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A REDESCRIPTION OF ACANTHONYX PETIVERII H.

MILNE EDWARDS,
1834, AND DESIGNATION OF A NEOTYPE FOR ACANTHONYX SIMPLEX
DANA, 1852 (BRACHYURA, MAJIDAE)
BY
ESTEBAN J. M. EMPARANZA1,3 ), GUILLERMO L. GUZMN1 ) and PETER K. L. NG2 )
1 ) Museo del Mar, Departamento de Ciencias del Mar, Universidad Arturo Prat, Casilla 121,
Iquique, Chile
2 ) Department of Biological Sciences, National University of Singapore, Kent Ridge, Singapore
119260, Republic of Singapore

ABSTRACT
The identity of the problematic spider crab species, Acanthonyx simplex Dana, 1852, originally
described from Hawaii and never reported since, is resolved. It is here shown to be identical to
Acanthonyx petiverii H. Milne Edwards, 1834, a species that is widely distributed on both sides
of the American continent. The supposed type locality of Acanthonyx simplex is almost certainly
wrong. Acanthonyx petiverii, nevertheless, is a highly variable species and, as such, is redescribed
in detail. To ensure a stable taxonomy for these species and the genus, a neotype for A. simplex is
designated.

RESUMEN
La problemtica identidad de la especie de cangrejo araa Acanthonyx simplex Dana, 1852,
originalmente descrita para las costas de Hawai y nunca estudiada desde entonces, ha sido
determinada. Aqu se demuestra que es idntica a Acanthonyx petiverii Milne Edwards, 1834,
especie que se distribuye en ambas costas del continente Americano. La supuesta localidad tipo
de Acanthonyx simplex es, casi con certeza, un dato errneo. Acanthonyx petiverii, no obstante, es
una especie altamente variable morfolgicamente y, en consecuencia, se redescribe en detalle. Para
estabilizar la taxonoma, se designa un neotipo para la especie A. simplex.

INTRODUCTION

The majid genus Acanthonyx Latreille, 1828 currently contains 17 species


(table I), most of which occur in the Atlantic and Indian Ocean. In the Pacific
3 ) Current address: Billund Aquaculture Chile S.A., Bernardino 1057, Mdulo 13, Parque Industrial
San Andrs, Puerto Montt, Chile; e-mail: akronos@yahoo.com

Koninklijke Brill NV, Leiden, 2007


Also available online: www.brill.nl/cr

Crustaceana 80 (5): 533-543

534

E. J. M. EMPARANZA, G. L. GUZMN & P. K. L. NG

TABLE I
List of the known species of Acanthonyx
Current name

Original genus
assigned

Subjective synonyms

Acanthonyx consobrinus
A. Milne-Edwards, 1862
Acanthonyx dentatus
H. Milne Edwards, 1834
Acanthonyx depressifrons
Manning & Holthuis, 1981
Acanthonyx dissimulatus Coelho, 1993
Acanthonyx elongatus Miers, 187?
Acanthonyx euryseroche
Griffin & Tranter, 1986
Acanthonyx formosa
Wu, Yu & Ng, 1999
Acanthonyx inglei
Tirmizi & Kazmi, 1988
Acanthonyx limbatus
A. Milne-Edwards, 1862
Acanthonyx lunulatus (Risso, 1816)

Acanthonyx

Acanthonyx

Dehaanius acanthopus MacLeay, 1838

Acanthonyx

Acanthonyx
Acanthonyx
Acanthonyx

Acanthonyx

Acanthonyx

Acanthonyx

Maia

Maia glabra Latreille, 1836


Acanthonyx viridis Costa, 1838
Gonosoma viridis Costa, 1844
Acanthonyx brevifrons A. MilneEdwards, 1869

Acanthonyx minor
Manning & Holthuis, 1981
Acanthonyx nodulosa (Dana, 1852)
Acanthonyx petiverii
H. Milne Edwards, 1834

Acanthonyx

Acanthonyx quadridentatus
Krauss, 1843
Acanthonyx sanctaehelenae
Chace, 1966
Acanthonyx scutellatus MacLeay, 1838
Acanthonyx undulatus Barnard, 1947

Acanthonyx

Acanthonyx simplex Dana, 1852


(present study)
Peltinia scutiformis Dana, 1851
Acanthonyx emarginatus H. Milne
Edwards & Lucas, 1843
Acanthonyx debilis Dana, 1851
Acanthonyx concamerata Kinahan,
1857

Acanthonyx

Acanthonyx
Acanthonyx

Acanthonyx macleaii Krauss, 1843

Peltinia

Ocean, there are only three known species, viz., A. petiverii H. Milne Edwards,
1834, A. simplex Dana, 1852, and A. formosa Wu, Yu & Ng, 1999 (cf. Dana,
1852; Manning & Holthuis, 1981; Griffin & Tranter, 1986; Wu et al., 1999).
Acanthonyx simplex was described from one specimen, supposedly originating

ACANTHONYX PETIVERII AND A. SIMPLEX

535

from Hawaii (Dana, 1852) and has never been recorded since its description
(Griffin & Tranter, 1986; Wu et al., 1999). Acanthonyx petiverii, on the other
hand, has been widely recorded in the Pacific from Baja California, Mexico, to
Valparaso, Chile, including Revillagigedo and the Galpagos Islands; and in the
Atlantic, from Miami, Florida, to Ro de Janeiro, Brazil (Garth, 1957, 1958, 1992;
Chirichigno, 1970; Wilson, 1987; Hendrickx, 1992; Retamal, 1994). The recently
described A. formosa is thus far only known from Taiwan (Wu et al., 1999).
During a study being carried out on the shallow water crabs of northern Chile,
examination of specimens of A. petiverii collected from different intertidal and
shallow water zones revealed considerable morphological variability, as had been
observed earlier by Garth (1958) and Chace (1966) with regard to the presence
or absence of tufts of setae on the carapace and legs, and also in the colour of
the carapace, since that depends on the colour of the host algae, be these green,
brown, or red (Wilson & Baeza, 1984; Wilson, 1987). Despite the fact that this
crab is found on both sides of the American continent (Rathbun, 1910), there are no
obvious characters that can be used to separate the Pacific and Atlantic populations
(Garth, 1958; Hendrickx, 1992) and, as such, these must be regarded as conspecific
for the time being.
Our comparisons of A. petiverii with the poorly known A. simplex Dana, 1852,
originally described from Hawaii, led us to the realization that these two taxa are in
fact conspecific, and that the type locality of the latter species was almost certainly
a mistake. In order to stabilize the taxonomy of these species, a neotype from
Chile is here designated for A. simplex. The neotype is described and A. petiverii
is recharacterized on the basis of the present material.
The following abbreviations are used: MNHNCL, Museo Nacional de Historia
Natural, Santiago, Chile; cl, total carapace length.

TAXONOMY

Acanthonyx petiverii H. Milne Edwards, 1834


(fig.1)
Cancer muricatus compressum Petiver, 1712, pl. 20 fig. 8.
Acanthonyx petiverii H. Milne Edwards, 1834: 343. Bell, 1835: 173; 1836: 62. De Haan, 1839,
pl. G. White, 1847: 11. Dana, 1852: 128; 1855, pl. 5 fig. 6a-d. Smith, 1869: 33. Miers, 1877: 654;
1886: 42. A. Milne-Edwards, 1878, pl. 27 fig. 7, 7a-f. Cano, 1889: 99, 100, 176. Rathbun, 1894: 72;
1901: 60; 1907: 72; 1910: 534, 571, pl. 46 fig. 4; 1925: 142, pl. 44, pl. 222 figs. 1-6. Boone, 1927:
137, fig. 38. Finnegan, 1931: 620. Hult, 1938: 11. Garth, 1946: 376, pl. 63 fig. 4. Crane, 1947: 71.
Buitendijk, 1950: 271.
Acanthonyx emarginatus H. Milne Edwards & Lucas, 1843: 9, pl. 5 fig. 2; type locality, near
Lima, Peru; type in Paris Museum.
Acanthonyx debilis Dana, 1851: 272; 1852: 127, pl. 5 fig. 5a, b; type locality, Valparaso, Chile;
type not extant.

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ACANTHONYX PETIVERII AND A. SIMPLEX

537

Peltinia scutiformis Dana, 1851: 273; 1852: 130; 1855, pl. 5 fig. 7a-c; type locality, Ro de
Janeiro; type not extant. Smith, 1869: 33.
Acanthonyx simplex Dana, 1852: 126; pl. 5 fig. 4a-d; type locality, Sandwich Islands; type not
extant.
Acanthonyx concamerata Kinahan, 1857: 334, pl. 14 fig. 1; type locality, North Chinchas Islands,
Peru; type in Royal Dublin Society Museum.
Acanthonyx petiveri Stimpson, 1871: 97. A. Milne-Edwards, 1878: 143 and synonymy. Garth,
1957: 22; 1958: 223.
Pugettia scultiformis Miers, 1886: 40, footnote. Moreira, 1901: 65, 138; 1920: 126. Lenz &
Strunck, 1914: 276, pl. 12 figs. 5-7.

Material examined. Five males (8.2-18.4 mm cl), 6 females (9.3-15.2 mm cl)


(MNHNCL D-N 11553), intertidal and shallow water zones of Campus Huayquique, Universidad Arturo Prat, Iquique, Chile, coll. G. Guzmn & E. Emparanza,
June 1998. Neotype of Acanthonyx simplex Dana, 1852, female (ovigerous,
22.8 mm cl) (MNHNCL D-N 11552), collected in a tide pool among leaves of
Glossophora kunthii C. Agardh, intertidal zone of playa Corazones, Arica, Chile,
coll. E. Emparanza, February 2000.
Description of neotype female of Acanthonyx simplex Dana, 1852. Carapace
longitudinally ovate, dorsal surface smooth with 2 tubercles on protogastric region,
each with tufts of stout, simple setae. Rostral sinus narrow, U-shaped, tips rounded,
densely covered with stout simple setae. Two strong preocular lobes densely
covered with stout bristles. Dorsal surface of rostrum with a row of hooked setae
on each side, between preocular lobe and tip of rostrum. Eyestalk short, relatively
slender, cornea terminal in position.
Hepatic margin produced into small lobe, with broad base and rounded tip;
length about half width of base; inner margin almost straight, posterior margin
straight, anterior margin about half length of posterior margin, posterior margin
almost parallel to midline of carapace. Branchial margin with small, rounded,
outwardly directed lobes on posterior part, about a third of size of hepatic lobe,
with stout straight setae, both lobes subequal in size. Lateral carapace margin with
very low ridge. Gastric and cardiac regions weakly elevated. Pterystogomial region
smooth.
Third maxilliped smooth; basis and ischium separated by shallow suture along
inner part only, ischium subrectangular, with slight median depression, anterointernal angle not produced, inner margin smooth with fine short bristles; merus
Fig. 1. a-f, j, Acanthonyx simplex Dana, 1852, neotype female (MNHNCL D-N 11552); g-i,
Acanthonyx petiverii H. Milne Edwards, 1834 (MNHNCL D-N 11553). a, carapace, dorsal view;
b, left third maxilliped; c, left cheliped; d, right fifth walking leg; e, rostrum, dorsal view, right area
with setae omitted; f, rostrum, ventral view, right area with setae omitted; g, abdomen, male, adult;
h, carapace, dorsal view (pink form); i, abdomen, female, juvenile; j, abdomen, ventral view. Scale
bars: a, e, f, h = 5 mm; b, c, d, g, i, j = 2 mm.

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E. J. M. EMPARANZA, G. L. GUZMN & P. K. L. NG

with slight median depression, anteroexternal angle slightly produced to rounded


lobe; exopod stout, basal two-thirds rectangular, outer margin straight, parallel
with lateral margins of ischium, distal one-third almost triangular with lateral margin straight, tip not reaching upper edge of merus. Basal antennal article smooth,
proximal part with shallow depression; basal and central parts broad, distal part
triangular without anterolateral projection, tip with long bristles.
Chelipeds elongate, subequal. Chelae slender, smooth; finger about 0.6 times
length of palm, fingers almost straight, slightly gaping when closed, tips gently
curved inward; dactylus with 8 submolar processes along cutting edge, pollex with
9 submolar processes along cutting edge; inner surface with 2 tufts of simple setae.
Ambulatory legs smooth, similarly shaped, first leg longest, fourth leg shortest.
Merus smooth. Carpus with very low median longitudinal ridge on outer surface.
Propodus with triangular median lobe on ventral margin, distal margin lined
with long stiff setae, forming subchelate structure with dactylus. Dactylus gently
hooked, inner margin with 2 rows of stout spines, varying in number from 7 to 11
along each row (decreasing from first to fourth leg).
Anterior thoracic sternum smooth, all sternites fused, sutures visible.
Abdomen with surface smooth, almost circular, telson almost semi-circular;
somites 2-6 fused, immovable. Somite 6 enlarged, almost trapezoidal, sutures
visible, somites 4 and 5 enlarged, sutures not visible, somites 2 and 3 depressed,
sutures visible, somite 1 almost dumb-bell shaped.
Additional characters for males and juveniles of Acanthonyx petiverii. Males
agree with females in all carapace and ambulatory leg features. However, male
chelipeds elongate, subequal; chelae swollen, smooth; finger about 0.6 times
length of palm, medially gaping when fingers closed, tips gently curved inward,
dactylus with submolar processes, submolar processes also present on pollex,
variable in number; merus smooth; carpus without blunt ridge along outer-dorsal
surface; male abdomen triangular, enlarged; telson triangular, enlarged, movable,
lateral margins almost straight; somite 6 relatively enlarged, with proximal twothirds of lateral margins broadly concave, distal third with lateral margins broadly
convex; somites 4 and 5 fused, immovable, enlarged, sutures not visible; twothirds of distal lateral margins straight, proximal third broadly trapezoidal; somites
2 and 3 trapezoidal, nearly equal in shape and size, except shape of lateral
margins; lateral margins of somite 2 following convex curves of somite 3; somite
1 dumb-bell-shaped, lateral margins almost straight; male first pleopod with
thickened tip, with triangular lobe at opposite side, covered with short spinules.
In juvenile female, abdomen almost rectangular, telson broadly triangular, somites
2-6 fused, immovable. Somite 6 broadly trapezoidal, sutures visible, somites 4 and
5 enlarged, sutures not visible, entire segment squarish, somites 1-3 depressed,
sutures visible, somite 1 dumb-bell shaped.

ACANTHONYX PETIVERII AND A. SIMPLEX

539

Variation. The carapace showed considerable variation among the specimens


studied, depending mainly on the type of algae that the crabs were taken from.
Pink specimens, living on Corallina officinalis var. chilensis (Dec.) Kutz., have
two tubercles on the protogastric region. In addition, some specimens possess
two tubercles on the lateral ends of the intestinal region, having tufts of stout
simple setae, and have two or three tubercles on the inner margin of the merus
of the chelae. Moreover, these crabs display a mosaic pattern for the carapace,
legs, and chelipeds, resembling the shape and colour of the host alga, which has
grayish-brown lines. Green specimens, occuring on Ulva lactuca L., often have
two tubercles on the protogastric region, all with tufts of stout but simple setae.
Brown specimens, which live on Glossophora kunthii C. Agardh, often do not
have tubercles on the carapace, although some may have two tubercles on the
protogastric region with tufts of setae, as well as tufts of stout, simple setae on
the lateral branchial lobes.
Remarks. The identity of A. simplex Dana, 1852 has always been problematic. Originally described from the Sandwich Islands (= present-day Hawaii), the
species has never been reported since Dana (1852), and its identity rests solely
on the relatively brief description and simple figures of the species provided. No
species of Acanthonyx is known from Hawaii or nearby waters, despite the many
intensive surveys of the late C. Edmondson in the area, and the many recent surveys
in Hawaiian waters. As Acanthonyx species are shallow-water inhabitants and associated with macroalgae, their presence cannot have been missed, especially since
the macroalgae of Hawaii and nearby waters have been studied in substantial depth
over the years. In addition, numerous surveys of the shallow-water fauna to examine the diversity of native and the impact of invasive species in recent years, have
not uncovered this species (L. Eldredge, pers. comm. to PKLN). The third author
has also examined numerous lots of identified and unidentified shallow-water majids in the Bishop Museum, Honolulu, and no specimen of Acanthonyx has been
uncovered as yet.
On the basis of Danas (1852) description and figures, the close resemblance
between A. simplex and A. petiverii is notable. Acanthonyx simplex is almost
identical to the smoother brown form of A. petiverii examined here from Chile,
no other differences being apparent. Considering that four other species, Peltinia
scutiformis Dana, 1851, A. emarginatus H. Milne Edwards & Lucas, 1843, A.
debilis Dana, 1851, and A. concamerata Kinahan, 1857 (see Garth, 1957), have
previously been synonymized with A. petiverii, it is rather surprising that its
similarity with A. simplex Dana, 1852 has escaped the attention of carcinologists.
The only possible explanation is, that workers who have studied A. petiverii
(and synonyms) have utilized literature and specimens from its known range, and
have not considered A. simplex at all since this was described from the much

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E. J. M. EMPARANZA, G. L. GUZMN & P. K. L. NG

more northerly region of Hawaii. We believe that the supposed type locality of
A. simplex, i.e., Hawaii, is erroneous, and the result of a mistake in labelling.
Considering that the specimens available to Dana came from all over the Pacific,
this mistake is quite possible and not too surprising. This is especially true, since
Dana himself had material from Chile. In fact, one of the species now considered
a synonym of A. petiverii, viz., Acanthonyx debilis Dana, 1851, was actually
described from Valparaso, Chile.
In view of the taxonomic problems that have been caused by A. simplex Dana,
1852, we believe a neotype should be designated to reflect our contention that
it is in fact a junior synonym of A. petiverii H. Milne Edwards, 1834, and that
its supposed type locality is wrong. As we have material at hand from Chile that
matches Danas (1852) description and figures of A. simplex very well, and we are
certain they are conspecific, we believe that selecting one of these as a neotype for
the species will stabilize the taxonomy of the species and facilitate future studies
of A. petiverii and related congeners. To this effect, we hereby select a recently
collected ovigerous female specimen (22.8 mm cl) from Chile as the neotype of A.
simplex Dana, 1852. This specimen is deposited in MNHNCL.
Male A. petiverii specimens from the southeastern Pacific have two abdominal
segments clearly fused and immovable, as has also been documented by Chace
(1966) for A. sanctaehelenae Chace, 1966 and by Wu et al. (1999) for A. formosa.
In the A. petiverii specimens examined, the male abdominal somites 3 and 4
were not fused. In the specimens of A. lunulatus Risso, 1816 examined by Chace
(1966), he reported that there was not always [a] completely functional suture
between abdominal somites 3 and 4. As for A. formosa, Wu et al. (1999) noted that
somites 3 to 5 were always fused and immovable, although the sutures between
them were still distinct. Griffin & Tranter (1986) stated that all the Atlantic
species of Acanthonyx have the two abdominal segments clearly fused, but in
the four Indian Ocean species examined by these authors, the sutures were often
indistinct.
In his work on the decapod crustaceans of St. Helena Island (South Atlantic),
Chace (1966) included drawings of A. sanctaehelenae, A. lunulatus, and A.
petiverii. His drawings of A. petiverii specimens showed median tubercles on the
gastric and cardiac regions. No tubercles were observed in these carapace regions
for the specimens of A. petiverii examined in the present study, and this could be
just a feature of the A. petiverii specimens inhabiting Martinique and the Virgin
Islands, the localities where Chaces specimens were taken from. Whether the two
populations are actually conspecific will need to be re-examined before anything
else can be said.

ACANTHONYX PETIVERII AND A. SIMPLEX

541

ACKNOWLEDGEMENTS

E. Emparanza wants to express his sincere gratitude to Dr. J. C. von Vaupel


Klein for bringing him into contact with the third author, thus giving him the
opportunity to generate this communication; to Mr. Luis Hidalgo, librarian of
the Museo Nacional de Historia Natural, Santiago de Chile, for providing help
to review some antique books; to Miss Ana Guzmn for helping to deliver the
neotype to the Museum in Santiago; to Mr. Patricio Molina and Mr. Freddy
Garca for providing help in the field work and transport to the collecting sites
in northern Chile, and to Mr. Mario Alcayaga and Ms. Adrienne in Clacton-onSea, Colchester, U.K., for additional logistic support and encouragement. P. K. L.
Ng thanks Lu Eldredge for help with his work in Hawaii, which led to a better
understanding of the problems associated with Danas Hawaiian species.

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First received 12 August 2006.


Final version accepted 29 December 2006.