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Collection Highlights

Oceanography and Marine Biology An Annual Review Volume 57

1st Edition S. J. Hawkins (Editor)

Marine Biology: - Function, Biodiversity, Ecology 1st

Edition Jeffrey S. Levintons

International Review of Cell and Molecular Biology First

Edition Kwang W. Jeon

Developmental Biology of the Sea Urchin and Other Marine

Invertebrates Methods and Protocol 2nd Edition David J.
Carroll
Biology and Ecology of Edible Marine Gastropod Molluscs
1st Edition Ramasamy Santhanam

Biology and Ecology of Pharmaceutical Marine Tunicates 1st

Edition Ramasamy Santhanam (Author)

Marine Organisms as Model Systems in Biology and Medicine

Malgorzata Kloc

Investigating Oceanography Raphael Kudela

Advances in Marine Biology 74 1st Edition Barbara E. Curry

(Eds.)
OCEANOGRAPHY
and
MARINE BIOLOGY
AN ANNUAL REVIEW
Volume 55
OCEANOGRAPHY
and
MARINE BIOLOGY
AN ANNUAL REVIEW
Volume 55

Editors
S. J. Hawkins
Ocean and Earth Science, University of Southampton,
National Oceanography Centre, UK
and
The Marine Biological Association of the UK, The Laboratory, Plymouth, UK

A. J. Evans
Ocean and Earth Science, University of Southampton,
National Oceanography Centre, UK
and
The Marine Biological Association of the UK, The Laboratory, Plymouth, UK

A. C. Dale
Scottish Association for Marine Science, Argyll, UK

L. B. Firth
School of Biological and Marine Sciences, Plymouth University, UK

D. J. Hughes
Scottish Association for Marine Science, Argyll, UK

I. P. Smith
School of Biological Sciences, University of Aberdeen, United Kingdom
International Standard Serial Number: 0078-3218

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Contents
Preface vii

Introduction to the Special Issue ix

Ally J. Evans, Louise B. Firth, Andrew Dale, David Hughes, I. Philip Smith
& Stephen J. Hawkins

Filling in the Grazing Puzzle: A Synthesis of Herbivory in Starfish 1

Aline S. Martinez, Maria Byrne & Ross A. Coleman

Intertidal Boulder-Fields: A Much Neglected, but Ecologically Important, Intertidal Habitat 35

M.G. Chapman

Ecological Dominance along Rocky Shores, with a Focus on Intertidal Ascidians 55

Marc Rius, Peter R. Teske, Patricio H. Manríquez, Rocío Suárez-Jiménez, Christopher D.
McQuaid & Juan Carlos Castilla

Giant Clams (Bivalvia: Cardiidae: Tridacninae): A Comprehensive Update of Species

and Their Distribution, Current Threats and Conservation Status 87
Mei Lin Neo, Colette C.C. Wabnitz, Richard D. Braley, Gerald A. Heslinga, Cécile Fauvelot,
Simon Van Wynsberge, Serge Andréfouët, Charles Waters, Aileen Shau-Hwai Tan, Edgardo
D. Gomez, Mark J. Costello & Peter A. Todd

How Anthropogenic Activities Affect the Establishment and Spread of Non-Indigenous

Species Post-Arrival 389
Emma L. Johnston, Katherine A. Dafforn, Graeme F. Clark, Marc Rius & Oliver Floerl

A Review of Herbivore Effects on Seaweed Invasions 421

Swantje Enge, Josefin Sagerman, Sofia A. Wikström & Henrik Pavia

Author Index 441

Systematic Index 471

Subject Index 475

v
Preface
The 55th volume of Oceanography and Marine Biology: An Annual Review (OMBAR) contains
six reviews that cover a range of topics, reflecting the wide readership of the series. A brief intro-
ductory comment outlines the inspiration for review articles in this special issue that stemmed
from contributions to and discussions at the 2015 Aquatic Biodiversity & Ecosystems conference
in Liverpool, UK.
OMBAR welcomes suggestions from potential authors for topics that could form the basis of
appropriate reviews. Contributions from physical, chemical and biological oceanographers that
seek to inform both oceanographers and marine biologists are especially welcome. Because the
annual publication schedule constrains the timetable for submission, evaluation and acceptance of
manuscripts, potential contributors are advised to contact the editors at an early stage of manuscript
preparation. Contact details are listed on the title page of this volume.
The editors gratefully acknowledge the willingness and speed with which authors complied
with the editors’ suggestions and requests and the efficiency of CRC Press, especially Jennifer
Blaise, Marsha Hecht and John Sulzycki, in ensuring the timely appearance of this volume.

vii
 Introduction to the Special Issue
ALLY J. EVANS1,2, LOUISE B. FIRTH3, ANDREW DALE4,
DAVID HUGHES4, I. PHILIP SMITH5 & STEPHEN J. HAWKINS1,2

1Ocean and Earth Science, National Oceanography Centre

Southampton, Southampton, SO14 3ZH, UK
2The Marine Biological Association of the UK, The Laboratory,

Citadel Hill, Plymouth, PL1 2PB, UK

3School of Biological and Marine Sciences, Plymouth University, Plymouth, PL4 8AA, UK
4Scottish Association for Marine Science, Scottish Marine

Institute, Oban, Argyll, Scotland, PA37 1QA, UK

5School of Biological Sciences, University of Aberdeen, Aberdeen, Scotland, AB24 2TZ, UK

In this 55th volume of Oceanography and Marine Biology: An Annual Review (OMBAR), we pres-
ent a special issue stemming directly and indirectly from contributions and discussions at the 2015
Aquatic Biodiversity & Ecosystems Conference in Liverpool, UK, plus related work from research
teams represented at that conference.
The Aquatic Biodiversity & Ecosystems Conference was organised by Dr Louise Firth at the
University of Liverpool in August–September 2015. The overarching theme was “evolution, inter-
actions and global change in aquatic ecosystems”, with a number of subthemes, within which over
250 papers were presented to delegates from 30 different countries. The meeting partially followed
up on the 1990 Plant–Animal Interactions in the Marine Benthos Conference hosted by Professor
Stephen Hawkins 25 years previously, which itself was prompted by a 1983 OMBAR review on
grazing of intertidal algae by marine invertebrates (Hawkins & Hartnoll 1983).
Through structured workshops and informal discussions at the 2015 conference, suggestions
were garnered for review articles for this special OMBAR issue, and for chapters in the forthcom-
ing book Interactions in the Marine Benthos—A Regional and Habitat Perspective, a follow-up
to Plant–Animal Interactions in the Marine Benthos (John et al. 1992) (which was inspired by the
1990 conference), to be published by Cambridge University Press. The neglected fields of herbivory
in starfish (Martinez et al. 2017) and intertidal boulder-fields (Chapman 2017) are synthesized to
stimulate further interest in these topics. Ecological dominance along rocky shores is discussed,
focussing on ascidians (Rius et al. 2017). Neo et al. (2017) review the ecology of giant clams.
Johnston et al. (2017) consider the spread of non-indigenous species. Enge et al. (2017) take this
theme further by discussing defences against herbivory of invasive seaweeds. The themes of the
conference— evolution, interactions and global change—are all explored in these contributions.
We hope that this special issue will be of value to oceanographers and marine biologists alike,
and will inspire further research where knowledge gaps have been identified. We look forward to
working with new contributors and welcome suggestions for reviews for future volumes. We would
especially value contributions from oceanographers for coming issues.
We sadly heard of the passing away of Professor Roger Hughes at the conference in August
2015. As the then Editor in Chief of OMBAR, this special issue was planned with Roger. We dedi-
cate this volume to his memory as a superb scientist, excellent editor, fine and fanatical fisherman
and great guitarist.

ix
 INTRODUCTION TO THE SPECIAL ISSUE

References
Chapman, M.G. 2017. Intertidal boulder-fields: a much neglected, but ecologically important, intertidal habi-
tat. Oceanography and Marine Biology: An Annual Review 55, 35–53.
Enge, S., Sagerman, J., Wikström, S.A. & Pavia, H. 2017. A review of herbivore effects on seaweed invasions.
Oceanography and Marine Biology: An Annual Review 55, 421–440.
Hawkins, S.J. & Hartnoll, R.G. 1983. Grazing of intertidal algae by marine invertebrates. Oceanography and
Marine Biology: An Annual Review 21, 195–282.
John, D.M., Hawkins, S.J. & Price, J.H. 1992. Plant-Animal Interactions in the Marine Benthos. Oxford:
Clarendon Press on behalf of the Systematics Association.
Johnston, E.L., Dafforn, K.A., Clark, G.F., Rius, M. & Floerl, O. 2017. How anthropogenic activities affect the
establishment and spread of non-indigenous species post-arrival. Oceanography and Marine Biology:
An Annual Review 55, 389–419.
Martinez, A.S., Byrne, M. & Coleman, R.A. 2017. Filling in the grazing puzzle: a synthesis of herbivory in
starfish. Oceanography and Marine Biology: An Annual Review 55, 1–34.
Neo, M.L., Wabnitz, C.C.C., Braley, R.D., Heslinga, G.A., Fauvelot, C., Van Wynsberge, S., Andréfouët,
S., Waters, C., Tan, A.S.-H., Gomez, E.D., Costello, M.J. & Todd, P.A. 2017. Giant clams (Bivalvia:
Cardiidae: Tridacninae): a comprehensive update of species and their distribution, current threats and
conservation status. Oceanography and Marine Biology: An Annual Review 55, 87–390.
Rius, M., Teske, P.R., Manríquez, P.H., Suárez-Jiménez, R., McQuaid, C.D. & Castilla, J.C. 2017. Ecological
dominance along rocky shores, with a focus on intertidal ascidians. Oceanography and Marine Biology:
An Annual Review 55, 55–84.

x
 Oceanography and Marine Biology: An Annual Review, 2017, 55, 1-34
© S. J. Hawkins, D. J. Hughes, I. P. Smith, A. C. Dale, L. B. Firth, and A. J. Evans, Editors
Taylor & Francis

FILLING IN THE GRAZING PUZZLE: A SYNTHESIS

OF HERBIVORY IN STARFISH

ALINE S. MARTINEZ1, MARIA BYRNE1 & ROSS A. COLEMAN1*

1Coastal and Marine Ecosystems Group, Marine Ecology Laboratories (A11),

School of Life and Environmental Sciences, The University of Sydney
New South Wales 2006, Australia
*Corresponding author: Ross A. Coleman
e-mail: [email protected]

Herbivory is an important ecological process controlling community structure and function in

almost all ecosystems. The effects of herbivores on algal assemblages depend primarily on con-
sumer and algal traits, but the strength of this interaction is contingent on physical and biologi-
cal processes. Marine herbivory is particularly intense, where grazers can remove around 70% of
primary production. Present understanding of marine herbivory is largely based on well-studied
groups including herbivorous fishes, gastropods, crustaceans and sea urchins. Herbivory in other
marine taxa is poorly understood, but nonetheless important. For instance, grazing by starfish has
the potential to strongly affect algal assemblages. Most starfish feed by extruding their stomach
and digesting their food externally. This feeding mechanism is distinctive and complex, and evolu-
tionarily advantageous as it allows individuals to explore many different food sources. Variation in
the feeding habits of herbivorous starfish is intriguing because some species are very specialized
whereas others are more generalist, and the reasons for those variations are not well understood.
Some herbivorous starfish are obligate herbivores while others vary from herbivory to carnivory
between life stages or between populations within the same species. The question that then arises is
how well we are able to predict grazing pressure from complex feeding habits on benthic systems?
This review provides a synthesis of herbivory in starfish showing that: the majority of species forage
on microalgae and soft tissue macroalgae; fidelity to an algal diet appears to be related to the size of
individuals; and, feeding habits are likely to change with variation in food availability. Directions
for future studies on the biology and ecology of herbivorous starfish are suggested to better under-
stand variation in species feeding behaviour. Elucidating the mechanisms that contribute to varia-
tion in the behaviour of herbivorous starfish is crucial to predict the effects that these species exert
on the structure of marine benthic communities. The influence of omnivorous species also warrants
more detailed study. Such investigations are important in the context of climate change, given the
potential for species invasions associated with range expansions.

Introduction
Herbivores play a key role in the world’s ecosystems by consuming primary production and altering
habitat structure. Mammalian herbivory, including inter alia grazing and browsing, is well known
to directly affect the distribution of plants in terrestrial systems (Hempson et al. 2015), whereas the
most important herbivores in structuring benthic communities in freshwater aquatic systems are

1
 ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN

snails, insects, crustaceans and small vertebrates (Huntly 1991, Steinman 1996). In marine ecosys-
tems, fish, molluscs, small crustaceans and sea urchins have been shown to dramatically affect the
structure of marine benthic communities (Hawkins & Hartnoll 1983, Hawkins et al. 1992, Coleman
et al. 2006, Poore et al. 2012, Poore et al. 2014). Thus far, the strongest known impact on a marine
habitat, in terms of change in marine algal/macrophyte canopy in the absence of a herbivore, is for
molluscs in rocky intertidal habitats (Poore et al. 2012).
The effects of herbivores on marine plants or microalgal assemblages depend primarily on the
feeding apparatus of the consumer and the interactions of these with the susceptibility of the pri-
mary producer to grazing (Hawkins et al. 1992). The feeding mechanism determines what resources
herbivores can exploit (Steneck & Watling 1982, Hawkins et al. 1989, Kennish & Williams 1997),
whereas food traits (e.g. detectability, toughness, palatability and digestibility etc.) will affect the
intake or intensity of consumption (Lubchenco & Cubit 1980, Lubchenco & Gaines 1981, Duffy &
Hay 1990, Norton et al. 1990, Paul et al. 2001, Dolecal & Long 2013). Variation in grazing pres-
sure is also associated with the size and mobility of herbivores. Small and/or sedentary herbivores
(e.g. amphipods, limpets) are likely to have local impacts on the assemblage while bigger and/or
more mobile species (e.g. fish, sea urchins) require a broader foraging range and can affect assem-
blages at larger scales (Lawrence 1975, Hay 1984, Norton et al. 1990, Duffy & Hay 1991, Vergés
et al. 2009). Apart from size and mobility of herbivores, abundance and composition of herbivores
will together play an important role in the magnitude of the effects of grazing, thereby affecting
the spatial heterogeneity and diversity of algal assemblages (Lubchenco & Gaines 1981, Benedetti-
Cecchi et al. 2005, Jenkins et al. 2008, Griffin et al. 2010).
Although there is a vast literature on the ecology of plant-animal interactions, knowledge from
marine systems is concentrated on a few groups of herbivores, i.e. fish, sea urchins, crabs, amphi-
pods and gastropod grazers (Poore et al. 2012). There are a few gaps that need to be addressed in
order to clarify grazing pressure by different herbivores. This is the case for starfish species, which
are abundant and distributed worldwide. Research on the feeding behaviour of starfish has largely
focused on predatory species (e.g. Brun 1972, Paine 1974, 1976, Town 1980, Keesing & Lucas
1992, Keesing 1995, Himmelman et al. 2005, Scheibling & Lauzon-Guay 2007, Estes et al. 2011,
Mueller et al. 2011, Menge & Sanford 2013). This is because numerous large and non-cryptic star-
fish species are carnivores and play an important role in controlling densities of prey (Chesher 1969,
Porter 1972, Paine 1974, Gaymer & Himmelman 2008, Kenyon & Aeby 2009, Pratchett et al. 2009,
Uthicke et al. 2009; Kayal et al. 2012, Baird et al. 2013, Pratchett et al. 2014). Several species of star-
fish have been reported to also feed on other resources such as detritus, dead animals, and macro-
and microalgae (for a review, see Sloan 1980, Jangoux 1982b). These species have been considered
to be omnivores, however their diet is poorly known because of the difficulty in identification of gut
contents, especially because of their peculiar extra-oral feeding mechanism.
Most starfish feed by everting their cardiac stomach and directly secreting enzymes from the
epithelium onto the food item and thereby digest their food externally. This feeding mechanism is
rare among other animals and allows starfish to utilize many different food types. Despite using the
same extra-oral feeding mechanism, diet varies among starfish species, ranging from carnivores
feeding on specific prey items to omnivorous feeders and herbivores (for a review, see Sloan 1980,
Jangoux 1982b).
The variation in feeding biology among starfish species is related to digestive tract anatomy and
tube foot morphology (Jangoux 1982a). There is, however, poor knowledge of the causes of varia-
tion in feeding habits among starfish species and populations. This is a particular concern for our
understanding of those starfish recorded as feeding on plant or microphyte resources because of the
lack of information on their ecological role as herbivores. Understanding the mechanisms involved
with a herbivorous diet in starfish is important to predict the potential impacts of grazing by these
animals on the structure of benthic assemblages. Therefore the aim of this review is to identify the
main findings and the fundamental gaps in knowledge on the ecology of herbivorous starfish, using

2
 FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH

a systematic review of the literature. We compile information of starfish species that forage on any
source of plant or microphytes and present a synthesis on the different aspects of the ecology of
herbivory in starfish. We describe the food source exploited by herbivorous starfish, the changes
in time and space on the use of food resource, and discuss the possible mechanisms driving these
variations. We also highlight the potential function of herbivorous starfish in controlling the distri-
bution of algal assemblages and the limited studies on interactions between herbivorous starfish and
other herbivores. We expect that this synthesis will encourage researchers to investigate in detail the
functional ecology of herbivorous starfish in marine benthic ecosystems.

Data collection and definition of terminology

Literature on starfish that forage on algae and/or marine macrophytes was compiled by searching
the ISI Web of Science database (Jan 1900–Jan 2015) using the following criteria: TOPIC: (starfish
or “sea star” or seastar or asteroidea) AND TOPIC: (herb* or graz*) AND TOPIC: (macrophyte* or
alga* or seagrass* or eelgrass* or seaweed*). We selected every article that cited or described algal
or vascular plant material in the feeding or diet of starfish species. The compilation was supple-
mented by subsequently searching material cited within those articles including published journal
articles and unpublished theses. We also gathered information from published reviews on star-
fish diet (Sloan 1980, Jangoux 1982b) and searched the references cited therein. The information
extracted from the literature included species name, food resource exploited, foraging behaviour,
distribution (habitat and region) and size of starfish. Any detailed or more relevant information on
feeding habits of a starfish was noted where appropriate. Starfish scientific names are presented
as the current accepted species name according to the world Asteroidea database (accessed at
http://www.marinespecies.org/asteroidea on Jan 7th, 2015). Previous species names in some of the
references cited in this review can be found in Appendix 1.
Hereafter true macrophytes are referred to as “plants”. Macroalgae and microalgae are used
to distinguish multi- and unicellular algae. We consider that microalgae correspond to any biofilm
assemblage (largely prokaryotes and microalgae). Algae are also considered in functional groups
according to the classification of Steneck & Dethier (1994). Finally, it is important to keep in mind
that all data of starfish presented as foraging on macroalgae or plant does not necessarily mean
that the starfish is feeding on the macroalgae or plant per se, but may be consuming associated
epiphytic organisms.

Composition of diet
Identifying the target plant or algal food source exploited by starfish species was used to detect com-
ponents of the benthos that are likely to be affected by asteroid grazing. We found 57 species whose
diet consisted entirely or partially of algae across a variety of habitats and ecosystems (Table 1). The
great majority of starfish (90%) that consume any source of plant/algae are found on hard-bottom
ecosystems, where 30% are exclusive to coral reefs and 32% to rocky shores (mostly temperate). The
remaining 28% of species are found in hard and soft bottom systems, including seagrass beds. Only
10% are exclusive to seagrass and sand/mud sediments. The information on the feeding habits of
starfish within different habitats is often descriptive and lacked accurate evidence of diet especially
for those species that are found on sediments. After compiling information from the literature, how-
ever, a clear pattern of diet emerged for starfish that forage on algal/plant resources.
The most frequently documented algal group exploited by starfish were those that comprise
biofilms. Biofilms usually contain a mix of diatoms, cyanobacteria, bacteria, spores of macroalgae
and other microorganisms embedded in a matrix of extra-polymer substances (Anderson 1995,
Decho 2000). The majority of the species (77%, 44 out of 57 spp.) were reported feeding on epi-
benthic biofilms on different substrata (Table 1). Thus, starfish feeding on biofilm were identified

3
 Table 1 Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References
Obligate herbivores

Asterinidae
Aquilonastra anomala 8 4.4 1.8 Biofilm (substrate feeding on algae and Coral reefs (Yamaguchi 1975)
(H.L. Clark, 1921) microbes)
Asterina gibbosa 16 9.4 1.7 Biofilm (diatoms/bacterial film), Rocky shores Intertidal rock pools, (Crump & Emson 1978, Sloan 1980,
(Pennant, 1777) macroalgae (Ulva) and decaying algae under boulders Jangoux 1982b, Crump & Emson
(Laminaria) 1983)
Asterina phylactica 7 4.4 1.6 Biofilm, epilithic organisms on algae Rocky shores Intertidal rocky pools, (Emson & Crump 1979, Crump &
(Emson & Crump, under boulders Emson 1983)

4
1979)
Cryptasterina hystera 12 7.5 1.6 Biofilm (bare rock) Rocky shores Mid intertidal under (Dartnall 1971; Martinez, A.S.,
(Dartnall & Byrne, small rocks unpublished, Dartnall et al. 2003)
2003)
Cryptasterina pacifica 12 9.2 1.3 Biofilm (bare rock) Coral reefs Intertidal rock pools, (Dartnall 1971, Dartnall et al. 2003)
(Hayashi, 1977) under coral rubble
Cryptasterina 11 5.6 2.0 Biofilm on coral rubble Coral reefs High intertidal rock (Dartnall 1971; Martinez A.S.,
pentagona (Muller pools, under coral unpublished)
& Troschel, 1842) rubble
Parvulastra exigua 10 6.7 1.5 Biofilm (diatoms, spores, detritus, Rocky shores, Intertidal rock pools, (Duyverman 1976, Branch & Branch
(Lamarck, 1816) bacteria), Macroalgae (Ulva, seagrass crevices, under 1980, Jangoux 1982b, Arrontes &
ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN

Enteromorpha, Ceramium, Corallina beds, soft boulders; seagrass and Underwood 1991, Stevenson 1992,
officinalis, Hormosira, encrusting bottom lagoonal sandflats Jackson et al. 2009, Pillay et al. 2010,
calcareous), detritus, shells. Shepherd 2013, Martinez et al. 2016)
Parvulastra 3 2.7 1.1 Biofilm Rocky shores Mid to low intertidal rock (Keough & Dartnall 1978, Wilson
parvivipara (Keough pools, under boulders 2004)
& Dartnall, 1978) and small rocks
Continued
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References
Parvulastra vivipara 15 12.0 1.3 Biofilm (diatoms, cyanobacteria) Rocky shores Intertidal rock pools and (Jangoux 1982b, Prestedge 1998,
(Dartnall, 1969) under boulders Polanowski 2002)
nomen dubiuma — — — Macroalgae and detritus Rocky shores (Jangoux 1982b)

Echinasteridae
Echinaster luzonicus 40 4.5 8.9 Biofilm (substrate feeding on algae and Coral reefs, Corals, seagrass, mud/ (Yamaguchi 1975, Jangoux 1982b)
(Gray, 1840) microbes) seagrass sand bottom, pebble,
beds, soft shell grits
bottom
Echinaster purpureus 80 8.3 9.6 Biofilm (epibenthic film of the hard Coral reefs, Corals, seagrass, sand (Thomassin 1976, Jangoux 1982b)
(Gray, 1840) substrate of the coral built formations) seagrass
beds, soft
bottom

5
Goniasteridae
Neoferdina cumingi 40 9.5 4.2 Biofilm (substrate feeding on algae and Coral reefs Seaward reef (Yamaguchi 1975, Jangoux 1982b)
(Gray, 1840) microbes)

Ophidiasteridae
Cistina columbiae 40 6.8 5.9 Biofilm (substrate feeding on algae and Coral reefs Coral and rock reefs (Yamaguchi 1975, Jangoux 1982b)
(Gray, 1840) microbes)
Dactylosaster 21 2.1 10.0 Biofilm (substrate feeding on algae and Coral reefs (Yamaguchi 1975, Jangoux 1982b)
cylindricus microbes)
(Lamarck, 1816)
Linckia guildingi 150 15.0 10 Biofilm (substrate feeding on algae and Coral reefs Reef flat and lagoon, (Yamaguchi 1975, Jangoux 1982b)
(Gray, 1840) microbes) seaward reef
Continued
FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References
Linckia laevigata 122 22.0 5.5 Biofilm (epibenthic film of seagrass and Coral reefs Reef flat and less (Laxton 1974, Yamaguchi 1975,
(Linnaeus, 1758) hard/dead coral substratum), common on coarse sand Thomassin 1976, Sloan 1980,
macroalgae (calcareous encrusting) flat, seaward reef Jangoux 1982b)
Linckia multifora 62 7.0 8.9 Biofilm (substrate feeding on algae and Coral reefs (Yamaguchi 1975, Jangoux 1982b)
(Lamarck, 1816) microbes)
Nardoa variolata 150* 26.8 5.6 Biofilm (epibenthic film of sea grass) Coral reefs, Coral reef flats and less (Thomassin 1976, Jangoux 1982b)
(Retzius, 1805) seagrass commonly on coarse
beds sandy flats
Ophidiaster 29 4.5 6.4 Biofilm (substrate feeding on algae and Coral reefs Reef flat and lagoon (Yamaguchi 1975, Jangoux 1982b)

6
cribrarius microbes)
(Lütken, 1871)
Ophidiaster granifer 41 11.4 3.6 Biofilm (substrate feeding on algae and Coral reefs Reef flats, under boulders (Yamaguchi 1975, Jangoux 1982b)
(Lütken, 1871) microbes) and coral rubble
Ophidiaster 120 11.3 10.7 Biofilm (substrate feeding on algae and Coral reefs Reef flat and lagoon, (Yamaguchi 1975, Jangoux 1982b)
hemprichi (Müller & microbes) seaward reef
Troschel, 1842)
Phataria unifascialis 70 10.3 6.8 Biofilm (film of algae and invertebrates Rocky shores Intertidal and subtidal (Morgan & Cowles 1997)
(Gray, 1840) on rock), macroalgae rocks

Oreasteridae
ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN

Pentaceraster 100 45.5 2.2 Macroalgae (bits of algae), calcareous Rocky shores, Rocks and reef flats (Jangoux 1982b, Dee et al. 2012);
cumingi encrusting algae and biofilm (rock) coral reefs
(Gray, 1840)
Pentaceraster 100 38.5 2.6 Biofilm (epibenthic felt of Coral reefs, Seagrass and reef flats (Thomassin 1976, Jangoux 1982b)
mammillatus microorganisms) seagrass
(Audouin, 1826) beds
Continued
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References
Pentaceraster regulus 100 28.6 3.5 Biofilm (epibenthic film; organic matter) Soft bottom Sand, mud and rubble (Coleman 1977, Jangoux 1982b)
(Müller & Troschel,
1842)
Protoreaster lincki 150* 62.5 2.4 Biofilm (substrate film-feeder, epibenthic Seagrass Seagrass, sand and cobble (Thomassin 1976, Coleman 1977,
(Blainville, 1830) felt of microorganisms), algae, detritus beds, soft Jangoux 1982b)
and dissolved organic material bottom
Protoreaster nodosus 150 50.0 3.0 Biofilm (epibenthic felt of microorgan- Seagrass Seagrass, sand and cobble (Thomassin 1976, Jangoux 1982b,
(Linnaeus, 1758) isms, film on Halimeda and on beds, soft Scheibling & Metaxas 2008)
calcareous encrusting), decomposing bottom
macroalgae, seagrass and sand
meiobenthos

Facultative herbivores (ontogenetic changes in diet)

7
Acanthasteridae
Acanthaster planci 20 16.7 1.2 Calcareous encrusting macroalgae Coral reefs, Reefs, cobble, sand (Yamaguchi 1973, Yamaguchi 1974,
(Linnaeus, 1758) (Poroliton) when juveniles; Other soft bottom 1975, Sloan 1980, De’ath & Moran
macroalgae as adults during outbreaks 1998)
(calcareous turfs, filamentous, foliose)

Echinasteridae
Echinaster (Othilia) — — — Biofilm (benthic diatoms) Coral reefs, Coral and rocky reefs, (Sloan 1980)
sentus (Say, 1825) seagrass seagrass and sand
beds, soft
bottom
Henricia leviuscula 5 1.7 2.9 Biofilm (polychaete tubes) Soft bottom Sand and mud (Birkeland et al. 1971, Sloan 1980)
(Stimpson, 1857)
Continued
FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References

Goniasteridae
Mediaster aequalis 10 4.2 2.4 Biofilm (tubes of polychaete, diatoms Rocky shores, Subtidal rocks, cobble, (Mauzey et al. 1968, Birkeland et al.
(Stimpson, 1857) mats on rock) soft bottom sand and mud 1971, Sloan 1980, Jangoux 1982b,
Sloan & Robinson 1983)

Luidiidae
Luidia foliolata 5 1.5 3.4 Biofilm (on tubes of polychaetes), Soft bottom Sand and mud (Birkeland et al. 1971, Sloan 1980)
(Grube, 1866) detritus and microorganisms
Oreasteridae
Culcita novaeguineae 7 5.4 1.3 Calcareous encrusting algae Coral reefs (Yamaguchi 1973, Yamaguchi 1975,
(Müller & Troschel, Yamaguchi 1977, Sloan 1980)

8
1842)

Pterasteridae
Pteraster tesselatus 5 4.5 1.1 Biofilm (polychaete tubes, films of Rocky shores Rocks in deep seas (Sloan 1980)
(Ives, 1888) detritus and microorganisms) and
sessile invertebrates (sponges)

Stichasteridae
Stichaster australis 10 8.3 1.2 Calcareous encrusting macroalgae Rocky shores Subtidal rocks (Barker 1977, 1979, Sloan 1980)
(Verrill, 1871) (Mesophyllum insigne)

Solasteridae
ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN

Crossaster papposus 5 2.8 1.8 Biofilm (polychaete tubes, films of Soft bottom Sand and mud (Birkeland et al. 1971, Sloan 1980)
(Linnaeus, 1767) detritus and microorganisms)
Solaster stimpsoni 5 2.0 2.5 Biofilm (polychaete tubes, films of Soft bottom Sand and mud (Birkeland et al. 1971, Sloan 1980)
(Verrill, 1880) detritus and microorganisms)
Solaster dawsoni 5 2.2 2.3 Biofilm (polychaete tubes, films of Soft bottom Sand and mud (Birkeland et al. 1971, Sloan 1980)
(Verrill, 1880) detritus and microorganisms)
Continued
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References

Facultative herbivores (spatial and temporal shifts in diet)

Asterinidae
Asterina stellifera 70 28.0 2 Biofilm (diatoms), red and green Rocky shores Subtidal boulders (Jangoux 1982b, Farias et al. 2012)
(Möbius, 1859) macroalgae, invertebrates (gastropods,
anemone, polychaetes, small crabs,
sponges, ascidians), and moribund/dead
invertebrates
Meridiastra calcar 60* 37.5 1.6 Macroalgae (Laurencia sp. and Gelidium Rocky shores Intertidal rock pools, Shepherd1968; Stevenson1992;
(Lamarck, 1816) sp.), biofilm (bare rock), invertebrates subtidal rocks Shepherd 2013
(gastropods, pelecipods), detritus and
moribund animals
Meridiastra gunnii 56* 43.1 1.3 Red macroalgae (Laurencia sp. and Rocky shore, Intertidal rock pools with (Shepherd 1968, Jangoux 1982b,
(Gray, 1840) Spyridia sp.), seagrass, sessile soft bottom boulders, subtidal rocks Shepherd 2013)

9
invertebrates (bryozoans, spirorbids and sand bottom
ascidians, sponges), carrion, detritus
and moribund animals
Meridiastra medius 38 26.6 1.4 Calcareous macroalgae, biofilm (bare Rocky shore, Intertidal rock pools with (Shepherd 2013)
(O’Loughlin, Waters rock) and sessile invertebrates soft bottom boulders, subtidal rocks
& Roy, 2003) (bryozoans and ascidians) and sand bottom
Patiria miniata 67 21.0 3.2 Microalgae, early life stages of kelps Rocky shores, Low intertidal rocks, (Araki 1964, Gerard 1976, Jangoux
(Brandt, 1835) (Macrocystis pyrifera), drift algae and kelp beds subtidal rocks and kelps 1982b, Dayton et al. 1984, Leonard
seagrass, macroalgae (Egregia, 1994)
Rhodymenia, Microcladia, Iridea,
Poryhyra, Prionitis, Gigartina,
Agardhiella, Laminaria, and
Delesseriaceae), sessile invertebrates
(ascidians, sponges and bryozoans) and
FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH

dead animals.
Continued
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References
Patiria pectinifera 50 31.3 1.6 Biofilm (periphyton films), macroalgae Rocky shores, Subtidal rocks, seagrass, (Bak 1981, Jangoux 1982b, Levin
(Muller & Troschel, (Ulva), calcareous turf algae seagrass shell grits, cobble and et al. 1987, Fujita 1999, Kurihara
1842) (Corallina), encrusting calcareous algae beds, soft sand 1999)
(Lithophyllum yessoense), seagrass, bottom
small invertebrates, dead animals and
detritus
Patiriella regularis 24 16.0 1.5 Biofilm (bare rock and diatom mats), Rocky shores, Intertidal under boulders, (Grace 1967, Crump 1969, Martin
(Verrill, 1867) encrusting calcareous and non- kelp beds, underwater rocks, 1970, Burgett 1982, Jangoux 1982b,
calcareous macroalgae, calcareous turf, soft bottom cobble, shell grit, sand Burgett 1988)
green filamentous algae, invertebrates and kelps
(gastropods, ascidians and crabs),
detritus and shell debris
Pseudonepanthia 85 9.3 9.1 Calcareous encrusting macroalgae Rocky shores, Subtidal rocks, under (Shepherd 1968, 2013)

10
troughtoni (Lithothamnia), calcareous turf and soft bottom boulders and sand
(Livingstone, 1934) sessile invertebrates (bryozoans,
ascidians and sponges)

Asteropseidae
Dermasterias 85 29.3 2.9 Green algae (Enteromorpha), sponges, Rocky shores, Intertidal and subtidal (Mauzey et al. 1968, Jangoux 1982b,
imbricata anemones, ascidians, hydroids, soft bottom rocks, cobble and sand Sloan & Robinson 1983)
(Grube, 1857) holothurians, detritus, moribund
animals

Goniasteridae
ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN

Fromia hemiopla 43 9.3 4.6 Biofilm (substrate feeding on algae and Coral reef (Yamaguchi 1975, Jangoux 1982b)
(Fisher, 1913) microbes) and sessile invertebrates
(sponges)
Nectria ocellata 97 34.6 2.8 Encrusting macroalgae, detritus and Rocky shores, Subtidal rocks, shell grit (Shepherd 1967, Coleman 1977,
(Perrier, 1875) sessile invertebrates (sponges, ascidians soft bottom and sand Jangoux 1982b, Shepherd 2013)
and bryozoans)
Continued
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References
Nectria saoria 66 24.4 2.7 Red and calcareous macroalgae, and Rocky shores Subtidal rocks (Shepherd 1967, Jangoux 1982b,
(Shepherd, 1967) sessile organisms (ascidians and Shepherd 2013)
bryozoans)
Nectria multispina 85 32.7 2.6 Macroalgae (red algae), rock encrusting Rocky shores Subtidal rocks (Shepherd 1967, Jangoux 1982b)
(H.L. Clark, 1928) organisms, sessile invertebrates
(sponges, ascidians and bryozoans) and
small bivalves
Mediaster aequalis 100* 33.3 3.0 Macroalgae (Ulva spp. and Porphyra Rocky shores, Subtidal rocks, cobble, (Mauzey et al. 1968, Birkeland et al.
(Stimpson, 1857) spp.), biofilm (tubes of polychaete, soft bottom sand and mud 1971, Sloan 1980, Jangoux 1982b,
diatom mats on rock) microalgae Sloan & Robinson 1983)
(planktonic diatoms), detritus, sessile
invertebrates (ascidians, sponges,
bryozoans) and moribund animals

11
Ophidiasteridae
Gomophia egyptiaca 50 7.4 6.8 Biofilm (substrate feeding on algae and Coral reef Reef flat and lagoon (Yamaguchi 1975, Jangoux 1982b)
(Gray, 1840) microbes) and sessile invertebrates
(sponges, ascidians) and moribund
animals

Oreasteridae
Oreaster reticulatus 165 126.9 1.3 Biofilm, green algae (Chaetomorpha, Seagrass Subtidal seagrass, sand (Scheibling 1979, Scheibling
(Linnaeus, 1758) Enteromorpha), epiphytes, seagrass, beds, soft and rubble 1980b,d, 1981, Jangoux 1982b,
invertebrates (sponges, sea urchins) bottoms Scheibling 1982, Martin et al. 2001,
Scheibling 2013)
Culcita schmideliana 120 109.1 1.1 Seagrass, sandy substratum, sponges, Seagrass Intertidal seagrass, sand (Thomassin 1976, Jangoux 1982b)
(Retzius, 1805) corals, film growing on dead corals beds, soft
bottoms
Continued
FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH
 Table 1 (Continued) Classification of starfish herbivory according to temporal and spatial fidelity of a species to a plant/algae resource
Size
Species R r R/r Food resource Ecosystem Habitat References

Stichasteridae
Granaster nutrix 70 25.0 2.8 Red macroalgae, small gastropods, Rocky shore Underwater rocks (Jangoux 1982b)
(Studer, 1885) ectoprots
a An uncertain starfish species from the family Asterinidae, formerly Marginaster littoralis Dartnall, 1970.
Note: Starfish size, food resource exploited, and distribution among ecosystems and habitats are presented for each species. Detailed information of food resource and habitat are presented
whenever available from the literature. The references cited in this table include the primary source of information for a given species and all other studies to date that recorded that
species feeding on plant/algal resources. Starfish metrics are shown for average sizes in mm (R – arm size, r – disc size) with exception for some species (maximum size instead) whose
average size was not found (indicated by an asterisk) and for those species classified with ontogenetic change in diet.

12
ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN
 FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH

as species feeding on “bare rock”, “diatoms”, “diatom mats on rock”, “films of dead corals, detri-
tus, tubes of polychaetes, shells, macroalgae, seagrass”, “microalgae”, “epibenthic film”, “bacterial
film”, “epilithic organisms on algae” and “epibenthic felt of microorganisms”; or characterized as
“substratal film-feeder” and “substratal feeding on algae and microbes”.
Biofilms are one of the main food resources that are also exploited by several grazing gastro-
pods. One of the reasons that gastropods forage on this food resource is that their feeding abilities
are limited by the physical characteristics of their radular feeding apparatus (Steneck & Watling
1982, Hawkins et al. 1989). Asteroids do not have any feeding apparatus such as radulae, Aristotle’s
lantern in sea urchins or teeth (e.g. fish) that can mechanically rasp, cut or bite the algae from the
substratum. The algal food resource utilized by herbivorous starfish relies on the digestive capac-
ity of their stomach. Feeding on biofilms could be a cost-efficient source of food for starfish, since
diatoms, bacteria and algal sporelings are protected only by individual cell walls, which are easier
to break down compared to the structures of multicellular macrophytes. Macrophytes not only have
thick walls, but also exhibit complex chemical defences (Hay 2009).
The enzymes produced in the pyloric ceca of some starfish (e.g. Oreaster reticulatus, Patiria
miniata, Patiriella regularis), are capable of digesting some oligosaccharides and polysaccharides
that are the sugar reserves of plants and algae (e.g. sucrose, trehalose, amylose, and laminarinose),
but they have a weak effect on, or failed to break down, the structural components of plant/algal cell
walls such as cellulose, alginate, agar, and carrageenan (Araki 1964, Araki & Giese 1970, Martin
1970, Scheibling 1980c). This suggests that starfish are potentially capable of digesting macroalgae,
but that feeding on these algae with thick cell walls might not be cost-effective. Still some starfish
seem to feed on macroalgae, but on less structured thalli forms. Soft-structured macrophytes such
as soft foliose (e.g. Ulva spp.) and filamentous algae (e.g. Ceramium, Chaetomorpha, etc.) appear
to constitute the main group of macroalgae in the diet of some herbivorous starfish. These algae
can be intensively grazed by several herbivores including snails, amphipods, isopods, crabs, and
fish (e.g. Choat 1982, Hawkins & Hartnoll 1983, Arrontes 1990, Poore 1994, Kennish & Williams
1997). This is because soft filamentous and foliose algae are generally easy to digest, have rapid
growth rates and energy intake, and do not offer strong resistance to herbivory (Littler & Littler
1980, Littler & Arnold 1982, Steneck & Dethier 1994). The cell walls of soft and thin thalli algae
are generally structured in a simple uni- or multiseriate cell configuration, which are not differenti-
ated and heavily corticated unlike those of thick leathery or calcareous or crustose algae (Steneck
& Dethier 1994). Thus, starfish could benefit from feeding on those soft algae by disrupting their
simple cell walls with their enzymes and accessing the digestible cell content (Kristensen 1972,
Scheibling 1980a). This hypothesis needs to be addressed through investigation of the digestive
capacity of herbivorous species.
The fact that starfish are not able to readily break down the cell walls of plants/macroalgae
raises the question of whether asteroids are able to feed on these more complex-structured pri-
mary producers. The diet of Asterina gibbosa, Patiria miniata and Protoreaster nodosus includes
macroalgal detritus (Gerard 1976, Crump & Emson 1978, Scheibling & Metaxas 2008), and it is
reasonable that these starfish species can eat decaying algae as the cell walls are already damaged.
Indeed, feeding on decomposing macroalgae by Patiria miniata enhanced the nutritional value of
the food source compared to intact layers of the algae (Gerard 1976). Some starfish species, how-
ever, were described as feeding on calcareous turf algae, calcareous and non-calcareous encrusting
algae, red corticated algae and seagrass (Table 1). There is no certainty whether those starfish were
actually feeding on the macroalgae itself or on the epiphytes or biofilm on those macrophytes. Some
authors argue that starfish probably eat the epilithic organisms that grow on the macroalgae, but
many authors noted that the plant/algae area on which a starfish had their stomach extruded became
discoloured (Araki 1964, Crump 1969, Yamaguchi 1973, Scheibling 1979, Scheibling 1982, Levin
et al. 1987, Leonard 1994, Farias et al. 2012). This is notably visible in calcareous algae, which
may look bright orange or white after starfish feeding (Figure 1A,B). The physical structure of

13
 ALINE S. MARTINEZ, MARIA BYRNE & ROSS A. COLEMAN

A B

30 mm 20 mm

C D

10 mm 50 mm

Figure 1 Discolouration of algae and scars caused by starfish herbivory: (A) Calcareous turf discoloured
(bright orange) by Meridiastra calcar feeding (Photo: Aline Martinez); (B-D) Feeding scars of Parvulastra
exigua on (B) encrusting calcareous algae (Photo: Ross Coleman), on (C) bare rock, in detail (Photo: Aline
Martinez), and (D) around rock pools on open rock (Photo: Aline Martinez).

those macrophytes, however, seems to not be affected by the starfish feeding. Indeed, studies of the
effects of starfish grazing on calcareous algae showed that the top layer of live tissue on the algae
was removed after being grazed, but the structure of the cell walls remained intact (Barker 1979,
Burgett 1982, Fujita 1999). Hence it is more likely that starfish are targeting epiphytic organisms
growing on the macroalgae than the macroalgae itself, although Bak (1981) showed that Patiria
pectinifera intensely consumed eelgrass.
In summary, it seems that the plant/algal resources exploited by starfish are limited by their
digestive enzymes. These enzymes can break down unicellular components of the biofilm and might
be able to disrupt the cell walls of soft filamentous and foliose algae to readily digest and assimilate
material from these macrophytes. Moreover, it is likely that starfish foraging on more robust mac-
roalgae are eating epiphytes and other organisms growing on the thalli. Whether or not the starfish
benefits from the apparently digested (discoloured) area on these macrophytes is not known.

14
 FILLING IN THE GRAZING PUZZLE: A SYNTHESIS OF HERBIVORY IN STARFISH

Categories of herbivory
The definition of trophic guilds (e.g. carnivores, herbivores, omnivores etc.) in asteroid echino-
derms is often convoluted and not clear for species that are not strictly carnivores. Trying to distin-
guish between herbivorous and omnivorous starfish becomes complicated because the behaviour
of herbivorous species can vary across time (amount or type of plant/algae consumed by individu-
als varies at different times) and space (individuals at different places have different diets). There
are also species that are herbivorous only during a specific life stage, usually the juvenile stage
(e.g. Yamaguchi 1973, Barker 1977, 1979, Sloan 1980; Kamya et al. 2016) whereas other species
might forage for their entire life on algae (e.g. Laxton 1974, Arrontes & Underwood 1991, Prestedge
1998, Wilson 2004, Jackson et al. 2009). Differentiating herbivorous behaviour becomes more com-
plicated when the same species displays different feeding habits between populations, locations
or seasons (e.g. Araki 1964, Mauzey et al. 1968, Shepherd 1968, Leonard 1994, Scheibling 2013).
Therefore, we approached this review by classifying starfish into different herbivore status depend-
ing on temporal and spatial fidelity of a species to an algal resource. Thus, starfish species were
classified into obligate and facultative herbivores (Table 1).

Obligate herbivores
Starfish species that primarily feed on algae were classified as obligate herbivores and are consid-
ered herbivorous starfish sensu stricto. Some species were noted to feed on detritus in addition to
algae and were also included in the obligate herbivorous category. While detritus may be a range
of decaying material, it is unlikely to include dead animals because starfish foraging on dead or
decomposing animals are clearly identified in the literature. Obligate herbivores also include some
species that, while foraging primarily on algae, on very rare occasions are reported to feed on ani-
mal-derived food resources, which was the case for Parvulastra exigua and Pentaceraster cumingi
(Branch & Branch 1980, Dee et al. 2012).
Most herbivorous starfish (70%) belong to the family Asterinidae and Ophidiasteridae. The
Asterinidae includes, Aquilonastra anomala, Asterina gibbosa, A. phylactica, Cryptasterina hys-
tera, C. pentagona, Parvulastra exigua, P. parvivipara and P. vivipara (Dartnall 1971, Yamaguchi
1975, Duyverman 1976, Crump & Emson 1978, Keough & Dartnall 1978, Emson & Crump 1979,
Jangoux 1982b, Chen & Chen 1992, Prestedge 1998; Dartnall et al. 2003). These herbivores are
small (average radius varies from 3 to 15 mm; see Figure 2A for example) and many occur in inter-
tidal pools on temperate rocky shores. There are also tropical species that occur in intertidal pools
of coral reefs (Aquilonastra anomola and Cryptasterina pentagona). These herbivorous asterinids
feed on surficial biofilms on rocks and coral rubble. Parvulastra exigua and Asterina gibbosa may
also feed on macroalgae (encrusting algae, soft filamentous and foliose algae, and decaying algae),
but these contribute a small portion of their diet (Crump & Emson 1978, Branch & Branch 1980).
The species from the Ophidiasteridae are nearly 10 orders of magnitude bigger (average radius
varies from 30 to 160 mm) than those in the Asterinidae. Herbivorous ophidiasterids include spe-
cies from the genera Cistina, Dactylosaster, Linckia, Nardoa, Ophidiaster and Phataria and occur
mostly in tropical coral reef flats and lagoons. Although they are bigger than the asterinids, the
extruded stomach (ca. 5–20 mm radius) of the biggest ophidiasterids (i.e. Linckia spp.; Figure 2B)
is similar in size to that of the small asterinids.
The other eight herbivorous starfish are distributed within the families Echinasteridae (2 spp.
from the genus Echinaster), Goniasteridae (Neoferdina cumingi) and Oreasteridae (2 spp. from the
genus Pentaceraster and 3 spp. from the genus Protoreaster). The Echinaster spp. and Neoferdina
cumingi are as big and similar in shape (i.e. long arms and small central disk) as Linckia spp.
These starfish inhabit coral reef flats and seagrass beds where they feed on the biofilms on hard
substrata and on epibenthic films of the sediment. The oreasterids are big starfish (average radius

15
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