Lobster larvae and their distribution_Johnson 1971.pdf

On Palinurid and Scyllarid Lobster Larvae and Their Distribution in the South China Sea
(Decapoda, Palinuridea)
Author(s): Martin W. Johnson
Source: Crustaceana , Nov., 1971, Vol. 21, No. 3 (Nov., 1971), pp. 247-282
Published by: Brill
Stable URL: https://www.jstor.org/stable/20101842
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ON PALINURID AND SCYLLARID LOBSTER LARVAE AND THEIR
DISTRIBUTION IN THE SOUTH CHINA SEA
(DECAPODA, PALINURIDEA)
BY
MARTIN W. JOHNSON
Scripps Institution of Oceanography, University of California San Diego, La Jolla,
California 92037, U.S.A.
INTRODUCTION
Among the marine animals of direct importance in the fisheries resources of
the sea are a number of species of lobsters especially the Palinuridae. Most resea
on these crustaceans has been concerned with the adult bottom-living phase. Stud
on the free-floating phyllosoma larval phase have received less attention because
taxonomic uncertainties resulting partly from inadequate collecting methods f
larvae from the sea and partly from difficulties in rearing the larvae experimentally
in the laboratory due to their fragile nature and long larval period.
The recent numerous oc?anographie expeditions have now collected large quan
tities of Zooplankton making it possible to contribute toward an understandin
of the complete life cycle and ecology of several important lobster species and t
dispersal of their larvae in the sea.
It is clear however that more research involving collecting and detailed morph
logical study is needed before a real comprehensive report on the systematics
the larvae occurring in the Indo-West Pacific can be given. The many species tha
occur in these waters from Japan to the Indian Ocean (see Holthuis, 1946), lead
to uncertainties in the specific identification of some larvae found in the plankto
Further difficulties arise from the dearth of information on the relative abunda
of the local species, their size range, and the relative number of eggs produced
by individual females.
The fifteen species of larvae dealt with here are from a section of the coa
not previously investigated to any extent for phyllosoma larvae and it is hoped
therefore that a description of larvae commonly found here will be of aid
furthering identification of larvae in this and other areas by a process of elimination
through comparisons and reference to the known geographic distribution of t
adult lobsters and their relative abundance in the areas concerned. Some of the
larvae dealt with here have been described in part previously in isolated stages o
partially complete developmental series. References to these will be made brief
under the species involved and only one or more of the developmental stag
illustrated here sufficiently to enable identification of the larvae in the later mo
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248 MARTIN W. JOHNSON
definitive stages. Although reference will be made to "stages" these suggest only
the relative position in a developmental series of unknown number of instars.
As a further aid to recognizing the developmental position of a given species the
length of the larva measured from midpoint between the eye stalks to the end of
the telson will be given. Irrespective of the number of stages or size involved,
the instar having a full compliment of well developed gills is considered the last
phyllosoma stage before metamorphosing to the bottom-seeking puerulus (pali
nurids) or nisto (scyllarids) stage.
SOURCE OF MATERIAL AND METHODS
The larvae dealt with were collected by the NAGA Expedition, 1959-1961,
sponsored by South Vietnam, Thailand, and the United States of America, and
was carried out mainly as the Southeast Asia Research Program of Scripps Institu
tion of Oceanography (see S.I.O. Reference 63-6, February, 1963, dealing with
"Ecology of the Gulf of Thailand and the South China Sea").
The net employed was a 2-meter stramin ring net about 7 meters long with
filtering mesh of about 1 to 2 mm aperture. The hauls were made both at the
surface and obliquely from various depths ranging to 400 or more meters during
dusk or dark hours, usually one to two hours or later after sunset.
All figures given here were made with the aid of a Wild drawing tube attach
ment to the binocular microscope.
The specimens dealt with are deposited in the U. S. National Museum as
indicated and duplicates in the Scripps Institution Invertebrate Collection when
available.
ACKNOWLEDGEMENTS
My sincere thanks are due to members of the NAGA Expedition, especially to
Dr. Edward Brinton for making available much of the plankton collection for
study and to Mr. H. George Snyder, curator of the collection. I am grateful to
Professor John D. Isaacs, Director of the Marine Life Research Group of Scripps
Institution of Oceanography for generous support in sponsoring this aspect of the
study and to Miss Margaret Knight for proof reading the manuscript and other
associated aids.
THE PHYLLOSOMA LARVAE OF PALINURIDAE (SPINY LOBSTERS)
Panulifus penicillatus (Olivier, 1791)
Stage VIII (figs. 1, 2). ? The cephalic shield (forebody) and the thorax
(hindbody) are equal or nearly equal in width in stages older than about VI or
VII. The posterior margin of the oval-shaped shield becomes progressively more
rounded in later stages but not fully rounded. Other distinguishing features are:
(1) anterior mouth parts situated well forward from the second maxillae; (2) the
second maxillipeds are long and slender and their exopods, when fully developed,
remain shorter than the first segment of the endopod (fig. 1); (3) there are no
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PHYLLOSOMA SOUTH CHINA SEA 249
Figs. 1-2. Panulirus penicillatus (Olivier). 1, stage VIII (21 mm); 2, first antenna.
subexopodal spines (a strong spine situated at the distal end of the second basal
segment adjacent to the exopod) on the legs; (4) the second and third segments
of the peduncle of the first antenna are about equal in length.
Remarks. ? A fuller description of different stages is given by Johnson (1968)
for Hawaiian specimens where the species occurs locally together with P. marginatus
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(Quoy & Gaimard, 182.4). An identical larva also occurs in the area around the
Galapagos Islands and the Revillagigedo Islands, where P. peni ci Hat us occurs
together with P. gracilis (Streets, 1871) and P. inflatus (Bouvier, 1895) (Johnson,
in press).
Specimen deposited in U. S. National Museum No. 137896.
Figs. 3-5. Panulirus longipes (A. Milne Edwards) (japonicus group). 3, stage IX (24.3 mm);
4, first and second antennae; 5, second maxilla, first and second maxillipeds.
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Panulirus longipes (A. Milne Edwards, 1868)
Stage IX, length 24.3 mm (figs. 3-5). ? The cephalic shield is symme
trically oval, 16.4 mm long and 9.7 mm wide. Thus it is conspicuously narrower
than the thorax which is 12.2 mm wide. The abdomen is 4.9 mm long
(fig. 3). As in P. penicillatus the anterior mouth parts are situated well forward
of the second maxillae and the second maxillipeds are long and slender and bear
short exopods (fig. 5). In contrast to that species, the third segment of the first
antennal peduncle is a little longer than the second segment (fig. 4). There are no
subexopodal spines and coxal spines are absent in the later stages.
Remarks. ? Stages V-IX and XI (35 mm, with gills) were present in the
South China Sea. Michel (1969) has described a larva referred to this species
Figs. 6-12. Panulirus ornatus (Fabricius). 6, stage IV? (7.1 mm), sp. = subexopodal spine; 7, first
antenna; 8, first maxilla; 9, stage VI (9.7 mm) ; 10, dactyl of first leg; 11, abdomen and fifth legs;
12, second maxilla, first and second maxillipeds.
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from the New Hebrides. These larvae are most likely produced by one of the
species belonging to the Panulirus japonicus group as defined by George & Holt
huis (1965) to include adults of P. japonicus (von Siebold), P. longipes (A. Milne
Edwards), P. marginatus (Quoy & Gaimard), P. pascuensis Reed, and P. cygnus
George (considered a subspecies of P. longipes by Chittelborough & Thomas, 1969).
Figs. 13-15. Panulirus ornatus (Fabricius). 13, stage VIII (14.7 mm); 14, abdomen and fifth legs;
15, first maxilla.
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It is probable that their larvae may be very similar or indistinguishable. This at
any rate appears to be true for the present larva and that of the Hawaiian species,
P. marginatus. It may also be true for the larva briefly described by George (1962)
assigned to P. c y gnus from western Australia.
Figs. 16-19. Panulirus ornatus (Fabricius). 16, stage IX (18.7 mm); 17, first antenna; 18, abdomen;
19, second maxilla, first and second maxillipeds.
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Panulirus omatus (Fabricius, 1798)
Stages V(?)-IX, XI (26.7 mm, with gills), (figs. 6-23). ? These larvae
appear not to have been described previously. The size range is 7.1 mm for stage
V(?) to 26.7 mm for XI, the gilled stage. The distinguishing features in the
series available include the following: (1) the cephalic shield is oval in outline
Figs. 20-23. Panulirus ornatus (Fabricius). 20, stage XI (26.7 mm); 21, uropods and telson;
22, second maxilla, first and second maxillipeds; 23, first and second antennae.
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but more tapered from about the mid-length toward the anterior end than toward
the posterior, and in the younger stages it is about equal in width to the thorax
but in the last stage it is distinctly wider, the shield being 17.9 mm long and
11.9 mm wide (fig. 20), while the thorax is 10.5 mm wide; (2) the mouth parts
are relatively closely spaced and the exopod of the second maxilliped when fully
developed is nearly as long as the first two segments of the endopod combined
(fig. 22); (3) subexopodal spines (fig. 6 sp.) are present on legs 1-3 and coxal
spines persist only on the third maxillipeds; (4) dorsal coxal spines are present
on the third legs in stages VIII and IX (and sometimes also on legs 2 and 4), and
on legs 3 and 4 in stage XI; (5) the second and third peduncular segments of the
first antennae are about equal in length (fig. 23).
Remarks. ? This larva, although distinct, appears to be rather closely related
to the following larva which is most likely Panulirus versicolor (Latreille), and
since that species is considered to be closely allied to P. ornatus, the present larva
is provisionally assigned to the latter species. P. ornatus is common in the Indo
West Pacific area.
Prasad & Tampi (1957) have described the first stage of P. ornatus hatched
from the egg. They do not mention the presence of dorsal coxal spines, but the
distinguishing features of several species appear not to be discernable until later
stages.
Specimens deposited in U. S. National Museum Nos. 137904 and 137907.
Panulirus versicolor (Latreille, 1804)
Stages IV-IX (19.5 mm), (figs. 24-26). ? In the later stages the cephalic
shield is egg-shaped in outline with a well rounded posterior margin. In stage IX
(fig. 24) it is 13.5 mm long and 10.2 mm wide and the thorax is 9.5 mm wide.
The mouth parts are closely spaced as in the preceding larva and the exopod of
the second maxilliped is also nearly as long as the first two segments of the endopod
combined (fig. 25). Subexopodal spines are present on legs 1-4 and in stage IX
coxal spines are present only on leg 1 and the third maxilliped. Dorsal coxal
spines are conspicuous on legs 2 and 3 in early stages (from stage IV) whereas
they were present only on legs 3 and 4 in the gilled stage which has a length of
26.1 mm.
Remarks. ? Michel (1969, fig. 2) has figured a larva from the New Hebrides
that agrees with the present larva and has referred it to P. versicolor, which is
followed here. Prasad & Tampi (1959) described a similar larva from the Lacca
dive Archipelago and referred it tentatively to P. penicillatus. But in light of more
recent work it seems more likely to belong to P. versicolor which occurs also in that
area and in the Indo-West Pacific. Chittelborough & Thomas (1969) report a
similar larva tentatively under the name P. penicillatus from the Indian Ocean
northwest of Australia.
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Figs. 24-26. Panulirus versicolor (Latreille). 24, stage IX (19.5 mm); 25, second maxilla, first and
second maxillipeds; 26, first antenna.
Panulirus (?) sp.
Stage IX, 26.8 mm (figs. 27-31). ? This larva of which two specimens were
obtained appears not to have been previously described and it cannot be referred
with certainty to Panulirus although its main features agree with that genus. The
cephalic shield is subcircular in outline, 20.5 mm long and 16 mm wide, with
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uniformly rounded posterior margin (fig. 27). The thorax is distinctly narrower
than the shield, being only 11.8 mm wide (a little wider in the 17.3 mm
specimen) and the relatively small abdomen is 2.9 mm long (fig. 28). Ventral
coxal spines are present on the third maxilliped and on legs 1-4, and subexopodal
spines are wanting. The mouth parts are very closely spaced and the exopod of
the second maxilliped is relatively long, reaching almost to the end of the endopod
(fig. 29). The proportional lengths of the peduncular segments 1 to 3 of the first
antenna are 2.1, 0.65, 1.35 mm respectively, giving unusually long first and third
Figs. 27-31. Panulirus} sp. 27, stage VIII? (26.8 mm); 28, abdomen and fifth legs; 29, second
maxilla, first and second maxillipeds; 30, first maxilla; 31, first and second antennae.
Crustaceana, 21 17
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segments and the peduncular segments 2 and 3 of the second antenna bear one
and two distal spines respectively (fig. 31). The anterior branch of the first
maxilla has only two strong masticatory spines, whereas three spines are commonly
found in most larvae at this stage of development, and no setae were observed
indicating the palp (fig. 30).
Linuparus White (?)
Stage IV (?), 6.2 mm (figs. 32-37). ? This larva also appears not to have
been described before. The cephalic shield is somewhat triangular with a rounded
posterior margin. It is 4.2 mm long and 3.85 mm wide with the greatest width at
about the level of the anterior mouth parts which are situated rather near the
second maxilla. The thorax is 2.7 mm wide (fig. 32). Coxal spines are present
on relatively short coxae. Subexopodal spines are present on legs 1-3. The dactyl
of leg 1 opposes a rather heavy spine, giving the appearance of a chelate appendage
(fig. 37). Leg 4 is still not fully developed but bears a two-segmented exopod.
Leg 5 is rudimentary but about as long as the abdomen. Rudimentary pleopods are
low buds, and bilobed uropods are present (fig. 33). The eyes are relatively long
and narrow. The third peduncular segment of the first antenna is about twice the
length of the second segment and the second antenna bears a strong outer process
at the base as in scyllarids (fig. 36). The first maxilla has two strong masticatory
spines on the anterior branch and the palp is indicated only by two short setae
(fig. 35). The distal segment of the second maxilla is fringed with setae and the
third maxilliped has a well developed exopod as in palinurids (figs. 32 and 34).
Remarks. ? A second specimen, a moult older than the one figured was taken
on the same cruise, S-6. Although the reference of these larvae to Linuparus
can be only tentative still it gains some credence from the fact that two species of
the genus are known to the area i.e., L. trigonus (von Siebold) of which numerous
specimens have been taken in trawls on the northern shelf region of South China
Sea (Bruce, 1965) and a rarer species, L. sordidus Bruce, 1965.
Puerulus angulatus (Bate, 1888)
Early stages, lengths 4.3 mm and 7.3 mm (figs. 38-46). ? The two larvae
here figured are early stages in the developmental series to which a 19.7 mm
larva from the South China Sea was described previously (Johnson, 1968a,
fig. 7).
As in the 19.7 mm specimen, the cephalic shield of the present larva is circular
with a truncated posterior margin. In the smaller larva the shield overlaps the
thorax to include the coxae of the first pair of legs and in the 7.3 mm larva it
overlaps also the second pair (figs. 38 and 43). This progressive backward expan
sion of the shield apparently continues in successive stages and in the 19.7 mm
larva it overlaps all of the thorax and part of the abdomen. The mouth parts are
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Figs, 32-37. Linuparus White (?). 32, stage IV? (6.2 mm); 33, abdomen and fifth legs; 34, second
maxilla, first and second maxillipeds; 35, first maxilla; 36, first and second antennae; 37, dactyl
first leg.
closely compacted, especially in the older stages. There are no subexopodal spines
but coxal spines are present on the legs and the third maxillipeds. The anterior
branch of the first maxilla has two masticatory spines and there is a well developed
palp with two terminal setae (fig. 42). The distal segment of the second maxilla
is fringed with setae (fig. 40). The first maxilliped is a rudimentary bud and the
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Figs. 38-42. Puerulus angulatus (Bate, 1888). 38, stage II? (4.3 mm); 39, abdomen, fourth and
fifth legs; 40, second maxilla, first and second maxillipeds; 41, first and second antennae;
42, first maxilla.
second maxilliped has a rather long rudimentary exopod without setae (fig. 46).
The first antenna is unsegmented and shorter than the second antenna which is two
segmented, the proximal segment bearing a strong lateral process at the base and
a spine at the distal end (fig. 41). The dactyl of leg 1 (as in legs 4 and 5 of the
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Figs. 43-46. Puerulus angulatus (Bate, 1888). 43, stage VII (7.3 mm); 44, tip of first leg; 45,
dactyl of third leg; 46, second maxilla, first and second maxillipeds.
19.7 mm larva) is not claw-like (fig. 44) and the dactyl of leg 2 (lost in the
19.7 mm larva) is long and prehensile and leg 3 has a short claw (fig. 45).
Legs 4 and 5 not yet fully developed.
Remarks. ? These larvae are of the kind early known as Phyllamphion (Rein
hardt, 1849) of which there are two types, one with a rectangular shield and the
other with a circular shield. The present larval type was called Phyllamphion
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santuccii by Belloc (1959) for a larva from the Gulf of Aden. His suggestion that
the larvae probably belong to the genera Puerulus and Palinurellus appears to be
well born out with increased knowledge of phyllosoma larvae of different genera.
Sims (1966) has concluded that a series of larvae with rectangular shield, and
occurring in the Caribbean area where adult Palinurellus occur, is of that genus.
Michel (1970) reports the larval type with rectangular shield from the New
Caledonia area and considers it Palinurellus wieneckii (De Man). The present
larva then appears to belong to Puerulus. The formula for the relative lengths of
the peduncular segments do agree well with that genus. Based on the presently
known distribution of Puerulus species mentioned by Holthuis (1966) and Berry
(1969) it seems likely that the present larva is P. angulatus. However, it must be
pointed out that the larva of the related genus Linuparus has not been identified
with certainty. Therefore Linuparus is still not ruled out as a possible alternate
in the identification of the present larva. A phyllamphion larva with a circular
shield has been reported from the Indian Ocean by Taisho (1966).
THE PHYLLOSOMA LARVAE OF SCYLLARIDAE (SLIPPER OR SHOVELNOSED LOBSTERS)
The following five larvae are believed to belong to the genus Scyllarus. The
main features separating them specifically are: (1) size in the gilled (final) stage;
(2) shape of cephalic shield; (3) shape of uropods; and (4) length of eyestalks.
Scyllarus sp. A
Four stages, 6.3 mm, 16.1 mm, 21 mm, and gilled stage 22 mm (figs. 47-56). ?
These larvae appear to fall into a developmental series not previously described.
For the 6.3 mm specimen (fig. 47) the shield is 4.4 mm long and 4.7 mm
wide; the thorax is 2.3 mm wide. In subsequent stages the ratio of shield to thorax
width remains about the same so that for the gilled stage the shield and thorax
are 19.7 mm and 8.9 mm wide respectively (fig. 54). The posterior margin of
the shield becomes progressively more truncated and bowed inward in successive
stages, and the anterior-lateral margins less sloping. In the last stage the shield
overlaps the coxae of the second legs. The telson in all stages examined bears two
strong terminal spines which become relatively shorter in the gilled stages (figs.
48, 52, 54). The uropods are not pointed. Subexopodal spines are present on legs
1-4. Leg 5 remains uniramous and small. Coxal spines are present on all legs and
the third maxillipeds. A dorsal thoracic spine is present near each of the coxae of
legs 1-4. The second and third peduncular segments of the first antenna are about
equal in length in the last stage. The second antenna bears a strong lateral process
and becomes very broad (figs. 50 and 56). Fringing setae are wanting on the
second maxilla (figs. 51 and 55). The exopods of the second and third maxillipeds
remain rudimentary buds. Each of the last two stages shown has a full complement
of gills, but the gills are relatively small especially in the younger specimen. This
would indicate that several moults may take place within the gilled stage.
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48 50 ^ 52
Figs. 47-52. Scyllarus sp.
5C, first and second anten
fifth legs.
Remarks. ? The oldest larva is somewhat similar to the gilled stage of Scyllarus
delfini (Bouvier, 1909) from north of Juan Fernandez Island but the shield is
relatively wider and overlaps more of the thorax, and the abdomen is shorter
(Johnson, 1971a). The youngest larva resembles somewhat a 6 mm larva from
Cape Verde Islands area that was referred to Scyllarides sp. by Gurney (1936,
fig. 36). Robertson (1969) has summarized the generic larval characteristics
of Scyllarus.
Scyllarus sp. B
Gilled stage, 29.2 mm (figs. 57-61). ? This is the largest Scyllarus larva en
countered in the collection. The cephalic shield is nearly circular but for a broadly
truncated posterior margin, and is 17.8 mm long and 20.5 mm wide. The thorax
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Figs. 53-56. Scyllarus sp. A. 53, larva (21 mm); 54, 22 mm larva; 55, second maxilla, first and
second maxillipeds; 56, first and second antennae.
is 10.6 mm wide and, as in the other Scyllarus larvae, is broadly confluent with
the abdomen. The distal ends of the uropods are smoothly rounded and the telson
bears only small lateral spines (fig. 57). Subexopodal spines are present on legs 1-4.
Leg 5 is short and uniramous. Small coxal spines are present on the third maxilliped
and all of the legs, and a dorsal thoracic spine is present adjacent to the coxa of
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Figs. 57-61. Scyllarus sp. B. 57, final stage (29.2 mm); 58, second maxilla, first and second
maxillipeds; 59, first maxilla; 60-61, first and second antennae.
legs 2, 3, and 4. The eyestalks are relatively short. The proportional lengths of
segments 1-3 of the first antennal peduncle are about 1.8 : 1.05 : 0.8 mm (fig. 60).
Other features, such as absence of fully developed exopods on the second and
third maxillipeds, the absence of fringing setae on the second maxilla (fig. 58),
and the presence of an exterior lateral process on short second antenna (fig. 61),
are as occurs in other Scyllarus larvae.
Scyllarus martensii Pfeffer, 1881
Gilled stage, 9.3 mm (figs. 62-64). ? This is the smallest final stage Scyllarus
larva encountered in the collection. The cephalic shield is more or less trapezoidal
in outline and is 5.3 mm long and 6.5 mm wide with a truncated posterior margin.
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The thorax is 4.5 mm wide (fig. 62). The distal ends of the uropods are pointed
and the telson bears small lateral spines (fig. 63). Subexopodal spines are present
on legs 1-4. Leg 5 is short and uniramous. Relatively large, broad coxal spines are
present on all of the legs. The proportional lengths of segments 1-3 of the first
antennal peduncle are about 1.7 : 1.0 : 1.0 mm (fig. 64). The eyes talks are short
(about 1 mm). Other features are characteristic of the genus.
Remarks. ? This larva is the second most abundant encountered. It is very
similar to, but about 3 or 4 mm shorter than the same stage referred to S. modestus
Figs. 62-64. Scyllarus martensii Pfeffer. 62, final stage (9.3 mm); 63, uropods and telson; 64, first
and second antennae.
Figs. 65-67. Scyllarus sp. C. 65, final stage (12.1 mm); 66, second maxilla, first and second
maxillipeds; 67, first and second antennae and eye.
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Holthuis, I960 from the Hawaiian Islands (Johnson, 1971), but that larva lacks
the lateral spines on the telson in the gilled stage. It is also near the larva referred
to Scyllarides sp. by Prasad & Tampi (I960, fig. 11), but that larva also lacks
the lateral spines on the telson. Larvae of this type have been erroneously referred
to Scyllarides in the earlier studies on phyllosomas, and are reported from South
Africa by Von Bonde (1932) and Gurney (1936), from the Gulf of Thailand
by Naiyanetr (1963), and from Japan by Shojima (1963).
Scyllarus sp. C
Gilled stage, 12.1 mm (figs. 65-67). ? The outline of the cephalic shield of this
larva is more circular than in the previous larva. It is 7 mm long and 7.8 mm wide.
The thorax is 4.3 mm wide. The uropods are only slightly pointed (fig. 65).
Subexopodal spines are present on legs 1-4. Leg 5 is short and uniramous. Coxal
spines are found on all of the legs and the third maxilliped but are less broad
than in the preceding larva. The eyestalks are short (1.6 mm) ; the first and second
antennae are about equal in length and the proportional lengths of the peduncular
segments 1-3 of the first antenna are about 0.6 : 0.49 : 0.35 mm (fig. 67). Other
features are characteristic of the genus but the second antennae appear to be some
what more advanced in growth than usual.
Specimen deposited in U.S. National Museum No. 137906.
Scyllarus sp. D
Gilled stage, 18.1 mm (figs. 68-71). ? In this intermediate sized larva the
cephalic shield is subcircular and only moderately truncated at the posterior margin.
It is 11.6 mm long and 11.1 mm wide. The thorax is 6.7 mm wide (fig. 68). The
distal ends of the uropods are smoothly rounded and the telson has small lateral
spines (fig. 69). The legs are relatively long and slender and subexopodal spines
are present on legs 1-4. Leg 5 is short and uniramous as in other Scyllarus species.
Coxal spines are present on the third maxilliped and on legs 1-5. The eyestalks
are relatively long (3.2 mm). The first antennae are longer than the second
antennae and the proportional lengths of peduncular segments 1-3 of the first
antenna are 0.95 : 0.75 : 0.65 mm (fig. 71). Other features are characteristic of the
genus. An early stage larva (4.8 mm) no doubt belonging to this species is shown
in figures 72-75.
Remarks. ? This larva was the most abundant phyllosoma taken in the area.
It is most likely the same specifically as the one called Scyllarus sp. II by Prasad &
Tampi (I960, fig. 9 A) from the Arabian Sea, and is similar to the larva from
South Africa figured by Von Bonde (1930, pi. 7).
Another larva that resembles very closely the present one in size was found at a
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Figs. 68-71. Scyllarus sp. D. 68, final stage (18.1 mm); 69, uropods and telson; 70, second maxilla,
first and second maxillipeds; 71, first and second antennae.
nearshore station just north of Saigon. It is 18.5 mm long in the gilled stage and
differs from the above larva especially in having short eyestalks and a less circular
shield with a distinctly truncated posterior margin. It is doubtless a separate species
but was discovered too late to be further described in this report.
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Figs. 72-75. Scyllarus sp. D. 72, larva (4.8 mm); 73, abdomen and fifth legs; 74, second maxilla
and second maxilliped; 75, first and second antennae.
Thenus orientalis (Lund, 1793)
Stage II, 3.7 mm (figs. 76-80). ? In this early stage larva, the cephalic shield
is subcircular, 2.45 mm long and 2.9 mm wide. The thorax is 1.5 mm wide
(fig. 76). The uropods are present as rudimentary buds (fig. 77). Subexopodal
spines are present on legs 1-4, and coxal spines on the third maxillipeds and
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Figs. 76-80. Thenus orientalis (Lund). 76, stage II (3.7 mm); 77, abdomen and fifth legs; 78, first
maxilla; 79, second maxilla and second maxilliped; 80, first and second antennae.
legs 1-4. Ventral thoracic spines are present at the margin adjacent to the coxae
of legs 1-4 (a feature commonly found in the larvae of palinurids). The eyes are
on long tapering stalks confluent with the eyes. The first antenna is unsegmented
and much longer than the second antenna, which is not yet provided with a lateral
process (fig. 80). A unique feature for such an early larva is the presence of three
strong masticatory spines on the anterior branch of the first maxilla (fig. 78).
The terminal segment of the second maxilla bears three setae (fig. 79). The
identification of this larva is based on comparisons with a similar larva hatched
from eggs of this species by Prasad & Tampi (1957). Naiyanetr (1963) has
reported similar larvae from the Gulf of Thailand.
In a later stage, 11.1 mm (figs. 81-84), the cephalic shield is bilobed
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with a truncated posterior margin having a small convexity at the midline.
It is 7.5 mm long and 10 mm wide. The posterior margin barely overlaps
the coxae of the first legs. The thorax is 4.7 mm wide. The abdomen is confluent
with the thorax, the uropods are biramous but not segmented, and the pleopods
are rudimentary buds. There is a small spine situated at the posterior lateral margin
of the telson, and the dorsal surface of the abdomen has a low, broad spine at the
posterior margin of each segment (fig. 82). Legs 1-4 have each a subexopodal
spine and coxal spines are present on the third maxilliped and legs 1-4. The fifth
Figs. 81-84. Thenus orientalis (Lund). 81, larva (11.1 mm); 82, abdomen, dorsal; 83, second
maxilla, first and second maxillipeds; 84, first and second antennae.
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pair of legs are relatively short, uniramous and consist of four segments and a
claw-like dactyl. The first antenna is longer than the second antenna and the
relative lengths of segments 1-3 are about 0.65 : 0.55 : 0.6 mm. The terminal seg
ment (not yet separated) of the second antenna has three inner marginal spines
(fig. 84).
Remarks. ? This stage is similar to a 17.8 mm larva from the Indian Ocean
figured by Prasad & Tampi (1957). Their larva was in a gilled stage but their
figures show the pleopods as being only rudimentary buds. It is important to note,
however, that, as in the present larva, leg 5 remains relatively short and without
an exopod.
In general, this larva shows some similarity to the following larva, but a marked
difference is especially the degree of development of the fifth pair of legs which
in the following larva are fully developed.
Ibacus novemdentatus Gibbes ?
Length 17.5 mm (figs. 85-88). ? The cephalic shield is bilobed in outline
with a deep depression at the anterior end from which the eyes and antennae
issue, and a similar deep inbowing of the posterior margin. The shield is 11.5 mm
long, measured along the mid axis, and 18 mm wide, and the lateral lobes overlap
the coxae of the third pair of legs (fig. 85). The thorax is 6.6 mm wide and the
abdomen is about 4 mm long and bears rudimentary biramous pleopods, biramous
uropods, and a telson with lateral spines (fig. 86). The third maxilliped has a
rudimentary exopod bud. Legs 1-5 are strongly developed and have subexopodal
and coxal spines. The eyestalks are 4.2 mm long. The first antennae are longer
than the second and the lengths of peduncular segments 1-3 are about 1.3 : 0.95 :
1.0 mm respectively, and the second antennae bear 5 spines on the inner edge
of the terminal segment (fig. 87). There are three strong masticatory spines on
the anterior branch of the first maxilla. The second maxilla is not fringed by setae.
The first maxilliped is rudimentary, and the second maxilliped has a small spine
on the coxa and a rudimentary exopod (fig. 88).
Remarks. ? A larva of this type was first described by Tokioka (1954) under
the larval name Phyllosoma utivaebi from Japan and has subsequently been
identified as belonging to Ibacus (Tokioka & Harada, 1963; Dotsu, Tanaka,
Shujima & Seno, 1966). The first stage has been reared from the eggs of Ibacus
ciliatus by Saisho & Nakahara (I960).
Dotsu et al. (1966) report finding two forms of larvae, one attaining a length
of about 43 mm and the other 30 mm. The first was assigned to I. ciliatus (Von
Siebold) and the second to /. novemdentatus Gibbes. Although the present larva
is not in the final stage, it has been referred to the latter species because of the
smaller size, for it seems highly unlikely that it would ever attain a size of 43 mm.
See also "Remarks" under the previous larva.
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Figs. 85-88. Ibacus novemdentatus Gibbes? 85, larva (17.5 mm); 86, abdomen; 87, first and second
antennae; 88, second maxilla, first and second maxillipeds.
Parribacus antarcticus (Lund, 1793)
Stage VII, length 16.6 mm (figs. 89-92). ? The cephalic shield is oval in
outline, 13.8 mm long and 10.1 mm wide. The thorax is 9 mm wide and the
abdomen small and situated within a deep thoracic concavity which begins at the
bases of the fifth pair of legs (fig. 89). There are no subexopodal or coxal spines.
The eyestalks are 3.9 mm long. The first antennae are over twice the length of the
second antennae and the peduncular segments 1-3 are about 0.6 : 0.5 : 0.45 mm
long respectively (fig. 92). No exopod buds are in evidence on either the second
or third maxillipeds.
Remarks. ? Only three specimens were taken. A fuller description of other
Crustaceana, 21 18
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Figs. 89-92. Parribacus antarcticus (Lund). 89, larva (16.6 mm); 90, abdomen; 91, second maxilla
and second maxilliped; 92, first and second antennae.
stages is given by Johnson (1971) for larvae from the Hawaiian Islands.
LARVAL ABUNDANCE AND DISTRIBUTION
Charts 1-3 give the locality records for phyllosoma larvae taken by the 2-meter
net during the main cruises S-4, S-6, and S-8 in the South China Sea between
February 15 and October 5, I960. Chart 4 covers two cruises in which S-2 occupied
91.139.102.157 on Tue, 28 Sep 2021 15:59:10 UTC

105? 106? 107? 108? 109? 110?
___-m
^rf
112? 113? i ?4?_115'
NAGA CRUISE S-4 %
Feb. 15-Mar. 20, 1960 :
2-m net station
? Panu?rus penicillatus
-o H longipes
-6- ii orna?us
o ?. ?
? Linuparus ?
0 Scyilarus martensii
? ? sp. A
4 h sp C
? sp. D
a Thenus orientalis
$ Parribacus antarcticus
J
0 8
$ 29
24
A 14
0. 2
6
? 2
0 68
62 -?- 1
0 1
14
A 2
J_1_L
n 2
? O I
$ i
o t
.4 4
11
<^ 4
? 1
* 1
? 2
1 -O?
4
O 1
? 2
1_L_L
106? 107? 108? IQ9?> IIIo 112? 113? " 114? 115?
Chart 1
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^kn
NAGA CRUISE S-6
May 26-June 23, 1960
2-m net station
O Panulirus ?ongipes
-6- ? ornatus
-?- " versicolor
o ? ?
? Linuparus ?
0 Scyllarus martensii
? sp. D
-0- ? sp. B
S ? early
A Thenus orientalis
Ibacus novemdentatus mA1
J>2
9- 1
15
0 2
100
0 8
1
* 0 24
0 75
* 440
S 10
9-1
81
l
A 1
0 1
? 1
oi
? Q 1
"9 2
0 2
-0- 1
S 3
O- 2
?- 2
17 0
14 #
1 A
0 77
4
A ?
114? 115?
M ?.S.H.O. CHART 5595
Chart 2
91.139.102.157 on Tue, 28 Sep 2021 15:59:10 UTC

^H#^
112? 113?
i i?r~
NAGA CRUISE S-8
Sept. 7-Oct. 5, 1960
1-m net station
XX Panulirus peniciliatus
o ? longipes
-<?>- ? ornatus
-9- ?? versicolor
? Scyllarus martensii
^ ? sp. A
$ " sp. C
? sp. D
S ? early
# Parribacus antarcticus
n l
?-?-2
?-3
-?- 55
0 8
50
$ 1
n 2
'-9- 5
0 1
7
O 2
? 0 2
0 1
'0 229
40
S 21
0 7
21
6
$ 1
-O 1
^ 3
? S 3
14
* 4
K 1
-6-4
.-9-6
l
0 2
0 15
103
S 6
Ml 1
Chart 3
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105? 106? 107? 108?_109?- 110?_IIIo 112?_113?_114? 115?
i?*xT?r
1?rT i?r~r i ' il
NAGA CRUISE S-2
Nov. 17 -Dec. 11, 1959
NAGA CRUISE S-10
Jan. 12 - Feb. 9, 1961
1-m net station
0 Scyllarus martensii
? ? sp. A
$ " sp. C
" sp. D
a Thenus orientalis
4
1*01
0 1
0 1
^ 3
CRUISE S-10
0 5
A 1 A 1
* 1
CRUISE S-2
2 0
105? 106? 10.7? 103? 109? 110? 111? 112? 113? 114? 115?
FROM U.S.H.O. CHART S595
Chart 4
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only 4 stations, using the 2-meter net, at the southern portion of the survey area
during November 18 to December 11, 1959 and S-10 occupied 7 stations with this
net at the northern portion during January 12 to February 9, 1961.
These five cruises together cover the alternating regimes of the southwest and
northeast monsoons each of which result in a corresponding alternating pattern of
surface water currents paralleling the coast of Vietnam. Characteristic features of
monsoon circulation are given by Wyrtki (1961). The alternating currents are
complicated but consist largely of surface currents. The homogeneous layer in
the China Sea is said to be about 30 to 40 m deep during the southwest monsoon
and 70-90 m in the northern portion during the strong northeast monsoon. The
surface flow is strongest along the coast of Vietnam where it may be 50 to 75
cm/sec as shown on Wyrtki's charts.
Table I
Summary of phyllosoma larvae taken with a 2-m net over certain depth ranges of
water in the South China Sea during three Naga Expedition cruises.
Cruise Range of Number Number Number Average Meters of wire
water of of of duration out for oblique
depths stations scyllarid palinurid of hauls hauls
(m) occupied larvae larvae (minutes)
taken taken
S-4
S-6
S-8
27- 60
80- 102
476-1254
over 2000
22- 53
70- 106
518-1957
over 2000
26- 57
68- 128
607-1372
over 2000
5
2
2
8
4
4
5
3
3
4
4
2
243
18
3
7
~271
640
222
32
1
~895~
849
97
5
1
952
1
0
2
5
9
0
1
11
0
~?2~
0
63
19
0
82~
38
45
72
73
73
44
79
77
37
34
114
313
75- 100
75- 100
800 and surface*
800 and surface*
75
100- 125
600-1500
800-1500
Northeast
monsoon
Southwest
monsoon
75- 100
100
800-1000
1000-3000
Changing
season
* One haul made at surface.
The long planktonic life of phyllosoma larvae coupled with prevailing
currents must result in a great deal of larval dispersal into and out of t
Nevertheless there is clearly a much greater concentration of larvae in t
lower water at the coastal stations off Vietnam and in the offshore shall
Sunda Shelf area to the south of Vietnam. These areas appear to maintai
centration of larvae that includes not only the young larvae, that may
produced locally or at some distance upstream on the coast, but also
stages found mingled with the young. Presumably there is a combination
91.139.102.157 on Tue, 28 Sep 2021 15:59:10 UTC

graphic mechanisms and larval behavior that enhances retention. Lateral dispersion
must, however, occur as indicated by the presence of a few larvae at the distant
stations over deep water where early as well as later stages were taken.
Recruitment to the bottom-living stage of lobsters is no doubt supplemented
by influx of larvae with water flowing from surrounding islands of the archipelago,
but the coastal concentration of larvae would suggest that the major recruitment
is probably within the China Sea itself. Reference to current charts show that in
addition to the seasonally alternating north and south currents there are concur
rently developed large eddies and countercurrents within the system such that
much of the water and its plankton content must be retained for a considerable
period of time. Thus although many larvae must be flushed out of the area, there
still remains a residue of locally produced individuals that maintains a favorable
balance.
Regarding the effect of the alternating monsoon seasons on larval occurrence,
only a few general remarks may be made. Relatively few larvae were found on
cruise S-4 during the northeast monsoon and a similar effect of the northeast
monsoon is suggested also by the short surveys taken during that season on cruise
S-2 and S-10. About three times as many larvae were taken during the southwest
monsoon season on cruise S-6, and more yet were taken on cruise S-8 when there
was apparently a changing of seasons (charts 1-4 and table I). By way of com
parison, in a survey of general Zooplankton volumes Brin ton (1963) found
generally higher volumes in the northern coastal waters of Vietnam during the
northeast monsoon, whereas the broad southern shelf and the offshore basin
yielded highest plankton volumes during the southwest monsoon.
As is characteristic of phyllosoma collections, the total number of larvae taken
(2241) in the area is not large; still it is sufficient to provide a rather consistent
picture of which the following six features stand out more or less clearly:
1. The predominantly coastal and shallow water distribution as already men
tioned.
2. A striking feature is the numerical predominance of scyllarid larvae over the
palinurids. They make up about 95% of the total number of larvae caught
(Table I).
3. Two species of scyllarids clearly dominate the catch, namely Scyllarus sp. D
and Scyllarus martensii in that order. The first species appears to be about equally
abundant in the north and south shallow water areas, whereas the second species
although occurring throughout the area is relatively more abundant in the southern
part.
4. Thenus orientalis was found only in the southern part but numbers were small.
5. Very few palinurid larvae were found and these appeared more frequently
in the northern part and to some degree offshore. In contrast to the present data
Prasad & Tampi (1965) working with the Dana Expedition collection from the
Indian Ocean found 1111 palinurids and 872 scyllarids. The population of
palinurid larvae was greater in the western half of the Indian Ocean and the
91.139.102.157 on Tue, 28 f on Thu, 01 Jan 1976 12:34:56 UTC

scyllarid larvae in the eastern part. This relative increase in scyllarids in the more
tropical area as the Indo-West Pacific area is approached is in keeping with the
predominance of scyllarids found in the Naga collection. Saisho (1966) found
202 palinurid and 138 scyllarid larvae during a cruise in the Indian Ocean.
However, most of his stations were also well offshore.
6. Judging from the regular occurrence of young stages, there appears to be
a fair amount of breeding throughout the year. However, very few larvae less than
3 mm in length and clearly recognized as stage I, were taken at any time, probably
due largely to the coarseness of the net employed and the considerable distance
offshore for most collecting stations. The 1-meter net samples taken during the
expedition have not as yet been analyzed for these larvae.
ZUSAMMENFASSUNG
In vorliegender Arbeit wurde eine Ausbeute von 2241 Phyllosoma-Larven untersucht, die w?hrend
der Naga-Expedition, zwischen November 1959 und Februar 1961, im S?dchinesischen Meer gesam
melt worden war. Auf Grund morphologischer Kennzeichen, vornehmlich der ?lteren Larvenstadien,
sind 7 Arten Palinuridae und 8 Arten Scyllaridae darin enthalten. Nicht alle Larven k?nnen auf ihre
Artzugeh?rigkeit bestimmt werden. Etwa 95 Prozent der Larven sind Scyllaridae, die vorwiegend der
Gattung Scyllarus angeh?ren. Beigef?gte Abbildungen dienen zum Bestimmen der Larven, und
Karten zeigen ihre Verbreitung, die sich haupts?chlich auf K?stenn?he und Regionen von relativ
flachem Wasser erstreckt. Die gr?sste Anzahl von Larven wurde w?hrend des S?dwestmonsuns und
w?hrend der darauf folgenden ?bergangszeit gefunden.
LITERATURE CITED
Belloc, G., 1959. Note sur un phyllamphion de la mer des Antilles. Bull. Inst. oc?anogr. Monaco,
1154: 1-10.
Berry, P. F., 1969. Rediscovery of the spiny lobster Puerulus carinatus Borradaile (Decapoda,
Palinuridae). Crustaceana, 17 (3): 239-252.
BrintoN, Edward, 1963. Ecology of the Gulf of Thailand and the South China Sea. Part IV,
Zooplankton abundance in the Gulf of Thailand and South China Sea. A report on the results
of the Naga Expedition, 1959-1961. Scripps Inst. Oceanog. Ref. No., 63-6: 53-58.
Bruce, A. J*., 1965. A new species of the genus Linuparus White, from the South China Sea
(Crustacea, Decapoda). Zool. Mededel. Leiden, 41 (1): 1-13.
Chittelborough, R. G. & L. R. Thomas, 1969. Larval ecology of the Western Australian marine
crayfish, with notes upon other panulirid larvae from the Eastern Indian Ocean. Aust. Jour.
Mar. Freshwater Res., 20: 199-223.
Dotsu, Yoshie, K. Seno & S. Inoue, 1966. Rearing experiments on early phyllosomas of Ibacus
ciliatus (von Siebold) and I. novemdentatus Gibbes (Crustacea: Reptantia). Bull. Fac. Fisher.
Nagasaki Univ., 21: 181-194.
George, R. W., 1962. Description of Panulirus cygnus sp. nov., the commercial crayfish (or spiny
lobster) of Western Australia. Jour. Royal Soc. West. Aust., 45 (4): 100-110.
George, R. W. & L. B. Holthuis, 1965. A revision of the Indo-West Pacific spiny lobsters of the
Panulirus japonicus group. Zool. Verh. Leiden, 72: 3-36.
Gurney, Robert, 1936. Larvae of decapod Crustacea, 3. Discovery Rep., 12: 400-440.
Holthuis, L. B., 1946. Biological results of the Snellius Expedition. 14. The Decapoda Macrura
of the Snellius Expedition. I. The Stenopodidae, Nephropsidae, Scyllaridae and Palinuridae.
Temminckia, 7: 1-178.
-, 1966. On spiny lobsters of the genera Palinurellus, Linuparus and Puerulus (Crustacea, Deca
poda, Palinuridae). Symp. Crust. Mar. Biol. Assoc. India, 1: 260-278.
Johnson, Martin W., 1968. Palinurid phyllosoma larvae from the Hawaiian Archipelago (Palinu
ridae). Crustaceana, suppl., 2: 59-79.
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282 JOHNSON, PHYLLOSOMA SOUTH CHINA SEA
-, 1968a. On phyllamphion larvae from the Hawaiian Islands and the South China Sea (Pali
nuridae). Crustaceana, suppl., 2: 38-46.
-, 1971. The phyllosoma larvae of slipper lobsters from the Hawaiian Islands and adjacent areas.
Crustaceana, 20 (l): 77-103.
-, 1971a. The phyllosoma larva of Scyllarus delfini (Bouvier) (Crustacea, Decapoda). Crusta
ceana, 21 (2): 161-164.
-, in press, a. The palinurid and scyllarid larvae of the tropical eastern Pacific and their distri
bution as related to the prevailing hydrography. Bull. Scripps Inst. Oceanog., Univ. California.
Michel, A., 1969. Les larves phyllosomes du genre Panulirus-Palinuridae (Crustac?s, D?capodes)
du Pacifique Tropical sud et Equatorial. Cah. O.R.S.T.O.M., (Oc?anogr.) 7 (4): 3-19.
-, 1970. Les larves phyllosomes du genre Palinurellus von Martens (Crustac?s, D?ca
podes: Palinuridae). Bull. Mus. nation. Hist. nat. Paris, (2) 41 (5): 1228-1237.
Naiyanetr, P., 1963. Scyllarid lobsters and their phyllosoma larvae in the Gulf of Thailand. Jour.
Nat. Res. Counc. Thailand, 4 (1-4) : 63-72. [in Thai with English abstract].
Prasad, R. R. & P. R. S. Tampi, 1957. On the Phyllosoma of Mandapam. Proc. Nat. Inst. Sei. India,
23 (B) (1-2): 48-67.
-, 1959. On a collection of palinurid phyllosomas from the Laccadive Seas. Jour. Mar. Biol.
Assoc. India, 1 (2): 143-164.
-, I960. Phyllosomas of scyllarid lobsters from the Arabian Sea. Jour. Mar. Biol. Assoc. India,
2 (2): 241-249.
-, 1965. A preliminary report on the phyllosomas of the Indian Ocean collected by the Dana
Expedition 1928-1930. Jour. Mar. Biol. Assoc. India, 7 (2): 277-283.
Reinhardt, J., 1849. Phyllamphion en ny Slaegt af Stomatopodernes Orden. Vidensk. Meddel.
Dansk naturhist. Foren. Copenhagen, 1: 2-6.
Robertson, Philip B., 1969. The early larval development of the scyllarid lobster Scyllarides
aequinoctialis (Lund) in the laboratory, with a revision of the larval characters of the genus.
Deep-Sea Res., 16: 557-586.
Saisho, Toshio, 1966. Studies on the phyllosoma larvae with reference to oceanographical conditions.
Mem. Fac. Fish. Kagoshima Univ., 5: 177-239.
Saisho, Toshio & Kantaro Nakahara, I960. On the early development of phyllosomas of Ibacus
ciliatus (von Siebold) and Panulirus longipes (A. Milne Edwards). Mem. Fac. Fish. Kagoshima
Univ., 9: 84-90.
Shojima, Yoichi, 1963. Scyllarid phyllosomas' habit of accompanying the jelly-fish (preliminary
report). Japan. Soc. Sei. Fish., 29 (4) : 349-353.
Sims, H. W., 1966. The phyllosoma larvae of the spiny lobster Palinurellus gundlachi von Martens
(Decapoda, Palinuridae). Crustaceana, 11 (2): 205-215.
Tokioka, Takasi, 1954. Droplets from the plankton net. Pub. Seto Mar. Biol. Lab., 3 (3): 360-368.
Tokioka, Takasi & Eiji Harada, 1963. Further notes on Phyllosoma utivaebi Tokioka. Pub. Seto
Mar. Biol. Lab., 11 (2): 425-434.
Von Bonde, W., 1930. Post-Brephalus development of some South African Macrura. Rep. Fish. mar.
biol. Survey South Africa, 8: 2-42.
Wyrtki, Klaus, 1961. Physical oceanography of the Southeast Asian waters. Naga Exped. Rep.,
2: 1-195.
Received for publication 29 March 1971.
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