Table Of ContentPROC. BIOL. SOC. WASH.
104(3), 1991, pp. 561-568
A LARGE BRACHYURAN-LIKE LARVA OF THE
HIPPIDAE (CRUSTACEA: DECAPODA: ANOMURA)
FROM THE BANDA SEA, INDONESIA:
THE LARGEST KNOWN ZOEA
Joel W. Martin and Bernard Ormsby
—A
Abstract. collection of planktonic Crustacea from the Banda Sea, In-
donesia, contained several specimens ofa large, late stage zoea belonging to
the anomuran family Hippidae. As is true for the few previous descriptions of
hippidlarvae, the zoea bears a remarkable (but superficial) similarity to larvae
oftrue crabs (Brachyura) in that the carapace is spherical and the lateral and
rostral spines are distinctly deflected ventrally. The zoea is also remarkable in
its size; the width across the lateral carapace spines is over 12.0 mm, making
this the largest known zoea ofthe Anomura or Brachyura. Characters ofthe
zoea are compared to previous descriptions ofhippid larvae.
Planktonic larval stages of the decapod largest known zoea in the Decapoda, but
crustacean infraorders Brachyura and An- because, like previously describedlarvae of
omura are usually easy to distinguish. The the Hippoidea, it is superficially similar to
two most obvious characters concern the zoeal stages ofthe Brachyura.
carapace: inanomuransthecarapaceislong-
er than broad, and the rostral spine is di- Materials and Methods
rected anteriorly, whereas in brachyuran
larvaethe carapaceis sphericalandthe ros- Zooplanktonsampleswere collecteddur-
tral spine (usually present) is directed ven- ing an expedition ofthe R/V Alpha Helix
trally (see Williamson 1982). Another dif- to the area ofthe Banda Sea, Indonesia, in
ferentiating character is that the sixth 1975. One lot contained 7 large, late stage
abdominal segment, in later stages, always zoeaethatwehereinassigntotheanomuran
bears setose uropods in the Anomura, family Hippidae; this lot was collected in
whereas uropods are absent in zoeal stages the south Ceram Sea,just northwest ofthe
of the Brachyura (except for raninids, ho- islandofCeram(sometimesspelledSeram),
AH
molids, and some other "primitive" crab at Station 19(3°55.7'S, 128°07.7'E,trawl
families; Rice 1980, Martin 1991). Also, RMT-8, 0-150 m). An additional lot con-
mostanomuranlarvaehaveahair-likepro- taining4 specimens ofthissamezoeal stage
AH
cess between the first and third posterior waslaterfoundincollections from Sta-
spinesonthetelson,althoughthiscondition tion 28, also in the Ceram Sea (3°14.8'S,
changeswithontogeny(Konishi 1989,Mar- 127°38.0'E, trawl RMT-8, 0-100 m). Zoo-
tin 1991). plankton was collected with an otter trawl
mm
The present paperdescribes a large zoeal fitted with a 0.5 mesh plankton net.
stage ofthe anomuran superfamily Hippoi- Most of the crustacean samples from this
dea. The zoea is ofinterest not only for its expedition, including those with the hippid
mm
greatsize(over6.0 carapacelength,and zoeae,driedoutcompletelybeforetheycame
over 12.0 mm in carapace width, including to our attention (early 1989). Some ofthe
the lateral carapace spines), making it the larvae were rehydrated in Trisodium Phos-
PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
562
Fig. L Large hippid zoea from the Banda Sea, Indonesia: carapace, abdomen, and telson. a, entire zoea,
frontalview;b,same,sideview;c,ventralviewofcarapaceandspineswithtelsonandabdomenremoved(note
howventralcarapace is shaped to conform to opercular-like telson; d, lateralview ofabdomen and telson; e.
VOLUME NUMBER
104, 3 563
phate for a period of 48 hours and then sharp spines separated by minute serra-
transferred to 70% ethanol. A few of the tions; spines becoming more dense ap-
larvaewereleftdry.Alldrawingsweremade proachingposteriorterminusoftelson. One
withaWildM5APOdissectingmicroscope immovable tooth (Fig. If) on either side of
and a Zeiss compound stereoscope, each lateral border at about two thirds length of
with camera lucida, from specimens col- telson.
AH
lectedat Station 19. Measurementswere Uropods (Fig. Ig, h).—Long, extending
made from the single largest zoea, which approximately half length of telson (more
wasamongthefourspecimenstaken atAH thanhalfifextendedfullyorifsetationcon-
Station 28. sidered), biramous, outer ramus bearing 8-
10longsimple setae, innerramusunarmed.
Description
First antenna (antennule) (Fig. 2a).—
mm
Size.—Carapacelength 6.5 (fromor- Stout, distal portion with 5 tiers containing
bittoposteriorborderofcarapace, allowing aesthetascs.
for slight curvature); carapace width (ex- Secondantenna(Fig. 2b).—Protopodwith
cluding lateral spines) 6.3 mm; length of three paired, evenly spaced spines, becom-
rostralspine 10.8 mm;lengthoflateralspine ing progressively shorter distally; terminus
6.8 mm; distance from tip to tip oflateral ofprotopod spine-like; single spine located
spines 12.3 mm. at articulation of endopod. Endopod ex-
Sffl^^.—Because ofthe large size and the tendingnearlytotipofprotopod,withthree
development ofthe pleopods and uropods, spines, tapering to acute distal tip; total
thisisprobablyalatestagezoea, equivalent length ofendopod approximately Vs length
to a stage 4 or later larva as compared to ofprotopod.
earlieraccountsofdevelopmentintheHip- Mandible(Fig. 2c).—Anteriorborderwith
pidae (e.g., Hanson 1969). 1 sharp and 1 blunt tooth, both of which
Carapace(Fig. la, b, c).—Cephalothorax exceed length of denticle row. Denticles
smooth, weakly calcified, globose, with ex- acute, sclerotized, subequal in length, num-
tremelylongdowncurvedrostralspine(lon- bering 7-8.
gerthancarapacelength)andslightlydown- Maxillule(Fig. 2d),—Endopodconsisting
curved lateral spines (less than rostral of a single segment with 1 long, minutely
length). No dorsal spine notedin this stage. serrulate, terminal seta. Exopod bilobed;
Abdomen (Fig. Id, g).—Five articulating basal endite with 3 stout serrate spines on
somites, with the sixth fused to the telson. distal margin; coxal endite with 1 short
Somites 2 through 5 with pleopods. Somite proximal seta, and 4 longer minutely ser-
length and width roughly equal. Fifth so- rulate setae distally, 3 ofwhich are in distal
mite with blunt ventrolateral protrusions group.
extending slightly posteriorly. Abdomen Maxilla (Fig. 2e).—Endopod with two
tightlyflexedforward, withtelsonexpanded simple and two plumose setae on margin
and covering pereiopods and mouthparts plusonelongsimplesetaondistalterminus.
ventrally. Endite not seen. Scaphognathite with nu-
Telson(Fig. le,f,g).—Longerthanbroad, merous plumose setae spaced as illustrated.
terminallysubtriangular.Dorsalsurfacewith Maxilliped 1 (Fig. 2f).—Basis bearing 5-
scattered short setae. Posterior border with 7 plumose setae usually arranged in pairs;
dorsal(functionalventral)surfaceoftelsonshowingindistinctfusionoftelsonwith6thabdominalsomite(light
stippling);f, highermagnificationoflateraltoothandspinesonrightsideoffiguree;g, ventral surfaceoftelson
with uropods and abdominal somites 5 and 6 in place (stippled area denotes removed abdominal somite); h,
righturopod. Scale barcorrespondsto figuresa-conly.
564 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig. 2. Large hippid zoea from the Banda Sea, Indonesia: antennae and mouthparts. a, first antenna; b,
secondantenna;c,innerviewofleftmandible;d, maxillule;e, maxilla; f,firstmaxilliped;g, secondmaxUliped.
Scalebarappliestoall figures.
endopod4-segmented, withsetation 3, 2, 1, tation 3, 1, 1,4 (proximaltodistal); exopod
4 (proximal to distal). Exopod uniramous with 9 to 11 natatory setae.
with 10-12 natatory setae distally. Maxilliped3.—Present only as small un-
Maxilliped 2 (Fig. 2g).—Basis with two developed bud.
plumose setae; endopod 4-segmented, se- Pleopods (Fig. Id).—Well developed, ex-
VOLUME NUMBER
104, 3 565
tending from somites 2 through 5, unira- third, or first and fourth, teeth on the pos-
mous. terior border of the telson are strong and
well developed. In contrast, the telsons in
hippid larvae tend to be terminally round-
Discussion
ed, with only the first posterior process
The Banda Sea zoea is probably a late modified as a strongimmovable tooth, like
stage larva. The large size, coupledwith the the telson ofthe larva described here. Our
well developed pleopods and uropods, in- larvae clearlybelongto the Hippidaerather
dicate to usthat this zoeais atleastin stage than to the Albuneidae.
4 or higher, ifdevelopment proceeds as in
othergeneraoftheHippoidea(e.g., Hanson
Assignment to Genus
1969).
Ofthetwofamilies,AlbuneidaeandHip- Assignment to a particular hippid genus
pidae, that comprise the anomuran super- is more difficult. There are currently three
familyHippoidea,larvaearebestknownfor accepted genera in the Hippidae: Emerita
the albuneids, with larval stages ofat least Scopoli, Hippa Fabricius, and Mastigochi-
four genera known (e.g., Johnson & Lewis rusMiers(seeHaig 1970). Larvaeareknown
1942, Knight 1970, Sandifer & Van Engel only for Hippa and Emerita, and the zoeal
1972, Gore & Van Dover 1980, and Stuck stages ofspecies in these two genera are ex-
& Truesdale 1986, for Lepidopa; Menon tremely similar. Hanson (1969), inthe only
1937, and Seridji 1988, forAlbunea; John- description ofa complete larval series ofa
son & Lewis 1942, Boschi et al. 1968, and species of Hippa {H. cubensis (De Saus-
Knight 1968, for Blepharipoda; Konishi sure)),mentionedthatlarvaeofthetwogen-
1987, forLophomastix). Seridji(1988:1299) eradifferonlyinthatzoeaeofEmeritabear
presented a table comparing larval charac- fewer aesthetascs on the antennule, fewer
teristics ofthe "three known genera ofthe spines on the telson, fewer setae on the sca-
Albuneidae." However, Seridji apparently phognathiteborder,andaregenerallysmall-
was unaware ofthe genusLophomastix, for er than zoeae of Hippa. Hanson's (1969)
which larvae ofone species have been de- description was actually oflarvae ofH. tes-
scribed (Konishi 1987), and the genera tudinaria (Herbst), rather than H. cubensis,
Leucolepidopa, Paralbunea, Zygopa, Aus- according to Haig (1970). Hanson's com-
trolepidopa, and Stemonopa (see Efford & parison with species ofEmerita was based
Haig 1968,Efford 1969,Haig 1974b,Serene on the work of Menon (1933), Johnson &
1979), for which no larvae have been de- Lewis (1942), Rees (1959), and Knight
scribed. The majordistinguishingcharacter (1967). The descriptions of Smith (1877)
separating hippid larvae and albuneid lar- and Faxon (1879) oflarvae ofHippa were
vae, according to Seridji (1988), is that al- actually of species in Emerita. Hanson
buneids have uniramous pleopods in later (1969) also mentioned that "The antenna
zoeal stages. This distinction does not al- ofEmerita develops a long flagellum in the
waysholdtrue,asStuck&Truesdale(1986) fourth or fifth zoeal stage" that persists
documentedbiramous pleopods (albeit mi- through the megalopa stage, whereas this
nutely biramous) in a fourth stage zoea of ffagellum "is entirely lacking in Hippo'''
analbuneid{Lepidopabenedicti). No known (Hanson 1969:155),andherelatedtheabove
albuneid larva has uropods as distinctly bi- morphological differences to differences in
ramousasinthezoeadescribedherein. Ad- feedingmodesbetweenthetwogenera, spe-
ditionally, all known albuneid larvae pos- cies ofHippa being scavengers rather than
sess a distinctive rhomboid-shaped telson filterfeeders. Ourzoeaiscloserto Hanson's
that is always wider than long and is pos- descriptionoflarvaeof//, testudinariathan
teriorly somewhat truncate. The first and to existing descriptions of Emerita larvae
566 PROCEEDINGSOFTHE BIOLOGICALSOCIETYOFWASHINGTON
(e.g., Knight 1967 for E. rathbunae). Con- adactyla, H. pacifica, and H. celaeno, are
sequently, we tentatively assign our large widely distributed in the Indian and Indo-
zoea to Hippa. West Pacific Oceans (Haig 1974a). There-
Anadditionalcharacterstrengtheningthe fore,knowngeographicdistributionsofspe-
assignmentofthe BandSealarvaetoHippa cies in the three hippid genera would also
is that in Hanson's description of H. tes- suggest the probability that the zoea de-
tudinaria larvae there is a noticeable ven- scribed herein belongs to Hippa.
trolateral projection extending from the We confirm, ifourlarvaeandthose ofH.
posterior border ofthe fifth abdominal so- testudinariaaretypicalforthegenusHippa,
mite, similar to the condition ofthe Banda and if our zoea does in fact belong to a
Sea zoea. This is also seen in plankton-col- species of Hippa rather than to Mastigo-
lected stages attributedto Hippa by Lebour chirus, that larval size is quite different be-
(1959: fig. 17) and to a lesser extent in the tween the genera Hippa and Emerita. Our
figures ofAl-Kholy (1959: figs. 40-48). We zoea,withacarapacelengthofover6.0mm
have not seen this characterin descriptions (excludingthe rostral spine) andwith a dis-
mm
of Emerita larvae. Our zoea differs from tance ofover 12.0 between the tips of
Hanson's (1969) description of the larvae thelateralspines,exceedsthatforanyknown
ofH. testudinaria mostly in size. Hanson's larval stage, includingthe megalopal stages,
largest zoea, at stage VI, had a carapace ofany species oiEmerita. Indeed, this zoea
length ofonly 3.87 mm, whereas the Banda islargerthananyzoea, andeven megalopa,
SeazoeahasaCLof6.5 mm, whichiseven ofthe Brachyura reported in the survey of
larger than the megalopa stage ofH. testu- Hines(1986). Tothebestofourknowledge,
dinaria. The two species also differ mor- the zoea described in this paper, while
phologically;theBandaSeazoeahasamore smaller than some ofthe unusual larvae of
strongly curved rostral spine, paired spines palinuridanderyonoidlobsters,whichhave
on the second antenna, and a single (un- phyllosoma and eryoneichus larvae some-
paired)ventraltoothonthedistalmandible times considered "zoeal" (e.g., Williamson
border (compare to Hanson's fig. 3a). 1969),isthelargestknownzoealstageamong
Because of the above-mentioned uncer- the decapod Crustacea.
tainty concerning generic placement ofour
larvae, the geographic distribution of all Acknowledgments
hippidgenerashouldbeconsideredintrying WeareverygratefultoJanetHaig, Frank
to assign the Banda Sea larvae. The geo-
Truesdale, and Pedro Baez R. forreview of
graphicallyclosestspeciesofEmeritawould
the manuscript and for assistance with the
beE. emeritus(Linnaeus),which, according literature. We thank Hans G. Kuck forrec-
to Effbrd (1976), ranges from thewestcoast
ognizing the unusual size ofthe specimens
ofIndia east to southern Sumatra. At least
andbringingthemtoourattention,andMs.
one species ofMastigochirus, M. quadrilo-
batus Miers, is known from western Aus- JenniferChristiansenforassistingwithsort-
ing and preliminary identification of the
tralia and Queensland and westward into
plankton. Specialthanksareextendedtothe
thecentral sectoroftheIndianOcean(Haig
Ichthyology Section ofthe Natural History
1974a, 1974b); the only other known spe- Museum ofLos Angeles County for donat-
cies in the genus, M. gracilis (Stimpson),
ing the specimens.
also inhabits the Indo-West Pacific (China
Sea and Sri Lanka, Haig 1974a). Hippa is
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