Chapter 1
A Brief History and Some
Fundamentals
Craig M. Young
This book documents the anatomy of representative
larval forms from all marine invertebrate phyla known
ro possess rhern, Ir is designed as a resource for students
and researchers who must work with larvae, identify
[arvae, and recognize their parts. Adapted to a drifting
existence in the relatively viscous medium of sea water,
the larvae of marine invertebrates often possess forms
that seem foreign to those of us who move actively and
freely through the relatively fluid medium of airo These
bizarre little creatures reeling about on microscope slides
have inspired creative works ranging from poetry
(Carstang, 1985) ro science fiction (sea urchin larvae
recently starred as alien life forms on a popular televi-
sion show), but are much more than biological
curiosities. Indeed, larval development is the most
common developmental pathway in the animal king-
domo Representing the furure generations of most
marine animals, larvae are, in a very real sense, the
lifeblood of the sea.
BIPHASIC LIFE CYCLES
Humans and most orher mammals are boro as minia-
ture adults, complete in virtually every particular except
rhe capacity ro reproduce. In this familiar process
known as direet development, babies become juveniles
and juveniles become adults by simple growth. Relative
proportions ofbody parts may change during this trans-
formation, but the changes are gradual and slow. Such
direct development is not the norm among animals rhat
live in the sea. Of the approxirnately 32 phyla in the
animal kingdom, only one, the Onychophora, lacks any
marine representatives. The majority of sorne 170 000
ATLAS OF MARINE INVERTEBRATE LARVAE
ISBN 0-12-373608-0
animal species in 23 of rhe 31 remaining phyla have indi-
rect development with larvae that may difrer from adults
in size, form, habitar, mode of nutrition, and/or ability
ro rnove. Thus, life cycies of most marine animals are
'biphasic', with the transition between phases being a
dramatic, often rapid, series of morphological, behav-
ioral, and physiological changes termed metamorphosis.
The evolution of life cycles in marine invertebrates,
inciuding the origin of larvae, has been a subject of
much discussion. Reviews by jagersten (1972), Nielsen
(1998) and Strathmann (1978, 1985, 1988, 1993) provide
good overviews of these ideas.
A SHORT HISTORY OF LARVAL BIOLOGY
The anatomical drawings and micrographs of larvae pre-
sented in this volume are best appreciated in the context
of the ciassical works produced by embryologists work-
ing more than a cenrury ago without the benefit of
electro n microscopy. Many of the nineteenth-century
renditions oflarval anaromy were beautifully composed
and replete with detail: they remain among the very
best illustrations oflarval form even roday. The ciassical
studies of invertebrate embryology are reviewed in sev-
eral older texts (MacBride, 1914; Dawydoff, 1928; Kume
and Dan, 1968) and a history of larval ecology has been
presented elsewhere (Young, 1990); here I highlight only
a few of the landmark discoveries important to the
understanding of larval morphology.
The larvae of polychaete worms, barnacle cyprids,
crustacean zoeas, and megalops were first drawn by
Martinus Slabber (1778), an early microscopist in
Holland around the time of the American and French
Copyright © 2006, EIsevier Lrd.
AH rights of reproduction in any farm reservcd.
2 Atlas o/ Marine l n u e r t e b ra t e Larvae
revolutions (Figure u) and nearly a century after
brooded oyster embryos were first described with rhe aid
of simple microscopes (Brach, 1689). Veliger larvae and
zoea larvae were probably seen by others over the fol-
lowing 70 years (Thompson, 1828; MacDonald, 1858),
though their significance as young snails and crabs was
not recognized. Then, in the 1820S, a well-traveled ama-
teur naturalist named J. Vaughan Thompson described
metamorphosis in the zoea larvae of crabs (Thompson,
1828) and the cyprid larvae of barnacles (Thompson,
1830), and also discovered the affinity of the dirninutive
penracrinus stage of unstalked feather stars (Thompson,
1835). His larvae (Figure 1.2) were collecred with a
makeshift 'muslin hoop net' in Cork Harbor, Ireland
where Thompson was stationed as a surgeon in the
Royal Navy. The discoveries ofThompson began a 70-
year heyday during which descriptions oflarval anatomy
and metamorphosis were at the forefront ofbiology and
resolved many of rhe pressing zoological and evolution-
ary questions of the day. Within the next 20 years,
bryozoan (Ehrenberg, 1834) and ascidian larvae (Milne-
Edwards, 1842) were discovered, and developrnent was
described in opisthobranchs (Nordman, 1846) and sea
stars (Sars, 1844; Desor, 1851). Veliger larvae were already
known from the plankton by this time, bur were
assumed to be miniscule adults until MacDonald (1858)
argued that they might be the young of benthic snails.
, In the mid-nineteenth century, many important dis-
coveries were made by the German zoologist and
physiologist Johannes Müller. Having no apparent
knowledge ofThompson's discoveries, Müller invented
the plankron net and discovered a wide assortrnent of
larval forms at Helgoland and Marseille, including
echinoderm plutei, bipinnariae and auriculariae,
phoronid actinotrochs, and the flatworm larvae that are
still known as Müller's larvae. Müller illustrated his
larvae in beautiful detail (e.g., Müller, 1846, 1847, 1850),
though often upside-down (Figure I.3) and was not
always able to attribute thern to the correct phylum.
Compound microscopes improved rapidly during this
period, and the establishment of seaside marine labora-
tories such as those in Naples, Marseille, Plyrnouth, and
Woods Hole facilitated rhe culture of embryos and
larvae obtained by artificial insernination. Such zoolog-
icalluminaries as Metchnikoff (1871), Hatschek (1883),
Kowalevsky (1867, 1883), Prouho (1892), Lacaze-
Duthiers (1856), Claparede (1863), Alexander Agassiz
(1877), and many others detailed the embryology, larval
development, and metamorphosis of representatives
from most marine phyla (Figures 1.4, I.5). Studies of
larval diversity were advanced by rhe collections made
during Henson's 1889 Plankton Expedition that tra-
versed an hourglass-shaped track through the North and
South Atlantic. Larvae collected during this expedition
were often illustrated with beautiful color woodcuts
(Figure 1.6), though the identities of many of these
larval forms were not known and the anatomical derails
were sornewhat stylized (Simroth, 1895; Hacker, 1898;
Mortensen, 1898).
While the broad outlines of animal phylogeny were
being formulated in the decades following publication
of On the Origin ofSpecies by Means ofNatural Selection
(Darwin, 1859), the various larval forms still being dis-
covered played a pivotal role in evolutionary arguments.
Ernst Haeckel's biogenetic law (Haeckel, 1866) was
based largely on larval anatorny (MacBride, 1914), as
were the arguments that eventually brought rhis influ-
ential but overly simplistic idea to its knees (Sedgwick,
1894; Garstang, 1922, 1929). One endeavor stimulated in
part by evolutionary questions was the study of inverte-
brate celllineage, which attained its zenith in the early
years of the rwentieth century at the Marine Biological
Laboratories, Woods Hole (reviewed by Lillie, 1944;
Gross, 1985). Other notable contributions included
detailed descriptions of a vast assortment of echinoderm
larvae by Mortensen (e.g., 1921), and descriptions of
early embryology and larval development of a large
number of species, mostly from temperare habitats
worldwide (see embryology books by Kume and Dan,
1968; Dawydoff, 1928; MacBride, 1914).
Douglas P. Wilson (1932) and Theodor Mortensen
(1921) discovered independently that larvae could choose
their settlernenr sites and delay metamorphosis. This
discovery stimulated research that would dominate
larval biology, particularly in the British Isles, for the
next half century (reviewed by Meadows and Campbell,
1972; Hadfield, 1998). Much of the anatomical work
during this period focused on the morphological basis of
habitar selection and serrlernent, though studies of mor-
phogenetic mechanisms continued. In Denmark,
Gunnar Thorson devoted his career to rhe description of
larval forms and distributions, first as a student in East
Greenland then in the Oresund between Sweden and
Denmark. Thorson's rhoughtful and far-reaching
reviews (Thorson, 1946, 1950, 1964) established the
broad outlines for the discipline of Larval Ecology,
during a time when experimental embryologists in
Europe (e.g., Horstadius, 1939), Japan (e.g., Kume,
1929), and North America were unraveling details of
larval morphogenesis and metamorphosis. The advent
of the transmission and scanning electron microscopes
in rhe mid-rwentieth century resulted in a second
heyday for studies of invertebrate larval anatorny, with
great emphasis being placed on mechanisms of meta-
morphosis (e.g., Chia and Rice, 1978). The micrographs
from this latter period form the nucleus of the present
volume.
During the second half of the rwentierh century, the
 A Brief History and Some Fundamentals
significance of oceanic larval dispersal was fully appre-
ciated (Scheltema, 1985) and theoretical studies of
life-history evolution and the evolution of larval form
became popular topics of discussion (see reviews in
McEdward, 1995). After rwo centuries of study, larvae
finally carne into the mainstream of marine ecology
with the realization that most marine 'populations' are
in fact open metapopulations, controlled in large meas-
ure by rhe dispersal and colonization oflarvae (reviewed
by Morgan, 2000; Underwood and Keough, 2000).
THE MANY DEFlNITIONS OF 'LARVA'
The term 'larva has been used by biologists in a number
of different ways. Disagreement about the definition
arises in part from diverse invertebrate life histories and
in part from differing perspectives of researchers. Thus,
definitions oflarva may focus on structure, evolutionary
history, developmental sequences, or ecological atrrib-
utes. Hickman (1999) has offered a complex structural
definition that is useful in evolutionary studies:
The larva is a structural state or series of states that
occurs berween the onset of the divergent morpho-
genesis following embryonic development (c1eavage,
blastula, gastrula) and metamorphosis to the adult
body plan.
This definition makes no assumption about the habitar
of the larva or its ability to rnove, unlike the various eco-
logical definitions that are useful when considering
larvaeas agents of dispersal, feeding, or habitat selection,
One such definition (Chia, 1974; Giese and Pearse, 1975;
Young, 1999) is:
A larva is a postembryonic stage of the life cyc1e
which differs from the adult morphologically and is
capable of independent locomotion.
From the standpoint of developmental biology, one may
define larva from a purely mechanistic point of view by
considering the fates of various cells that form adult
and pre-adult structures (Davidson, 1991; Hickman,
1999):
A larva is the premetamorphic consequence ofType
1 embryogenesis, a type of specification of cell fates
that is fundamentally different in invertebrate larvae
until metamorphosis.
No single definition is useful for all approaches to the
study of larvae. The ecological definitions have been
criticized because sorne encapsulated or brooded stages
resemble larvae morphologically but do not disperse.
3
This is particularly common in gastropod mollusks,
where encapsulated stages may inc1ude trochophore-like
or veliger-like stages that rotate within their capsules
but hatch as crawl-away juveniles. A worker interested
in the evolution of form would regard these encapsu-
lated stages as larvae because their genes express larval
structures. However, one interested in life-history strat-
egies or dispersal might regard these stages as
functionally equivalent to direct development, since
they do not disperse.
Although we limit our definition of larvae in the
present volume to free-living, unencapsulated forms, we
also have inc1uded for the sake of completeness a few
groups that have direct development but whose juven-
iles are sufficiently different from the adult in form,
size, or habit to have been called larvae at sorne time in
the pasto These inc1ude the ctenophores, whose tiny
cydippid 'larvae' have the same basic body plan as the
adults, and the cephalopods whose 'paralarvae' are often
endowed with juvenile features that adapt thern ro a
planktonic existence and are either lost or transformed in
the adult stage. Likewise, many polychaete 'larvae' dis-
play no dramatic metamorphosis, making the transition
from larva to juvenile by the simple addition of seg-
ments. The megalops 'larva ofbrachyuran crabs and the
puerulus 'larva of spiny lobsters are today regarded as the
first juvenile instars of the life cycle, despite their free-
swimming habits and their role in definitive habitar
selection. We inc1ude them here because they have trad-
itionally been known as larvae. Sorne parasitic animals,
such as acanthocephalans, have larval stages that do not
disperse freely in the water column, but live within an
animal host. Because me life cyc1es of mese organisms are
biphasic or even triphasic, these internal stages are legit-
imately regarded as larvae. Sorne workers also regard the
asexually propagating polyps ofbenthic hydrozoans and
the large physonects of siphonophores as larval forms in
life cyc1es that alternate between sexual and asexually
propagating phases. For the purposes of this volume, we
have not accepted this very broad definition,
How LARVAE ARE FORMED
Most larvae originate from gametes, the products of
sexual reproduction. There are exceptions, inc1uding the
larvae of sorne bryozoans that are produced by polyem-
bryony (asexual mulriplication of embryos similar to
the formation of identical rwins in humans) and the
miracidia larvae of parasitic flarworrns that arise asexu-
ally from rhe redia stage. Ir has recently been discovered
that the oceanic larvae of sorne asteroids can propagare
by budding; the relative importance of sexual and
asexual reproduction in the production of these larvae is
unknown.
4 Atlas 01 Marine l n u e r t e b r a t e Larvae
Larvae produced sexually may orrgmate by
oviparous, ovoviviparous or viviparous development. In
oviparous development, gametes are spawned into the
sea, where embryos and larvae develop without any
parental careo This form of development occurs in most
phyla (Arthropoda is a notable exception). In ovovivip-
arous development, embryos are brooded by the parent
but the parent provides no nutrients for development
beyond that invested initially in the egg; the larva is the
first free-living stage. Ovoviviparity is particularly
common in clonal animals such as bryozoans, hydro-
zoans, and colonial ascidians, but is also the major mode
of development in crustaceans and probably sponges. In
viviparous development, nutrients are transferred from
the rnother to the embryo through a direct tissue con-
nection. Sorne directly developing ascidians and
echinoderms have viviparous development, but this is
the most rare developmental mode in the marine envi-
ronment. A fourth mode of developmenr, known as
mixed development, occurs in the mollusks, flatworms,
phoronids, and polychaetes. In mixed developrnenr,
eggs are deposited in capsules or gelatinous egg masses
and embryos hatch as larvae. Thus, early developmenral
stages are protected, but the larval stages disperse freely.
Mixed development differs from ovoviviparity because
in the former, embryos are not brooded within or under
rhe parent.
LARVAL FüRMS AND DIVERSITY
The body forms of invertebrate larvae are spectacularly
diverse (reviewed by Levin and Bridges, 1995) and a
large number of larval forms have been given specific
names (Table 1). Larvae of crustaceans are unciliated
and move by muscular locomotion, as do the tadpole
larvae of ascidians and the miracidia larvae of digenean
trernatodes (Chia et al., 1984). The majority of larval
forrns, however, are ciliated. The ultrastructure oflarval
ciliary bands and the associated nervous system has been
studied for a number oflarval forms (e.g., Lacalli, 1986)
and this Íiterature is reviewed by Nielsen (1987). Yolky
larvae with uniform ciliation, termed atrochal larvae,
are found in most phyla and are the most common
larval forms among the cnidaria, porifera, and plary-
helminthes. Sorne, including the planula larvae of
cnidarians, are simple spheres or spheroids, but others,
including the Müller's larvae of flatworms, are more
complex in shape and may have organized ciliary fields
with cilia of different lengths. Sorne atrochal larvae,
including the amphiblastula larvae of sponges and sorne
polychaete larvae, are ciliated on only a portion of the
body.
Trochallarvae, which have cilia organized into dis-
crete bands, are found in most phyla. Indeed, it has
been suggested (Nielsen, 1995) that an ancestral form
resembling a trochophore larva may have given rise to
the metazoan phyla. Trochal larvae may generally be
assigned to one of rwo larval types, each of which is
characteristic of a major subdivision of the coelomate
metazoans. The spiralian eucoelomates or protostornes,
induding rhe annelids and mollusks, characteristically
have trochophores or trochophore-like (e.g., veliger)
larvae. Trochophores and veligers have two parallel
bands of compound cilia, the prototroch and the
mesotroch. Additional ciliary bands (the meratroch,
neurotroch, and telotroch) may also be present, partic-
ularly in later larval stages. The deuterostome phyla,
incIuding the echinoderms and the hemichordates,
characteristically have larvae with a single ciliary band,
though the shape of this band may be complex and con-
voluted. In its basic form, this kind of larva has been
called rhe dipleurula.
Ir is nor uncommon to find larvae with several dif-
ferent names in the Iife cycle of a single species.
Polychaetes may be classified as trochophores early in
larvallife and as nectochaetes [ater on, as segments are
added. Sipunculans have rwo metamorphoses, one that
transforms the trochophore to a pelagosphera larva and
a second one at the time ofsettlemenr. Mollusk trocho-
phores become veligers with the development of the
larval shell. Bipinnaria larvae ofstarfish become brachio-
laria larvae, with the addition of rhe attachrnent organs
used at settlernenr. In holothurians, there is commonly
a progression from an atrochal virellaria larva to a dolio-
laria larva, wirh discrete bands of cilia, to an auricularia,
with a single convoluted band of cilia, and finally to a
pentactula, which develops tentacles and tube feet, loses
the ciliary bands, and takes up a benthic existence.
The larval stages of crustaceans are instars separated
by molting events and designated with individual names
or numbers. The early instars of brachyuran crabs are
known as zoeal stages and the terminal instar, or mega-
lops, is the juvenile that selects a habitar for settlement.
In similar fashion, barnacles have several naupliar instars
before molting to the non-feeding cyprid stage prior to
metamorphosis.
Parasites often have definitive and intermediate hosts,
each infected initially by a particular larval formo In a
typical marine digenean fluke, for example, the ciliated
miricidium larva infects a mollusk intermediare host by
burrowing into ir. Within rhe intermediate host, the
miricidium transforms itself and propagates asexually
to form a second larval form, the cercaria, which leaves
the host and propels itselfby means of a muscular rail to
burrow into the definitive host or to locate a setrlernenr
site where it is likely to be eaten. Cycliophorans have
very complex life cycles with pandora larvae and chor-
doid larvae. Rhombozoans, parasitic in the kidneys of
6 Atlas o/ Marine Invertebrate Larvae
cephalopods, have internal vermiform larvae and dis­
persive, ciliated infusoriform larvae.
MOOE OF NUTRITION
Larvae have been dassified by nutritional mode, site of
development, dispersal potential, and morphogenesis
(Levin and Bridges, 1995), with mode of nutrition
(Thorson, 1946) being the most often used. The litera­
ture on larval feeding and nutrition has been reviewed
thoroughly by Strathmann (1987), Han and Strathmann
(1995), and Boidron-Metairon (1995). The basic con­
cepts are introduced here ro familiarize the reader with
terminology common ro many phyla. A few larval
forms, induding sorne cnidarian planulae and ascidian
tadpoles, carry symbiotic algae from which they may
derive sorne of their nutrition. Most larvae, however,
are either plankrotrophic or lecithotrophic.
Lecithotrophic larvae, by definition, rely entirely on
yolk placed in the egg by the mother, whereas plank­
totrophic larvae acquire food during theirdeveloprnent,
either by collecting and concentrating partides or by
absorbing organic molecules from sea water.
Lecithotrophic larvae tend ro be large, yolky, opaque,
and colorful, whereas plankrotrophic larvae are generally
smaller and transparento The traditionally dear distinc­
tion between planktotrophy and lecithotrophy has
recently been blurred by the discovery that sorne
lecithotrophic larvae take up dissolved organic matter.
Moreover, sorne species, the so-called facultative plank­
totrophs, are capable of collecting food and will do so if
food is available, but have sufficient yolk to complete
development wirhout feeding if necessary. lf we
recognize thar all species are lecithotrophic at least
during the embryonic period, larval nutrition may be
regarded as a continuum along which the relative dura­
tions of the feeding and non-feeding periods vary.
Most planktotrophic larvae feed on phytoplankton,
though sorne crustacean and polychaete Íarvae are
predatory. Ciliated larvae use at least three different
mechanisms for collecting food particles. Planula larvae
of sorne cnidarians and a few polychaete trochophores
secrete a strand of mucus which is pulled behind the
larva like a fishing lineo The strand itself is ingested
along with any adherent particles. Trochophores,
veligers, and related larval forms collect food with paired
bands of cilia and transport the food in a ciliared food
groove that lies between the two bands. The anterior
prototroch is comprised of large, compound cilia that
funcrion simultaneously in locomotion and particle col­
lection. Particles are deílecred into the food gro ove,
apparently with the aid of a secondary band, the
mesotroch, whose function remains poorly understood.
Echinoderm and hemichordate larvae collect food with
unpaired convoluted bands of simple cilia. When indi­
vidual particles strike cilia of the band, individual cilia
reverse the direction of their beat, deflecting partides
roward the mouth and causing them to concentrare in
the circumoral field. They also capture sorne partides
without ciliary reversal, apparently by directing the flow
of partides into the oral region. Cyphonaures larvae of
anascan bryozoans use stiff cilia of the locomorory band
as a sieve to capture particles, Herbivorous crustacean
larvae capture particles by means of fine spines and hairs
on their locomotory appendages, whereas raptorial crus­
tacean larvae such as megalopae feed on individual prey
items in much the same way as adult crabs, using the
mourhparts to sort the food and chelipeds and peri­
opods to hold the prey and disassemble it.
LARVAL ECOLOGY, DISPERSAL, SETTLEMENT,
ANO METAMORPHOSIS
Ecological and behavioral aspects of larval biology gen­
erally fall outside the scope of the present volume,
though many of the chapters give brief descriptions of
certain aspects of larval life. The interested reader is
referred ro reviews in McEdward (1995) and Giese et al.
(1987 and other volumes in this series) and to the recent
review oflarval ecology by Morgan (2000) for conveni­
ent portals into this extensive literature.
The behaviors and cues involved in larval habitat
selection and settlernent have been studied extensively
and reviewed many times in the literature (e.g.,
Meadows and Campbell, 1972; Crisp, 1974; Pawlik,
1992). Hadfield (1998) has provided a recent overview of
the development of thought in this field.
Metarnorphosis involves multiple physiological
processes and morphogenetic changes including cell
death, resorption of tissues, reorganization of tissues
and organs, activation of organ systems, etc. Sorne of
these processes take place at dramatic speed, whereas
others, particularly those that require growth and re­
organization of tissues, may take longer ro complete.
Details are summarized for many groups by the authors
in Chia and Rice (1978) and many examples are given in
the individual chapters of the preseot volume.
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Zool. Haruard5, 1-131.
Boidron-Metairon, LE (1995). Larval nutrition. In Ecology o[
Marine Inuertebrate Laruae(ed. L.R. McEdward), pp. 223-248.
CRC Press, Boca Raton, Florida.
Brach, J. (1689). De ovis ostreorum. Ephemer. Acad. Nat. Curo
Dec. 2 Ann. 8, 506-508.
Chía, ES. (1974). Classification and adaptive significance of
developmental patterns in marine invertebrates. Tbalassia
[ugoslauica 10, 267-284.
 A Brief History and Some Fundamentals
Chia, ES. and Rice, M.E. (eds) (1978). Settlement and metamor­
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Chia, ES., Buckland Nicks, ]. and Young, e.M. (1984).
Locomotion of marine invertebrate larvae: a review. Can. J
Zool 62, 1205-1222.
Claparedc, A.R.E. (1863). Beobachtungen uber Anatomie und
Entwicklungsgeschichte wirbelloser Thiere an der Knste von
Normandie angesstellt. Wilhelm Engelmann Verlag, Leipzig.
Crisp, D.]. (¡974). Facrors influencing the setrlernenr of marine
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P.T. Grane, and A.M. Mackie (eds), pp. 177-265. Academic
Press, New York.
Darwin, C. (1859). On the Origin o[Species by Means o[Natural
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Davidson,E.H. (¡991). Spatial mechanisms of gene regulation in
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Dawydoff, e. (¡928). Traite d'Embryologie Comparee des
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Desor,P.].E. (1851). Unrirled notes. Proc. Boston Soco Nat. Hist. 3,
1l,13-14·
Ehrenberg, c.G. (¡834). Drirter Beitrag zur Erkenntnis grosser
Organization in der Richtung des kleinsten Raumes. Adhandl.
Konig. Akad. Wissensch. Berlin. 1833, 145-336.
Garstang, W (¡922). The rheory of recapitulation: a critical re­
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Garstang, W (¡985). Larval Forms and Otber Zoological Verses.
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Giese, A.c. and Pearse,] .5. (eds) (1975). Reproducrion of marine
invertebrates, Vol. 1. Academic Press, San Diego.
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Blackwell, Palo Alto and Boxwood, Paeific Grove, California.
Gross, P. (1985). Laying rhe ghost: embryonic development, in
plain words. Biol Bull. 168, 62-79.
Hacker, V. (r898). Die pelagischen polychaeten und achaeten
Larven der Plankton-Expedition. Ergeb. Atlantic Ocean
Plankton Exped. 2, 1-50.
Hadfield, M.G. (r998). The D.P. Wilson lecrure. Research on
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Haeckel, E. (1866). Generelle Morphologie der Organismen:
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Descendenz- Theorie. Reimer, Berlin.
Hart, M.W and Strathmann, R.R. (1995). Mechanisms and rates of
suspensionfeeding. In Ecology o[MarineInvertebrate Laruae (ed,
L.R. McEdward), pp. 193-22!. CRC Press, Boca Raton, Florida.
Harschek, B. (1878). Studien uber Enrwicklungsgeschichte der
Anneliden. Arb. Zool Inst. Wien 1, 1-128.
Hatschek, B. (1883). Uber die Enrwicklung von Sipunculus nudus.
Arb. Zool Inst. Univ. Wien. Zool. Sta. Triest 5, 61-14°.
Hickrnan, C.S. (1999). Larvae in invertebrate development and
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7
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Academic Press, London and San Diego.

FIGURE 1.1

Larval forms depicted by Slabber (1778) in one of the earliest lmown works
on microscopic marine animals
Thisfigure is reproduced in color between pages 606 and 607
The hand-colored plares were drawn by M. Slabber and P.M. Brasser and engraved by R. Mays.
The chapters were originally published in German as 18 separare works berween 1769 and 1778.
A. Larvae of a syllid polychaere being released frorn their morher's brood, from Slabber's Piare X.
B. Magnified view of a syllid larva (A), from Piare X.
C. Serigerous polychaere larva írorn Slabber's Piare XVII.

D Zoea larva of a brachyuran crab frorn Slabber's Piare V Compare wirh Figure 17.2.

E. Nauplius larva of a balanomorph barnacle from Slabber's Piare VI. Compare wirh figure 17.4G
F. Megalops larva of a brachyuran crab frorn Slabber's Piare XVIII. Compare with figure 17.12
G. Nauplius larva of a lepadomorph barnacle frorn Slabber's Piare VIII. Compare wirh figure
17-4E
H. Shrimp zoea frorn Slabber's Piare V Compare with Figure 17.6
10 Atlas 01 Marine Invertebrate Larvae

FIGURE 1.2

Plates aeeompanying the original descriptions oí barnacle and crab

metamorphosis by Thompson (1828, 1830)
A. Cirripede cyprid larva (top middle) and newly metamorphosed barnacles (bottom figures) from
Thompson (1830). Compare with Figures 12.9 D,E.
B. Zoea larva, with detailed drawings of its appendages and mouthparts, from Thompson (1828).
Compare with Figure 12.7.
A

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• :lE. s.
ZOEA_. _
 Chapter 17
Phylum Ar t h r op o d a : Crustacea
Alan W Haruey, joel W Martín and Regina
INTRODUCTION
Ruppert and Barnes (1996: 682) begin their introduc­
tion ro rhe subphylum Crustacea with the observation
rhat "... crustacean diversity is so great that a descrip­
tion of a 'rypical' form is impossible," and this is as true
of the larvae as it is of the adults. Furtherrnore, larval
development is known for only a small percentage of
species in five of rhe six recognized elasses of crustaceans,
and not at all for the elass Rernipedia, which suggests
that biologists have only begun ro sample the existing
¡: diversity of crustacean larval types. Thus, the present
,
1~
t:
chapter is in no way an exhaustive review of the fasci­
nating larval forms present in the Crustacea. Ir is meant
to be, instead, merely a glimpse oflarval diversity in the
world's most diverse taxon (Martin and Davis, in press),
Crustacean development can be direct, in which the
egghatches into a fully formed but miniature version of
me adult (as in most of the superorder Peracarida), or
entirely anarnorphic, in which change berween succes­
sive molts consists essentially of increasing body size,
adding segments and limbs, and developing existing
limbs. Usually rhere is sorne merarnorphosis, and at
times this can be striking.
The larval phase of marine crustaceans typically
ineludes a pelagic phase that usually lasts several weeks.
In sorne species, however, this planktonic larval phase
may last over a year. In many other taxa, particularly
those with abbreviated developrnent, the larvae may be
exclusively benthic, or spend only a very brief time in
che plankron (Figures 17.2A,B) (johnson, 1974; Serfling
and Ford, 1975; Rabalais and Gore, I985). Many crus­
tacean larvae are initially lecithotrophic, but most are
ATLAS OF MARINE INVERTEBRATE LARVAE
ISBN 0-12-373608-0
~tzer
planktivorous at sorne point in their pelagic life.
Lecithotrophy is commonly associated with abbreviated
development, and in lecithotrophic species, at least
sorne of the cephalic appendages are often reduced or
absent (Figures q.2C-E) (Rabalais and Gore, I985).
In those taxa with planktonic larvae and benrhic
adults, there is usually a morphologically distinct tran­
sitional phase that settles out of the plankton and
metamorphoses into rhe benthic post-larval phase
(Figures I7.9-12). Cornpetenr larvae use a wide variety
of chemica! and ractical cues to evaluate potential set­
tlernenr sites (Figure I7.uF), and are commonly able ro
delay metamorphosis in the absence of such cues
(Figure 17.uH) (Crisp, 1974; Harvey, 1996; O'Connor
and Judge, 1999). Curiously, in a number of taxa (such
as cirripedes and pagurid herrnit crabs), the settlernent
phase does not feed, which appears to set definite limits
on the ability of these taxa to delay metamorphosis
(Lucas et al., 1979; Dawirs, 1981; Harvey and
Colasurdo,1993).
The literature contains a vast number of descriptive
names for crustacean larval forms. This confusing situ­
ation arose in part because taxonomic names were
assigned to larval forms before ir was known to which
group they belonged, or often that they were even larvae
(e.g., Nauplius, Zoea, Megalopa, Glaucothoe, Cerataspis,
Eryoneicus, Erichthus, Alima, Phyllosoma, and Amphion).
Once these organisms were recognized as the larval
stages of other taxa, the now-defuncr generic narne often
became the descriptive name ofthe transitional stage for
thar group. This pracrice is fa!ling out of favor, alrhough
names for particularly distincrive or well-known larvae
persisto
Copyright © 2006, Elsevier Ud.
AlI rights of reproducrion in any forro reserved.

 Atlas of Marine Invertebrate Larvae
Typically, crustacean larvae have been grouped broadly
into three main types, which are identified by the
appendages primarily responsible for swimming; a 'nau­
plius' swims with its cephalic appendages (Figure 17.rA), a
'wea with its rhoracic appendages (Figure q.1C), and a
'megalopa' with its abdominal appendages (Figure 17.1D)
(Williamson, 1969; 1982a). Ecologica!ly, the nauplius and
zoea are usually dispersive phases and rhe megalopa is the
transitional settlernent phase (bur see Harvey, 1993).
The 'nauplius-wea-megalopa' series represents a
generalized developmental sequence as well, although
most crustaceans do not pass through all three phases.
Thus, carcinologists refer ro these as phases as well as
types. Within a given developmenral phase, successive
rnolts in which changes are essenrially anamorphic in
nature are referred ro as stages (e.g., ZI-Z5 in Figure
17.2A). The number of stages within a phase varies
widely among taxa, and is often but not always fairly
consistent wirhin species (Williamson, 1982a; Gore,
1985). The addition and development of segments and
appendages during anamorphic development within a
phase usually occurs in an anterior ro posterior direction
(Williamson, 1982a; Schram, 1986).
A free-swimming naupliar phase is known from at
leasr sorne species in all classes for which developmental
information is available. Zoeae and megalopae, as they
are commonly understood, are restricted ro rhe c1ass
Malacostraca. In other c1asses, developmental stages rhar
swim wirh thoracic appendages typically differ from
adulrs only in anamorphic derails and are thus consid­
ered to be juveniles rather rhan zoea larvae (e.g.,
copepodids in rhe Copepoda, although Williamson,
1982a classifies rhese as protozoeae). Megalopae are only
found in the Malacosrraca because true pleopods are
only found in this raxon (Williamson, 1982a).
NAUPLIUS
In terms of sheer nurnbers, the crustacean nauplius
(Figures 17.1A, 17.3, 17.4) has been called "the most
abundant type of multicellular animal on earth" (Fryer,
1987). The nauplius is the most prirnirive crusracean
larval type found in extant crustaceans, and the earliesr
free-swimming phase in crustacean development. By far
the most taxonomically widely disrributed type of crus­
tacean larva, rhe nauplius has been used as a key feature
that unites rhe entire subphylum Crustacea (Cisne,
1982; Schram, 1986; Walossek and Müller, 1990).
However, many crusraceans have lost this larval stage, or
pass through ir prior to eclosion from rhe egg.
Across the Crustacea, the nauplius exhibits a surpris­
ingly conservarive morphology (Figure 17.rA) (Dahrns,
2000). The body is typically covered wirh a dorsal cara­
pace, or cephalic shield, which is typically widesr
anteriorly, A carapace may be present at hatching, or
may develop in later stages; its appearance and develop­
ment can be either gradual or abrupto In rhe
orthonauplius (Figure 17.3), which is rhe typica! form of
newly hatched nauplius, there are initially only three pairs
of cephalic appendages: antennules, antennae, and
mandibles. The antennules are uniramous and lack a fla­
gellum, whereas the antennae and mandibles are typically
biramous. Initially, the rrunk exhibirs no sign of segmen­
tarion. A single median eye, visible near rhe anterior
margin of rhe carapace, is typical bur not universal (e.g.,
it is lacking in euphausiids; Mauchline and Fisher, 1969).
During subsequent rnolrs, rhe trunk develops seg­
rnentation, and rypically rhe remaining cephalic
appendages and the rhoracic appendages of rhe adults
are added and developed. Once a nauplius develops
appendages in addition to the three typical pairs of
cephalic appendages, it is called a meranauplius (Figure
17.4). These additional appendages, not used in loco­
motion, may inc1ude additional head appendages,
which may be rudimentary or functional, as well as
rudimentary thoracic appendages. In many groups, eggs
hatch as orthonauplii and beco me metanauplii after one
ro several molts: in sorne groups, (e.g., cephalocarids,
Figure 17.4A), eggs hatch directly as metanauplii. Late
naupliar stages often develop a pair of compound eyesin
addition to rhe naupliar eye.
Swimming and feeding in rhe nauplius are accom­
plished wirh the cephalic appendages. Feeding behaviors
seem to be triggered by chemosensory cues, at leasr in
barnac1es (Anderson, 1994). Nauplii feed on small
phytoflagellates, diaroms, and other microplankron
(Moyse, 1963; Barker, 1976; Srone, 1989). In a grear
many taxa, however, rhe nauplius is a non-feeding
phase, especially in rhe early stages. All nauplii in mala­
costracan taxa (e.g., euphausiids and dendrobranchs)
are lecithotrophic (Gurney, 1942).
There are many variations on this relatively simple
therne (Figures 17.3, 17-4). The relative size and degree of
setation of the cephalic appendages varies considerably
across taxa, as does the shape and armature of rhe cara­
pace. There are also more emphatically modified
naupliar forms. The ostracod nauplius, for exarnple,
possesses a late rally compressed, bivalved carapace like
that of the adult (Figure 17.3C). In fact, Cohen and
Morin (1990) argue that ostracods have juveniles, not
larvae, because there are no fundamental differences in
rhe morphology, behavior, or habitar of 'larval' and
adult ostracods. Hansen's (1899) 'Y-Iarvae' are peculiar
nauplii and cyprids (see below) rhat have been known
for more than 100 years, but their raxonomic idenriry is
still unknown (Figure 17.3F). The well-developed
carapace of a Y-nauplius is divided into a set of
symmetrical plates. There are one or rhree large spines
 Phylum Arthropoda: Cr u s t a c e a
on me posterior margin of the carapace, and a single
median spine that sits anterior to a unique structure
called the dorsal caudal gland. The function of this
gland, whose presence in the Y-nauplius is marked by a
slightly raised oval median plate in the carapace, is
unknown (Schram, 1970a, 1972).
Most nauplii are free-swirnrning, but sorne taxa
have benthic naupliar stages (e.g., Mystacocarida,
Cephalocarida, Ostracoda, and sorne harpacticoid
copepods; Dahrns, 2000). Many barnacles reta in
lecirhotrophic nauplii in the parent's mantle cavity,
releasing thern as cyprids (Anderson, 1986).
ZOEA
(Figures 17.1c' 17.5-8.) The genus Zoea (Bosc, 1802) was
created for what turned out to be the planktonic larval
stages of several species of brachyuran crabs. fu biolo­
gists began to better understand the Iife-history patterns
of crustaceans, the term zoea eventually carne ro refer to
any crustacean larva with functional thoracic
appendages, replacing numerous taxon-specific names
for the same stage (see Gurney, 1942 and Williamson,
1969, 1982a for a more complete discussion of these
larval names). Within this broad ter m lies a bewildering
diversity of shapes, sizes, and ornamentation, for the
zoeal body plan is most assuredly not subject to the
same conservatism as that of the nauplius. Here we can
do little more than summarize the general features of
zoea larvae, and mention sorne common or striking
variations.
AII zoeae possess a carapace, which covers the head
and the anterior portion of the thorax, and a pair of
compound eyes, which are almost always sessile in the
first stage and stalked in subsequent stages. A naupliar
eye is also found in the early zoeal stages of the
Amphionidacea, Caridea, and Palinuroidea, as well as of
those malacostracans that hatch as nauplii, hereafter
referred to as 'naupliar malacostracans.'
In early-stage zoeae, the antennular pedunde lacks
flagellae and is unsegmented, except in stomatopods
and 'naupliar malacostracans', where it has two or three
segments (Figures 17.5C,0,F, 17.8C); the basal segment
is annulated in protozoeal dendrobranchs (e.g., Oshiro
and Omori, 1996). Later zoeal stages may subdivide the
pedunde and add a dorsal flagellum, ventral flagellum,
or both. The antenna is biramous except in storn­
atopods (Figures 17.6B,E, 17.8C). The antennal exopod
is typically a flattened scale with setose margins (Figures
17.6B,E); major exceptions are found in the Brachyura,
where it is normally rod-like (Figure 17.7), and the unre­
lated taxa Palinura, Porcellanidae, and Hippidae, where
it is a simple spine (Figure 17.80). The mandibles are
usually at least slightly asymmetrical, usually lack a palp
339
in early stages (exceptions are rnost likely to be seen in
speeies with abbreviated developrnent), and often pos­
sess incisor and molar regions that are discernible but
much less developed than in adults, The maxillule and
maxilla are always present in zoeae, though their degree
of development varies among taxa (Figures 17.2C-E)
(Williamson,1982a).
Zoeal thoracic appendages are typically biramous.
Of rhe eight pairs found in adults (the first three pairs
correspond to maxillipeds, the last five to pereopods, or
legs), stage 1 zoeae may have as few as only the first pair
(e.g., in euphausiids) or as many as all eight, although
this latter is usually only seen in speeies with abbreviated
development. In most taxa, all eight pairs appear by the
final zoeal stage (Figure 17.1C).
The abdomen of stage 1zoeae typically consists of five
segments plus a terminal telson (Figures 17.6C,0, 17.70),
and the last segment subdivides in the next molt or two
(Figures 17.50, 17.6A). In sorne groups, the abdomen is
unsegmented through the first (e.g., 'naupliar malacos­
tracans') or most (e.g., Palinuroidea; Figure 17.80) zoeal
stages; in a few cases, first-stage zoeae have all six seg­
ments, and in a few brachyurans, the last abdominal
segment is fused ro the telson (Lucas, 1971). The shape of
the telson varies widely across the Crustácea, often fol­
lowing phylogenetic lines. The terminal margin of the
telson possesses several pairs of 'processes', and often a
single median process. These may be setae, articulated
spines, or fused spines, and rheir number, type, and size
are often surprisingly constant within a zoeal stage across
crustacean families or even orders. Uropods usually
appear in the third zoeal stages (Figure 17.50), developing
into fully articulated biramous structures over the next
few molts (Figures 17.6B,E), although there are numerous
exceptions. Their appearance and development may be
accelerated in taxa with abbreviated development (Figure
17.18); they appear more slowly in the Palinuroidea and
the Sromatopoda (Figure 17.8B), and are completely
absent in most brachyurans, which lack uropods as adults
(Figure 17.7).
Under the general heading of zoea Íarvae, several dis­
tinctive names remain in active use in the literature,
either for functional or historical reasons. Several of
these are briefly considered below.
Protozoea and mysis
(Figures 17.5C,E,F.) In dendrobranchiate shrirnps, the
first three zoeal stages are suffieiently different from the
subsequent zoeal stages that they have been given sepa­
rate names, protozoea and mysis. Unlike the archetypal
zoeae, protozoeae swim with the combined efforts of
the antennae and the first two pairs of thoracic
appendages; the remaining thoracic appendages are
absent or relatively undeveloped. In mysis larvae, the
 Atlas o/ Marine l n u e r t e b ra t e Larvae
antennae are no longer used for swimming and most ro
all of the rhoracic appendages are well developed. Mysis
larvae are not found in rhe Mysidacea; rhe name reflects
the apparent similariry between lare-srage dendrobranch
zoeae and mysid juveniles, in the same way that rhe
cypris larvae ofbarnacles are so named for their similar­
iry ro the osrracod genus Cypris (Williamson, 1982a).
Elaphocaris and acanthosoma
In sergesrid shrirnps, rhe prorozoea and the mysis srages
have a prominenr1y spinose carapace and are called
elaphocaris and acanthosorna, respecrively.
Calyptopis and furcilia
(Figures 17.5A,B,D.) As m dendrobranchs, rhe
euphausid meranauplius is followed by rwo disrincr
zoeal rypes, the calypropis and the furcilia. The calyp­
ropis is characrerized by a carapace rhar covers rhe sessile
eyes, an elongared trunk (compared wirh earlier srages),
and progressive segmentarion. There are normally sev­
eral calypropis srages in euphausids. In rhe furcilia,
which likewise persisrs rhrough several rnolts, rhe eyes
are movable and no longer covered by rhe carapace, and
rhe antennules, thoracic, and abdominal appendages
develop roward rhe adulr formo Unlike the rnysis, rhe
furcilia conrinues ro swim wirh borh rhe antennae and
the rhoracic appendages (Mauchline and Fisher, 1969).
Amphion
The amphion is rhe larva of Amphionides reynaudi, rhe
sole species in the order Amphionidacea (Williamson,
1973). In mosr respects, rhe amphion is similar ro a
caridean zoea, differing primarily in that irs relson is
spatulate in early srages, pointed and possessing only
rwo rerminal spines in larer srages. Ir has hepatic caecae,
and lacks chelae at any srage.
Phyllosoma
One of the largesr and most unusual decapod larval
forms is rhe phyllosoma (Figures 17.8D,E), a unique
srage shared by members of the relared Palinuridae
(spiny lobsters) and Scyllaridae (slipper lobsters), Sorne
giant phyllosomas have been reponed ro reach 8 cm in
rorallengrh (Roberrson, 1968a). Bur rhis size is decep­
rive; rhey are leaflike, flattened, and nearly paper-rhin,
possiblyan adapration for riding on the medusae of jel­
Iyfishes (Thornas, 1963; Herrnkind et al., 1976).
MEGALOPA
(Figures 17.1D; 17.10-12.) Many crusracean groups pos­
sess an interesring and somewhar intermediare larval
form that serves as a morphological and ecological tran­
sirion from rhe plankronic zoea ro the benrhic adult.
The rerminology of rhis transirional form has been quite
confusing. Commonly referred ro as posr-Iarvae
(Gurney, 1942), this rerm has fallen into disfavor because
rhey are clearly disrincr from subsequent juvenile srages.
Like zoea larvae, many of rhese rransirional forms were
originally described as new species or even genera befare
rheir developmenral status was recognized; once rhese
links were established, rhe now-defuncr generic name
ofren became rhe name of the rransirional srage for that
group. The earliesr of these, Megalopa (Leach, 1814), is
rhe transirional form of a brachyuran crab, and me rerm
megalopa has beco me (not wirhour conrroversy; see
Felder et al., 1985) the general rerm across the Crustacea
for the transitional larval srages where swimming is
achieved rhrough pleopods (Williamson, 1982a). As
mentioned earlier, megalopae are only found in the
Malacostraca, Anorher name describing rhis phase in
decapods is 'decapodid' (Kaesrner, 1970).
Megalopae are also morphologically intermediare
berween rhe zoeal and juvenile stages, often possessing
features characrerisric ofborh in addition to sorne char­
acrers unique ro the megalopa (Rice, 1981). In 'naupliar
rnalacostracans', there is a gradual transition from larval
ro adulr morphology from the lare mysid rhrough rhe
numerous megalopal srages and into the juvenile phases
(Gurney, 1942; Mauchline and Fisher, 1969). Likewise, in
sorne carideans, the transition from megalopa ro juvenile
is gradual enough that sorne aurhors repon two or rhree
megalopal srages while orhers question whether the con­
cepr of a megalopal phase is even applicable (e.g.,
Haynes, 1976; Rorhlisberg, 1980). In mosr orher groups,
however, many morphological structures acquire rhe
basic adulr characrerisric in rhe rnolt ro rhe megalopa,
which rhen represents a dramatic meramorphosis from
rhe last zoeal srage. In rhese groups, rhere is normally
only a single megalopal srage, alrhough eryonids have
several (see discussion under eryoneicus below).
Wirh a variable mix of zoeal and juvenile characters,
rhe megalopal phase shows even grearer diversity than
does rhe zoeal phase across the Malacosrraca. Generally,
megalopae have proponionarely larger eyes (hence the
narne) and smaller pereopods rhan do adults, and the
carapace is commonly narrower and smoorher. In the
megalopa, rhe abdomen projecrs posreriorly even when ir
is tucked under rhe rhorax in adulrs (i.e., in 'crabs')
(Figure 17.12). Megalopae rypically exhibir complete bilat­
eral syrnmetry, even in rhose raxa wirh pronounced
asymmetries as adulrs (e.g., chela asymmerry in homarid
lobsrers, fiddler crabs, and hermir crabs; abdominal asym­
rnetry in hermir crabs), Paguro id hermir crabs (families
Paguridae, Parapaguridae, and Lirhodidae) are un usual
in rhis respecr, as their megalopae often have asyrnmetri­
cal claws and uropods (e.g., McLaughlin et al., 1988).
In mosr crusraceans, sexual rnaturity is nor reached
 PhyLum Ar t b r op o d a : Cr u s t a c e a

until post-juvenile stages. However, ir appears that in
rhe Amphionidaceae, males matute as megalopae, and
females reach maturity at the next molt (Williamson,
1973), and one shrimp (Discias atLanticus) also apparently
reaches sexual maturity as a megalopa (Williamson,
1970). Sexual characrers sometimes develop even prior ro
the megalopal phase in a few groups (e.g., Palinura).
Several of the numerous names that have been used for
megalopae at lower taxonomic levels still persist in the lit­
erature today, in pan, beca use the organisms bear little
resemblance ro the archerypal brachyuran megalopa.
of a rype of pelagic lobster, owing ro their huge size and
the common presence in larger specimens of developing
male reproductive structures (Bouvier, 1905; Bernard,
1953). Early eryoneicus larvae swim with thoracic
appendages, and are thus referable ro zoeae; Íater stages
have well-developed pleopods (e.g., Figures 17.roC,D)
and fit the definition of megalopae in this respect
(Williamson, 1982a). However, the transitional phase
(i.e., megalopa in the ecological sense) has not yet been
idenrified for any eryonid, and in fact no eryoneicus
larva has been conclusively identified ro species.

Cyrtopia
Antizoea and pseudozoea
In the Euphausiacea, the eyrropia differs from the preced­
(Figures 17.8A-e.) The well-developed raptorial claws of
ing furcilia mainly in that swimming is now done with the
larval stomatopods (mantis shrimps) make thern imme­
pleopods. Thus, the cyrtopia fits the definirion of a mega­
diarely recognizable members of the plankton; although
lopae, bur most carcinologists now consider rhis to be a
newly hatched larvae are lecithotrophic and initially lack
stage in the furcilia phase (e.g., McLaughlin, 1980).
raptorial claws (e.g., Figure 17.8C), these rypically
remain in the parent's burrow (Manning and
Mastigopus
Provenzano, 1963). Like rypical zoeae, early larvae of the
These megalopae of sergestid shrimps are noteworthy
family Lysiosquillidae lack pleopods, have uniramous
mostly in that rhey have Íost all traces of the spines that
antennules, as well as five pairs of biramous thoracic
characterize the earlier zoeal phase. One un usual feature
appendages, and are called antizoeae. Other storn­
that characterizes both rhe megalopae and adults of this
atopods hatch as pseudozoeae, which initially possess
group is rhat the fourrh and fifrh pereopods are lost or
only rwo pairs of uniramous thoracopods but do have
greatly reduced.
functional pleopods. Antizoeae and pseudozoeae
develop anamorphically, and Íater stages are called
Puerulus
erichthus larvae in the superfamilies Lysiosquilloidea
(Figures 17.roA,B.) The unique phyllosoma larvae of
and Gonodactyloidea and alima larvae in rhe
Palinuroidea metamorphose into an un usual form of
Squilloidea (Provenzano and Manning, 1978;
megalopa termed rhe puerulus (or nisto, or pseudibacus,
Williamson, 1982a). The funcrional pleopods of the
in the family Scyllaridae). This phase is similar in form
pelagic erichthus and alima stages technically qualify
to rhe benthic adult, bur is transparent, with a srnooth,
thern as megalopae. However, borh erichthus and alima
decalcified cara pace and large, serose pleopods.
larvae are followed by a disrincr transitional form that is
the ecological and morphological equivalent of the
megalopal phase in other malacostracans (Schram,
AMBIGUOUS LARVAL TYPES
1986).
As Schram (1986) observes, not all known rypes of crus­ The naupliar phase of the subclass Thecostraca (bar­
tacean larvae can be easily placed in a functional nacles and rheir relatives) concludes with a
'nauplius-zoea-megalopa' classification. Most arnbigu­ rnetamorphic molt ro a cyprid phase (Figures 17.1B,
ous cases involve gradual transitions berween phases, or 17.9), which firs the broad definirion of a zoea but is
else reflect conflicts between the functional and ecolog­ functionally more analogous to the megalopa. The
ical components (e.g., a larva that swims with its cephalic appendages of the nauplius are lacking except
thoracic appendages but represents rhe transitional set­ for well-developed antennules, which the cyprid uses ro
dement stage). Several examples are described below. walk on the substrate in search of a suitable attachrnent
site. In addition, the cyprid possesses six pairs of tho­
Eryoneicus racic appendages. The fusiform cephalic shield now
Eryoneicus (Figures 17.roC,D) is the remarkable larva of encloses the body, but normally lacks a middorsal hinge,
an eryonid lobster (family Polychelidae). With a maxi­ unlike its ostracod namesake. A pair of compound eyes
mum-recorded size of over 6 cm, these crustacean larvae accompanies rhe naupliar eye. The cyprid does not feed,
ate exceeded in length only by sorne of the giant phyl­ and swims using irs thoracic appendages, as do zoeae
losomes, but rheir nearly spherical cara pace makes them (Anderson, 1994).
far more massive than any paper-thin phyllosoma. However, like a megalopa, the cyprid represents a
Eryoneicus larvae were long considered ro be rhe adults transition between the free-swimming larva and the
 Atlas o/ Marine Invertebrate Larvae
benthic adult, and is responsible for locating suirable
habitar for rhe sessile post-larval phase (Figure 17.90)
(Anderson, 1994). Cyprids vary widely among species in
rhe nature and precision of the cues they use to assess
potential settlernenr sites (Crisp and Meadows, 1963;
Lewis, 1978; Chabot and Bourget, 1988;Raimondi, 1988;
Young, 1991). The inability of non-feeding cyprids ro
replenish their energy reserves may consrrain the time
available to search for a suitable setrlement site; in sorne
sessile barnacles (Balanomorpha), it appears rhar older
cyprids become less discriminating regarding settlernent
sites (Crisp and Meadows, 1963).
In most barnacles, a newly settled cyprid undergoes a
rapid and profound metarnorphosis into a sessile juven­
ile barnacle. Ouring rhis metamorphosis, the animal
changes its orientation from dorsal to head-down, the
carapace and compound eyes are lost, and rudimenrs of
rhe capitular platcs appear. In those rhizocephalan bar­
nades known as kentrogonids, however, settlernent of
rhe cyprid on a suitable hosr leads not to a juvenile bar­
nacle, but to one of two unique larval forrns, the
kentrogon and the trichogon.
Kentrogon
An individual femal e kentrogonid rhizocephalan can
produce either small or large eggs, which develop into
small or large larvae. Cyprids that develop from small
eggs settle on a suitable host and develop into en dopar­
asitic females, whereas cyprids from large eggs setrle on
juvenile externae as males. Once a female cyprid settles
on a host, it molts into a unique attachment stage,
known as a kentrogon. In sorne species, the kentrogon
retains nearly all cypris organs except the thorax, but in
others rhe kentrogon keeps only those structures specif­
ically needed to perform its function. The kentrogon
rapidly develops a piercing sryler rhrough which the
kentrogon injects yet another instar called the vermigon.
This motile, worrn-shaped stage has an exceedingly
simple structure that includes only a few types of cells,
primarily an epidermis, large cells that will develop into
somatic tissues such as muscles, and rhe germlinecells
(Ritchie and Hoeg, 1981; Heeg, 1985, Glenner et al.,
2000). The vermigon is enelosed within a rhin cuticle
thar later develops into the nutrient-absorbing cuticle of
the internal parasite.
Trichogon
Male rhizocephalan cyprids take another path,
Following setrlement (onro immature fernales), they
rneramorphose into a trichogon, which like the ver­
migon is elongate and worm-shaped, and may be motile
as well. The trichogon enters the aperture of the externa
and migrares through the fernale's mantle cavity toward
the seminal receptades. The spinose cutide of the tri­
chogon is shed, forming a plug thar prevents rhe entry
of subsequent trichogons into the same receptacle. The
trichogon eventually reaches the lumen of the receptade
where irs germinative cells mature into spermatogonia
(it is unclear whether male sornatic tissue also survives
and functions wirhin the receptacle). The trichogon
remains with the female for rhe remainder of her life­
time (Hoeg, 1987).
A few 'Ycyprids' have been found developing within
Y-nauplii or collecred from the plankton (McMurrich,
1917; Schram, 1970b; Ito, 1989). These distinctive
cyprids (Figure 17.9B) have a plated carapace that does
not cover the abdomen, which is also covered with
plates posteriorly and possesses a pair of morpholcgi­
cally complex caudal ramio fu in cirripedes, the Y-cypris
has six pairs of biramous thoracic appendages, and the
only cephalic appendages present are rhe antennules,
although these differ considerably between the twO
types of cyprids (Figures 17·1B, 17.9B).
ACKNOWLEDGEMENTS
We rhank Craig Young for inviting us to contribute to
rhis atlas, and Shane Ahyong, Roy Caldwell, Anne
Cohen, Jens Hoeg, and Dieter Walossek for helpful dis­
cussions. Jens Hoeg improved the section on cyprids.
This work was supported in part by the National
Science Foundation via the following grants:
OEB-9996I62, from the Systernatic Biology program,
to A.W Harvey; OEB 9972IOO, from the Biotic Surveys
and Inventories program, to T.L. Zimmerman and].W
Martin: OEB 9978193, from the PEET initiative of me
Systernatic Biology program, to J.W Martin and D.K.
Jacobs.
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in decapods. In Crustacean Issues II: Larva! Growth (ed. A.M.
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Raimondi, P.T (r988). Serrlernenr cues and determination of the
verrical limir of an interridal barnac1e. Eco!ogy 69, 400--407.
Rice, A.L. (r981). The megalopa stage in brachyuran crabs, the
Podotrernata Guinor. j. Natl Hist. 15, I003-IOIl.
Rirchie, L.G. and Heeg, J.T. (r981). The life history of
Lernaeodiscus porcellanae (Cirripedia Rhizocephala) and coevo­
lution with irs porcellanid host, j. Crust. Biol. 1, 334-337.
Roberrson, P.B. (r968a). The complete larval developmenr of rhe
sand lobsrer, Scyllarus americanus (Srnirh), (Decapoda,
Scyllaridae) in rhe laborarory with notes on larvae frorn rhe
plankton. Bull. Mar. Sci. 12, 294-342.
Robertson, P.B. (r968b). A giant scyllarid phyllosoma larva frorn
the Caribbean Sea, wirh notes on smaller specirnens
(Palinuridae). Crustaceana Supp!. 2, 83-97.
Rodrigues, S.l. and Manning, R.B. (1992). First stage larva of
Coronisscolopendra Latreille (Stomaropoda: Nannosquillidae).
j. Crust. Biol. 12, 79-82.
Rorhlisberg, p.e. (r980). A complete larval descriprion of Pandalus
jordani Rathbun (Decapoda, Pandalidae) and irs relarion ro
other members of the genus Pandalus. Crustaceana 38, 19-48.
Rupperr. E.E. and Barnes, R.D. (r996). [nuertebrate Zoo!ogy, 6th
edn. pp. 682. Saunders College Publishing, Fort Worth.
Sanders, H.L. (1963). The Cephalocarida: functional morphology,
larval developmenr, comparative external anatomy. Mem.
Conn. Acad. Sci. 15, 1-80.
Schram, TA. (1970a). On rhe enigrnaticallarva nauplius Y type 1.
Sarsia 45, 53- 68.
Schram, TA. (r970b). Marine biological investigations in the
Bahamas 14. Cypris Y, a later developmenral stage of nauplius
Y. Sarsia44, 9- 24.
Schram, T.A. (r972). Further records of nauplius Y type I~ from
Scandinavian warers, Sarsia 50, 1-24.
Schram, ER. (1986). Crustacea. Oxford University Press, Oxford.
Serfling, SAo and Ford, R.E (1975). Ecological studies of the
pueruli larval stage of rhe California spiny lobster, Panuliru:
tnterruptus. Fish. Bull. 73, 360--377.
Seridji, R. (r988). Sorne plankronic larval stages of Albunea
carabus (L., 1758) (Crustacea, Decapoda, Anomura). j. Nati
Hist. 22, IZ93-1300.
Srone, C,], (r989). A comparison of algal diets for cirripede nau­
plii. j. Exp. Mar. Biol. Eco!. 132, 17-40.
Strasser, K.M. and Felder, D.L. (r999). Larval developmenr in
two populations of rhe ghost shrimp Callichirus major
(Decapoda: Thalassinidea) under laboratory condirions. J
Crust. Bio!. 19, 844-878.
Stuck, K.e. and Truesdale, EM. (1986). Larval and early postlar­
val developrnent of Lepidopa benedicti Schmitt, 1935
(Anomura: A1buneidae) reared in the laborarory. j. Crust. Biol.
6,89-Il O.
Subrahmanyam, e.B. (1971). Descriptions of shrimp larvae
(family Penaeidae) off rhe Mississippi coast. Gu/fRes. Reports3.
241-258.
Subrahmanyam, e.B. and Gunrer, G. (r970). New penaeid
shrimp larvae frorn the Gulf of Mexico (Decapoda, Penaeidea).
Crustaceana 19, 94-98.
Suh, H.-L., Soh, H.Y. and Hong, S.Y. (r993). Larval dcvelopmenr
of the euphausiid Euphausiapacifica in the Yellow Sea. Mar.
Biol. 115, 625-633.
Thomas, L.R. (r963). Phyllosoma larvae associared with medusae.
Nature (Lond.) 198, 202.
Walley, L.J. (1969). Studies on rhe larval structure and metarnor­
phosis of Balanus balanoides (L.). Phi!. Trans. R. SOCo B 256,
237- 280 .
Walossek, D. and Muller, K,J. (r990). Upper Cambrian srem-lin­
eage crusraceans and rheir bearing upon rhe rnonophyletic
origin of Crustacea and the posirion of Agnostus. Lethaia 23,
409-427.
Wear, R.G. and Fielder, D.R. (r985). The marine fauna of New
Zealand: larvae of the Brachyura (Crusracea, Decapoda). New
Zea!. Oceanogr. Inst. Mem. 92, 1-90.
Webber, WR. and Booth, J.D. (r988). Projasus parkeri (Stebbing,
1902) (Crustacea, Decapoda, Palinuridae) in New Zealand and
descriprion of a Projasus puerulus from Ausrralia. Nat. Mus.
New Zea! Rec. 3, 81-92.
Wehrtrnann, !.S., Veliz, D., Pardo, L.M. and Albornoz, L. (r996).
Early developmenral stages, inc1uding rhe first crab, of
Allopetro!isthes angu!osus (Decapoda: Anornura: Porcellanidae)
from Chile, reared in rhe laborarory. j. Crust. Bio!. 16,
730--747·
Williamson, D.l. (r969). Names of larvae in rhe Decapoda and
Euphausiacea. Crustaceana 16, 2IO-213.
Williamson, D.!. (1970). On a collecrion of plankronic Decapada
and Srornaropoda (Crusracea) from the easr coast of the Sinai
Peninsula, norrhern Red Sea. Bull. Sea Fish. Res. Stn Haift 56,
1-48.
Williamson, D.l. (r973). Amphionides reynaudii (H. Milne
Edwards), represenrarive of a proposed new order of eucaridan
Malacostraca. Crustaceana 25, 35-50.
Williamson, D.!. (r982a). Larval morphology and diversity. In
The Bio!ogy ofCrustacea, Vo!. 2. Embryo!ogy, Morpho!ogy, and
Genetics (ed. L.G. Abele), pp. 43-IlO. Academic Prcss, New
York.
Williarnson, D.!. (1982b). The larval characters of Dorhynchus
thomsoniThomson (Crustacea, Brachyura, Majoidea) and their
evoluciono j. Natl Hist. 16, 727-744.
Young, B.L. (r991). Spartina axil zones: preferred setdernenr sites
of barnac1es. j. Exp. Mar. Bio!. Eco!. 151, 71-82.
 Atlas 01 Marine Invertebrate Larvae

FIGURE 17.1
Examples of primary types of crustacean larvae

A. Generalized nauplius, ventral view. (Reproduced with permission from Dahrns, 2000.)
B. Cyprid of Semibalanus balanoides (family Archaeobalanidae), lateral view. Scale bar: 0.25 mm.
(Reproduced with permission from Walley, 1969.)
C. Zoea of Coenobita compressus (family Coenobitidae), lateral view. Scale bar: 0.5 mm.
(Reproduced with permission fróftl Brodie and Harvey, 2001.)
O. Megalopa of Notomithrax minor (family Majidae), lateral view. Scale bar: 0.5 mm. (Reproduced
with permission from Wear and Fielder, 1985.)
A
\ Carapace Naupliar Eye

\ '", y -,,, - - - - - ;/
B
Naupliar Eye
Carapace

Antennule
Thoracic
Appendages

Rostrum
e Antennule
Abdomen
~ Antenna

Maxilliped 3 Maxilliped 1

Telson
Telson
o
Antennule

Uropod
Rostrum

'T". '
Antenna -­1/
{ Pleopods

~~
Pereopods 2-5

 Atlas o/ Marine l n u e r t e b ra t e La r u a e

FIGURE 17.2

Typical and abbreviated development in congeneric crustaceans (family

Coenobitidae; A,B) and examples of reduced feeding structures

in lecithotrophic larvae (C-E)

A. Planktotrophic zoeal developrnenr in Coenobita compressus, dorsal view. Scale bar: 1.0 mm.
ZI-Z5: Zoea stage 1-5. (Reproduced with permission from Brodie and Harvey, 2001.)
B. Lecithotrophic zoeal development in Coenobita uariabilis, dorsal view. The two species are
similar as megalopae (shown here) and as adults. Scale bar: 1.0 mm. (Reproduced with
permission from Harvey, 1992.)
C. Maxillule in first stage zoea of C. compressus. Scale bar: 0.25 mm. (Reproduced with permission
from Brodie and Harvey, 2001.)
D. Maxillule in fifth zoea of C. compressus. Scale bar: 0.25 mm. (Reproduced with permission from
Brodie and Harvey, 2001.)
E. Maxillule in first stage zoea of C. uariabilis. Scale bar: 0.25 mm. (Reproduced with permission
from Harvey, 1992.)
A

~~

\ l'

i I

I '
1/ ,1"
I
" .;"
35° Atlas o/ Marine l n u e r t e b ra t e Larvae

FIGURE 17.3
Examples of orthonauplius larvae

A. First stage nauplius of Hemicyclops japonicus (family Clausidiidae), ventral view. Scale bar: 30
flm. (Reproduced with permission from Itoh and Nishida, 1997.)
B. Second stage nauplius of Paramphiascella jUlvofasciata (family Harpacticoida), ventral view.
Scale bar: 30 flm. (Reproduced with permission írom Dahrns, 2000.)
C. First stage nauplius of Cypris fasciata (family Cyprididae), lateral view. Scale bar: 30 flm.
(Reproduced with permission from Dahrns, 2000.)
D. First stage nauplius of Trachypenaeus (family Penaeidae), dorsal view. Scale bar: 0.3 mm.
(Reproduced with permission from Subrahmanyam, 1971.)
E. Sixth stage larva of Baccalaureus falsiramus (family Ascorhoracidae), ventral view, Scale bar: 0.1
mm. (Reproduced with permission from Dahrns, 2000.)
F. 'Type VI' Y-nauplius from Greenland (infraclass Facetotecta), dorsal (left) and ventral view.
Scale bar: 0.1 mm. (Reproduced with permission from Grygier, 1987.)
352 Atlas o/ Marine Ln u e r t e b r a t e Larvae

FIGURE 17.4

Examples of nauplius larvae

A. Third stage larva (metanauplius) of Hutchinsoniella macracantba (family Hutchinsoniellidae),

ventral view. Scale bar: 0.1 mm. (Reproduced with permission from Sanders, 1963.)
B. Larva (metanauplius) of Thysanopoda tricuspidata (family Euphausiidae), dorsal view. Scale bar:
0.25 mm. (Reproduced with permission from Knight, 1973.)
C. Larva (rnetanauplius) of Euphausia pacifica (family Euphausiidae), dorsal view. Scale bar: 0.2
mm. (Reproduced with permission from Suh et al., 1993.)
D. First stage larva (metanauplius) of the Upper Cambrian fossil maxillopod Bredocaris
admirabilis, ventral view. Scale bar: 50 ~m. (Scanning electron micrograph courtesy of Dieter
Walossek.)
E. Larva (orthonauplius) of an unidentified barnacle collected from plankton in the Bahamas,
ventral view. Magnification unknown. (Photograph by C.M. Young.)
F. First stage larva (metanauplius) of Derocheilocaris typicus (family Derocheilocarididae), ventral
view. Scale bar: 50 ~m. (Reproduced with permission from Hessler and Sanders, 1966.)
G. Larva (orthonauplius) of Pollicipes polymerus (family Pollicipedidae) from the San Juan Islands,
Washington, ventral view. Nauplii in this species range in length from approximately 0.2-0.6
mm (Lewis, 1975). (Phorograph by C.M. Young.)
B \J e

\ /

?1 ­
rr­
354 Atlas o/ Marine Ln u e r t e b r a t e Larvae

FIGURE 17.5
Euphausiid and dendrobranch zoeae

A. Calyptopis of euphausiid (family Euphausiidae), dorsal view. Scale bar: 0.25 mm. (Photograph
by Jocelyne Martin/IFREMER.)
B. Fureilia of euphausiid (family Euphausiidae), lateral view. Scale bar: 0.25 mm. (Photograph by
Jocelyne Martin/IFREMER.)
C. Third stage protozoea of Gennadas (family Penaeidae), dorsal view. Scale bar: 0.5 mm.
(Reproduced with permission from Subrahmanyam and Gunter, 1970.)
D. Third stage calyptopis of Thysanopoda tricuspidata (family Euphausiidae), dorsal view. Scale
bar: 0.25 mm. (Reproduced wirh permission from Knight, 1973.)
E. Mysis of penaeoid shrimp (order Dendrobranchia), dorsal view. Body length approxirnarely 3
mm. (Photograph by Peter Parks/Drawing Quest 3-D.)
F. Third stage mysis of Solenocera (family Penaeidae), dorsal view. Scale bar: 0.5 mm. (Drawing
reproduced with permission after Subrahmanyam, 1971.)
 Atlas o/ Marine Invertebrate Larvae

FIGURE 17.6
Zoeae of Caridea, Thalassinidea, and Anomura
A. Third stage zoea of Crangon crangon (family Crangonidae), lateral view. Scale bar: 0.5 mm.
(Photograph by jocelyne Martin/IFREMER.)
B. Fourth stage zoea of Callichirus major (family Callianassidae), dorsal view. Scale bar: 0.5 mm.
(Reproduced with permission fram Strasser and Felder, 1999.)
C. First stage zoea of Galathea dispersa (family Galatheidae), dorsal view. Scale bar: 0.5 mm.
(Photograph by jocelyne Martin/IFREMER.)
D. First stage zoea of Albunea carabis (family Albuneidae), dorsal view. Scale bar: 0.5 mm.
(Repraduced with permission fram Seridji, 1988.)
E. Ninth stage zoea of Latreutcs laminirostris (family Hippolytidae), dorsal (upper) and lateral
view. Scale bar: 0.5 mm. (Reproduced with permission fram Kim and Hong, 1999.)
F. First stage zoea of Pisidia longicornis (family Porcellanidae), lateral view. Scale bar: 0.5 mm.
(Photograph by Jocelyne Martin/IFREMER.)
 Atlas o/ Marine Invertebrate Larvae

FIGURE 17.7
Zoeae of brachyuran crabs

A. First stage zoea of Dromia wilsoni (family Drorniidae), dorsal view. Scale bar: 0.25 mm.
(Reproduced with perrnission fram Wear and Fielder, 1985.)
B. Second stage zoea of Philyra platychira (family Leucosiidae), lateral view. Scale bar: 0.2 mm.
(Repraduced with permission fram Ko, 2000.)
C. First zoeal stage of Notomithrax minor (family Majidae), lateral view. Scale bar: 0.5 mm.
(Repraduced with permission fram Wear and Fielder, 1985.)
D. Zoea of Tetralia (family Xanthidae), dorsal view. Length fram tip of rastrum to tip of dorsal
spine: 6.4 mm (George and John, 1975).
E. First zoeal stage of Homola barbata (family Homolidae), lateral view. Scale bar: 0.25 mm.
(Reproduced fram Williamson, 1982b.)
F. Second zoeal stage of Dorhynchus thomsoni (family Majidae), lateral view. Scale bar: 0.5 mm.
(Repraduced with permission fram Williamson, 1982b.)
G. Third stage zoea of Thia scutellata (family Thiidae), anterolateral view. Scale bar: 0.5 mm.
(Photograph by Jocelyne Martin/IFREMER.)
H and I. Unidentified brachyuran zoeae fram the Bahamian plankron. Magnifications unknown.
(Photographs by C.M. Young.)
J. Unidentified brachyuran zoea fram the plankton at Friday Harbor, Washington, frantal view,
showing carapace spines. Magnification unknown. (Photograph by C.M. Young.)
B

Dorsal
Carapace
( Spine

Lateral
Carapace

SPT

.:

Rostral Spine
 Atlas o/ Marine l n u e r t e b r a t e Larvae

FIGURE 17.8

Atypical zoeal forms

A. Pseudozoea of a stornatopod (superfamily SquiIloidea), ventral view. Body lengrh

approximately 3 mm. (Phorograph by Peter Parks/Orawing Quest 3-D.)
B. Fifth pseudozoeal stage (second pelagic stage) of Gonodactylus oerstedii (family
Conodacrylidae), lateral view. Scale bar: LO mm. (Reproduced with permission from
Provenzano and Manning, 1978.)
C. Antizoea of Coronis scolopendra (family Lysiosquillidae), lateral view. Scale bar: 0.25 mm.
(Reproduced with permission from Rodrigues and Manning, 1992).
D. Fourth stage phyIlosoma of Tbenus orientalis (family SeyIlaridae), ventral view. Scale bar: 0.25
mm. (Reproduced with permission from Barnett et al., 1984.)
E. Early stage phyllosorna collected from plankton in the Baharnas, probably of Panulirus argus
(family Palinuridae), dorsal view. Body length 3mm. (Photograph by C.M. Young.)
 Atlas 01 Marine l n u e r t e b r a t e Larvae

FIGURE 17.9
Cyprids

A. Typical balanomorph cyprids (superorderThoracica.) Scale bar: 0.25 mm. (Photograph by

Jocelyne Martin/IFREMER.)
B. Cypris Y-larvaof Hansenocaris ftrcifera (infraclass Facetotecra), in dorsal (Ieft) and lateral view.
Scale bar: 0.1 mm. (Reproduced with permission from Ito, 1989.)
C. Dendrogaster defOrmator (family Dendrogasteridae), lateral view. Collected from adult
parasitizing the asreroid Novodinea antillensis at bathyal depths in the Bahamas. Scale bar: 0.25
mm. (Photograph by CM. Young.)
O. Cyprid larva of the barnacle Balanus ampbitrite, with thoracic appendages extended. Scale bar:
0.25 mm. (Photograph by CM. Young.)
E. Dense aggregation of settling balanomorph cyprids (probably Balanus glandula) (family
Balanidae) near Friday Harbor, Washington. Carapace length of cyprids in B. glandula is
0.6-0.7 mm (Brown and Roughgarden, 1985.) (Photograph by CM. Young.)
 Atlas 01 Marine Invertebrate Larvae

FIGURE 17.10

Megalopae of the infraorder Palinura

A. Puerulus of Scyllarus americanus (family Scyllaridae), lateral view. Scale bar: 1.0 mm.
(Reproduced with permission from Robertson, 1968b.)
B. Puerulus of Projasus (family Palinuridae), lateral view. Scale bar: 10.0 mm. (Reproduced with
permission from Webber and Booth, 1988.)
C. Eryoneicus larva of polychelid lobster (family Polychelidae), lateral view, Scale bar: 10.0 mm.
(Photograph by R. Meier, from a speeimen in the collections of the Natural History Museum
ofLos Angeles County.)
D. Same specimen as in C, dorsal view. Scale bar: 10.0 mm.
 Atlas o/ Marine Ln u e r t e b r a t e Larvae

FIGURE 17.II

Megalopae ofThalassinidea and Anomura (A-e) and settlement by

hermit crab megalopae (D-G)
A. Callichirus major (family Callianassidae), dorsal view. Scale bar: 0.5 mm. (Reproduced with
permission from Srrasser and Felder, 1999.)
B. Lepidopa benedicti (family Albuneidae), dorsal view. Scale bar: LO mm. (Reproduced with
permission from Stuck and Truesdale, 1986.)
C. Allopetrolisthes angulosus (family Porcellanidae), dorsal view. Scale bar: 0.5 mm. (Reproduced
with permission from Wehrtmann et al., 1996.)
D. Swimming megalopa of Pagurus brevidactylus (family Paguridae), lateral view. Scale bar: LO
mm. (Phorograph by A.W Harvey.)
E. Megalopa P. brevidactylus walking on the substrate, dorsal view. Megalopae commonly make a
gradual transition from being primarily pelagic ro primarily benthic. Scale bar: LO mm.
(Phorograph by A.W Harvey.)
F. Initial investigation of a gasrropod shell by a megalopa P. brevidactylus. Shells are essential to
post-larval survival in most hermit crabs. Scale bar: 0.5 mm. (Photograph byA.W Harvey.)
G. Megalopa of Coenobita variabilis (family Coenobitidae), occupying a gastropod shell. Scale bar:
LO mm. (Phorograph by A.W. Harvey.)

H. Delayed metamorphosis of megalopae in the absence of required post-larval cues. Two same­
age siblings of Clibanarius longitarsus (family Diogenidae), 6 weeks aíter becoming megalopae.
The large specimen on rhe right received a gastropod shell 5 weeks earlier, metamorphosed a
week later, and is a sixth stage juvenile. The small specimen on the left has not received a shell,
and is still a swimming megalopa. Scale bar: LO mm. (Photograph by A.W Harvey.)
 Atlas o/ Marine I n u e r t e b r a t e Larvae

FIGURE 17.12

Megalopae of brachyuran crabs

A. Dromia wilsoni (Dromiidae), dorsal view, Scale bar: 0.25 mm. (Reproduced with permission
from Wear and Fielder, 1985.)
B. Cancer nouaezelandiae (Cancridae), dorsal view. Scale bar: 0.5 mm. (Reproduced with
permission from Wear and Fielder, 1985.)
e and D. Menippe adina (family Xanthidae), in dorsal (C) and lateral (D) view. Scale bar: 0.5 mm.
(Reproduced wirh permission from Martin et al., 1985.)
E. Paramolapetterdi (family Homolidae), lateral view. Scale bar: 2.0 mm. (Reproduced with
permission from Wear and Fielder, 1985.)
F. Sesarmaguttatum (family Grapsidae.) Scale bar: 0.2 mm. (Reproduced with permission from
Pereyra Lago, 1993.)
G. Pisa sp. (family Majidae.) Scale bar: 0.5 mm. (Photograph by Jocelyne Martin/IFREMER.)
A