RAYMOND B. MANNING A Review of the
Genus Harpiosquilla
r (Crustacea, Stomatopoda),
r with Descriptions of
Three Mew Species
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? 1969 NUMBER 36
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SMITHSONIAN CONTRIBUTIONS TO
ZOOLOGY
NUMBER 36
Raymond B. Manning A R e v i e w of t h e
Genus Harpiosquilla
(Crustacea, Stomatopoda),
with Descriptions of
Three New Species
SMITHSONIAN INSTITUTION PRESS
CITY OF WASHINGTON
1969
ABSTRACT
Manning, Raymond B. A Review of the Genus Harpiosquilla (Crustacea, Stomato-
poda), with Descriptions of Three New Species. Smithsonian Contributions to
Zoology, 36:1-41. 1969.?Available specimens of the large squillid Harpiosquilla
from the collections of the Australian Museum, Sydney, and the Division of Crustacea,
National Museum of Natural History, Smithsonian Institution, show that the genus
comprises seven species; three of these species are newly described. The species of
Harpiosquilla occur throughout the Indo-West Pacific region, from Japan and
Australia westward to the Red Sea and South Africa; some species, particularly
H. annandalei, H. harpax, and H. raphidea are widely distributed in the region
whereas others, including H. indica, new species (India), H. japonica, new species
(Japan), and H. stephensoni, new species (Australia), exhibit more limited distribution
patterns. All available literature is summarized, and the descriptive accounts are
accompanied by notes on biology, development, and distribution.
Official publication date is handstamped in a limited number of initial copies and is recorded
in the Institution's annual report, Smithsonian Year
UNITED STATES GOVERNMENT PRINTING OFFICE
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Contents
Page
Introduction 1
Acknowledgments 1
Systematic Account 1
Genus Harpiosquilla Holthuis, 1964 1
Key to Species of Harpiosquilla 5
1. Harpiosquilla annandalei (Kemp, 1911) 6
2. Harpiosquilla raphidea (Fabricius, 1798) 9
3. Harpiosquilla japonica, new species 15
4. Harpiosquilla stephensoni, new species 17
5. Harpiosquilla melanoura Manning, 1968 21
6. Harpiosquilla harpax (de Haan, 1844) 25
7. Harpiosquilla indica, new species 33
General Remarks 36
Bibliography 37

Raymond B. Manning A R e v i e w o f t h e
Genus Harpiosquilla
(Crustacea, Stomatopoda),
with Descriptions of
Three New Species
Introduction
Examination of two species of Harpiosquilla for a re-
port on the stomatopods of Madagascar (Manning,
1968b) revealed that a single specimen of Harpio-
squilla from Australia identified by me in 1966 as H.
harpax was actually an undescribed species. Through
the kindness of John C. Yaldwyn, formerly of the
Australian Museum, I was able to examine all ma-
terial of Harpiosquilla in the collection of that institu-
tion. The series included H. harpax, H. melanoura,
which was described from Madagascar in 1968, and
an undescribed species, named H. stephensoni below.
Although Australian specimens of both H. harpax and
H. raphidea had been recorded in the collections of the
Australian Museum, the latter species apparently does
not occur in Australian waters. The surprising absence
of H. raphidea, supposedly the most widely distributed
species of the genus, in Australian waters, led me to
examine the series of Harpiosquilla in the Division of
Crustacea, National Museum of Natural History,
Smithsonian Institution (SI Crust.) and the collec-
tions made by the International Indian Ocean Ex-
pedition (HOE). Two other undescribed species were
found in these collections. The present review of the
species of Harpiosquilla results from the study of these
three collections.
Raymond B. Manning, Chairman, Department of Inverte-
brate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.
Terms and measurements used herein have been ex-
plained in detail in earlier papers (Manning, 1966,
1968b).
Acknowledgments
I am indebted to John Yaldwyn for the loan of the
extensive series of Harpiosquilla from the Australian
Museum (AM) and for promoting the exchange of
several specimens from that series with the Division
of Crustacea, Smithsonian Institution. L. B. Holthuis,
Rijksmuseum van Natuurlijke Historie, Leiden
(RMNH), made available the lectotype of Harpio-
squilla harpax, without which the identity of several
of the species recognized below would have remained
in doubt. The line drawings were made by my wife
Lilly with the support of the Office of Oceanography
and Limnology, Smithsonian Institution. I also thank
Fenner A. Chace, Jr. and David L. Pawson for their
comments on the manuscript.
Systematic Account
Genus Harpiosquilla Holthuis, 1964
Alimerichthus Claus, 1871 p. 147 [p. 39 on separate; a junior
homonym of Alimerichthus Guerin-M6neville, 1855].?
Holthuis and Manning, 1964, pp. 138, 143.?Evans and
China, 1966, p. 205 [luted] [name no. 1822 on Official
Index of Rejected and Invalid Generic Names in
Zoology].
Harpiosquilla Holthuis, 1964, p. 140.?Manning, 1968a, p.
121.
1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DIAGNOSIS.?Eye large, T-shaped, cornea bilobed
and much broader than stalk; ocular scales separate;
carapace usually with median, intermediate, lateral
and reflected marginal carinae; median carina not bi-
furcate at either end; posterolateral margins of
carapace with deep excavations, anterior margin of
each excavation angled; mandibular palp present; 5
epipods present; dactylus of raptorial claw with teeth,
outer margin of dactylus usually with prominent
angled or rounded lobe in adult males; opposable mar-
gin of propodus of raptorial claw with spaced erect
spines rather than closely set pectinations; lateral proc-
ess of fifth thoracic somite usually an inconspicuous
lobe, fifth somite usually also with ventral pair of
spines; lateral processes of sixth and seventh thoracic
somites not strongly bilobed, usually acute and sharp
posterolaterally; abdomen broad, with 8 pairs of
carinae on first 5 somites, 6 on the sixth, submedians
usually low, absent in some species; telson without sup-
plementary dorsal ornamentation other than median
carina and carinae of marginal teeth; 3 pairs of mar-
ginal teeth present on telson, submedians with fixed
apices, prelateral lobes present; inner spine of basal
prolongation of uropod the longer, inner margin
crenulate or serrate.
TYPE-SPECIES.?Squilla harpax de Haan, 1844, by
original designation. The gender is feminine.
NUMBER OF SPECIES.?Seven, of which three are
described as new herein.
RANGE.?Indo-West Pacific region, from Japan and
Australia in the Pacific to the Red Sea and South
Africa. Also see General Remarks (p. 36).
REMARKS.?Until 1952, only two of the species cur-
rently assigned to Harpiosquilla, H. annandalei and H.
raphidea, were recognized by students of the group.
In that year K. K. Tiwari and S. Biswas, studying the
collections of the Zoological Survey of India, pointed
out that raphidea comprised two species, raphidea
sensu stricto and harpax, a species described from
Japan by W. de Haan in 1844. They gave the diagnos-
tic features of the two species and included a list of
localities based on materials in the Zoological Survey.
Another species, H. melanoura, was described by me
in 1968(b), and three additional new species are de-
scribed below. The genus now comprises seven species.
Most accounts in the literature are too brief to allow
us to determine which of the seven species might have
been reported by earlier workers as Squilla raphidea.
Where possible, however, I have tried to identify rec-
ords in the literature and assign them to one of the
species recorded here. Sometimes the accounts give
information, either in the text or in the figures, which
makes this possible; in other cases I have been able to
examine specimens previously recorded, as in the col-
lections of the Australian Museum which proved to
include three species, all incorrectly identified as H.
raphidea; and in some cases, notably records of H.
raphidea from China and Japan, corrections can be
made on the basis of geography. Harpiosquilla raph-
idea does not occur in the Red Sea and is not known to
occur north of the Philippine Islands in the Pacific
Ocean, so records from those areas must be referable
to other species. Under the accounts of both H. harpax
and H. raphidea I have included a section on "Synon-
ymy" to explain my identification of the old records.
Early records which have not been verified by re-
examination of the material must be accepted with
caution.
Harpiosquilla includes the largest known stomato-
pods, the largest species being H. raphidea (Fabricius),
which attains a total length of 335 mm. Their large
size and the deep posterolateral excavations on the
margins of the carapace make adult members of the
genus very easy to recognize. Another characteristic
of species of Harpiosquilla is the row of spaced, erect
spines on the opposable margin of the propodus of the
claw. These spines apparently replace the closely set
pectinations found on the propodus of all other mem-
bers of the family Squillidae. Similar spines are also
found on the propodus of the two species of Bathy-
squilla, the only genus in the Bathysquillidae. Bathy-
squilla and Harpiosquilla, however, were derived from
separate stocks and apparently have developed the re-
markable armature of the claw independently. The
function of the spines on the propodus is unknown;
they may aid in holding prey. Their number and ar-
rangement may have some value as a specific character
for H. raphidea, for in that species, as pointed out by
Tirmizi and Manning (1968), there is usually no more
than one smaller spinule or denticle between the major
erect spines. In the other species there are usually
several smaller spinules and denticles between the
major spines.
Adults of Harpiosquilla show no apparent close re-
lationship to any other genus in the family Squillidae;
the genus occupies an isolated position in the family.
On the basis of his studies on the larval development
of stomatopods, K. H. Alikunhi (1952, p. 313) made
NUMBER 3 6
the following observations: "The larvae of S.[quilla]
raphidea occupy a peculiar position. Though described
as belonging to the Alimerichthus group, they show
some conspicuous differences from the typical Alimer-
ichthus of the 'Chloridella' group, viz.; larger size;
broad shield-like carapace and absence of free teeth
other than the terminal on the dactylus. In the num-
ber of marginal spinules on the carapace (4 + 3) these
larvae perhaps occupy an intermediate position be-
tween Alimerichthii of the 'Chloridella' and Scorpio
groups on the one hand and the typical Alima larvae
on the other." Since that was written, each of these
groups referred to above have been recognized as dis-
tinct genera. Thus Harpiosquilla, on the basis of
larvae, may occupy a position intermediate between
(a) Clorida and Cloridopsis, and other small-eyed
genera, and (b) Oratosquilla and its allies. Its precise
relationship to any of these genera cannot be deter-
mined with available information.
There is no fossil record of the genus.
That H. harpax is generically distinct from Squilla
was recognized perhaps as long as 100 years ago by
the zoologist William Stimpson; he used the generic
name Prolepta on the label of the specimen collected
by the North Pacific Exploring Expedition. As far as
I can determine, that name has not been published.
Inasmuch as we know relatively little about the
biology of stomatopods in general, we are fortunate in
having some basic information available on the early
ontogeny and postlarval development of Harpio-
squilla, primarily as a result of the work of K. H.
Alikunhi and K. B. Nair in India. Possibly this is due
to the easily recognizable facies of Harpiosquilla
adults. It is ironic that we cannot determine with cer-
tainty which of the species of Harpiosquilla they
studied. Nair (1941) reported on extrusion of eggs and
embryological development in a Harpiosquilla from
Madras, and Alikunhi, in a long series of studies on
larvae culminating in his extensive work on postlarval
development of stomatopods published in 1967, exam-
ined material from Madras and other localities. Both
H. harpax and H. raphidea could occur there, so I
have here referred their papers to whichever name was
used in their works.
The characteristic larvae of Harpiosquilla, first re-
ported by Claus (1871) as Alimerichthus, are dis-
cussed in some detail by Alikunhi (1952, 1959). They
are easily recognized by the broad carapace with one
or two strong, laterally directed lateral spines. Ali-
kunhi (1967) has also given a very detailed account
of growth changes in the postlarvae. He was able to
hold one specimen in the laboratory for 282 days; the
specimen underwent 14 molts.
All of the descriptions given here are based on sub-
adults or adults. I have seen no very young specimens
of any species and can only predict some possible dif-
ferences in major morphological features between ju-
veniles and adults; Alikunhi (1967) has noted some
characteristics of juveniles. The eyes of juveniles are
undoubtedly larger than those of adults; within the
size range of the specimens examined the corneal in-
dices of smaller specimens are always smaller than those
of larger specimens (Table 1) ; this is the normal pat-
tern found also in other squillids. The body carinae of
young specimens are not so well developed as in adults;
this could cause a problem in recognizing small speci-
mens of H. harpax, for example, in which the subme-
dian carinae of the abdomen of adults are at best low
and poorly developed. In H. stephensoni, new species,
the lateral process of the fifth thoracic somite of the
smallest specimens examined is not as markedly angled
nor does it project as far laterally as in the largest speci-
mens; thus the lateral spine on that somite, character-
istic of both H. annandalei and H. raphidea, may be
but a blunt lobe in juveniles.
Alikunhi (1967, p. 903) made the following ob-
servations on postlarvae and early juveniles of
Harpiosquilla:
. . . the early post-larva is characterised by the absence of
anterolateral spines on the carapace and the presence of
terminally articulated submedian spines on the telson. Though
rudimentary spines appear on the anterolateral corners of
the carapace during the second or third post-larval stage, the
terminal articulation of the submedian spines of the telson
continues to be present even after the ninth post-larval moult.
In the species of the nepa group the terminal articulation
of the submedian spines disappears with the first post-lai.al
moult; i.e., it is present only during the first post-larval stage.
In this feature also H. raphidea differs from S. nepa or S.
woodmasoni but resembles S. latreiUei (Chloridella group).
Adults are easily recognized by the presence of an
inflated median carina on the telson; often the sub-
median carinae of the sixth abdominal somite are also
inflated in adults of both sexes. Adult males of most
species may be distinguished by the presence of a prom-
inent angled or rounded projecting lobe on the outer
margin of the dactylus of the raptorial claw; the bases
of some of the teeth of the claw may also be enlarged.
The chelae and telsons of both sexes have been figured
where possible.
The color pattern of preserved specimens is a good
specific character, particularly the coloration of the
uropod. In H. melanoma the distal segment of the uro-
podal exopod is entirely black. In H. annandalei the
same segment is black with a white midrib and this
species also has distinctive paired submedian spots on
the telson. In H. harpax, H. indica, new species and H.
stephensoni, new species, only the inner half of the
distal segment of the uropodal exopod is dark, but
there are other less distinctive differences in color pat-
tern. I have not been able to determine with certainty
the color pattern of the uropod in either H. japonica,
new species, or H. raphidea.
The armature of the abdominal carinae appears to
be a very good specific character. In H. annandalei, for
example, the submedian carinae of both the fifth and
sixth abdominal somites are armed posteriorly; in all
other species only the submedians of the sixth somite
are provided with spines. Similarly, in H. raphidea and
H. annandalei the intermediate carinae of all six ab-
dominal somites are spined posteriorly, but in H. indica
and H. japonica, the intermediate carinae of the first
and second somites are unarmed and different patterns
of abdominal armature are present in the other three
species. In general, the number of armed carinae in-
creases with age.
The relative distinctness of the submedian carinae
of the last three thoracic and first five abdominal
somites also may be of help in the recognition of species.
In H. raphidea these carinae are strongly developed,
more so than in any of the other species. The sub-
median carinae of the body in both H. japonica and
H. stephensoni are not so well developed, being lower
and less well defined, but they are still readily dis-
tinguishable. In H. harpax the submedian carinae are
very low, poorly defined, and often can be detected
only when the dorsal surface is dried. The submedian
carinae are completely absent in both H. indica and
H. melanoura; the latter species is also the only one
which lacks a median carina on the carapace.
The marginal denticles of the telson are sharper and
better defined in H. annandalei, H. harpax, H. indica,
new species, and H. melanoura than in the remainder
of the species.
The eyes of each species are of different sizes and
these differences are reflected in the corneal indices,
summarized for all species in Table 1. Because of the
change in the index with size I have separated the
indices in the table by arbitrary subgroups based on
rarapace length. The overall range given at the bottom
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Corneal Indices of species of Harpiosquilla
Carapace
length
in (mm)
11-15
16-20
21-25
26-30
31-35
36-40
41-45
46-50
51-55
^6-60
Overall
range
H. annandalei
Range
280-291
260-287
-
-
-
-
-
-
-
240-291
No.
1
3
6
-
-
-
-
-
-
-
?
Mean
262
285
278
-
-
-
-
-
-
-
?
H. raphidea
Range No.
-
-
-
_
-
1
1
1
475-504 3
380-530 -
Mean
-
-
-
-
-
410
486
469
489
?
Carapace
length
in (mm)
11-15
16-20
21-25
26-30
31-35
36-40
41-45
46-50
51-55
56-60
Overall
range
H. japonica
Range No. Mean
_ _ _
_ _ _
1 328
325-337 2 331
_
_ _
_
_
_
325-337 -
H. stephensoni
Range
_
-
-
-
349-392
-
396-420
-
410-425
336-434
No.
_
-
1
3
1
2
1
2
?
Mean
_
-
-
336
364
366
408
434
418
?
Carapace
length
in (mm)
11-15
16-20
21-25
26-30
31-35
36-40
41-45
46-50
51-55
56-60
Overall
range
H. melanoura
Range No.
1
237-246 3
1
1
-
-
-
_ _
- _
229-271 -
Mean
229
240
252
271
-
-
-
_
_
?
H. harpax
Range
251-292
263-301
269-311
269-320
305-358
294-344
-
-
_
_
251-388
No.
9
4
10
18
16
5
1
1
_
_
?
Mean
275
283
290
306
327
323
329
356
_
-
?
H.
indica
_
-
-
-
290
-
-
-
-
-
290
NUMBER 3 6
of each column includes records of eye size given in
the literature.
One feature of telson morphology is helpful in
species recognition, particularly when more than one
species is present in a sample. The carinae of the mar-
ginal teeth are dorsally tuberculate or nodulose in all
species, and the known species show a wide range of
size and prominence of this feature. In H. raphidea
the tubercles are large, inflated, and few in number;
the opposite extreme can be seen in H. harpax and H.
melanoura in which the tubercles are small, often
sharp, and very numerous. Harpiosquilla annandalei,
H. indica, new species, H. japonica, new species, and
H. stephensoni, new species, fall between these two ex-
tremes; in H. annandalei the tubercles are large, low,
and sparse. Specimens of H. stephensoni, new species,
and H. harpax (or any of those with very fine tubercu-
lation) can always be separated on the basis of the
tuberculation of the marginal carinae. This feature is
shown in the plates in which the telsons of each species
have been illustrated. The species, if placed in a series
based on the number of tubercles, would be arranged
as follows: H. raphidea (few tubercles), H. annan-
dalei, H. japonica, H. stephensoni, H. indica, H. har-
pax, and H. melanoura (many tubercles). The mar-
ginal carinae of the telson are sharpest in H. annan-
dalei, H. harpax, and H. melanoura.
Another feature which has not been used previously
as a specific character is the relative length of the
lateral and marginal carinae of the telson. In adults of
H. raphidea both carinae are inflated and may fuse
completely. In the remainder of the species they are
sharper and distinct throughout their length. The mar-
ginal carina of H. melanoura is almost three times as
long as the lateral carina; in H. harpax it is more than
twice but less than three times as long as the lateral
carina, and in the remainder of the species it is less
than twice as long. It seems to be shortest in H. indica,
for in the only known specimen of that species the
marginal carina is no more than one and one-half
times as long as the lateral carina. The species with the
longest marginal carinae, H. harpax and H. mela-
noura, can be distinguished from all other species on
the basis of this character.
 k
The ventral keel of the eighth thoracic somite seems
to have limited value as a specific character; it is sharp
in annandalei, rounded in the remainder of the species.
Two of the species, H. annandalei and H. raphidea,
differ from the other five in having the intermediate
carinae of the thoracic somites armed posteriorly; the
remaining species in the genus have unarmed inter-
mediate carinae on the thoracic somites.
The species seem to fall into two natural groups
based on the shape of the rostral plate. In H. harpax,
H. indica, new species, H. melanoura, and H.
raphidea the rostral plate is elongate, anteriorly taper-
ing to a slender median projection. In H. annandalei,
H. japonica, new species, and H. stephensoni new
species, the plate is short, with the length and width
subequal or the width greater, and the apex is rounded
or obtusely angled, lacking a noticeable median
projection.
The key to species is based on adults.
Key to Species of Harpiosquilla
1. Submedian carinae distinct on posterior 3 thoracic and first 5 abdominal somites. (Carapace
with median carina.) 2
1'. Submedian carinae indistinct or lacking on posterior 3 thoracic and first 5 abdominal somites.
(Rostral plate longer than broad, with apical projection; raptorial dactylus of male with
prominent angular projection on outer margin; fifth thoracic somite laterally acute; ven-
tral keel of eighth thoracic somite rounded; intermediate carinae of thoracic somites
unarmed posteriorly; submedian carinae of fifth abdominal somite unarmed posteriorly. ).5
2(1). Intermediate carinae of thoracic somites and first abdominal somite spinous posteriorly.
(Raptorial dactylus with 8 teeth; fifth thoracic somite laterally acute; intermediate
carinae of second abdominal somite spinous posteriorly.) 3
2'. Intermediate carinae of thoracic somites and first abdominal somite unarmed posteriorly.
(Rostral plate as broad as long, without apical projection; ventral keel of eighth thoracic
somite rounded; submedian carinae of fifth abdominal somite unarmed posteriorly; mar-
ginal carina of telson less than twice as long as lateral carina.) 4
3(2). Rostral plate as broad as long, without apical projection; raptorial dactylus of adult male
without angular projection on outer margin; ventral keel of eighth thoracic somite sharp;
submedian carinae of fifth abdominal somite spinous posteriorly; size moderate, total
length less than 150 mm. (Marginal carina of telson less than twice as long as lateral
carina; distal segment of uropodal exopod black with white midrib.). . . . 1. H. annandalei
356-607 O - 69 - 2
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
3'. Rostral plate longer than broad, with long apical projection; raptorial dactylus of adult
male with prominent angular projection on outer margin; ventral keel of eighth thoracic
somite rounded; submedian carinae of fifth abdominal somite unarmed posteriorly;
size large, total length up to 335 mm 2. H. raphidea
4(2')- Raptorial dactylus with 8 teeth; intermediate carinae of second abdominal somite un-
armed posteriorly; size moderate, total length less than 175 mm. (Fifth thoracic somite
rounded laterally.) 3. H. japonica, new species
4'. Raptorial dactylus with 7 teeth; intermediate carinae of second abdominal somite spinous
posteriorly; size large, total length up to 315 mm. (Raptorial dactylus of adult male
with prominent angular projection on outer margin; distal segment of uropodal exopod
dark colored on inner half.) 4. H. stephensoni, new species
5(1'). Carapace without median carina; distal segment of uropodal exopod entirely black. (Rap-
torial dactylus with 8 teeth; no submedian carinae on thoracic and first 5 abdominal
somites; intermediate carinae of first abdominal somite unarmed posteriorly; marginal
carina of telson more than twice as long as lateral carina; size moderate, total length
less than 170 mm.) 5. H. melanoma
5'. Carapace with median carina; distal segment of uropodal exopod dark colored on inner
half only 6
6(5'). Raptorial dactylus with 8 teeth; indistinct submedian carinae on thoracic and first 5
abdominal somites; intermediate carinae on second abdominal somite spinous posteriorly;
marginal carina of telson more than twice as long as lateral carina; size moderate to
large, total length up to 248 mm 6. H. harpax
6'. Raptorial dactylus with 9 teeth; no submedian carinae on thoracic and first 5 abdominal
somites; intermediate carinae on second abdominal somite unarmed posteriorly; mar-
ginal carina of telson less than twice as long as lateral carina; size moderate, total length
of mature individuals 150 mm or more 7. H. indica, new species
1. Harpiosquilla annandalei (Kemp, 1911)
FIGURES 1-3
Squilla annandalei Kemp, 1911, p. 99; 1913, p. 92, pi. 7 (figs.
78-80).?Sunier, 1918, p. 71.?Kemp and Chopra, 1921,
pp. 298 [listed], 307.?Chopra, 1934, p. 27, fig. 2; 1939,
p. 159, fig. 7.?Foxon, 1939, p. 259 [discussion of larva].?
Serene, 1954, pp. 6, 8 [listed].?Stephenson and McNeill,
1955, p. 255 [key].?Holthuis, 1964, p. 140 [listed].?
Manning, 1968a, p. 121 [listed].
Squilla annendalei.?Alikunhi, 1952, pp. 267, 269 [discussion
of larvae; erroneous spelling].
Harpiosquilla annandalei.?Manning, 1965, p. 250, pi. lla.?
Lee and Wu, 1966, p. 51, figs. 6A-B.?Manning, 1968b,
p. 14 [key].?Tirmizi and Manning, 1968, p. 31 [discussion].
MATERIAL.?1 $ , 71 mm; Tung Kang, Taiwan;
74-93 m; S. Lee, S. K. Wu; 5 March 1965; SI Crust.
113649.?1 4 , 107 mm; 2 $ , 109-117 mm; Andaman
Sea, Mergui Archipelago; 09?13'N, 97?51'E; 58-60
m; ANTON BRUUN Cruise 1, Sta. 20; IIOE; 23 March
1963; SI Crust. 125350.?3 9, 72-93 mm; Bay of
Bengal, off Burma; 19?32'N, 92?52'E; 53 m; ANTON
BRUUN Cruise 1, Sta. 49; IIOE; 6 April 1963; SI
Crust. 125348.?1 $ , 99 mm; Gulf of Oman, off Iran;
25?45-50'N, 57?O7'E; 92-95 m; clay, mud, sand,
minute gastropod shells; ANTON BRUUN Cruise 4B,
Sta. 255A; IIOE; 30 November 1963; SI Crust.
125349.-1$, 97 mm; Gulf of Oman, off Iran;
26?10-13'N, 57?02'E; 55-64 m; green mud; ANTON
BRUUN Cruise 4B, Sta. 256A; IIOE; 30 November
1963; AM.?2^, 129-135 mm; Gulf of Oman, off
Arabia; 25?12'N, 56?47-51'E; 206 m; grey soft mud;
ANTON BRUUN Cruise 4B, Sta. 263A; IIOE; 2 Decem-
ber 1963; SI Crust. 125716.
DIAGNOSIS.?Size moderate, adults with TL less
than 150 mm; antennular peduncle longer than cara-
pace and rostral plate combined; corneal indices rang-
ing between 240 and 291 (Table 1); rostral plate (Fig-
ure 2) with length and width subequal or width
greater, apex rounded or obtusely angled, lacking
slender anterior projection; carapace with median
carina; opposable margin of propodus of claw with
smaller spines and denticles between largest spines;
dactylus of claw with 8 teeth, outer margin of dactylus
lacking prominent angular projection in adult males:
fifth thoracic somite (Figure 3) acutely angled or
spined laterally; posterior 3 thoracic somites with
submedian and intermediate carinae, intermediates
with posterior spines; ventral keel of eighth thoracic
NUMBER 3 6
somite sharp; all six abdominal somites with sub-
median carinae, abdominal carinae spined as follows:
submedian 5-6, intermediate 1-6, lateral 1-6, margi-
nal 1-5; denticles 5-7, 10-11, 1, inner submedians
largest, rounded, remainder spiniform; marginal
carina less than twice as long as carina of lateral tooth;
postanal carina short, not extending halfway between
anus and posterior margin; distal segment of uropodal
exopod black with white midrib.
COLOR.?Second and third segments of antennular
peduncle with proximal dorsal black spot and distal
black ring, proximal spot fainter on third segment;
dark transverse bar present dorsally between ophthal-
mic and antennular somites; medan portion of rostral
plate outlined in dark pigment; carapace with antero-
lateral margins, posterior median margin, grooves, and
carinae outlined with dark pigment; merus of claw
with inner subdistal black spot and with proximal and
distal dark spot on dorsal depression; posterior 3 tho-
racic and all 6 abdominal somites with dark posterior
line; submedian and intermediate carinae of carapace
darker than adjacent surface; second abdominal
somite with transverse dark bar middorsally; first and
third to fifth abdominal somites each with traces of
broken transverse bar; telson with pair of proximal,
submedian dark circles, margin darker than center;
pits, arranged in concentric rows on dorsum of telson,
dark; distal segment of uropodal exopod black with
white midrib; distal half of uropodal endopod with
diffuse dark pigment. The overall color pattern is
shown on Figure 1.
SIZE.?Males, TL 71-135 mm; females, TL 72-117
mm. Males of TL 106-123 mm and females of TL
43-115 mm have been recorded in the literature.
Other measurements of a male, TL 107 mm: cara-
pace length 22.1; cornea width 8.5; antennular pe-
duncle length 26.5; rostral plate length 2.7, width
3.3; raptorial propodus length 27.9; fifth abdominal
somite width 20.0; telson length 17.8, width 16.8.
DISCUSSION.?Harpiosquilla annandalei can be
distinguished readily from all other species in the genus
by a variety of characters, including the length of the
antennular peduncle (longer than the carapace and
rostral plate combined), the sharp ventral keel on the
eighth thoracic somite, the armed submedian carinae
of the fifth abdominal somite, and the color pattern
of the telson and uropod (Figure 1). None of these
features is found in any other species of Harpiosquilla
now known.
FIGURE 1.?Harpiosquilla annandalei (Kemp). Male, TL
135 mm, Gulf of Oman: dorsal view.
8Harpiosquilla stephensoni, new species, and H. ja-
ponica, new species, also have a short rostral plate but
in neither of those species are the intermediate carinae
of the thorax or the submedian carinae of the fifth
abdominal somite armed posteriorly. Harpiosquilla ste-
phensoni, new species, and H. raphidea also have an
acute or even spiniform lateral process on the fifth
thoracic somite; H. raphidea is the only other species
of the genus in which the intermediate carinae of the
last three thoracic somites are armed posteriorly and
in which all intermediate carinae of the abdomen also
are spined.
FIGURE 2.?Harpiosquilla annandaUi (Kemp). Male, TL
135 mm, Gulf of Oman: anterior portion of carapace.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Corneal indices of H. annandalei are summarized
in Table 1. The eyes of the specimens reported herein
are similar to those of H. harpax of comparable size,
but the overall range of corneal indices, 240-291, in
contrast to that observed in H. harpax, 251-358, sug-
gests that the eyes of H. annandalei are larger than
those of H. harpax.
Chopra (1934, 1939) has pointed out that the an-
terior lobes of the lateral processes of the sixth and
seventh thoracic somites are variable in shape and in
some specimens may be subdivided secondarily. Simi-
lar variation in the shape of the anterior lobes of these
somites is evident in the specimens reported herein.
The lateral processes in H. annandalei (Figure 3) are
more strongly bilobed than in any other species of the
genus.
Harpiosquilla annandalei also has a higher and
sharper median carina on the telson than any other
species of Harpiosquilla; Lee and Wu (1966) com-
mented on the shape of the carina in their specimens
from Taiwan, and they noted that the carina, in lateral
view, is convex anteriorly, concave posteriorly. The
relative thinness of the median carina does not seem to
be affected by age, and the median carina is not par-
ticularly inflated in either large males or large females.
One specimen, from ANTON BRUUN Sta. 49, a female
TL 72 mm, had nine teeth on the dactylus of one
claw, the normal complement of eight teeth on the
other dactylus.
In all specimens examined the intermediate carinae
of the last three thoracic somites and of all six ab-
dominal somites were armed posteriorly.
SEXUAL DIMORPHISM.?This species does not exhibit
secondary sexual dimorphism to the extent found in
other species such as H. harpax and H. raphidea. The
two males reported herein from ANTON BRUUN Sta.
263A are the largest recorded for the species and they
show no marked sexual dimorphism. The outer edge
of the dactylus of the claw in the largest male, TL 135
mm, is faintly sinuous but not markedly different in
shape from the claw of the females. The teeth on the
raptorial claw of these two males are slightly more
inflated basally than in smaller males or in any females,
but the difference in shape is relatively minor.
BIOLOGY.?Virtually nothing is known about the
biology of this species. In general, it occurs in deeper
water, 15-206 meters, than either H. harpax or H.
raphidea. Specimens collected from the ANTON BRUUN
were taken on mud or grey mud and sand. Serene
NUMBER 3 6 9
(1954) recorded the species from sandy mud, in 15-25
meters, in the Baie de Cauda, Viet-Nam. The specimen
reported by Chopra (1939) was taken on green mud
in 201 meters in the Gulf of Oman. Harpiosquilla
annandalei was taken together with H. melanoura al
ANTON BRUUN Sta. 20, in 5&-60 meters off the Mergui
Archipelago and with H. harpax at ANTON BRUUN
Sta. 256A, in 55-64 meters in the Gulf of Oman.
DEVELOPMENT.?Larval stages of H. annandalei
FIGURE 3.?Harpiosquilla annandalei (Kemp). Male, TL
135 nun, Gulf of Oman: lateral processes, of fifth, sixth,
and seventh thoracic somites.
have not been identified with certainty. Foxon (1939)
identified larvae from the Red Sea, Arabian Sea, and
Gulf of Oman with Alima multispinus (Claus) She
noted that the larvae could be separated into two
groups, referred to by her as typical and atypical, and
suggested that they might be the larvae of H. raphidea
and H. annandalei (note that neither of these species
is known to occur in the Red Sea proper and H. raph-
idea is not known with certainty to occur in the Gulf
of Oman.)
Alikunhi (1952, p. 269) pointed out that "while
the atypical form is perhaps identical with S. {Alima)
raphidea, the typical larva might well belong to the
very closely related S. annendalei [sic]." Ingle and
Delia Croce (1967) corroborate Alikunhi's identifica-
tion of Foxon's atypical larvae with H. raphidea. This
seems highly unlikely if the atypical larvae came from
the Red Sea rather than the Arabian Sea; however, the
larvae identified with H. raphidea by Alikunhi could
be the larvae of H. harpax, and this might explain the
presence of this larval form in the Red Sea,
TYPE.?The four syntypes are in the Zoological
Survey of India [Indian Museum], Calcutta.
TYPE-LOCALITY.?Gulf of Martaban, Burma, in 53-
67 fathoms.
DISTRIBUTION.?Indo-West Pacific region, from
scattered localities between Japan and the Gulf of
Oman in depths between 15 and 206 meters. Records
for adults in the literature are as follows: JAPAN:
Hayama, Sagami Bay (Manning, 1965). TAIWAN:
Tungkang market, 40-50 fms (Lee and Wu, 1966).
VIET-NAM: Baie de Cauda, 15-25 m (Serene, 1954).
JAVA SEA: 30-35 fms (Sunier, 1918). BAY OF BENGAL:
off Kabusa Island, Mergui Archipelago, 33 fms (Kemp
and Chopra, 1921); Gulf of Martaban, Burma, 53-67
fms (Kemp, 1911,1913); Sandheads, mouth of Hughli
River [Hooghly], India, less than 20 fms (Chopra,
1934). ARABIAN SEA; Gulf of Oman, 201 m (Chopra,
1939). Some of the larvae reported from the Red Sea,
Arabian Sea, and Gulf of Oman by Foxon (1939) may
be referable to this species.
2. Harpiosquilla raphidea (Fabricius, 1798)
FIGURES 4-9
Squilla raphidea Fabricius, 1798, p. 416.?Latreille, 1802, p.
279; 1828, p. 471 .?Milne-Edwards, 1837, p. 524.?Gibbes,
1850, p. 199 (discussion; p. 35 on separate].?Miers,
1880, p. 27 [part].?Preudhomme de Borre, 1882, p. cxi.?
de Man, 1888, p. 296.?Thallwitz, 1892, p. 55.?Hender-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
son, 1893, p. 453.?Sharp, 1893, p. 108.?Thurston, 1895,
p. 120.?de Man, 1898, p. 694.?Lanchester, 1900, p. 264;
1901, p. 553.?Nobili, 1903, p. 38.?Lanchester, 1906, p.
133.?Kemp, 1913, p. 88 [part]; 1915, p. 172; 1918, p.
297.?Sunier, 1918, p. 70, fig. 3.?Hansen, 1921, p. 79 ?
Kemp and Chopra, 1921, p. 298 [listed].?Smedley, 1927,
p. 231.?Roxas and Estampador, 1930, p. 101.?Chopra,
1934, p. 27 [part].?Tweedie, 1934, p. 40.?Foxon, 1939,
p. 259 [discussion of larvae].?Gravely, 1941, p. 74.?
Holthuis, 1941, p. 256 [part].?Nair, 1941, p. 544, figs. 3 -
10, 21, 23-30, pi. 29 (figs. 34-35), pi. 30 (figs. 37-38, 40)
[embryology].?Chacko, 1942, p. 404.? Alikunhi and
Aiyar, 1942, p. 56, figs. 3-4 [larvae]; 1943, p. 81 [growth].
?Alikunhi, 1950, p. 103 [discussion; larvae]; 1952, p. 265,
fig. 9 [larvae].?Tiwari and Biswas, 1952, p. 356, figs. 3a,
c, e.?Ingle, 1963, p. 14 [key].?Zimsen, 1964, p. 653
[listed].?Holthuis, 1964, p. 140 [listed].?Alikunhi, 1965,
p. 35 [postlarva; listed].?Manning 1968a, p. 121 [listed].
Squilla raphidia.?Bosc, 1801-1802, p. 122 [erroneous
spelling].
Squilla mantis (B) var. major Lamarck, 1818, p. 187 [in
synonymy].?Deshayes and Milne-Edwards, 1838, p. 322;
1839, p. 373 [in synonymy].
Squilla Mantis.?Latreille, 1818, p. 6, pi. 324 [not Squilla
mantis (Linnaeus)].
Squilla Raphidia.?Bosc, 1830, p. 95 [erroneous spelling].
Squilla Raphidea.?Herklots, 1861, p. 152 [p. 39 on separate;
listed].
Alimerichthus Claus, 1871, p. 147, pi. 8 (fig. 30) [larva; p. 39
on separate].??Brooks, 1886, p. 96 [discussion].
Alimerichthus pyramidalis Lanchester, 1903, p. 457 [larva].?
Gurney, 1946, p. 161 [references].
Squilla raphidea var. africana Balss, 1910, p. 8, fig. 2b.
Alima pyramidalis.?Gurney, 1946, p. 159 [references].
Squilla (Alima) raphidea.?Alikunhi, 1952, p.269 [discussion
of larvae].
?Harpiosquilla raphidea.?Alikunhi, 1967, p. 894, figs. 114-
125, pi. 2 [larvae and young; part?].
Harpiosquilla raphidea.?Chhapgar and Sane, 1968, p. 45
[key.]?Manning, 1968b, p. 14 [key].?Tirmiri and Mann-
ing, 1968, p. 31, fig. 12.
not Squilla raphidea.?Berthold, 1845, p. 47; 1847, p. 29 ?
White, 1847, p. 84.?Bigelow, 1894, pp. 511 [key], 535 ?
Tattersall, 1906, p. 166.?Balss, 1910, p. 8, fig. 2a.?Parisi,
1922, p. 103.?Komai, 1927, p. 323.?Komai, Akatsuka,
and Ikari, 1927, p. 295.?Komai and Ikari, 1929, p. 121.?
Gravier, 1930, p. 525.?Serene, 1937, p. 68 [listed].?
Komai, 1938, p. 268.?Suvatti, 1938, p. 52.?Chopra, 1939,
p. 158?Liu, 1949, p. 43, pi. 6 (figs. 15-17).?Anony-
mous, 1949, p. 843, fig. 2421.?Barnard, 1950, p. 851, figs,
lc, g.?Suvatti, 1950, p. 132.?Dawydoff, 1952, p. 145.?
Serene, 1953, p. 507; 1954, pp. 6, 8, 62, pi. 4 (figs. 1-6).?
Millard and Harrison, 1954, p. 176 [listed].?Day and Mor-
gans, 1956, p. 306 [listed].?Utinomi, 1956, p. 91, pi. 46
(fig. 2) [colored figure].?Yamaji, 1959, fig. on p. 68.?
Utinomi, 1960, p. 114, pi. 57 (fig. 7) [colored figure].?
Chuang, 1961, pp. 181, 206, pi. 81 (fig. 4), lowest fig. on
pi. 82.?Stephenson, 1962, p. 34.?Crosnier, 1965, p. 61
[listed]. [All = H. harpax (de Haan).]
not Squilla raphidea.??Tate, 1883, p. 48.?Stephenson,
1952, p. 4; 1953, p. 43. [All probably = H. stephensoni,
new species.]
not Chloridella raphidea.?Rathbun, 1902, p. 55. [ = / / .
japonica, new species.]
not Squilla raphidea.?Fukuda, 1913, pp. 70, 72, fig. on p. 71.
[=H. japonica, new species.]
not Squilla raphidia.?Torralbas, 1917, p. 621 [p. 81 on sep-
arate] [=Squilla empusa Say, 1818].
not Squilla raphidoea.?Serene, 1939, p. 349 [erroneous spell-
ing; ?=/ / . harpax].
not Squilla raphidea.?Stephenson and McNeill, 1955, p. 239
[a mixture of H. harpax (de Haan), H. melanoura Man-
ning, and H. stephensoni, new species].
MATERIAL.?1 9 , 216 mm; Sandakan, North Bor-
neo, Indonesia; Herre, col.; 30 June 1929; SI Crust.
125360.?29, 250-277 mm; Thailand; Hugh M.
FIGURE 4.?Harpiosquilla raphidea (Fabricius). Female,
TL 250 mm, Thailand: anterior portion of body.
NUMBER 36 11
FIGURE 5.?Harpiosquilla raphidea (Fabricius). Male, TL
310 mm, off Bombay: raptorial claw (pro pod us length
72.5 mm).
FIGURE 6.?Harpiosquilla raphidea (Fabricius). Female,
TL 250 mm, Thailand: raptorial claw (propodus length
69.7 mm).
Smith; SI Crust. 93736.?1 9, 270 mm; Pak Poon
[Pakpoon, western side of Gulf of Siam, Nakon Srita-
marat Province], Thailand; Hugh M. Smith; 19 Octo-
ber 1923; SI Crust. 69530.?1 S , CL 56.0 mm; same;
SI Crust. 106388.?1 $, 310 mm; taken with a 60
mile radius of Bombay, India; C. L. Kaufmann; SI
Crust. 104363.
DIAGNOSIS.?Size large, adults with TL to 335 mm;
antennular peduncle shorter than carapace; corneal
indices ranging between 380 and 530 (Table 1);
rostral plate (Figure 4) longer than broad, with
slender anterior projection; carapace with median
carina; upper margin of propodus of claw usually with
1 smaller spine or denticle between largest spines;
dactylus of claw with 8 teeth, outer margin with prom-
inent angular projection in adult males (Figure 5);
fifth thoracic somite (Figure 7) with lateral spine;
posterior 3 thoracic somites with submedian and inter-
mediate carinae, intermediates spined posteriorly;
ventral keel of eighth thoracic somite rounded; all 6
abdominal somites with submedian carinae, abdominal
carinae spined as follows: submedian 6, intermediate
1-6, lateral 1-6, marginal 1-5; denticles rounded,
4-5, 8, 1; marginal carina about twice as long as carina
of lateral tooth, usually fused with it in large speci-
mens; postanal carina extending about halfway be-
tween anus and posterior margin; proximal portion
of inner half of distal segment of uropodal exopod with
dark pigment.
COLOR.?The color pattern is not well marked in
any of the specimens examined. Posterior 3 thoracic
and first 5 abdominal somites each with dark posterior
line; fifth thoracic somite with dark anterior line;
telson with proximal pair of dark spots; uropodal
exopod with dark spot, primarily on inner surface,
on distal third of proximal segment; some dark pig-
ment on inner proximal third of distal segment of
uropodal exopod, but inner half of distal segment not
completely dark.
Tirmizi and Manning (1968, p. 33) noted that a
fresh specimen from Karachi, West Pakistan was col-
ored as follows:
. . . ocular peduncles light pink; posterior border of cara-
pace with a black band; antennal scale yellowish, outlined
with dark pigment; claw with merus pink, also marked with a
greenish yellow patch; distal end of propodus with bright
yellow streak; thorax and abdomen appearing speckled; last
3 thoracic and first 3 abdominal somites pink, last 3 abdomi-
nal somites more cream-colored; tips of spines of last 4
abdominal somites yellow; posterior margin of first 4 abdomi-
nal somites black; carinae of telson bluish, apices of teeth
yellow; telson with pair of submedian yellow-brown spots;
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
uropod lightly marked with yellow and black, inner half of
distal segment of exopod grayish, outer half yellow.
SIZE.?Only intact male examined, TL 310 mm;
females, TL 216-277 mm. Males ranging between
197-275 mm TL and females ranging between 160-
288 mm TL have been recorded in the literature.
Kemp (1913) recorded a specimen of TL 335 mm but
did not specify its sex.
Other measurements of a female, TL 277 mm:
carapace length 57.0; cornea width 12.0; antennular
peduncle length 42.0; rostral plate length 13.6, width
8.2; raptorial propodus length 81.4; fifth abdominal
somite width 59.8; telson length 52.4, width 54.3.
DISCUSSION.?Several of the features of H. raphi-
dea allow its ready separation from the other known
species of the genus. It shares with H. annandalei the
sharp lateral process on the fifth thoracic somite
(Figure 7), and the armed intermediate carinae of
the posterior three thoracic and all six abdominal
somites; these two features will distinguish it from
FIGURE 7.?Harpiosquilla raphidea (Fabricius). Female
TL 250 mm, Thailand: lateral processes of fifth, sixth, and
seventh thoracic somites.
the five other species. It differs from H. annandalei in
several characters over and above color pattern and
size of adults, for the submedian carinae of the fifth
abdominal somite are unarmed, the rostral plate is
elongate, with a slender median projection (Figure 4) ,
and adult males of H. raphidea have the large angled
prominence on the outer margin of the dactylus of the
claw (Figure 5). As noted under the Discussion of
H. annandalei (p. 7), the color pattern of that species
is quite distinctive.
Harpiosquilla raphidea is the largest known stoma-
topod; Kemp (1913) recorded a specimen from Bom-
bay with a total length of 335 mm. In very large
specimens, as in the male, TL 310 mm, recorded here
from Bombay, the marginal teeth of the telson are
markedly compressed and inflated. The lobe between
the spines of the basal prolongation of the uropod
becomes comparatively smaller with increasing size.
Inflation of the carinae of the marginal teeth of the
telson in adults smooths out the normal dorsal rugosity
of these carinae.
Two authors have recorded specimens with nine
rather than eight teeth on the dactylus of the raptorial
claw. Lanchester (1906) noted that a male from
Patani Bay, Halmahera, had nine spines on the left
dactylus and Roxas and Estampador (1930) reported
that their specimens had nine, rarely eight teeth, on
the claw. The latter authors may have been dealing
with another species, for their specimens were under
160 mm TL.
The structure of the dorsal organ of this species
was discussed by Hansen (1921).
SYNONYMY.?Inasmuch as H. raphidea was the
only species in the genus which was recognized be-
tween 1798 and 1911, when Kemp described annan-
dalei, and which was considered to include all speci-
mens distinct from annandalei between 1911 and 1952,
it is not surprising to find that the literature is quite
confused; many of the records of raphidea in the liter-
ature could refer to one or more of the six other
species recognized herein. Some of the records here
referred to raphidea may include that species and other
species as well.
Many of Kemp's (1913) records were verified by
Tiwari and Biswas (1952) in their account in which
H. harpax (de Haan) was shown to be distinct from
H. raphidea; the latter authors also verified identifi-
cations of some material reported on by Chopra
(1934). These verified records have been used to com-
NUMBER 3 6 13
pile the distribution of the species as recorded in the
literature.
Balss (1910) distinguished his variety africana from
raphidea sensu stricto by the shape of the rostral plate
(which he illustrated), the presence of well-developed
keels on the abdomen and telson, and the well-devel-
oped teeth of the telson. His raphidea sensu stricto is
apparently H. harpax whereas his africana is raphidea
sensu stricto.
Sunier (1918) has clearly illustrated a specimen of
raphidea; the distinctive lateral process of the fifth
thoracic somite and telson are easily discernible.
In his account of postlarval development and growth
of raphidea, Alikunhi (1967) notes that according to
the features discussed by Tiwari and Biswas (1952) his
juveniles should be referred to harpax. He mentions,
however, that at one stage the juveniles have all of the
intermediate carinae of the abdomen provided with
spines. This is a feature always found in H. raphidea
and rarely found in H. harpax, for the intermediate
carinae of the first abdominal somite are usually un-
armed in H. harpax. It seems likely that Alikunhi was
dealing with specimens of both species. Adults of both
species occur on the eastern coast of India.
As noted under Distribution (p. 15), most records
in the literature require verification.
SEXUAL DIMORPHISM.?As in most other species of
Harpiosquilla, large males exhibit marked changes in
the morphology of the raptorial claw and the telson; to
a lesser extent the carinae of the sixth abdominal
somite are also affected. The outer margin of the dac-
tylus of the claw in the male (Figure 5) has a large,
angled protuberance and the bases of the teeth on the
inner margin of the dactylus, particularly the second
through the fifth tooth, have basal inflations. The
carinae of the sixth abdominal somite, especially the
submedians, are inflated in large adults of both sexes,
but they are more inflated in the males; in some speci-
mens the posterior spines of the submedian teeth are
obsolete. In adult males (Figure 8*), the median carina
of the telson is so inflated that it may obliterate the
distal dorsal spine. The carinae of the marginal teeth
are also inflated; usually the marginal and lateral
carinae of the telson are completely fused. Claws (Fig-
ures 5, 6) and telsons (Figures 8, 9) of both males and
females have been illustrated. Although the carinae of
the telson are also inflated in adult females, as in the
specimen from Thailand, TL 250 mm, illustrated here,
the extent of inflation is never so great as in males of
similar size.
There is no information available on the size at
which males begin to exhibit dimorphism, but Alikunhi
(1967, p. 903) noted that: "Sexual maturity is not
FIGURE 8.?Harpiosquilla raphidea (Fabricius). Male, TL
310 mm, off Bombay: posterior portion of body (telson width
55.8 mm).
FIGURE 9.?Harpiosquilla raphidea (Fabricius). Female,
TL 250 mm, Thailand: posterior portion of body (telson
width 48.1 mm).
356-607 O - 69 - 3
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
attained even after the 15th moult after metamor-
phosis. Other specimens up to 137.0 mm in length
were also found immature in March."
Kemp (1918) noted that one male from Tale Sap,
Thailand, TL 200 mm, lacked an angular projection
on the claw.
BIOLOGY.?As in the other species of Harpiosquilla,
very little is known about the biology of H. raphidea.
It occurs in relatively shallow water; the few depth
records in the literature indicate that it has been taken
in depths between 1 and 28 fathoms. Kemp (1918)
noted that it was taken in brackish water, specific
gravity 1.0085, at Tale Sap, Thailand.
Several authors have reported specimens from local
fish markets, so it may have economic importance in
some areas and it may be collected in connection with
commercial fishing operations. It has been recorded
from markets in Karachi, West Pakistan (Tirmizi and
Manning, 1968), Singapore (Tweedie, 1934), Sara-
wak (Tiwari and Biswas, 1952), Divisoria Market,
Manila (Kemp, 1915), and markets in Manila (Roxas
and Estampador, 1930) ; the specimens from Thailand
reported by Kemp (1918) were taken in fishermen's
nets.
Smedley (1927) commented that raphidea was the
commonest of local mantis shrimps at Morib, on the
west coast of the Malay peninsula.
Sunier (1918) found fragments of the shrimp
Penaeus indicus Milne-Edwards in the stomachs of
two specimens and Brachyurum [Brachyura?] in the
stomach of a third.
There is no information available on other stoma-
topods or invertebrates associated with this species.
Postlarval development and changes with age have
been discussed in some detail by Alikunhi (1967).
DEVELOPMENT.?Nair (1941) discussed egg extru-
sion and embrological development in a species from
Madras, India, identified by him as Squilla raphidea;
he also made observations on the embryology of Ora-
tosquilla holoschista (Kemp), O. nepa (Latreille),
and O. woodmasoni (Kemp).
Alikunhi (1952) has given a detailed account of a
larval form identified by him as raphidea; Alikunhi
and Aiyar (1942, 1943) and Alikunhi (1950) had
previously reported the metamorphosis of a larva into
postlarval and juvenile raphidea. In an abstract of a
paper presented at the Symposium on Crustacea spon-
sored by The Marine Biological Association of India,
Alikunhi (1965) noted that raphidea was among eight-
een species from the Madras coast, the postlarval de-
velopment of which was well known.
In the paper presented at the Symposium and pub-
lished in Part II of the Proceedings of the Symposium
on Crustacea, Alikunhi (1967) has presented results of
the most detailed study of growth in stomatopods ever
made. Aspects of postlarval growth are reported in
varying detail for eighteen species, including raphidea.
Of particular interest is the fact that the movable sub-
median teeth of the postlarva are retained beyond that
stage in Harpiosquilla. Although Alikunhi identifies his
material with raphidea and mentions that by the
twelfth molt the intermediate carinae of the first and
second abdominal somites are armed posteriorly, a
characteristic of adult raphidea, he notes on p. 904
that according to the diagnostic characters of harpax
and raphidea the reared material actually belongs to
harpax. It seems highly likely that both species were
represented in his material.
Alikunhi (1952) identified his larva with Alimerich-
thus sp. Claus (1871) from the Indian Ocean and with
Alimerichthus pyramidalis Lanchester, 1903 from
North Male Atoll, Maldive Islands; Lanchester had
identified his larva with Claus' species and with an
Alimerichthus reported by Brooks (1886, pp. 95-98).
Brook's larvae, however, were from the Atlantic and
are not likely to be the larvae of an Indo-West Pacific
species.
Alikunhi noted that his series of larvae from Madras
included two distinct forms. He identified his typical
larva with H. raphidea and stated (1952, p. 267) :
The post-larva obtained by moult from one such larva more
or less agrees with S. raphidea. However, since the early post-
larvae in most cases do not show the full adult characters and
since S. raphidea and S. annendalei [sic] are very closely
related, it is a possibility that the atypical larva probably
belongs to the latter species. This is, however, just a surmise
which is not based on any positive evidence.
Alikunhi (1952, p. 269) also noted that Erichthus
multispinus Claus, identified with H. raphidea by
Foxon (1939), "appears to be a very different species,
probably not related to S. raphidea, even though it also
possesses laterally directed spinules on the carapace.
Borradaile's [1907] specimens are perhaps identical
with the specimen figured by Claus."
TYPE.?Probably not extant. Zimsen (1964) did not
locate the type in her comprehensive survey of
Fabrician types.
TYPE-LOCALITY.?Oceano Indico D. Daldorff. I. K.
Daldorff collected material for Fabricius at Fredericks-
NUMBER 3 6 15
nagore, on the eastern coast of India. Fredericksnagore,
located on the Hooghly River, north of Calcutta, is
now known as Serampore (Webster's Geographical
Dictionary, Rev. Ed., 1966).
DISTRIBUTION.?Indo-West Pacific region, from
Indonesia and the Gulf of Siam westward to East
Africa. Most of the records in the literature probably
refer to H. harpax, H. indica, new species, or H.
melanoura as well as H. raphidea; from most accounts
it is not possible to determine which species was ac-
tually being recorded. The following records are
probably referable to H. raphidea: SOUTH CHINA
SEA: NE of Tegal, 06?53'N, 110?21'E, 6.5-9 fms
(Sunier, 1918). INDONESIA: Indischen Archipel (de
Man, 1898); Java Sea, between Singapore and Sura-
baya [Surabaja, Java] (? part; de Man, 1898); Borneo
Bank, Makassar Strait, 15 fms; Madura Strait, ca. 16
fms; Java Sea, N. of western Madura, 7.5 fms; vicinity
of Madura; near Tandjong Priok, Batavia [Djakarta],
less than 1 fm; same, 6-7 fms; beyond western entrance
of Banka Strait, Sumatra, 5-8 fms; outside of estuary
of Rokan River, Sumatra, 18-20 fms; Bagan Si Api
Api, eastern Sumatra; east coast of Sumatra (all
Sunier, 1918). MALAYSIA : Singapore (Kemp, 1913;
Tiwari and Biswas, 1952); Penang, 11 fms (Sunier,
1918); Kuching Fish Market, Sarawak (Tiwari and
Biswas, 1952). THAILAND: (?) Tale Sap (Kemp,
1918; Tiwari and Biswas, 1952). INDIAN OCEAN:
(Fabricius, 1798). BAY OF BENGAL: north and south
of Eastern Channel (Tiwari and Biswas, 1952). EAST
PAKISTAN: COX'S Bazaar (Tirmizi and Manning,
1968). INDIA: Sunderbans [Sundarbans] (Tiwari
and Biswas, 1952); Sandheads, mouth of River Hughli
[Hooghly; including Saugor Island] (Kemp, 1913;
Chopra, 1934; Tiwari and Biswas, 1952); Calcutta
Bazaar (Tiwari and Biswas, 1952); Chandipur, Bala-
sore (Tiwari and Biswas, 1952); offPuri (Kemp, 1913;
Tiwari and Biswas, 1952); Bombay (Kemp, 1913;
Chhapgar and Sane, 1968). WEST PAKISTAN:
Karachi (Tirmizi and Manning, 1968). EAST
AFRICA: (Balss, 1910).
The following records need to be verified; all fall
within the known range of H. raphidea but it is not pos-
sible to determine with certainty the identity of the spe-
cies reported by each author. PHILIPPINE ISLANDS:
Philippines (Holthuis, 1941); Manila Bay (Roxas and
Estampador, 1930) ; Divisoria Market, Manila (Kemp
1915) ; Cebu (Roxas and Estampador, 1930) ; Davao
Gulf, Mindanao (Roxas and Estampador, 1930). IN-
DONESIA: Dutch East Indies; Skroe, Dutch New
Guinea; Amboina; Haroekoe (all Holthuis, 1941);
Patani Bay, Halmahera (Lanchester, 1906); Moluccas
(Holthuis, 1941); Kisar, near Timor (Holthuis,
1941); Borneo (Miers, 1880); Sandakan, northeast
Borneo; Balikpapan, east Borneo; Java; Java Sea
(all Holthuis, 1941); Madura (Thallwitz, 1892);
Batavia [DJakarta] (Preudhomme de Borre, 1882) ;
Bay of Batavia [Djakarta]; Tandjong Priok, near Ba-
tavia [Djakarta]; Cheribon, Java; Deli, east Sumatra;
mouth of Djambi River, east coast of Sumatra; Padang,
west coast of Sumatra; coast of Atyeh, Sumatra; Telok
Dalam, Nias (all Holthuis, 1941). MALAYASIA:
Singapore (Nobili, 1903); Siglap, Singapore
(Tweedie, 1934); Morib (Smedley, 1927); Kota
Bharu (Lanchester, 1901); [?] Moratabas (Lanchester,
1900); Malacca (Sharp, 1893). THAILAND: Singora
(Lanchester, 1901). INDIAN OCEAN: (BOSC, 1801?
1802, 1830; Latreille, 1802; Lamarck, 1818; Milne-
Edwards, 1837; Deshayes and Milne-Edwards, 1838,
1839). ANDAMAN ISLANDS: Port Blair (Kemp,
1913). BURMA : Mergui Archipelago (de Man, 1888;
Kemp, 1913); off Irrawaddy Delta, 15?20'N, 94?55'E,
20 fms (Kemp, 1913); Rangoon (Kemp, 1913);
Akyab, Tenasserim (Kemp, 1913). INDIA: no spe-=
cific locality (Sharp, 1893) ; Sunderbunds [Sundar-
bans] (Henderson, 1893); off Gopalpur, Ganjan coast,
25-28 fms (Kemp, 1913); Indes Orientales, Pondi-
chery (Latreille, 1828); Madras (Henderson, 1893;
Nair, 1941; Gravely, 1941); Pamban, Gulf of Manaar
[Mannar] (Thurston, 1895); Krusadai Island, Gulf of
Mannar (Chacko, 1942); Bombay Harbour, Bombay
[part?] (Kemp, 1913). CEYLON: (Kemp, 1913).
WEST PAKISTAN: Karachi (Kemp, 1913). PERSIAN
GULF: 28?59'N, 50?05'E, 25 fms (Kemp, 1913).
Larvae attributed to H. raphidea have been recorded
from the following localities: INDIAN OCEAN: (Claus,
1871). INDIA: Madras (Alikunhi and Aiyar, 1942,
1943; Alikunhi, 1950, 1952, 1965, 1967. MALDIVE
ISLANDS: North Male Atoll, 27-35 fms (Lanchester,
1903). EAST AFRICA: Mozambique Channel (Ingle
and Delia Croce, 1967).
3. Harpiosquilla japonica, new species
FIGURES 10-11
Chloridella raphidea.?Rathbun, 1902, p. 55 [not Squilla
raphidea Fabricius].
Squilla raphidea.?Fukuda, 1913, pp. 70, 72, figure on p. 71
[not S. raphidea Fabricius] [in Japanese].
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
HOLOTYPE.?1 9 , 172 mm; Wakanoura, Kii, Japan;
Jordan and Snyder, col.; 1900; SI Crust. 26340.
PARATYPES.?1 3 , 165 mm; 1 $ , 135 mm; data as
in holotype; SI Crust. 125718.
DIAGNOSIS.?Size moderate, adults with TL less
than 175 mm; antennular peduncle not so long as
carapace; corneal indices ranging between 325 and
337 (Table 1) ; rostral plate (Figure 10) with length
and width subequal, apex rounded or obtusely angled,
lacking slender anterior projection; carapace with
median carina; upper margin of propodus of claw with
smaller spines and denticles between largest spines;
dactylus of claw with 8 teeth; fifth thoracic somite
(Figure 11) rounded laterally; posterior 3 thoracic
somites with submedian and intermediate carinae, none
armed posteriorly; ventral keel of eighth thoracic
somite rounded; all 6 abdominal somites with sub-
FIGURE 11.?Harpiosquil-
la japonica, new species.
Female holotype, TL 172
mm, Wakanoura: lateral
processes of fifth, sixth,
and seventh thoracic
somites.
MOURE 1U.?tiarpiosquilla japonica, new species. Female
holotype, TL 172 mm, Wakanoura: anterior portion of
body.
median carinae, abdominal carinae spined as follows:
submedian 6, intermediate 3-6, lateral 1-6, marginal
1-5; denticles 4-6, 10-11, 1; marginal carina of telson
less than twice as long as carina of lateral tooth; post-
anal keel extending about halfway between anus and
posterior margin; color of uropodal exopod unknown.
COLOR.?Completely faded in the type-specimens.
SIZE.?Only male examined, TL 165 mm; females
TL 135-172 mm.
Other measurements (in mm) of male: carapace
length 34.1; cornea width 10.5; antennular peduncle
length 32.4; rostral plate length 5.4, width 5.4; fifth
abdominal somite width 35.3; telson length 31.3, width
30.3.
DISCUSSION.?Harpiosquilla japonica, new species,
resembles both H. annandalei and H. stephensoni, new
species, and differs from the remainder of the species
in the genus in having a short rostral plate, as broad
as long, without an anterior projection. Harpiosquilla
annandalei differs in numerous features, as noted un-
der the discussion of that species, and it can be dis-
tinguished immediately from H. japonica by the
presence of armed submedian carinae on the fifth ab-
dominal somite. Harpiosquilla stephensoni and H.
japonica are very similar in general facies, but they
can be distinguished readily by the characters used in
the key, namely (1) the presence of eight teeth on the
claw of H. japonica, seven on H. stephensoni; (2) the
intermediate carinae of the second abdominal somite
are unarmed in H. japonica, spined posteriorly in H.
stephensoni, and (3) the lateral process of the fifth
NUMBER 3 6 17
abdominal somite is rounded in H. japonica, angled
or spined in H. stephensoni. The last feature may be
of limited value in distinguishing specimens of H.
japonica from smaller specimens of H. stephensoni,
for, as noted under the Discussion (p. 19) of the latter
species, small adults may not have the projecting lat-
eral process well developed.
Although the corneal indices of H. japonica, new
species, overlap those of the smallest specimen of H.
stephensoni, new species, examined for this study, the
eyes of the two species differ in shape when two speci-
mens of similar size are compared. The lobes of the
cornea are more inflated in H. stephensoni and the
eye appears smaller in that species.
I have previously assumed that records of H. raphi-
dea from Japanese waters are referable to H. harpax,
for H. raphidea is not known to occur that far north-
ward. However, some of the specimens reported in
the literature may well be H. japonica, new species.
Fukuda (1913) clearly shows a short rostral plate,
short marginal carinae on the telson, and relatively
strong submedian carinae on the abdomen, all of
which are characteristic of H. japonica rather than
H. harpax.
Although Rathbun (1902) reported five specimens
in the lot collected by Jordan and Snyder, only three
are now in the collections of the Division of Crustacea,
National Museum of Natural History, Smithsonian
Institution. The other two specimens probably were
returned to Stanford University and may now be
housed in the collections of the California Academy of
Sciences.
SEXUAL DEMORPHISM.?The male paratype lacks
raptorial claws, so no comparison of that appendage
can be made with the claw of the female (only one
female, the holotype, has claws). The median carina
of the telson in the male is slightly more inflated than
that of each female.
BIOLOGY.?Unknown.
DEVELOPMENT.?Unknown.
TYPES.?The holotype and two paratypes are in
the Division of Crustacea, National Museum of Nat-
ural History, Smithsonian Institution.
TYPE-LOCALITY.?Wakanoura, Kii, Japan.
NAME.?The name refers to the occurrence of the
species off Japan.
DISTRIBUTION.?Known only from Japan. Fukuda's
article is in Japanese so I cannot determine whether
he mentioned any specific localities.
4. Harpiosquilla stephensoni, new species
FIGURES 12-17
?Squilla sp. [Alima pyramidalis].?Foxon, 1932, p. 381 [larva;
? several species; identification questioned by Ingle and
Delia Croce, 1967].
Squilla raphidea.?Tate, 1883, p. 48 [?].?Stephenson, 1952,
p. 4; 1953, p. 43 [part?].?Stephenson and McNeill, 1955,
p. 239 [part] [not S. raphidea Fabricius].
Harpiosquilla harpax.?Manning, 1966, p. 87, fig. 1 [not S.
harpax de Haan].
Harpiosquilla sp.?Manning, 1968a, fig. 4.
HOLOTYPE.?1 9,315 mm; Port Curtis, Queensland,
Australia; seine net over flat; no other data; AM P -
9664.
PARATYPES.?1 $ , 220 mm; near Emery Point, Dar-
win Harbour, Northern Territory; taken in seine net
FIGURE 12.?Harpiosquilla stephensoni, new species.
Male paratype, TL 272 mm, Port Curtis: anterior portion
of body.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 13.?Harpiosquilla stephensoni, new species. Male
paratype, TL 272 mm, Port Curtis: raptorial claw (propodus
length 63.0 mm).
FIGURE 14.?Harpiosquilla stephensoni, new species. Fe-
male holotype TL 315 mm, Port Curtis: raptorial claw (pro-
podus length 79.2 mm).
off beach; Capt. E. F. Wells, col.; early 1954; AM
P-12382.?1 5 , 170 mm; same data; SI Crust. 111354.
?1 9 , 168 mm; Darwin Harbour, Northern Territory;
no other data; AM P-12770.?1 9 , 240 mm; same; in
60 ft; C. W. Holman, col.; 2 July 1962; AM P-14924.
?4 $ , 136-202 mm; 1 9 , 135 mm; southern Gulf of
Carpentaria, Queensland; 14 fms or less; CSIRO
Prawn Survey; December 1963; AM P-15503 to P -
15507.?1 $, 208 mm; southeastern Gulf of Carpen-
taria, Queensland; 14 fms or less; CSIRO Prawn Sur-
vey; December 1963; AM P-15509.?1 broken 9,
CL 51.6 mm; Cairns Inlet, northern Queensland; AM
P-9685.?1 5 , 245 mm; Port Curtis area, Queens-
land; M. Ward and W. Boardman, col.; July 1929;
AM P-14925.?1 9 , 277 mm; same; AM P-14926.?
1 $ , 272 mm; Port Curtis, Queensland; SI Crust.
125720.
DIAGNOSIS.?Size large, adults with TL to 315 mm;
antennular peduncle shorter than carapace; corneal
indices ranging between 336 and 434 (Table 1) ; rostral
plate (Figure 12) as long as or slightly longer than
broad, apex obtusely angled or rounded, not projecting
anteriorly; carapace with median carina; upper mar-
gin of propodus of claw with smaller spines and den-
ticles between largest spines; dactylus of claw with 7
teeth, outer margin of dactylus (Figure 13) with
prominent obtusely rounded projection in adult males;
fifth thoracic somite (Figure 15) angled laterally or
with lateral acute projection; last 3 thoracic somites
with submedian and intermediate carinae, intermedi-
ates unarmed; ventral keel of eighth thoracic somite
rounded; submedian carinae present on all 6 abdomi-
nal somites, abdominal carinae armed as follows: sub-
median 6, intermediate 2-6, lateral 1-6, marginal 1-5;
denticles 4-6, 6-11, 1, usually 5-6, 7-8, 1; marginal
carina less than twice as long as carina of lateral tooth;
postanal keel extending slightly more than halfway
between anus and posterior margin; at most inner half
of distal segment of uropodal exopod dark.
COLOR.?Black transverse line present on dorsum
between antennular and ophthalmic somites; antennal
scale lacking any marked concentration of dark pig-
ment; carinae, grooves, and both anterior and posterior
margins of carapace dark; display patch on dorsal sur-
face of merus yellow, with proximal dark oval bar;
last 3 thoracic and first 5 abdominal somites with dark
posterior line and irregular dark area on dorsum; telson
with pair of oval or elongate dark spots on anterior
dorsal surface; proximal portion of anterior margin of
uropod dark; proximal segment of uropodal exopod
with inner dark patch, distal segment with dark, diffuse
pigment on inner half or third; distal half of endopod
darker than proximal, but apex light
J. C. Yaldwyn made available a color slide taken by
D. F. McMichael of one specimen, AM P-15507.
Most of the pattern mentioned above is visible. The
background color of the body is ivory. The major dif-
ference from the pattern on the same specimen in pre-
servative is that there is but one anterior, elongate
patch of dark pigment on the telson; it is not broken
into two submedian patches.
NUMBER 36 19
SEE.?Males TL 136-272 mm; females TL 135-315
mm. Only H. raphidea, among the currently known
species of Harpiosquilla, grows to such a large size; it
is known to reach 335 mm in total length.
Other measurements of female holotype, TL 315
mm: carapace length 59.1; cornea width 14.4; anten-
nular peduncle length 46.7; rostral plate length 10.9,
width 9.9; raptorial propodus length 79.2; fifth ab-
dominal somite width 61.4; telson length 57.2, width
55.6.
DISCUSSION.?The short rostral plate of this spe-
cies (Figure 12), a feature which it shares with H.
annandalei and H. japonica, new species, will distin-
guish it from H. harpax, H. indica, new species, H.
melanoura, and raphidea. H. stephensoni, new species,
differs from H. annandalei in many features, includ-
ing size of adults and color pattern, but the absence of
spines on the submedian carinae of the fifth abdominal
? " . ? ? ? ? . - ? ? . . ? .
FIGURE 15.?Harpiosquilla stephensoni, new species. Male
paratype, TL 272 mm, Port Curtis: lateral processes of fifth,
sixth, and seventh thoracic somites.
somite in H. stephensoni is the best way to separate the
two species.
Harpiosquilla stephensoni, is very similar to H.
japonica, described above. However, H. japonica dif-
fers in having eight teeth on the claw, unarmed inter-
mediate carinae on the abdomen anterior to the third
somite, and the lateral process of the fifth thoracic
somite is rounded, not angled posterolaterally. The
submedian carinae of the abdomen are more promi-
nent in H. stephensoni than in H. japonica.
Although the corneal indices of H. stephensoni and
H. japonica overlap, the eyes of H. japonica appear to
be larger when specimens of similar size are compared.
Harpiosquilla japonica is probably a smaller species,
for the single male examined, TL 165 mm, shows signs
of inflation of the median carina of the telson, a sec-
ondary sexual feature characteristic of adult males.
See also the remarks under the account of H. japonica,
new species (p. 15).
Adult specimens (TL 200-225 mm or more) differ
from younger ones in having the lateral process of the
fifth thoracic somite shaped differently. In adults (Fig-
ure 15) the edge of the lateral process slopes postero-
laterally to an acute angle, the apex of which may
form a blunt tubercle or spine. The spine, if present, is
never so well developed as in H. annandalei or H.
raphidea. The lateral process of subadult specimens
is not so markedly angled posterolaterally. However,
juveniles of H. stephensoni can be distinguished by the
other features cited above.
Corneal indices for H. stephensoni, new species, are
summarized in Table 1. They overlap those of H.
harpax to some extent, but the eyes of the largest speci-
mens of H. harpax are about the same size as those of
the smallest specimens of H. stephensoni, and the lobes
of the cornea are not as inflated as in the latter species.
The eyes of the smaller specimens of H. raphidea re-
ported herein are similar in size to those of the largest
specimen of H. stephensoni, but the eyes of large speci-
mens of H. raphidea are much smaller.
Specimens recorded as S. raphidea by Stephenson
and McNeill (1955) from Darwin Harbour (2 9 ) ,
to belong to H. stephensoni. Most of the remainder of
the specimens reported by Stephenson and McNeill
are referred herein to H. harpax; one specimen proved
to be H. melanoura. It is likely that some of the
specimens from Queensland recorded as S. raphidea by
Stephenson (1952, 1953) belong to H. harpax, but I
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 16.?HarpiosquiUa stephensoni, new species. Male
paratype, TL 272 mm, Port Curtis: posterior portion of body
(telson width 45.3 mm). FIGURE 17.?Harpiosquilla stephensoni, new species. Female
holotype, TL 315 mm, Port Curtis: posterior portion of body
(telson width 55.6 mm).
include his records here because of the apparent
abundance of H. stephensoni in Queensland waters.
In 1966 I reported on a specimen of H. stephensoni
as H. harpax and noted that it differed from other
specimens available for study in several features. Two
of these, the convex lobe on the basal prolongation of
the uropod and the elongate dark spots on the telson,
have proved to be of little value. The two specimens
from off Emery Point, Darwin Harbour, show both
of these features but in other respects they appear to
be typical subadult specimens of H. stephensoni. No
other specimens have elongate spots on the telson but
several show variation in shape of the lobe between
the spines of the basal prolongation. I can only con-
clude that these features vary in this species.
The carinae of the marginal teeth of the telson in
H. stephensoni are lined dorsally with large tubercles;
the carinae appear nodulose in lateral view (Figures
16, 17). These tubercles are well defined in large speci-
mens are are not obliterated in them as in H. raphidea.
The carinae are inflated and the apices of the teeth
are upturned. In these features H. stephensoni resem-
bles H. raphidea, but the tubercles are much larger in
the latter species.
The marginal denticles of the telson are less numer-
ous, in general, than in other species of Harpiosquilla.
They are also sharper and more distinct in smaller
specimens than in the largest ones examined.
SEXUAL DIMORPHISM.?Males of TL 180 mm or
more have secondary sexual characteristics similar to
those found in some other species. There is a pro-
nounced lobe, often irregular in outline, on the outer
margin of the dactylus of the claw (Figure 13); it is
not so sharply angled as in males of H. raphidea. The
bases of the second to fifth dactylar teeth are inflated.
The submedian carinae of the sixth abdominal somite
and the median carina of the telson are also inflated in
adult males (Figure 16), but the median carina of the
telson in large females may also be very swollen. Claws
(Figures 13, 14) and telsons (Figures 16,17) of a large
male and female have been illustrated.
BIOLOGY.?Relatively little information is available.
Specimens have been taken in shallow water, 25 meters
or less; three specimens were taken in seine nets. The
range of the species in Australia overlaps that of H.
harpax but whether both species occur together is not
known.
One of the specimens from the southern Gulf of
Carpentaria, AM P-15507, has numerous spherical
egg cases, approximately 1.5 mm in diameter, attached
to both the dorsal and ventral surfaces of the body;
they are most numerous on the posterior portion. Many
of the capsules had ruptured, but one was found to
contain three small gastropods, each about 1.1 mm
long. Commensals have not been recorded from mem-
bers of this genus, and egg cases of other organisms are
NUMBER 3 6 21
rarely found on stomatopods. At the time of this writ-
ing, the egg cases had been identified tentatively by Dr.
Joseph Rosewater, Division of Mollusks, Smithsonian
Institution, as those of marginellid gastropods. Both
marginellids and Harpiosquilla are burrowers, and
possibly the adult gastropod used the Harpiosquilla in
its burrow (?) as a substrate for attachment of its egg
capsules.
DEVELOPMENT.?Larvae of this species are not
known with certainty. Foxon (1932) reported on
larvae from the Great Barrier Reef which she identi-
fied with Alimerichthus pyramidalis Lanchester; she
also synonymized A. unidens Lanchester, 1903, and
Alimerichthus a Tattersall, 1906, with A. pyramidalis.
A. pyramidalis was subsequently identified with H.
raphidea by Alikunhi (1952) who also pointed out that
A. unidens and Tattersall's larva as well could be at-
tributed to other species. Alikunhi (1952, p. 268) also
noted: "That Foxon was dealing with a collection of
species under the name A. pyramidalis is, therefore,
obvious. Since larvae measuring 18 mm in length are
reported in this collection, it is likely that S. raphidea
is also represented in it."
However, Ingle and Delia Croce (1967), who re-
examined Foxon's material, stated (p. 67) :
Alikunhi (1952, p. 265) attributes Foxon's (1932, p. 381)
material of Alima pyramidalis (Lanchester) to the adult
species Squilla raphidea Fabricius. These particular speci-
mens in the B.M. (N.H.) Collections do not quite correspond
to Alikunhi's description and figures as most of them have
spinule no. 5 . . . poorly developed and one or more teeth on
the inner margin of the raptorial dactylus.
The possibility remains that Foxon was dealing with
some larvae of either H. harpax or H. stephensoni, new
species, both of which are common on the coast of
Australia.
TYPE.?The holotype and most of the paratypes are
in the Australian Museum, Sydney. One male and one
female paratype are in the collections of the Division
of Crustacea, National Museum of Natural History,
Smithsonian Institution.
TYPE-LOCALITY.?Port Curtis, Queensland, Aus-
tralia.
NAME.?The species is named for Professor W.
Stephenson of the University of Queensland who has
been instrumental in the development of our knowl-
edge of Australian stomatopods.
DISTRIBUTION.?Known only from the coastal
waters of Australia, where it has been taken from
scattered localities between Darwin, Northern Terri-
tory, and Port Curtis, Queensland; shallow water to
14 fathoms. The following records in the literature
may be based on material of this species. AUSTRALIA:
Darwin, Northern Territory (Manning, 1966); prob-
ably Port Darwin, Northern Territory (Tate, 1883) ;
Queensland (Stephenson, 1952); Cooktown, Queens-
land (Stephenson, 1953); Tambian Beach, Keppel
Bay, Queensland (Stephenson and McNeill, 1955) ;
Peel Island, Moreton Bay, Queensland (Stephenson,
1953); Woody Point Pier, Moreton Bay, Queensland,
10-11 m (Stephenson, 1952); 3 miles south of Woody
Point Pier, Moreton Bay, Queensland, 6 m (Stephen-
son, 1952); 4 miles east of Scarborough Pier, Moreton
Bay, Queensland, 10 m (Stephenson, 1952); and
Southport, Queensland (Stephenson, 1953).
Stephenson (1952, 1953) may also have had H.
harpax represented in his collections from Queensland.
Larvae of this species from the Great Barrier Reef
may have been reported by Foxon (1932) as Alima
pryramidalis.
5. Harpiosquilla melanoura Manning, 1968
FIGURES 18-27
Squilla raphidea.?Stephenson and McNeill, 1955, p. 239
[part] [not S. raphidea Fabricius].
Harpiosquilla melanoura Manning, 1968b, pp. 14 [key] 18,
fig. 5.
MATERIAL.?1 $ , 161 mm; Rose Bay, Port Jackson,
New South Wales, Australia; in prawn net; 10 March
1925; AM P-7929.?1 9 , 151 mm; Andaman Sea,
Mergui Archipelago; 09?13'N, 97?51'E; 58-60 m;
ANTON BRUUN Cruise 1, Sta. 20; IIOE; 23 March
1963; SI Crust. 125719.?19, 128 mm; Andaman
Sea, off Burma; 14?07'N, 97?05'E; 69-73 m; ANTON
BRUUN Cruise 1, Sta. 38; IIOE; 30 March 1963; SI
Crust. 125358.
In addition to these specimens I have examined the
type-series, 2 S , TL 102-129 mm, 2 9 , TL 126-136
mm, from the Bane de Pracel, western coast of
Madagascar.
DIAGNOSIS.?Size moderate, adults with TL less
than 170 mm; antennular peduncle not so long as cara-
pace and rostral plate combined; corneal indices rang-
ing between 229 and 271 (Table 1) ; rostral plate
(Figure 19) longer than broad, lateral margins taper-
ing to slender median projection; carapace without
median carina; upper margin of propodus of claw with
smaller spines and denticles between largest spines;
22
FIGURE 18.?Harpiosquilla melanoura Manning. Female,
TL 151 mm, Mergui Archipelago: dorsal view.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
dactylus of claw with 8 teeth, outer margin of dac-
tylus with prominent angular projection in adult males;
fifth thoracic somite (Figure 21) rounded laterally;
posterior 3 thoracic somites lacking submedian carinae;
intermediate carinae of thoracic somites unarmed;
ventral keel of eighth thoracic somite rounded; first to
fifth abdominal somites lacking submedian carinae,
abdominal carinae spined as follows: submedian 6,
intermediate (2) 3-6, lateral 1-6, marginal 1-5; denti-
cles 5-7, 9-12, 1; marginal carina of telson almost 3
times as long as carina of lateral tooth; postanal keel
not extending halfway between anus and posterior
margin; distal segment of uropodal jet black.
COLOR.?Inner and outer surfaces of antennular
peduncles with distal black spot on each segment; an-
terolateral angle of antennal scale dark, anterointernal
angle dark in some specimens; basal portion of rostral
plate with oval area outlined with dark pigment; an-
terolateral margins, gastric grooves, and posterior
median margin of carapace dark, median area, anterior
to cervical groove, with dark oval patch; merus of claw
with 3 dark spots, 1 at inner distal margin, 1 on proxi-
mal edge of dorsal depression, and 1 subdistally on
outer surface; propodus of claw with inner and outer
dark spot; posterior 3 thoracic and anterior 5 ab-
dominal somites (sixth in some specimens) with pos-
terior black lines; second abdominal somite with thin,
transverse dark bar, traces of dark pigment also present
medially on third to fifth somites; dorsal surface of
telson with proximal submedian pair of oval dark spots,
surface pits also dark; ventral surface of telson dark
along midline, with dark patch at base of each tooth;
anterior edge of basal segment of uropod dark; distal
portion of proximal segment of uropodal exopod dark,
distal segment of exopod jet black; distal half or two-
thirds of endopod dark. The overall color pattern is
shown in Figure 18.
SIZE.?Only male examined, TL 161 mm; females
TL 128-151 mm. Including the type-series from Mada-
gascar, males are known to range between TL 102-161
mm, females between TL 126-151 mm.
Other measurements of a female, TL 151 mm:
carapace length 31.2; cornea width 11.5; antennular
peduncle length 31.7; rostral plate length 6.4, width
5.0; raptorial propodus length 43.4; fifth abdominal
somite width 32.2; telson length 28.5, width 27.6.
DISCUSSION.?Harpiosquilla melanoura is the only
species in the genus lacking both the median carina of
the carapace and the submedian carinae of the last
NUMBER 3 6 23
FIGURES 19-23.?Harpiosquilla melanoura Manning. Male holotype, TL 129 mm, Bane de
Pracel, Madagascar: 19, anterior portion of body; 20, eye; 21, lateral processes of exposed
thoracic somites; 22, last two abdominal somites, telson, and uropod; 23, uropod, ventral view
(setae omitted; from Manning, 1968b).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
three thoracic and first five abdominal somites. It fur-
ther differs from all other species of Harpiosquilla now
known in having the entire surface of the distal seg-
ment of the uropodal exopod jet black in color.
Harpiosquilla melanoura most closely resembles H.
indica, new species, described herein, but the latter
species has a well-developed median carina on the
carapace. Other differences are discussed under H.
indica (p. 33).
Corneal indices for H. melanoura are summarized
in Table 1. The eyes of this species are similar in size
to those of H. harpax but are slightly larger and more
inflated.
The marginal carinae of the telson are compara-
tively longer in H. melanoura than in other species of
Harpiosquilla for each is almost three times as long
as the carina of the lateral tooth. H. harpax is the
only other species in which the marginal carina is
more than twice as long as that of the lateral tooth.
Of the seven specimens of this species now known,
only one has the intermediate carinae of the second
abdominal somite armed posteriorly. In the other
specimens the intermediate carinae of the third to sixth
somites only are provided with posterior spines.
SEXUAL DIMORPHISM.?Large males (TL 129 mm
or more) (Figure 24) have a well-developed obtuse
projection on the outer margin of the dactylus of the
claw and the bases of individual teeth on the claw
are noticeably inflated. Claws of a male and a female
are shown in Figures 24 and 25. Males also differ from
females in having the submedian carinae of the sixth
abdominal somite and the median carina of the tel-
son more inflated (Figures 26,27).
BIOLOGY.?Essentially unknown. Most specimens
have been taken in depths between 55 and 73 meters;
they have not been taken in the littoral habitat as have
both H. harpax and H. raphidea. Specimens from
Madagascar were taken on sand or muddy sand.
Harpiosquilla melanoura was taken along with H.
annandalei at ANTON BRUUN Sta. 20 off the Mergui
Archipelago.
DEVELOPMENT.?Unknown.
TYPES.?Division of Crustacea, National Museum
of Natural History, Smithsonian Institution. Paratypes
are in the Smithsonian Institution and the Museum
National d'Histoire Naturelle, Paris.
TYPE-LOCALITY.?Bane de Pracel, western coast of
Madagascar.
DISTRIBUTION.?Indo-West Pacific region, from
Rose Bay, Port Jackson, New South Wales, Australia
(Stephenson and McNeill, 1955, as S. raphidea);
Mergui Archipelago; off Burma; and the Bane de
Pracel, off the western coast of Madagascar (Manning,
1968b). It has not been recorded previously east of
Madagascar.
FIGURE 24.?Harpiosquilla melanoura Manning. Male, TL
161 mm, Port Jackson, New South Wales: raptorial claw
(propodus length 39.0 mm).
FIGURE 25.?Harpiosquilla melanoura Manning. Female,
TL 151 mm, Mergui Archipelago: raptorial claw (propodus
length 43.4 mm).
NUMBER 3 6 25
FIGURE 26.?Harpiosquilla melanoura Manning. Male, TL
161 mm, Port Jackson, New South Wales: posterior portion
of body (telson width 30.0 mm).
6. Harpiosquilla harpax (de Haan, 1844)
FIGURES 28-38
Squilla harpax de Haan, 1844, atlas, pi. 51 (fig. 1); 1849,
text, p. 222.?Tiwari and Biswas, 1952, p. 358, figs. 3b,
d, f.?Barnard, 1955, p. 49?Alikunhi, 1959, p. 132, fig-
13 [larva].?Ingle, 1963, pp. 14, 18, figs. 9, 59.?Holthuis,
1964, p. 140 [listed].?Manning, 1968a, p. 121 [listed],
fig. 3a.
Squilla raphidea.?Berthold, 1845, p. 47; 1847, p. 29.?
White, 1847, p. 84.?Miers, 1880, p. 27 [part].?Bigelow,
1894, pp. 511 [key], 535.?de Man, 1898, p. 694 [part ?].?
Tattersall, 1906, p. 166.?Balss, 1910, p. 8, fig. 2a.?Kemp,
1913, p. 88, pi. 7 (fig. 77) [part].?Parisi, 1922, p. 103.?
Komai, 1927, p. 323.?Komai, Akatsuka, and Ikari, 1927,
p. 295.?Komai and Ikari, 1929, p. 121.?Gravier, 1930,
p. 525.?Chopra, 1934, p. 27 [part].?Serene, 1937, p. 68
[listed].?Komai, 1938, p. 268.?Suvatti, 1938, p. 52.?
Chopra, 1939, p. 158.?Holthuis, 1941, p. 256 [part].?
Liu, 1949, p. 43, pi. 6 (figs. 15-17).?Anonymous, 1949,
p. 843, fig. 2421.?Barnard, 1950, p. 851, figs, lc, g.?
Suvatti, 1950, p. 132.?Dawydoff, 1952, p. 145.?Serene,
1953, p. 507; 1954, pp. 6, 8, 62, pi. 4 (figs. 1-6).?
Millard and Harrison, 1954, p. 176 [listed].?Stephenson
and McNeill, 1955, pp. 239 [part], 255 [key].?Day and
FIGURE 27.?Harpiosquilla melanoura Manning. Female,
TL 151 mm, Mergui Archipelago: posterior portion of body
(telson width 27.6 mm).
Morgans, 1956, p. 306 [listed].?Utinomi, 1956, p. 91,
pi. 46 (fig. 2) [color figure].?Yamaji, 1959, fig. on p.
68.?Utinomi, 1960, p. 114, pi. 57 (fig. 7) [color figure].?
Chuang, 1961, pp. 181, 206, pi. 81 (fig. 4), lowest fig. on
pi. 82.?Stephenson, 1962, p. 34.?Crosnier, 1965, p. 61
[listed]. [All not Squilla rap hide a Fabricius.]
Squilla obsoleta White, 1847, p. 84 [nomen nudum; in
synonymy].
Squilla Harpax.?Herklots, 1861, p. 152 [p. 39 on separate].
Squilla raphidea van?Gravier, 1937, p. 186, figs. 8-10.
Squilla gilesi.?Boone, 1938, pp. 201, 203, pi. 71 [not S.
gilesi Kemp].
Squilla raphidoea.?Serene, 1939, p. 349 [erroneous spelling].
?Squilla sp. [Alima multispinus].?Foxon, 1939, p. 258, fig.
2B [larva] [not Erichthus multispinus Claus, 1871].
?Squilla sp.?Gohar and Al-Kholy, 1957, p. 106 [larvae; part,
several species mixed ?].
Squilla hanpax.?Alikunhi, 1959, p. 132 [discussion; erro-
neous spelling].
Harpiosquilla harpax.?Lee and Wu, 1966, p. 51, figs.
7A-F.?Manning, 1967, p. 103.?Holthuis, 1967, p. 15.?
Chhapgar and Sane, 1968, p. 45 [key].?Manning, 1968b,
pp. 14 [key], 15, fig. 4.?Tirmizi and Manning, 1968, p.
33, fig. 13.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
?Alima stages of Harpiosquilla raphidea-harpax group.?
Ingle and Delia Croce, 1967, p. 6, figs. 19, 24 [larvae].
not Harpiosquilla harpax.?Manning, 1966, p. 87, fig. 1
[=H. stephensoni, new species].
MATERIAL.?1 3 , 152.5 mm; Japan; lectotype of S.
harpax; RMNH 28H.?1 3,175 mm; China or Japan;
u.s.s. PALOS; SI Crust. 5146.?1 3 , CL 20.2 mm; be-
tween Kobe and Yokahama, Japan; 34?06'15"N,
134?57'50"E; 40-37 fms; fine gray shell mud; ALBA-
TROSS Sta. 4963; 27 August 1906; SI Crust. 76058.?
1 3 , 85 mm; Takao, Taiwan; F. Baker, col.; December
1914; SI Crust. 47935.?1 3 ,109 mm; same; SI Crust.
47928.?13, 146 mm; Tungkang market, Taiwan;
40-50 fms; S. Lee and S. K. Wu; 5 March 1965; SI
Crust. 113645.-1$, 215 mm; Hong Kong; North
Pacific Exploring Expedition; SI Crust. 2108.?13,
137 mm; Makung, Penghu Island, Pescadores Islands,
China Sea; 19 April 1961; SI Crust. 124762.?1 3 ,
132 mm; Nhatrang Bay, Viet-Nam; DE LANESSAN,
col.; 1930; SI Crust. 124721.?2 3 (1 broken), 132
mm; 2 9 (1 broken), 109 mm; Sandakan Fish Market,
North Borneo; Chin Phui Kong, col.; 5 November
1956; AM P-14923.?1 broken 9, CL 39.5 mm; off
North East Island, near Groote Eylandt, western Gulf
of Carpentaria, Northern Territory, Australia; 25 fms;
CSIRO Prawn Survey; 4 September 1963; AM P -
15508.?19, 248 mm; Tin Can Bay, near Gympie,
Queensland, Australia; taken in net by C. Lea; 7 July
1952; AM P-12256.?1 broken 3 , CL 36.2 mm; Cape
Flattery, Queensland, Australia; AM G-1792.?19,
225 mm; The Basin, Newcastle Harbour, New South
Wales, Australia; from prawn trawl; A. A. Racek, col.;
27 May 1954; AM P-12512.?13, CL 29.6 mm;
Hawkesbury River, N.S.W., Australia; hooked by line
fisherman; March 1952; AM P-12230?1 broken S,
CL 27.5 mm; Bobbin Head, near Jerusalem Bay,
Hawkesbury River, N.S.W., Australia; taken from
stomach of fish, flathead; May 1952; AM P-12229.?
IS, 188 mm; Jerusalem Bay, Hawkesbury River,
N.S.W., Australia; hooked by line fisherman; 18 May
1952; SI Crust. 124765.?19, 143 mm; Jerusalem
Bay, Hawkesbury River, near Sydney, N.S.W., Austra-
lia; R. Woodger, col.; AM P-15399.?19, 180 mm;
Jerusalem Bay, Hawkesbury River, near Sydney,
N.S.W., Australia; handline from 30 ft of water on
hook baited with small fish; A. B. Iverach, col.; 24
January 1965; AM P-15402.?19, 175 mm; Cowan
Creek, Hawkesbury River, N.S.W., Australia; 1 Feb-
ruary 1953; SI Crust 124766.?13, 165 mm; same,
from stomach of 71 lb flathead fish; March 1960; AM
P-13324.?1 9 , 190 mm, same; handline and baited
hook; L. Kelly, col.; 23 April 1960; AM P-15400.?
1 9 , 185 mm; same; handline and baited hook;
A Leslie, col.; 18 March 1961; AM P-15401.?
19, 195 mm; same; handline from 15 ft water on
small hook baited with prawn; 26 April 1965; AM
P-15403.?IS, 110 mm; Lem Sing, Thailand; 10
January 1924; SI Crust. 124761.?1 9 , 168 mm;
Tachalom, Gulf of Siam; trap; Hugh M. Smith; 21
July 1923; SI Crust. 69533.?19, 157 mm; South
Kang Kradan, Thailand; Hugh M. Smith; 1923; SI
Crust. 69534.?19, 153 mm; beach north of Singora,
Thailand; Hugh M. Smith; 5 October 1923; SI Crust.
69535.?IS, 87 mm; Bay of Bengal; 15?04'N,
95?51'E; 29?33 m; ANTON BRUUN Cruise 1, Sta. 41A;
IIOE; 31 March 1963; SI Crust. 125356.?1 9 , 153
mm; Sandheads, mouth of River Hooghly [Hughli],
India; LADY FRASER; SI Crust. 119312.?IS, 120 mm;
2 9 , 129-173 mm; India; 17?54-57'N, 72?23-27'E;
46-55 m; green mud, shells; ANTON BRUUN Cruise 4B,
Sta. 201A; IIOE; 13 November 1963; SI Crust.
125352.?23, 71-146 mm; 29 , 122-164 mm; off
Bombay, India; 18?47.3-49/N, 72?36-38'E; 12.5-14
fms; M.V.F. JANJIRA; Field No. 9682; I IOE; 12 May
1964; SI Crust. 125354.?73, 72-120 mm; 5 9 ,
64-140 mm; India; 20?30/N, 70?50-54/E; 33 m;
brown sticky mud; ANTON BRUUN Cruise 4B, Sta 204A;
IIOE; 15 November 1963; SI Crust. 125355.?1 9 ,
165 mm; northwestern India; 22?52-54'N, 68?34-
36'E; 15 m; soft mud; ANTON BRUUN Cruise 4B, Sta.
223A; IIOE; 19 November 1963; SI Crust. 125353.?
19, 148 mm; Gulf of Oman; 26?10-13'N, 57?02'E;
55-64 m; green mud; ANTON BRUUN Cruise 4B, Sta.
256A; IIOE; 30 November 1963; SI Crust. 125351.?
5 3 , 139-166 mm; 8 9 , 134-185 mm (in 2 lots);
Archico Bay, south of Massawa, Ethiopia, Red Sea;
Israel South Red Sea Exped., Sta. E62/4118; 8 April
1962; SI Crust. 124763,124764.
DIAGNOSIS.?Size moderate to large, TL to 248 mm;
antennular peduncle usually shorter than carapace;
corneal indices ranging between 251 and 388 (Table
1); rostral plate longer than broad (Figure 31), with
slender anterior projection; carapace with median
carina; opposable margin of propodus of claw with
smaller spines and denticles between largest spines;
dactylus of claw with 8 teeth, outer margin of dactylus
with prominent angular projection in adult males;
NUMBER 3 6 27
fifth thoracic somite (Figure 32) rounded laterally;
posterior 3 thoracic somites with submedian and inter-
mediate carinae, none armed posteriorly; ventral keel
of eighth thoracic somite rounded; abdomen with sub-
median carinae on all somites, abdominal carinae
spined as follows: submedian 6, intermediate (1)
2-6, lateral 1-6, marginal 1-5; denticles 4-6, 10-12
(14), 1; marginal carina of telson more than twice as
long as carina of lateral tooth; postanal keel extending
about halfway between anus and posterior margin;
distal segment of uropodal exopod with inner half
black.
COLOR.?In preservative, black transverse line on
dorsum between antennular and ophthalmic somites;
antennal 6cale with diffuse dark pigment, concentrated
along anterior margin; carinae, grooves, and anterior
and posterior margin of carapace dark, pair of dark
spots also present on lateral plates of carapace at level
of cervical groove; display patch on dorsal surface of
merus yellow, with proximal oval dark patch and
distal dark spot; last 3 thoracic and first 5 abdominal
somites with dark posterior line; telson with pair of
dark circles or square spots on anterior dorsal surface;
distal segment of uropodal exopod with diffuse dark
chromatophores on inner half, distal third of proximal
segment also dark; distal half of endopod dark, with
light midrib in some specimens; apices of teeth of
uropods and telson yellow in some specimens. The
overall color pattern is shown in Figure 28.
The notes on color in life given below from Komai
(1927, p. 323) and Utinomi (1956, 1960) may refer
to this species but may also refer to H. japonica, new
species.
When fresh, the entire surface is pale and slightly ochrace-
ous, besprinkled with minute dark spots. A pinkish color is
found on the carapace, raptorial limbs and on the thoracic
and abdominal somites. A conspicuous round patch of chest-
nut-brown color occurs on either side of the base of the mid-
dorsal carina of the telson.
According to Utinomi (1956) this species is pinkish,
setae of scale and uropod yellow, telson with pair of
proximal red spots. He (1960, p. 114) also reports:
Antennal scale blue, setae yellow; propodus yellow distally,
apices of teeth of chela yellow; submedian carinae of first
to fifth abdominal somites and sixth to eighth thoracic somites
yellow, lateral carinae on body with more red; submedian
carinae of sixth abdominal somite green; median carina of
telson and pits green; carinae of marginal teeth green, apices
yellow; most of uropod yellow.
Tirmizi and Manning (1968, p. 33) reported the
following color pattern in fresh specimens from Ka-
rachi, West Pakistan:
FIOURE 28.?Harpiosquilla harpax (de Haan). Male, TL
188 mm, Jerusalem Bay, New South Wales: dorsal view.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURES 29-34.?Harpiosquilla harpax (de Haan). Female, TL 118 mm, Ambaro Bay, Mada-
gascar: 29, anterior portion of body; 30, outline of eye; 31, outline of rostral plate; 32, outline
of lateral processes of exposed thoracic somites; 33, last two abdominal somites, telson, and
uropod; 34, uropod, ventral view. (Setae omitted; from Manning, 1968b).
NUMBER 36 29
. . . eyestalks bright yellow; antennal peduncle and scale
with pink patches; carinae of carapace lined with dark spots;
merus and carpus of raptorial claw with pink patches; pos-
terior margin of each of thoracic and abdominal somites
lined with black spots; lateral portions of abdominal somites
pink; sixth abdominal somite pink between submedian cari-
nae; spines on abdomen, telson, and uropod yellow; telson
with grey pits and carinae except for red median carina;
proximal submedian patches of telson maroon (black in
preservative); base of uropod pink.
SIZE.?Males, TL 71-188 mm; females, TL 64-248
mm. Males ranging in TL between 80 and 195 mm and
females TL 112 to 215 mm have been recorded in the
literature (size data from corroborated distribution
records only).
Other measurements of male lectotype, TL 152.5
mm: carapace length 29.1; cornea width 9.2; anten-
nular peduncle length 28.1; rostral plate length 5.0,
width 4.8; raptorial propodus length 38.4; fifth ab-
dominal somite width 30.6; telson length 29.4, width
26.2.
DISCUSSION.?Harpiosquilla harpax, the common-
est and most widely distributed species of the genus,
can be distinguished with ease from the other species.
The apical median projection of the rostral plate at
once separates it from H. annandalei, H. japonica,
new species, and H. stephensoni, new species; in all
of those species the rostral plate is blunt apically, either
broadly rounded or obtusely rounded. Harpiosquilla
harpax is a smaller species than H. raphidea, and the
carinae on the body are not nearly so well developed
as in that species; in H. harpax the intermediate cari-
nae of the last three thoracic somites are unarmed
whereas they are provided with spines in H. raphidea,
and the lateral process of the fifth thoracic somite is
rounded in H. harpax, spined laterally in H. raphidea.
The faint submedian carinae on the abdomen will dis-
tinguish H. harpax from both H. indica, new species,
and H. melanoura; the latter species lacks the median
carina on the carapace as well. In H. indica, the in-
termediate carinae of the first and second abdominal
somites are unarmed; the carinae of the second ab-
dominal somite are always armed in adults of H.
harpax.
The color pattern of H. harpax, in preserved speci-
mens, is very similar to that of both H. indica and
H. stephensoni, but very distinct from those of H.
annandalei and H. melanoura. In H. harpax, the inner
half of the distal segment of the uropodal exopod is
covered with diffuse dark pigment. In H. annandalei
that segment is black with a white midrib and in H.
melanoura it is entirely black. The color pattern of the
uropod is not nearly so well marked in H. raphidea as
in any of these species. The color pattern of H. japon-
ica, is not known.
The shape and length of the marginal carinae of
the telson are also distinctive in H. harpax which has
a slender marginal carina over twice as long as the
carina of the intermediate tooth. In H. melanoura the
marginal carina is three times as long as the intermedi-
ate, and in the remainder of the species it is less than
twice as long.
Harpiosquilla harpax is not n small species but it
apparently does not get nearly so large as do either
H. raphidea or H. stephensoni. The maximum ob-
served size in H. harpax is 248 mm in contrast with
335 mm for H. raphidea and 315 mm for H.
stephensoni.
The intermediate carinae of the first abdominal
somite were armed in only three of these specimens,
a female from Luzon, a female from India, and a
male from Viet-Nam. In addition, the small specimen
(CL 20.2 mm) from Japan lacked spines on the inter-
mediate carinae of the first two abdominal somites. All
but one of the specimens from the collections of the
Australian Museum had both intermediate carinae of
the first abdominal somite unarmed posteriorly. In all
other respects the specimens from Australia agree well
with the remainder of the specimens from other local-
ities available for study.
Holthuis (1967) noted that the anterior projection
of the rostral plate was a variable feature in his speci-
mens from the Red Sea. There is also some variation
in relative length of the anterior projection in speci-
mens from Australia. The plate in H. harpax, however,
never terminates so bluntly as it does in H. annandalei,
H. stephensoni, and H. japonica.
All available specimens of H. harpax were examined
to determine the extent of variation of the corneal
indices, if any. Although the samples from any area
were small, no obvious variation by sex, size, or loca-
tion could be detected. The eye is remarkably uni-
form in size and shape over a wide geographic area.
Corneal indices of all species are summarized in
Table 1.
All specimens examined by me had eight teeth on
the dactyl us of the claw. Parisi (1922) recorded one
specimen from Suruga Gulf, Japan, with but seven
teeth on one claw.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 35.?Harpiosquilla harpax (de Haan). Male, TL
188 mm. Jerusalem Bay, New South Wales: raptorial claw
(propodus length 42.9 mm).
FIGURE 36.?Harpiosquilla harpax (de Haan). Female, TL
175 mm, Cowan Creek, New South Wales: raptorial claw
(propodus length 45.9 mm).
It has been pointed out already that Alikunhi's
(1967) excellent account of growth is probably based
on both H. harpax and H. raphidea. Certainly some
of the observations made by Alikunhi on changes with
age apply to this species as well as to H. raphidea.
SYNONYMY.?It has been very difficult to determine
which of the early records of S. raphidea actually be-
longed to H. harpax. Several criteria have been used
but only a few records could be checked so most of the
references referred herein to H. harpax require
verification.
Geography has been the basis for assigning some
records of H. harpax rather than to H. raphidea. The
latter species is not known to occur with certainty as
far north as China or Japan, as far south as Australia,
or as far west as South Africa. Most locality records
from Japan for H. raphidea probably are referable to
H. harpax, but some may be referable to H. japonica,
new species. Most records from Japan require
verification.
In some cases I have examined material previously
recorded in the literature, although only in the case
of the collections from the Australian Museum and
the Smithsonian Institution have I been able to ex-
amine the bulk of the collection. Some of the material
recorded by the following authors has been examined:
de Haan, 1844, 1849; Holthuis, 1941, 1967; Lee and
Wu, 1966; Boone, 1938; Gravier, 1930; Millard and
Harrison, 1954; Stephenson and McNeill, 1955. I
have accepted the records of Tiwari and Biswas (1952)
and those of Kemp (1913) and Chopra (1934) which
they corroborated. The records of Ingle (1963) and
the earlier records corrected by him have also been
assigned to H. harpax.
The numerous records from Viet-Nam by Gravier
(1930), Serene (1937, 1939, 1953, 1954) and
Dawydoff (1952) are assigned here for two reasons.
Their accounts concur with my observations on H.
harpax rather than H. raphidea, and we have no au-
thenticated records of H. raphidea from east of Borneo
and north of Thailand. Serene's (1954) observations
on sexual dimorphism agree well with what I have
observed in H. harpax. All of these authors, however,
could have been dealing with both species.
Gravier (1937) recorded three specimens from the
Baie d'Along [Tonkin], Viet-Nam, as S. raphidea var.
NUMBER 3 6 31
His illustrations clearly show that his specimens are
referable to H. harpax, for the thorax and telson of
the illustrated specimen are typical of that species.
De Man (1898) noted that a small male, TL 11 cm,
from the Java Sea had poorly developed submedian
carinae on the abdomen and lacked the spine on the
fifth thoracic somite; he attributed these features to
individual aberration. He was almost certainly deal-
ing with H. harpax in this case; he may also have
examined some H. raphidea.
Gravier (1937) also commented that another speci-
men collected by Dawydoff differed from his material
in several features. I cannot determine from his ac-
count the distinguishing features of DawydofFs
material.
Tattersall (1906) reported that a male from Ceylon
had an obtuse lateral process on the fifth thoracic
somite; he, too, was probably dealing with H. harpax.
Judging from the size of specimens reported from
Queensland, Australia, TL to 222 mm, some of the
specimens identified by Stephenson (1952, 1953) as
S. raphidea may actually be referable to H. stephen-
soni, new species, and H. harpax.
Suvatti's records of S. raphidea from Thailand
(1938, 1950) are apparently based on collections made
there by Hugh M. Smith. Material from the localities
given by Suvatti is in the Smithsonian Instituion and
is clearly referable to H. harpax. Although Smith also
collected H. raphidea in Thailand, those specimens
were not recorded by Suvatti.
The specimens from Singapore reported by Boone
(1938) as S. gilesi are almost certainly H. harpax;
although a lateral spine is shown on the fifth thoracic
somite by Boone (op. cit., pi. 72), the specimens ac-
tually lack lateral spines. It seems likely that the ven-
tral spines on that somite are exaggerated in the figure.
Boone's records have been corrected by both Holthuis
(1941) and Manning (1967).
The specimen figured by Chuang (1961) on his
plate 82 shows no trace of submedian carinae on the
abdomen or of a lateral spine on the fifth thoracic
somite. For these reasons I have assigned his records to
H. harpax, which has very low submedian carinae on
the abdomen. Chuang (1961, p. 181) commented on
the presence of middorsal carinae on the last two tho-
racic somites as a distinguishing feature of S. raphidea;
none of the species of Harpiosquilla are ornamented
with middorsal carinae on either the thorax or
abdomen.
FIGURE 37.?Harpiosquilla harpax (de Haan). Male, TL
188 mm, Jerusalem Bay, New South Wales: posterior portion
of body (telson width 33.9 mm).
FIGURE 38.?Harpiosquilla harpax (de Haan). Female, TL
175 mm, Cowan Creek, New South Wales: posterior portion
of body (telson width 32.2 mm).
32
Liu (1949), on the other hand, shows strong sub-
median carinae on the thorax and abdomen and short
marginal carinae on the telson, but mentions that the
fifth thoracic somite has no lateral process; an apical
projection is shown on the rostral plate. It seems likely
that the submedian carinae of the body and the carinae
of the telson were exaggerated in the illustration and
that he was dealing with H. harpax.
SEXUAL DIMORPHISM.?Males of H. harpax exhibit
the secondary sexual characters also found in most
other species of Harpiosquilla. The obtuse prominence
on the outer margin of the claw in adult males can be
observed at a size of about 120 mm TL (CL ca. 22
mm) (Figure 35). It becomes larger and more angu-
lar with increasing age. It is not so well developed in
H. harpax at TL 150 mm as it is in H. indica, new
species. The bases of the second through the sixth dac-
tylar teeth are inflated, as illustrated by Serene (1954,
pi. 4, fig. 4). The submedian carinae of the sixth ab-
dominal somite and the median carina of the telson
may be inflated in adults of both sexes; often the me-
dian carina of the telson is more inflated in females
than in males (Figures 37, 38). The carinae of the
marginal teeth of the telson are not markedly inflated
in adults of either sex.
BIOLOGY.?As is the case in the other species of
Harpiosquilla, we can glean relatively little informa-
tion on the biology of H. harpax from the literature and
from specimen-associated data. It is a widely distrib-
uted species in the Indo-West Pacific region, and it
occurs in shallow water, between 2 and 93 meters.
Chuang (1961) recorded it from the sublittoral zone
in Singapore. It apparently has a preference for mud
bottom, for recorded bottom types include green mud,
shell; brown sand; grey sand; brown sticky mud; soft
mud; green mud; muddy sand; and sandy shore. It
was taken together with H. annandalei in 55-64 meters
in the Gulf of Oman by the ANTON BRUUN ; in general,
it occurs in shallower water than does H. annandalei.
Harpiosquilla harpax has been recorded from fish
markets from three areas, the Tungkang Market, Tai-
wan (Lee and Wu, 1966), the Sandakan Fish Market,
Borneo, and Fish Harbour, Karachi, West Pakistan
(Tirmizi and Manning, 1968). In Pakistan, at least,
it is not sought for food but is brought to shore along
with the trash from commercial shrimping operations.
The specimens from Australia were taken in prawn
nets, on hook and line by fishermen, and from the
stomach of a fish known locally as flathead. It may
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
enter water of lowered salinity, for it seems to be abun-
dant in different sections of the Hawkesbury River,
New South Wales.
DEVELOPMENT.?The larvae of H. harpax are not
known with certainty. Foxon (1939) reported larvae
from the Red Sea, the northern Arabian Sea, and the
Gulf of Oman; she noted that the larvae were of two
types, with one (typical form) or two (atypical form)
laterally directed spines on the lateral margin of the
carapace. Both forms occurred in all three areas. She
suggested that these might be the larvae of raphidea
and annandalei, and pointed out the similarity of the
larva reported by her to Erichthus multispinus Claus.
In his discussion of the development of raphidea,
Alikunhi (1952, p. 269) noted that "The atypical form
of Alima multispinous described by Foxon (1939) also
shows close similarity to the final pelagic larva of
S. raphidea," differing in having one rather than two
laterally directed spinules on the carapace. Alikunhi
further pointed out that the typical form reported by
Foxon differed from known Harpiosquilla larvae in
having four free teeth on the dactylus of the claw.
Alikunhi then suggested that the atypical form
might be identified with raphidea whereas the typical
form might be the larva of annandalei. Ingle and Delia
Croce (1967, p. 67) reexamined Foxon's specimens
and stated, "The specimens identified by Foxon (1939,
p. 258) as atypical forms of Alima multispinosa
(Claus) agree with Alikunhi's figures and description
of late S. raphidea larvae."
Inasmuch as H. raphidea is not known to occur in
the Red Sea and H. harpax is the only Harpiosquilla
which occurs in the Red Sea, northern Arabian Sea,
and the Gulf of Oman, it seems likely that Foxon's
atypical larva may be the larva of H. harpax. Whether
or not the other larva reported by Foxon is even iden-
tifiable with Harpiosquilla remains to be seen.
Some of the larval forms from the Red Sea identi-
fied as Squilla sp. by Gohar and Al-Kholy (1957) may
be identifiable with H. harpax (see their figs. 5-7, pi.
xu). Judging from the illustrations, their Squilla sp.
includes the larvae of more than one species.
Alikunhi (1959) recorded young larvae of harpax
from off the Mahanadi Estuary, India.
The larval stages of Harpiosquilla reported from
the Mozambique Channel by Ingle and Delia Croce
(1967) may be referable to H. harpax or H. melanoura,
both of which occur as adults off Madagascar, or to H.
raphidea, which occurs on the east coast of Africa.
NUMBER 3 6 33
The alima larva from Japan figured by Yamaji
(1959) adjacent to his figure of raphidea does not
resemble the larva of Harpiosquilla.
Some of the observations made by Alikunhi (1967)
on Harpiosquilla postlarvae may apply to H. harpax
as well as to H. raphidea.
TYPE.?The lectotype and paralectotypes are in the
Rijksmuseum van Natuurlijke Historie, Leiden, The
Netherlands.
TYPE-LOCALITY.?Japan.
DISTRIBUTION.?Indo-West Pacific region, from
Japan and Australia in the Pacific to the Red Sea and
South Africa in the Indian Ocean. Shallow water to
93 m. The following records in the literature are prob-
ably correct: JAPAN: NO specific locality (de Haan,
1844, 1849; Herklots, 1861; Holthuis, 1941; Manning,
1968b).?TAIWAN: Tungkang Fish Market, 40-50
fms (Lee and Wu, 1966).?CHINA: Hongkong (Bige-
low, 1894; Kemp, 1913; Tiwari and Biswas, 1952).?
VIET-NAM: Baie d'Along [Tonkin] (Gravier, 1937);
Baie Nhatrang, 50 m (Gravier, 1930).?PHILIPPPINE
ISLANDS: NO specific locality (Miers, 1880; Ingle,
1963).?MALAYSIA: Singapore (Boone, 1938; Tiwari
and Biswas, 1952; Manning, 1967); Siglap, Singapore
(Chuang, 1961); Morib (Chuang, 1961).?INDO-
NESIA: Java Sea, between Singapore and Surabaya
[Surabaja] (de Man, 1898).?AUSTRALIA (all Stephen-
son and McNeill, 1955) : Cape Flattery and Tin Can
Bay, Queensland; Cowan Creek, Jerusalem Bay, and
Hawk River, New South Wales.?THAILAND: Tach-
alom and Gulf of Siam, north of Singora (Suvatti,
1938,1950).?ANDAMAN ISLANDS: Port Blair (Tiwari
and Biswas, 1952).?INDIAN OCEAN: (White, 1847;
Miers, 1880; Ingle, 1963).?INDIA: mouth of River
Hughli [Hooghly, including Sandheads] (Kemp, 1913;
Tiwari and Biswas, 1952); off Madras coast (Kemp,
1913); Madras Presidency (Tiwari and Biswas, 1952);
Bombay (Chhapgar and Sane, 1968) ; off Bombay,
20-25 fms (Tiwari and Biswas, 1952).?CEYLON:
southwest part of Palk Bay, off Adam's Bridge and
Rameswaram Island, 7-9 fms (Tattersall, 1906).?
WEST PAKISTAN: Karachi (Tirmizi and Manning,
1968).?GULF OF OMAN: (Chopra, 1939; Ingle,
1963).?RED SEA: southern Red Sea (Ingle, 1963);
Archico Bay, south of Massawa, Ethiopia, 8?17 fms
(Holthuis, 1967).?ZANZIBAR: (Miers, 1880; Ingle,
1963).?MADAGASCAR: Ambaro Bay, Nosy Be, 2-5 m
(Manning, 1968b) ; Baie de Narendry, 6 m (Manning,
1968b).?SOUTH AFRICA: Richards Bay (Millard and
Harrison, 1954); Durban (Barnard, 1950, 1955; Day
and Morgans, 1956).
The following records, listed in the literature as
raphidea, may be referable to H. harpax; records for
Japan might be dealing with H. japonica, new species,
and those from Australia may be referable to H. ste-
phensoni, new species. JAPAN: NO specific locality
(Anonymous, 1949; Utinomi, 1956, 1960; Yamaji,
1959); Suruga Gulf (Kemp, 1913; Parisi, 1922); Kii
Peninsula (Komai, Akatsuka, and Ikari, 1927); Seto,
Kii Province (Komai, 1927, 1938); Tanabe Bay, Kii
Peninsula (Komai and Ikari, 1929); Tanabe (Komai,
1938); Wakanoura (Komai, 1938); Kasiwazima, Tosa
Province (Komai, 1927); Misaki (Komai, 1927);
Nagasaki (Balss, 1910; Parisi, 1922; Komai, 1927).?
TAIWAN: northern Formosa (Komai, 1927); Takao
(Balss, 1910).?CHINA: no specific locality (Berthold,
1845, 1847); Hongkong (Liu, 1949) .?VIET-NAM:
Indo-China (Gravier, 1930; Serene, 1937, 1953); Baie
de Cauda, 15-25 m (Serene, 1954); Baie de Nhatrang,
8-12 m (Serene, 1939); Baie d'Along [Tonkin] (Dawy-
doff, 1952); Baie de Tourane, 25-30 m; Embouchure
du Bassac; Cap Saint Jacques; Baie de Hone Coke,
entree de Port Dayot; Poulo Condore Islands (all
Gravier, 1930).?SUBLICHES OSTASIENS: (Balss,
1910).?AUSTRALIA: Wyndham, Western Australia
(Stephenson, 1962).?INDIA: Sandheads (Chopra,
1934) .?MADAGASCAR : (Crosnier, 1965).
Larvae which may prove to be the larvae of H. har-
pax have been reported from the Red Sea by Foxon
(1939) and Gohar and Al-Kholy (1957), the northern
Arabian Sea and Gulf of Oman by Foxon (1939), the
Mozambique Channel by Ingle and Delia Croce
(1967), and perhaps, from the east coast of India by
Alikunhi (1952, 1959). See also the section on larvae
for both H. harpax (p. 25) and H. raphidea (p. 9) .
7. Harpiosquilla indica, new species
FIGURES 39-43
HOLOTYPE.?1 <J , 150 mm; 10 miles due south of
Mandapam Camp, South India; 12 fms; otter trawl;
L. P. Woods on Norwegian trawler M-2; Sta. LW-20
(GA 64-7); HOE; 14 February 1964; SI Crust.
125717.
DIAGNOSIS.?Size moderate, male with adult char-
acters at TL 150 mm; antennular peduncle shorter
than carapace; corneal index 290; rostral plate (Figure
40) longer than broad, lateral margins tapering to
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
slender median projection; carapace with median ca-
rina; upper margin of propodus of claw with smaller
spines and denticles between largest spines; dactylus of
claw with 9 teeth, outer margin with prominent angu-
lar projection in adult male; fifth thoracic somite
rounded laterally (Figure 42); posterior 3 thoracic
somites lacking submedian carinae; intermediate
carinae of thoracic somites unarmed; ventral keel of
eighth thoracic somite rounded; first to fifth abdominal
somites lacking submedian carinae, abdominal carinae
spined as follows: submedian 6, intermediate 3-6, lat-
eral 1-6, marginal 1-5; denticles 5, 11, 1; marginal
carina of telson not more than 1.5 times as long as
carina of lateral tooth; postanal keel short, not extend-
ing halfway between anus and posterior margin; at
most inner half of distal segment of uropodal exopod
dark.
COLOR.?Inner and outer surfaces of antennular
peduncles lined with dark pigment; antennal scale out-
lined with dark chromatophores; dark transverse bar
present dorsally between antennular and ophthalmic
somites; rostral plate outlined with dark pigment, basal
portion of plate with oval ring of dark chromato-
phores; margins, carinae, and grooves of carapace out-
lined with dark pigment; meral depression of claw with
proximal and distal dark spots; posterior 3 thoracic
FIGURE 39.?Harpiosquilla indica, new species,
holotype, TL 150 mm: dorsal view.
Male FIGURE 40.?Harpiosquilla indica, new species. Male
holotype, TL 150 mm: anterior portion of carapace.
NUMBER 36 35
and all abdominal somites with dark posterior line and
intermediate carinae lined with dark pigment; sub-
median area of thoracic and abdominal somites with
diffuse dark pigment; telson with marginal carinae and
dorsal pits darker than remaining surface; ventral sur-
face of telson with bases of teeth and submedian distal
area darker than remainder of surface; anterior edge
of basal segment and proximal segment of uropodal
exopod with diffuse dark spot; distal fourth of proximal
segment of uropodal exopod with diffuse dark pigment,
inner half of distal segment black; distal half of endo-
pod dark with apex lighter along midline. The overall
color pattern is shown in Figure 39.
SIZE.?Only specimen known, the male holotype,
TL 150 mm. Other measurements (in mm) : carapace
length 30.5; cornea width 10.5; antennular peduncle
length 27.0; rostral plate length 6.8, width 5.4; rap-
torial propodus length 39.4; fifth abdominal somite
width 33.0; telson length 29.6, width 28.0.
DISCUSSION.?Harpiosquilla indica, new species,
most closely resembles H. melanoura, differing from
the latter species as follows: (1) there are nine teeth
on the dactylus of the claw rather than eight; (2) there
is a well-developed median carina on the carapace;
(3) only the inner half of the distal segment of the
uropodal exopod is black; and (4) the marginal carina
of the telson is comparatively shorter, being not more
than one and one-half times rather than nearly three
times as long as the carina of the lateral tooth.
The lack of submedian carinae on the posterior three
thoracic and anterior five abdominal somites, the un-
armed intermediate carinae anterior to the third ab-
dominal somite, and the shorter marginal carina of
the telson will distinguish H. indica from H. harpax,
which it otherwise closely resembles. In H. indica the
projection on the outer margin of the dactylus of the
male is much more angular than that found in H.
harpax, and the median carina of the telson in the
holotype of H. indica is more inflated than in males of
either H. harpax or H. melanoura of similar size.
Harpiosquilla japonica differs from H. indica in
lacking the median projection on the rostral plate and
in having submedian carinae on the posterior three
thoracic and all six abdominal somites.
The eye of the holotype of H. indica is similar in
size to that of H. melanoura and noticeably larger than
FIGURE 41.?Harpiosquilla indica, new species. Male
holotype, TL 150 mm: raptorial claw (propodus length
39.4 mm).
FIGURE 42.?Harpiosquilla indica, new species. Male
holotype, TL 150 mm: lateral processes of fifth, sixth, and
seventh thoracic somites.
36
FIGURE 43.?Harpiosquilla indica, new species. Male
holotype, TL 150 mm: posterior portion of body (telson
width 28.0 mm).
the eyes of either H. stephensoni or H. harpax of simi-
lar size.
The color pattern of H. indica is very similar to that
of H. harpax. The holotype of H. indica lacks the
paired submedian spots on the telson but that may be
an artifact of preservation, and it also lacks the con-
centration of dark chromatophores on the anterolateral
margin of the antennal scale which is characteristic
of H. harpax.
SEXUAL DIMORPHISM.?The holotype has a pro-
nounced projection on the dactylus of the claw (Figure
41) which is much more angular than that found in
H. harpax. The basal inflation of the teeth of the claw,
swollen submedian carinae of the sixth abdominal so-
mite, and inflated median carina of the telson (Figure
43) are well developed in the single male specimen
available for study.
BIOLOGY.?Unknown.
DEVELOPMENT.?Unknown.
TYPE.?The holotype is in the Division of Crustacea,
National Museum of Natural History, Smithsonian
Institution.
TYPE-LOCALITY.?Ten miles due south of Manda-
pam Camp, South India, in 12 fathoms.
NAME.?The name refers to the occurrence of the
species off India.
DISTRIBUTION.?Known only from the type-locality.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
General Remarks
The species of Harpiosquilla are large squillids which
are found in relatively shallow water throughout most
of the Indo-West Pacific region. All species occur on
the shelf; the maximum depth recorded is 206 meters
for H. annandalei. That species, which has been taken
in depths between 15 and 206 meters, also shows the
widest depth range. H. melanoura has been taken in
depths between 55 and 73 meters. The remaining
species, H. harpax, H. indica, new species, H. japonica,
new species, H. stephensoni, new species, and H. raphi-
dea, generally occur in depths of 51 meters or less,
although H. harpax has been collected as deep as 91
meters. At least three species, H. harpax, H. stephen-
soni, and H. raphidea, may occur in the sublittoral
zone.
Judging from the limited information available both
in the literature and from data associated with speci-
mens reported here, the species of Harpiosquilla all
prefer a soft, level bottom; they do not appear to be
associated with coral reefs, or with rough, rocky bot-
tom. At least one species, H. raphidea, is tolerant of
lowered salinities.
The genus is not known to occur in Oceania. This
may be a result of the paucity of collections from that
area but more likely reflects the lack of suitable habitat
there.
In a study of certain penaeid shrimps from the Indo-
Malayan area, Hall (1962) commented on the exist-
ence of an apparent distributional barrier in the Ma-
lacca Straits. On the basis of this he cast suspicion
on the validity of all littoral penaeid species reported
to range throughout the Indo-West Pacific region. This
does not seem to be true in Harpiosquilla, for four of
the seven known species occur on both sides of the
Indo-Malayan area, and two of these four species have
broad distributional patterns in the Indo-West Pacific
region. H. harpax, for example, extends from Japan to
Australia in the western Pacific, and from there west-
ward to the Red Sea and South Africa. H. annandalei
has a similar distribution pattern, based on many fewer
samples, but it does not occur in Australia and has
not been recorded with certainty west or south of the
Gulf of Oman. H. raphidea is known to occur from
Borneo and Thailand to India, West Pakistan, and
East Africa; it is not known with certainty to occur
north of Borneo and Thailand in the Pacific and there
are no apparent reasons for its relatively limited dis-
tribution in the western Pacific or its absence in Aus-
NUMBER 36 37
tralian waters. H. melanoura occurs in New South
Wales, Australia, the Andaman Sea, and Madagascar.
The remaining species of Harpiosquilla may be in-
digenous to the areas from which they are now known.
H. indica, new species, is known only from the type
locality off southern India, H. japonica, new species,
only from Japan, and H. stephensoni, new species, from
the Northern Territory and Queensland, Australia.
The latter two species are very similar and possibly
represent the terminal portions of a more widely dis-
tributed ancestral stock. Among the stomatopods, a
similar Japan-Australia link is shown by two species of
Heterosquilla, H. latifrons from Japan and H. brazieri
from Australia (Manning, 1966).
Within Australia the two most abundant species,
H. harpax and H. stephensoni, have an interesting dis-
tributional patern. H. stephensoni is known with cer-
tainty from Darwin Harbour, Northern Territory to
Port Curtis, Queensland; i.e., it is known only from
the tropical waters of the eastern and northeastern
coast on the Solanderian Province (Knox, 1963). If,
as I have assumed, Stephenson's (1952, 1953) records
for Squilla raphidea prove to be based on H. stephen-
soni rather than H. harpax, its distribution on the east-
ern Australian coast will be extended to Southport,
Queensland, in the northern edge of the Peronian
Province. H. harpax, however, extends from the Gulf
of Carpentaria, on the Queensland side, south to the
vicinity of Sydney in the cooler Peronian Province,
is much more abundant in the southern portion of its
range. It is nevertheless surprising to find H. harpax,
the most abundant species of the genus, all of which
otherwise are tropical, in the cooler, warm-temperate
environs of Sydney. Perhaps its ability to adapt to a
wider range of conditions in part helps to explain its
overall wide distribution.
Bibliography
Alikunhi, K. H.
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titles will carry a final line indicating the higher categories to which the taxon is referable:
"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify
the author on the first page of text with an unnumbered footnote that includes his professional
mailing address. A table of contents is optional. An index, if required, may be supplied by the
author when he returns page proof.
Two headings are used: (1) text heads (boldface in print) for major sections and chapters
and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings
may be worked out with the editor.
In taxonomic keys, number only the first item of each couplet; if there is only one couplet,
omit the number. For easy reference, number also the taxa and their corresponding headings
throughout the text; do not incorporate page references in the key.
In synonymy, use the short form (taxon, author, date, page) with a full reference at the end
of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent
lines indented about three spaces. Within a taxon, use a period-dash (.?) to separate each
reference. Enclose with square brackets any annotation in or at the end of the taxon. For
references within the text, use the author-date system: "(Jones, 1910)" or "Jones (1910)!" If
the reference is expanded, abbreviate the data: "Jones (1910, p. 122, pi. 20: fig. 1)."
Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they
should not contain rules. Formal tables must be submitted as pages separate from the text, and
each table, no matter how large, should be pasted up as a single sheet of copy.
Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed
throughout the printed text. They will be termed Figures and should be numbered consecutively;
however, if a group of figures is treated as a single figure, the individual components should be
indicated by lowercase italic letters on the illustration, in the legend, and in text references:
"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as
full pages on a different stock of paper, they will be termed Plates, and individual components
should be lettered (Plate 96) but may be numbered (Plate 9: figure 2). Never combine the
numbering system of text illustrations with that of plate illustrations. Submit all legends on pages
separate from the text and not attached to the artwork.
In the bibliography (usually called "Literature Cited"), spell out book, journal, and article
titles, using initial caps with all words except minor terms such as "and, of, the." (For capital-
ization of titles in foreign languages, follow the national practice of each language.) Underscore
(for italics) book and journal titles. Use the colon-parentheses system for volume, number, and
page citations: "10(2) :5-9." Spell out such words as "figures" and "plates" (or "pages" when
used alone).
For free copies of his own paper, a Smithsonian author should indicate his requirements on
"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive
50 free copies; order forms for quantities above this amount with instructions for payment will be
supplied when page proof is forwarded.