, distal one lobes obtuse, median lobe scarcely extend-
usually largest. ing beyond lateral lobes; frontal margin
Uropods and telson (Fig. 2H) fringed with row of upwardly directed short setae.
with long plumose setae; males with pair of Orbits shallow; eyes large, completely vis-
pleopods. ible from above.
—
Habitat. The only known specimens Basal segments of antennae triangular,
were collected by hand from the shore, but with transverse crest, partly visible from
no commensalism was reported. above; segments of peduncle cylindrical,
—
Etymology. The specific name is de- smooth, flagellum large, reaching nearly
rived from the type locality, Tulear, Mada- twice length ofcarapace, with short stiffse-
tae throughout length.
gascar. —
Remarks. Polyonyx tulearis is most Third thoracic stemite narrow, trilobate
anteriorly; median lobe broadly triangular,
similar to congeners that have a prominent
crested lobe on the merus of the chelipeds, lateral lobes with rounded tip. Ischium of
third maxilliped (Fig. 4C) about as long as
more than three terminal spines on the pro-
broad with scattered long setae on outer
podus of the walking legs, and lack spinu-
face.
lation on the inner side of all walking legs.
Chelipeds (Fig. 3) unequal, merus with
Polyonyx quadratus is distinct because of
well developed cristate lobe extending dis-
the tubercles on the outer side ofthe chelae.
tally beyond proximal border ofmerus, up-
The remaining species, P. maccullochi, P.
per surface with some rugae, lateralborders
senegalensis, P. vermicola, and the second
fringed with plumose setae. Carpus about
new species described here, P. thai, have
1.5 times longer than wide, upper surface
the fingers of the larger cheliped strongly
evenly rounded, smooth; inner border ta-
curved outward, a character that easily dis- pering proximally, broadening to large
tinguishes them from P. tulearis. rounded lobe fringed with line oflarge plu-
mose setae; external portion of carpus with
Polyonyx thai, new species scattered setae. Larger chela (Fig. 4F) swol-
Figs. 3, 4 len, surface smooth, thickly setose near ex-
— ternal margin; similar setation on internal
Material. Holotype: male, CL 2.7 mm, margin extending into gape of fingers. Fin-
CW
4.0 mm, Thailand; Chorburi Province, gers curved outward, gaping, dactylus con-
Bang Saen Beach, Gulf of Thailand, siderably shorter than pollex; cutting edge
13°20'N 100°55'E, 15 Jan 1959; exposed of pollex with large conical tooth proxi-
sand beach, commensal with polychaete mally, dactylus with large rounded tooth
worms (Chaetopteridae), leg. Dr. G. M. nearbase and a smallertuberculate one sub-
USNM
Moore, 296463. Paratype: oviger- medially. Chela of smaller cheliped (Figs.
VOLUME 114, NUMBER 1 113
3mm
Fig. 3. Polyonyx thai, new species. Male holotype. Bang Saen Beach, Chorburi Province, GulfofThailand,
USNM 296463: dorsal view ofbody and chelipeds.
4D, E) slender, fingers slightly curved out- male paratype bears a faint dorsal ridge
ward; palm with faint ridge starting from which joins the dorsal articulation of the
articulation ofmovable finger; outer border dactylus and which continues, more pro-
ofpalm and fingers covered with long, plu- nounced, on the dorsal margin of the dac-
mose setae, movable finger with long ele- tylus, and becomes tuberculate towards the
vated ridge along outer margin formed by tip ofthe finger. The three left walking legs
row of flattened rounded tubercles; fingers of the paratype female also have more
closing completely, cutting edges with fine spines on the lower margin ofthe propodus
denticulation. than the holotype, numbering eight, five
Ambulatory legs (Figs. 4G-I, K) de- and seven respectively in addition to the
creasing in size from first to third pair, three terminal spines, from first to third leg,
thickly setose. Merus and carpus unarmed; versus five, three and four respectively for
propodus 2 times as long as wide, armed the male ho—lotype.
on lower margin with three to five conical Habitat. The new species was collected
spines in addition to three terminal spines. from chaetopterid worm tubes togetherwith
Dactylus compact, on lowermarginwithbi- a series of P. sinensis (three males and two
fid curved tip and two strong conical spi- females). —
nules, distal one largest. Etymology. The species name is de-
Uropods and telson fringed with long rived from the country of the type locality,
plumose setae; first pair ofpleopods present Thailand, us—ed as a noun in apposition.
in males. — Remarks. Polyonyx thai has the fingers
Variations. The setation of the frontal of the larger cheliped strongly curved out-
margin ofthe female paratype is denser and ward, a character shared only with three
not confined in a row as in the male holo- other species of the group. From P. vermi-
type. In contrast to the smooth surface of cola, it is most easily distinguished by the
the larger chela of the male, that of the fe- form of the carpus of the larger cheliped.
114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
2mm -• A, D, E, F, G
1mm -t B, C, H, I, K
Fig. 4. Polyonyx thai, new species. Male holotype Bang Saen Beach, Chorburi Province, GulfofThailand:
A. front, anterior view; B. left lateral wall ofcarapace; C. third left maxilliped, outer view; D. right (smaller)
chela, ventral view; E. same, dorsal view; F. left (greater) chela, dorsal view; G. first right ambulatory leg; H.
same, detail of propodus and dactylus; I. second right ambulatory leg, detail of spinulation of propodus and
dactylus; K. third right ambulatory leg, detail ofspinulation ofpropodus and dactylus.
VOLUME 114, NUMBER 1 115
the inner margin being concave over the which was placed in the P. biunguiculatus
proximal half in that species, whereas P. group by Sankolli (1965).
thai has only a short concavity on the much As a result, the P. biunguiculatus group
shorter carpus of both chelipeds. The cut- now comprises just three species that are
ting edge of the pollex of the larger chela restricted to the Indo-West Pacific. All of
is entire in P. vermicola, whereas there is a those three species are free living, and
very prominent rounded tooth in P. thai. while they are sometimes associated with
Polyonyx senegalensis can be distinguished corals and sponges, they are never found in
by the convex frontal carpal lobe of the worm tubes, as are most of the remaining
chelipeds, and the absence of prominent Polyonyx species.
teeth on the cutting edges ofthe larger che- By far the largestPolyonyx species group
la. Polyonyx maccullochi differs in the sub- is the P. sinensis group. Johnson (1958)
ovate carapace which is only slightlybroad- considered only the species from the Indo-
er than long, the narrow front with a strong West Pacific, and subsequent workers (e.g.,
median lobe and the lack of a concavity on Sankolli 1965, Ng & Sasekumar 1993,
the proximal inner margin of the carpus of Hsueh & Huang 1998) never compared all
the chelipeds. The new species and the species in a worldwide context. For the
specimens of P. sinensis included in the eastern Pacific, Haig (1960) recognized P.
same sample represent the first finding of nitidus Lockington, 1878 and P. quadriun-
members of the P. sinensis group in Thai- gulatus Glassell, 1935. Haig (1960) consid-
land and increase the number of species re- ered the warm temperate outer Baja Cali-
ported for Thailand by Yang & Naiyanetr forniapopulation ofP. quadriungulatus dif-
ferent from the Gulf of California P. niti-
(1997) to 17.
dus. The range of P. nitidus was later
extended southward to Panama (Haig 1962)
Discussion and Isla Gorgona, Colombia (Werding &
Haig 1982). The discovery of P. quadriun-
When Johnson (1958) divided the genus gulatus by Kudenow & Haig (1974) in the
Polyonyx he characterized the P. biungui-
Gulf of California led those authors to ad-
culatus group as ". the central andlargest
. . mit that both taxa might be conspecific.
group of the genus" and assigned P. biun- Comparison ofthe basic characters used by
guiculatus (Dana, 1852), P. obesulus Miers, Haig (1960) to separate the two species
1884, P. parvidens NobiH, 1905, P. triun- (spinulation of the propodi and dactyli of
guiculatus Zehntner, 1894 and, with some the walking legs) indicate that these char-
doubts, P. hendersoni Southwell, 1909 to acters are not constant. Therefore, P. quad-
that group. Later, Haig (1979) included P. riungulatus is considered ajunior synonym
parvidens in the synonymy of P. obesulus. ofP. nitidus. Additionally, a second species
Actually, the differences discussedby John- P. confinis Haig, 1960 is present in the east-
son (1958), and some other morphological em Pacific. This species has been found
features which can be seen in the descrip- only once, and an association with a host
tion of P. hendersoni by Tirmizi et al. species was not observed. The description
(1989), and additional ones which were re- of P. bella by Hsueh & Huang (1998) and
vealed in a recent study ofHendersons ma- a pair of paratypes from the USNM was
terial from the London Museum set this compared with a series of P. sinensis from
species apart from typical Polyonyx. Poly- Thailand, found together with the type se-
onyx hendersoni does not fit at all with any ries of P. thai, new species, in the USNM.
group of Polyonyx and will be assigned to Polyonyx bella is not distinguishable from
a separate genus in a forthcoming paper, to- P. sinensis, and is therefore considered a
gether with P. splendidus Sankolli, 1963 junior synonym of that species.
116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Polyonyx cometes is here considered a the respective value given by Ng & Naka-
member of the P. sinensis group since the sone for P. cometes is "about 1 : 1.8". Fi-
characters revealed by Ng & Nakasone nally, the insertion ofthe telson setae ofthe
(1993) for the genus Eulenaios Ng & Na- first zoea as appears in Fig. 5, I in the paper
kasone, 1993, do not withstand a critical re- ofNg & Nakasone (1993) shows apparently
view on a worldwide level. The authors an incompletely developed telson as it is
consider the setal pattern as perhaps the typically present in prezoeae. On the other
"most distinctive feature of the species". hand, the P. sinensis group including P.
However, the setation is a character which cometes exhibit some key characters which
exhibits a wide variation in the different make advisable to maintain the group as a
species, and seems to be inadequate for the systematic category. The common features
establishment of a separate genus. The are in the unique form ofthe chelipeds and
same is true for the shape of the carapace the dactyls of the walking legs. An even
and the form of the front and the carapace more significant feature may be seen in the
regions. The ridge of granules observed on adaptation of living in tubes of annelid
the dactylus of the smaller cheliped is used worms which is developed only in the
by Ng & Nakasone (1993) as an additional members of the P. sinensis group.
character defining Eulenaios. But similar
ridges are present in P. senegalensis and in Key to species of the Polyonyx sinensis
both new species described herein. Further, group
the authors mention the membranous artic-
ulation between cheliped carpus, merus and 1. Walking legs with merus unarmed ven-
trally 2
propodus as "seeming more extensive than - At least third pair of walking legs
in other species". This obviously quantita- armed with spines ventrally on merus
tive feature appears in a very similardegree 14
in P. gibbesi where it was checkedforcom- 2. Frontal and lateral parts of carapace
parison. Another argument is the size ofthe andchelipeds densely coatedwith setae
species which is described as "larger than which obscure the outline of the struc-
any other known Polyonyx species". But ture (Singapore)
the largest P. cometes dealt with is 11.2 X Polyonyx cometes Walker, 1887
8.2 mm. The holotype of P. quadriungu- - Setation of carapace and chelipeds dif-
latus (= P. nitidus) measures 13.5 X 9.1 ferent 3
mm 10X7 mm 3. Propodus of walking legs without
and that ofP. nitidus (Haig
spines except for terminal triplet .... 4
1960), and Williams (1984) indicates for P. - Propodus of walking legs with at least
gibbesi a maximum breadth of 16 mm. For one spine in addition to terminal triplet
the australian P. transversus, Baker (1905) 6
and Haig (1965) record a size of 11 X 8 4. Frontal lobe acute in frontal view, ex-
mm. As a distinguishing larval property Ng tending well beyond obtuse lateral an-
& Nakasone (1993) mention the form ofthe gles (western Africa)
telson ofzoea I "being more proportionally P. bouvieri Saint-Joseph, 1900
elongate than in any other known species". - Frontal lobe scarcely produced, not ex-
Probably, the authors were not aware ofthe tending beyond lateral angles 5
larval descriptions given by Shenoy & San- 5. Carapace margins with thickly matted
setae; meral lobe of chelipeds slightly
kolli (1973) forP. loimicola and those from
produced (western coast of India) ....
Shepherd (1979) for P. transversus. As all P. loimicola Sankolli, 1965
other known Polyonyx larvae, the two spe- - Carapace margins not setose; meral
cies exhibit an elongate telson in Zoea I. lobe of chelipeds largely produced (Ja-
Shepherd records a broads: length ration of pan) P. utinomii Miyake, 1943
1 : 1.75 for the larvae of P. transversus and 6. Outer surface of chelae with scattered
VOLUME 114, NUMBER 1 117
tubercles (western Africa) ting edge (peninsular Malaysia)
. . .
P. quadratus Chace, 1959 .... P. vermicola Ng & Sasekumar, 1993
- Outer surface of chelae without tuber- 14. Proximal angle of carpal lobe of cheli-
cles 7 peds obtuse (Indian Ocean)
7. Dorsal surface of carapace with scat- P. pedalis Nobili, 1905
tered long setae; Propodus of walking - Proximal angle of carpal lobe of cheli-
legs with numerous (>60) minute spi- peds strongly pronounced, subrectan-
nules at lower margin (western Pacific) gular (American species) 15
P. haigae McNeil, 15. Longitudinal crest on the manus—of the
1968 (Syn. P. plumatus Yang & Xu. 1994) mayor cheliped (eastern Pacific Nic-
- Dorsal surface of carapace not setose; aragua) P. confinis Haig, 1960
propodus of walking legs without nu- - Manus of chelipeds without longitudi-
merous minute spinules (<20) at lower nal crest 16
margin 8 16. Propodus ofwalking legs without mov-
able spinules additional to the poster-
8. Irmer margin of carpus of both cheli-
peds evenly convex 9 odistal triplet, manus ofminor cheliped
- Innermargin ofat least mayor cheliped markedly outcurved (western Atlantic)
P. gibbesi Haig, 1956
convcave proximally, convex distally
- Propodus of walking legs with one or
12
two movable spinules additional to the
9. Propodus of walking legs with more posterodistal triplet, manus of minor
than 12 spinules on lowermargin (Aus-
cheliped not markedly outcurved (east-
tralia) .... P. transversus (Haswell, 1882) em Pacific) P. nitidus Lockington, 1878
- Propodus ofwalking legs with lessthan . .
10 spinules on lower margin 10
Acknowledgments
10. Chelipeds slender, fingers of both che-
lae straight (Madagascar) I am very grateful to R. Lemaitre for his
P. tulearis, new species help and orientation during my visit to the
- Chelipeds stout, fingers of larger chela National Museum of Natural History,
bent outward distally 11 Washington, D.C., and to him, P. K. L. Ng
11. Front narrow, forming a prominent
and anonymous reviewers for the revision
acute median lobe; proximal inner an-
of the manuscript. Peter K. L. Ng also pro-
gle ofcarpus ofchelipeds softly round-
ed (Australia) P. maccullochi Haig, 1965 vided me with some literature that I was not
- Front broadly obtuse, median lobe aware of. To the Smithsonian Institution's
blunt; proximal innerangle ofcarpus of Office of Fellowships and Grants, I ac-
chelipeds forming a roimded nearly knowledge support in the form of a short-
rectangular angle (western Africa) . . . term visitor grant. I am grateful to Mrs.
P. senegalensis Chace, 1959 Helga Schmitt for kindly inking the draw-
12. Inner border of carpus of chelipeds ta- ings of the type specimens.
pering proximally forming a deep short
concavity; carpus of larger cheliped Literature Cited
about 1.5 times broaderthan long (Gulf
of Thailand) P. thai new species Baker,H.W. 1905. Noteson SouthAustraliandecapod
—
- Inner border of carpus of larger cheli- Crustaceans. PartIII. TransactionsoftheRoy-
ped concave over the proximal half; al Society of South Australia 29:2—52-269.
carpus oflarger cheliped about 2 times Chace, E A. 1959. Porcellanid crabs. Expedition
broader than long 13 Oceanographique Beige dans les Eaux Cotieres
Africaines de I'Atlantique Sud (1948-1949),
13. Pollex of larger cheliped with large
Resultats Scientifiques III (5):1-45.
blunt subproximal tooth on cutting edge
Dana, J. D. 1852. United States Exploring Expedition
(1w8e5s8te(rSnynP.aciPf.icb)el.l.a.H.sPu.esi&nenHsuiasnSgt,im1p9s9o8n), duunrdienrgthtehecyoemamrsan1d83o8,fC1h8a3r9l,es18W4i0l,ke1s8.4—1,U.1S8.4N2.,
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——
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