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Larval development, metamorphosis and early

growth of the gumboot chiton Cryptochiton
stelleri (Middendorff, 1847) (Polyplacophora:
Mopaliidae) on the Oregon coast

Article · April 2011

DOI: 10.1093/mollus/eyr004

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 LARVAL DEVELOPMENT, METAMORPHOSIS AND EARLY
GROWTH OF THE GUMBOOT CHITON CRYPTOCHITON STELLERI
(MIDDENDORFF, 1847) (POLYPLACOPHORA: MOPALIIDAE)
ON THE OREGON COAST
JOSHUA P. LORD
Oregon Institute of Marine Biology, PO Box 5389, Charleston, OR 97420, USA

Correspondence: J.P. Lord; e-mail: lord@uoregon.edu or jlordjosh@gmail.com

(Received 18 August 2010; accepted 7 January 2011)

ABSTRACT
Cryptochiton stelleri is the largest herbivore in the intertidal and subtidal zone throughout its North
Pacific range, but its larval development and metamorphosis have not been well documented. A
description of larval development for specimens in Hokkaido, Japan, has been used in multiple text-
books yet shows many features atypical of chiton development. In the present study in Oregon, C.
stelleri larvae were raised in culture and displayed developmental stages similar to other chitons, very
different from the previous description. Plate development began 3 days after hatching. Larvae were
competent beginning 3 days posthatching and metamorphosed in response to extract from encrusting

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coralline algae. Larvae survived for over a month without metamorphosing and did not metamor-
phose in response to increased temperature, presence of adults or the addition of algal foods of adults
or juveniles. Juvenile C. stelleri were discovered in the field and grew c. 4 mm per month in captivity.
Juveniles had exposed shell plates and fed on the red alga Cryptopleura.

INTRODUCTION MATERIAL AND METHODS

Cryptochiton stelleri (Middendorff, 1847) is a common interti-
dal inhabitat on the western coast of North America and, at
up to 36 cm long, is the largest chiton species in the world.
Its range stretches from central California to Alaska and
westward to northern Japan and its habitat is from the mid-
intertidal zone down to a depth of 60 m. Despite the abun-
dance of this species, small individuals (smaller than 15 cm
long) are rarely found (Palmer & Frank, 1974; Yates, 1989)
and juveniles (Heath, 1897) and larvae (Okuda, 1947) have
each been described in only a single publication (besides
brief mentions by Eernisse, 2004, and Vendrasco et al.,
2008). The life cycle of C. stelleri is therefore largely
unknown. Larval development and metamorphosis have,
however, been described for several species of chitons, includ-
ing descriptions by Cowden (1961), Thorpe (1962), Barnes &
Gonor (1973), Watanabe & Cox (1975), Kniprath (1980),
Rumrill & Cameron (1983) and Voronozhskaya, Tyurin &
Nezlin (2002).
The timing of spawning in C. stelleri has been described in
several papers, but these reports differ, even at the same
location (Heath, 1905; Okuda, 1947; Tucker & Giese, 1962;
Lawrence & Lawrence, 1965; Palmer & Frank, 1974; Yates,
1989). During spawning, release of eggs by females triggers the
release of sperm by males (Tucker & Giese, 1962). Eggs are
green and are released in a loosely connected gelatinous mass
(Yates, 1989). Sperm are released freely and are activated
(become mobile) when they contact seawater.
The only description of C. stelleri development, by Okuda
(1947), differs from other literature reports with regard to egg
colour and spawning behaviour in this species, as discussed
later. No photographs have been published of the development
of C. stelleri, and the only illustrations are drawings by Okuda
(1947) that contain some inconsistencies.
The present study seeks to describe the spawning, larval devel-
opment, metamorphosis and juvenile growth rate of C. stelleri.
Adult Cryptochiton stelleri were observed spawning naturally
(without artificial cues) in the spring of both 2009 and 2010.
Since different experiments were done each year, the methods
are described separately.
Larval cultures in 2009
Adults were collected from the rocky intertidal zone at South
Cove, Cape Arago, Oregon (43818.1910 N, 124823.1980 W), on
April 11, 2009. The low tide on that day was 20.21 m
(actual) relative to MLLW, the sky was overcast, air tempera-
ture was 88C and sea surface temperature 10.18C. While con-
ducting intertidal surveys, four chitons were observed with
released eggs near the paired gonopores in the pallial groove.
They were discovered c. 30 min before an 8:16 a.m. low tide,
after a night with a full moon. These individuals were brought
to the laboratory and eggs were isolated in culture dishes filled
with 45-mm filtered seawater (not autoclaved). After a subset
of the eggs was photographed under a compound light micro-
scope, they were fertilized with sperm from a male that had
released sperm in a running-seawater table that morning.
Some were left unfertilized and were observed in order to
determine that eggs had not already been fertilized. Cultures
were changed via reverse filtration with 45-mm Nytex mesh
and then refilled with 0.45-mm filtered seawater every day for
the first week and then every other day thereafter.
Cultures of C. stelleri were maintained at 118C (+18C),
roughly the ambient Oregon seawater temperature, and kept
in natural light. Photographs were taken regularly with a Sony
3CCD ExwaveHADw microscope camera on a DIC compound
microscope and observations were made throughout
development.
In an attempt to induce settlement and metamorphosis,
beginning 2 weeks after fertilization several potential cues were
introduced into the culture dishes. One centimetre square

Journal of Molluscan Studies (2011) 77: 182–188. Advance Access Publication: 22 March 2011 doi:10.1093/mollus/eyr004
# The Author 2011. Published by Oxford University Press on behalf of The Malacological Society of London, all rights reserved
 LARVAL DEVELOPMENT OF CRYPTOCHITON STELLERI

Table 1. List of potential metamorphosis cues tested on larvae of
Cryptochiton stelleri.
Treatment ( potential cue)
1 cm2 pieces of alga
Mazzaella splendens
1 cm2 pieces of alga Ulva
lactuca
Rocks covered in
encrusting corallines
Lithothamnion and
Clathromorphum
Shavings of encrusting
corallines Lithothamnion
and Clathromorphum
Increased phytoplankton
concentrations
(Rhodomonas,
culture dishes each of 50 larvae for each treatment. The first
was a control, with nothing added to the filtered seawater.
Treatment 2 had a small rock covered with encrusting coralline
Duration (h) Replication Sources
algae in each dish. Treatment 3 had encrusting coralline algal
(no. of extract added; this was prepared by scraping the coralline
dishes) algae off rocks, grinding it with a mortar and pestle, adding fil-
tered seawater and then centrifuging in 15-ml vials to obtain
48 5 Adult food
the supernatant. Treatment 4 had extract of Cryptopleura (a
(Yates,
delicate leafy red alga eaten by juvenile C. stelleri), prepared as
1989) for coralline algal extract. Larvae were examined under a dis-
48 5 Adult food secting microscope once a day for 1 month to observe
(Yates, metamorphosis.
1989)
48 5 Barnes &
Juveniles
Gonor
(1973) Searches in the intertidal zone for juveniles were performed at
Sunset Bay, Cape Arago and Cape Blanco throughout the
48 5 Barnes &
spring and summer. Juveniles collected at Cape Blanco on July
22, 2009 were raised in a container with mesh sides (20 cm 
Gonor
20 cm by 15 cm high) in a flowing seawater table at the
(1973)
Oregon Institute of Marine Biology. The length and weight of
48 5 the juveniles were measured once a week during March 2010
and time-lapse images were taken every 5 min during August
2009 in order to determine movement patterns.

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Dunaliella, Isochrysis sp.)
Water from tank with adult 48 5 Burke
C. stelleri (1984)
Temperature increased 48 5 Boettcher
to 258C (2005)
Treatments were done consecutively on the same culture dishes and are
listed in the order in which they were tested.
pieces of the green alga Ulva and red alga Mazzaella, the food
of adult C. stelleri, were placed in culture dishes with the
larvae. Since the chemical g-aminobutyric acid from coralline
algae has been shown to induce settlement in the chitons
Katharina tunicata (Rumrill & Cameron, 1983), both shavings of
and rocks covered with encrusting coralline algae
(Lithothamnion and Clathromorphum) were added. The phyto-
plankton species that are commonly used to feed larvae,
Rhodomonas lens (division Cryptophyta), Dunaliella tertiolecta
(division Chlorophyta) and Isochrysis galbana (division
Haptophyta), were also added from laboratory cultures to
larval culture dishes as potential cues for metamorphosis. This
was done in case the spring phytoplankton bloom that occurs
off the Oregon coast served as a cue.
In order to determine if the presence of adult C. stelleri was a
cue for metamorphosis, adults were kept in 3 l filtered seawater
for 1 h after which 5 ml of this water was added to larval culture
dishes. As another potential cue, the temperature of the cultures
was raised to 258C. All of these cues were added sequentially to
five 11.4-cm wide glass culture dishes, with each dish containing
c. 15 larvae. After 2 days, the water was changed and another
potential cue was tested in the order listed in Table 1.
Larval cultures in 2010
Male and female C. stelleri collected at South Cove, Cape
Arago and kept in flowing seawater tanks at the Oregon
Institute of Marine Biology spawned on May 6, 2010 and con-
tinued to release gametes intermittently for 2 days. Eggs were
fertilized with sperm from males that also spawned in the lab-
oratory. Larvae were again raised in 11.4-cm culture dishes in
flowing seawater at c. 128C and were kept in 45-mm filtered
seawater. After the larvae hatched out of the egg hulls (about
2 days postfertilization), four treatments were set up, with four
RESULTS
Eggs and spawning behaviour
The four female Cryptochiton stelleri from which eggs were col-
lected in April 2009 were not spawning at the time of capture,
so egg release was not observed. Some females raised their pos-
terior ends while spawning in May 2010, while others lay
entirely upside down to release or showed no visible difference
in behaviour while spawning. The green eggs were stuck in the
mucous covering the gills, towards the posterior end of the
pallial groove. Both the eggs and sperm sank in seawater.
Unfertilized ova of C. stelleri averaged 301 mm in diameter
(SD ¼ 3.16, n ¼ 15) with a hull diameter of c. 600 mm (SD ¼
18.1, n ¼ 15). The eggs were yolky, indicated by their opaque
appearance, but the hull obstructed a clear view of the
embryos.
Larval cultures in 2009
Fertilization was observed shortly after gametes were mixed
together in culture (Fig. 1A). The first holoblastic (total div-
ision) cleavage occurred 4 h 40 min postfertilization, and sub-
sequent divisions occurred approximately every 40 min
(Table 2). By 27 h postfertilization, four distinct patches of
cells had developed cilia. The eggs remained on the bottom of
the dishes until hatching. At 46 h the early trochophore
hatched and was characterized by an apical tuft and proto-
troch, which propelled the bright green larvae rapidly around
the culture dish (Fig. 2). A large amount of yolk remained and
there was no indication of plate formation or any bumps that
could be precursors of shell plates (Fig. 2).
Over the next 2 days the post-trochal region elongated, a
pair of red larval eyes appeared behind the prototroch and
dorso-ventral flattening occurred as the foot developed
(Fig. 3A). At about 5 days after fertilization (3 days posthatch-
ing), the larvae developed seven shell plate rudiments and
settled to the bottom of the culture dishes where they began
creeping (Fig. 3B). In controls with just filtered seawater,
development went no further; loss of the prototroch and plate
formation did not occur, suggesting that the larvae were await-
ing some cue to undergo metamorphosis. The addition of
different kinds of seaweeds, high phytoplankton concentrations,

183
 J. P. LORD

Figure 2. Two-day-old trochophore larva of Cryptochiton stelleri just after

hatching. Note the lack of dorsal plates or bumps that are the
precursors to plate formation. There is no elongation of the post-trochal
region, and the eyes and foot have yet to develop. Scale bar ¼ 100 mm.

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Figure 3. Late trochophores of Cryptochiton stelleri. Dorsal side is facing
the upper right corner of photographs. A. Elongation of post-trochal
region is visible as well as some dorso-ventral flattening; foot and seven
plate precursors are evident underneath body surface; yolk is still
visible. B. The seven bumps that are plate precursors are visible on
outer surface of larval body; as in (A) eyes and prototroch are present.
Figure 1. Early development of Cryptochiton stelleri. A. Fertilized egg Scale bars ¼ 100 mm.
with surrounding hull. B. 16-cell stage with spiral cleavage evident.
Scale bar ¼ 100 mm.
water that had contained conspecific adults and increased
water temperatures all failed to induce metamorphosis. The
larvae in these dishes and in controls died in culture c. 8 weeks
Table 2. Developmental timetable for Cryptochiton stelleri. after hatching, having failed to metamorphose.
Time Stage
postfertilization
Larval cultures in 2010
0 Gametes mixed The embryos cultured in May 2010 developed on a similar
,1 min Fertilization envelope forms timeline, beginning to hatch 2 days after fertilization. The tro-
4 h, 40 min First cleavage: 2-cell stage chophores exposed to coralline algal extract and coralline
5 h, 20 min Second cleavage: 4-cell stage algae-covered rock began metamorphosing 3 days after hatch-
5 h, 55 min Third cleavage: 8-cell stage ing (5 days postfertilization). Larvae were competent to meta-
6 h, 40 min Fourth cleavage: 16-cell stage morphose after the post-trochal region had elongated and
11 h, 10 min Fifth cleavage: 32-cell stage
seven shell-plate rudiments had formed, when the larvae spent
most of their time on the bottom of the dishes (Fig. 4). In the
17 h, 10 min Blastula
control and Cryptopleura-extract treatments, larvae stayed in
26 h, 55 min Ciliated cells form, 4 patches of cilia
this stage for over 2 months and metamorphosis could still
46 h Hatching from hull occur at this time. Metamorphosis involved the loss of the pro-
70 h Prototroch formed from ciliary patches totroch and apical tuft and the beginning of the formation of
90 h Elongation of post-trochal region, eyes form seven shell plates. The plates began as thin lines (Fig. 5A) and
118 h Settle to bottom, shell-plate precursors appear on slowly expanded until they covered the body of the juvenile.
dorsal side Postmetamorphosis, the larvae were observed feeding on
5 days Metamorphosis; loss of prototroch and apical tuft; diatoms and cyanobacteria that were present on the bottom of
beginning of shell-plate calcification the culture dishes. Faecal pellets were discovered, some com-
posed entirely of cyanobacteria and others containing largely
Eggs were collected from the South Cove of Cape Arago and gametes were diatoms with silica skeletons that were not broken down by the
mixed on April 11, 2009. addition of bleach. Grazing trails were formed on the bottoms

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Figure 4. Contrasting descriptions of Cryptochiton stelleri development. A. Development in Hokkaido, Japan (modified from Okuda, 1947): just after
hatching (a); just before settlement (b); recently settled (c). B. Development observed in Oregon in this study: just after hatching (no plate
formation or post-trochal elongation) (a); after settlement (shell plate rudiments are plate precursors as no calcification has occurred) (b). Scale
bars ¼ 100 mm.

of the dishes but did not show any pattern and repeatedly
crossed over other trails. Shell plates continued to develop and
widen as juveniles grew (Fig. 6A).
Juveniles
Four juvenile C. stelleri were discovered on the red alga
Cryptopleura at Cape Blanco on July 22, 2009. They measured
7.6– 12.6 mm long and, unlike the adults, their dorsal plates
were still exposed (Fig. 6B). Juveniles were yellow with tufts
of red spicules partially covering the girdle, giving them an
orange appearance. The mouth, foot and other anatomical
features appeared to be fully developed and the juveniles fed
on Cryptopleura. Feeding was captured on time-lapse video
and the red alga was visible in the gut through the juvenile
foot.
By time-lapse photography juvenile C. stelleri were found to
move more at night (3.60 mm/h) than during the day
(1.9 mm/h), but this difference was not significant (ANOVA,
df ¼ 4, F ¼ 2.42, P ¼ 0.13). In January 2010, 6 months after
the juveniles were captured and estimated 9 months after they
metamorphosed, they began to feed on Ulva lactuca as well as
Cryptopleura sp. on which they had been feeding since capture.
By 1 year juveniles could feed on Mazzaella splendens, a
common adult food source. After July 2009, juveniles grew an
average of 1.96 mm or 0.14 g per month and were c. 30 mm
long by March 2010 (Fig. 7). This growth rate appears to be
similar to that in the field; three additional juveniles collected
in the field were in the same size range as those kept in the lab-
oratory; one was found on November 11, 2009, at Middle
Cove of Cape Arago (13 mm), another on December 5, 2009,
at Cape Blanco (15 mm), and the third on February 3, 2010,
at Cape Blanco (19 mm).
DISCUSSION
The short larval period, developmental timetable and non-
feeding larvae of Cryptochiton stelleri described in this study are
similar to those of most other described species of non-brooding
chitons (Pearse, 1979; Rumrill & Cameron, 1983; Shanks,
2001). The trochophore larva looks similar to that of
other chitons, but is slightly larger, 300 mm at hatching
(Fig. 2). The appearance of only seven plates is consistent with
other chiton species, since many chitons do not develop the
eighth plate until well after metamorphosis (Voronozhskaya
et al., 2002).
Metamorphosis was triggered solely by the addition of coral-
line algal extract, which has also been shown to induce species
such as Tonicella lineata (Barnes & Gonor, 1973), Katharina
tunicata (Rumrill & Cameron, 1983), Haliotis diversicolor (Bryan

185
 J. P. LORD

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Figure 5. Photographs of newly metamorphosed juveniles of Cryptochiton stelleri. A. Dorsal view of 1-day post-metamorphosic juvenile with valves
beginning to form. B. Ventral view of 1-day post-metamorphosic juvenile with two red eyes and foot visible. C. Dorsal view of juvenile 5 days after
metamorphosis; valves have developed further. D. Dorsal view of juvenile 14 days after metamorphosis with valves completely covering mantle and
foot. Scale bar ¼ 100 mm.

Figure 7. Growth rate of juvenile Cryptochiton stelleri raised in flowing

seawater tables (n ¼ 4, with SE). Growth rate is in terms of wet
weight, measured to 0.01 g.

or could have evolved convergently. Why C. stelleri uses this

Figure 6. Photographs of juvenile Cryptochiton stelleri. A. Valves of
4-week-old juvenile. Scale bar ¼ 100 mm. B. Four-month-old juvenile
discovered at Cape Blanco, Oregon. Eight exposed plates are visible,
not yet overgrown by mantle, and red spicules are starting to cover
surface of mantle. Scale bar ¼ 1 mm.
& Qian, 1998) and other species of molluscs to metamorphose
(Hayakawa et al., 2008; Steller & Caceres-Martinez, 2009;
Roberts, Barker & Mladenov, 2010). This feature of a diverse
array of molluscs could be a highly conserved ancestral trait,
cue is unclear, because encrusting coralline algae are present in
many locations with high wave action in which C. stelleri would
be unable to survive and because at no stage of its life cycle
does C. stelleri feed on coralline algae. It is possible that
recruitment occurs mainly within highly localized populations;
C. stelleri is most abundant in small coves (personal obser-
vation) and a ubiquitous cue for metamorphosis may be
suitable if larvae generally remain in these protected habitats.
It is also possible that a coralline algal cue ensures that meta-
morphosis occurs at a low tidal level, where encrusting coral-
line algae are found. Since newly metamorphosed juveniles fed
on diatoms and bluegreen algae, it is possible that the response
to encrusting coralline algae because of the biofilm commonly
found among coralline algae. However, the coralline algae
were cleaned thoroughly before the experiments, so the biofilm

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 LARVAL DEVELOPMENT OF CRYPTOCHITON STELLERI
itself could not be the cue. Since older juveniles feed on the
leafy red alga Cryptopleura and this did not induce metamor-
phosis, the cue may not be directly related to diet.
The developmental stages of C. stelleri were compared with
those described by Okuda (1947) in the only other study on
the development of this species. There are several differences
between the observations of Okuda and those reported here.
He described eggs as red or cinnamon-coloured (not green)
and in an egg mass (not released freely). His illustrations
show a thin hull, unlike the wide hull found here (Fig. 1).
Okuda stated that plate formation and post-trochal elongation
occurred before hatching (Fig. 4A). In the present study post-
trochal elongation commenced 2 days after hatching and
plate precursors appeared 3 days posthatching (Figs 2, 3).
Plates did not begin to calcify until metamorphosis, which
could occur as soon as 3 days posthatching. Okuda’s (1947)
description of C. stelleri eggs with a small hull, laid in an egg
mass, and of development of shell rudiments before hatching
and without a metamorphosis cue are all characteristics of
brooding chiton species or those that lay benthic egg masses
(Eernisse, 1988; Buckland-Nicks, 1993). These are inconsistent
with existing descriptions of C. stelleri from North America
(Tucker & Giese, 1962; Pearse, 1979) and with the present
observations. Furthermore, brooding chitons are usually small
as adults (Strathmann & Strathmann, 1982; Creese, 1986;
Eernisse, 1988) and brooding is associated with small adult
size in organisms such as sea stars, bivalves, calyptraeid gas-
tropods and sabelliform polychaetes (Strathmann, 1985).
Therefore, it is unlikely that Japanese members of the largest
chiton species in the world would brood, and instead they
may develop in benthic egg masses and then crawl away. The
existence of two such contrasting modes of development
within a single molluscan species is extremely rare (e.g.
Ellingson & Krug, 2006). It is therefore possible that
Cryptochiton ‘stelleri’ in Japan and North America are distinct
species, and there is some molecular evidence for this (D.
Eernisse, personal communication). Since the type locality of
C. stelleri is the Kamchatka Peninsula, Russia (Middendorff,
1847), the name should be applied to the northwestern
Pacific form.
ACKNOWLEDGEMENTS
This manuscript was based upon work supported by NSF GK-
12 grant number 0638731. I thank Dr Richard Emlet and Dr
Craig Young for advice on larval culture. I also thank Dr Alan
Shanks and Dr Cynthia Trowbridge for assistance with the
manuscript.
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