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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
CRC Press
Taylor & Francis Group
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© 2017 by S.J. Hawkins, A.J. Evans, A.C. Dale, L.B. Firth, D.J. Hughes, and I.P. Smith
CRC Press is an imprint of Taylor & Francis Group, an Informa business
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Version Date: 20170206
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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: ross.coleman@sydney.edu.au
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 &
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
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
Size
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
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
Size
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
Size
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
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
Size
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
dead animals.
Continued
Size
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
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
Size
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
Size
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
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
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
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
A B
10 mm 50 mm
C D
5 mm 100 mm
Figure 2 Aboral view of different categories of herbivorous starfish: Obligate herbivores. (A) Parvulastra
exigua in a feeding mode shape (round; Photo: Aline Martinez), (B) Linckia laevigata (Maria Byrne); and
facultative herbivores, (C) Acanthaster planci at juvenile stage (Photo: Phil Mercurio) and (D) Oreaster retic-
ulatus (Photo: Marcela Rosa).
100–150 mm) and typically inhabit soft bottoms; foraging on seagrass, sand, and among cobbles.
Pentaceraster cumingi is the only oreasterid species restricted to hard bottoms where it feeds mostly
on biofilms and calcareous encrusting algae (Dee et al. 2012). Oreasterids exploit a broad range of
plant/algal resources including biofilm, macroalgae, decomposing algae and seagrass.
It appears that eating plant/algal resources by obligate herbivores is associated with the lack
of ability to catch or manipulate animal resources as predatory starfish do. Pentaceraster cumingi
consumes small sea urchins (corresponding to 1% of their diet), but feeding was observed on teth-
ered sea urchins (Dee et al. 2012). Whether the starfish actively captured live sea urchins or was
attracted by a moribund sea urchin already attacked by other predators (e.g. fish) is not clear. Other
herbivorous species such as Protoreaster nodosus might not primarily forage on animal-derived
resources, but benefit from ingesting small invertebrates (i.e. meiofauna) when foraging on plant/
algal food (Thomassin 1976, Scheibling 1982, Scheibling & Metaxas 2008). Complementing an
16
algal diet with animal resource can benefit animal growth as demonstrated for herbivorous crabs
(Wolcott & O’Connor 1992). An explanation for this is that animal food is much richer in essential
amino acids, vitamins and sterols compared to plant/algal resource (Phillips 1984).
Interestingly, obligate herbivorous starfish do not forage, or rarely forage, on soft sessile inverte-
brates such as sponges, bryozoans or ascidians, even though there is no need to handle these organ-
isms. Possible causes for this feeding behaviour include feeding preferences, chemical defences
of sessile invertebrates, especially sponges and ascidians, physical prey defences and/or diges-
tive capabilities of starfish. This is an unexplored subject in herbivorous starfish that needs to be
addressed to better understand their feeding behaviour.
Facultative herbivores
Many starfish species that forage on algae also feed on animal-derived material and this is quite
variable among and within species, individual lifespan, seasons and location of populations. These
species are classified as facultative herbivores, as their algal/plant diet changes in time or space.
Ontogenetic changes in diet

Herbivory in starfish that display ontogenetic changes in diet is reported from the early life stages
of carnivorous or omnivorous species (e.g. Yamaguchi 1973, Barker 1977, 1979, Barker & Nichols
1983). Only a few species have been described as herbivores when juvenile and carnivores when
adult (Table 1). The lack of information on the feeding behaviour of juvenile starfish is due to the
difficulty in finding newly-settled or post-metamorphosed starfish in situ (Yamaguchi 1975, 1977).
Juveniles are small and often cryptic. Despite the lack of detailed studies, there is evidence that
some predatory starfish are herbivores when young (for a review, see Sloan 1980), for example
Acanthaster planci (Yamaguchi 1973, Yamaguchi 1975; Kamya et al. 2016), Mediaster aequalis
(Birkeland et al. 1971) and Stichaster australis (Birkeland et al. 1971, Yamaguchi 1973, Barker
1977, Sloan 1980).
The feeding behaviour of juveniles of the predatory starfish Acanthaster planci is well docu-
mented. Encrusting calcareous and coral rubble substrata induce settlement and metamorphosis of
A. planci (Johnson et al. 1991). After metamorphosis, juveniles feed on calcareous encrusting algae
for around 18 weeks (radius ca. 20 mm; Figure 2C), after which, they start foraging on small coral
polyps (Henderson & Lucas 1971, Yamaguchi 1973, Yamaguchi 1975). Similarly, the coral reef
starfish Culcita novaeguineae settles on encrusting calcareous algae and spends at least 12 months
on this substratum (radius reaches ca. 3.5 mm) (Yamaguchi 1973, Yamaguchi 1975), but whether
further feeding and growth rely on such algae is not known. Adult C. novaeguineae forage on ses-
sile invertebrates including coral, but still eat algae. Similar to these tropical species, the temperate
starfish Stichaster australis uses the coralline alga Mesophyllum insigne as a nursery area. Juvenile
Stichaster australis remain in this habitat and feed on the algae for 15 to 18 months (until ca.
6.5 mm radius) and then switch to juvenile mussels (Perna canaliculus) (Barker 1977, 1979). The
larvae of Mediaster aequalis settle and feed on the biofilm that grows on tubes of the polychaete
Phyllochaetopterus prolifica, until the juvenile reaches around 5 to 10 mm radius and then feeds on
animal prey and algae (Birkeland et al. 1971). In a survey of the feeding behaviour of juvenile star-
fish, Birkeland et al. (1971) noted that individuals smaller than 5 mm radius (Henricia leviuscula,
Luidia foliolata, Pteraster tesselatus, Crossaster papposus, Solaster stimpsoni and S. dawsoni)
feed on the tubes of Phyllochaetopterus prolifica. It appears that a period of herbivory in the early
juvenile stage may be a common feature of carnivorous starfish, although not all juveniles studied
were observed feeding on biofilm. Juvenile Asterias rubens and Marthasterias glacialis eat small
encrusting invertebrates as soon as metamorphosis occurs, although these starfish settle in a variety
of substrata covered with biofilm (Barker & Nichols 1983).
17
The extent to which herbivory occurs in young starfish appears to be influenced by the growth
patterns of a given species. Newly-settled starfish only have one to two pairs of tube feet and so
feeding depends on extruding the stomach (Sloan 1980). Juveniles of predatory starfish pass through
a phase during which young starfish start foraging on small animals as soon as they are big enough
to catch prey. At this stage the juveniles stop feeding on algae. When adult, many predatory species
use their disc-ending tube feet to hold prey and open shells (Hennebert et al. 2013). These starfish
may also have extensive batteries of specialized pedicellariae (jaw-like structures) that the adults
use to ensnare large prey, including fishes (e.g. Stylasterias forreri, Chia & Amerongen 1975). For
predatory starfish, it appears that morphological changes are required as a juvenile develops into a
competent predator. However, ecological–morphological studies of diet change through growth of
juveniles are necessary to test this hypothesis.
In addition to the intrinsic biology of asteroids, external factors such as availability of resource
and predation risk might influence diet shifts in starfish, as has been shown for crabs. Juvenile crabs
inhabit macrophyte habitats where they are safe from predators and forage on the available food
(i.e. plant/algal resources) until they achieve a certain size at which they are less vulnerable to pre-
dation and then move to open areas (Laughlin 1982, Alexander 1986, Orth & Vanmontfrans 1987,
Williams et al. 1990). In a similar way, Yamaguchi (1975) argued that the cryptic behaviour of tropi-
cal juvenile starfish is associated with the risk of predation, which has been shown to be high in the
early life stage (Keesing & Halford 1992). Foraging on encrusting algae exploits a highly abundant
algal resource available within sheltered-cryptic habitat in coral reefs. When animals become big
enough to escape predation, starfish would then start foraging on open areas on the reefs, as dem-
onstrated for Acanthaster planci (Yamaguchi 1973, Yamaguchi 1975). These observations suggest
that herbivorous behaviour may have evolved in these species through a predator–prey interaction
with the juvenile stages remaining in cryptic algal dominated habitat until they grow to a size where
they are less vulnerable to predation.
It is clear that the ontogenetic change in starfish diet is related to starfish size (Birkeland 1989).
Thus, investigations of feeding habits associated with morphological changes in post-metamor-
phosed juvenile starfish, as well as feeding behaviour associated with available food resources and
predation risk, would be of great interest to understand the shifts from a herbivorous to a carnivo-
rous diet in asteroids. This may help in understanding the mechanisms that drive different feeding
behaviours in starfish, a taxon where feeding is not constrained by possession of a complex feed-
ing apparatus.
Spatial and seasonal changes in diet

Most asteroids are considered to be opportunistic with feeding habits reflecting the availability of
resources (for a review, see Sloan 1980, Jangoux 1982b). A more detailed examination of the litera-
ture indicates that several species classified as omnivores or even carnivores, display herbivorous
behaviours linked to the variability of food between locations and seasons. Species are referred
to here as facultative herbivores. Eighteen species were included in this category belonging to the
families Asterinidae (8 spp.), Goniasteridae (5 spp.), Oreasteridae (2 spp.), Asteropseidae (1 sp.),
Ophidiasteridae (1 sp.) and Stichasteridae (1 sp.). The most evident aspect of the feeding behaviour
of facultative herbivorous starfish is that the proportion of algae eaten by a population varies accord-
ing to spatial and temporal fluctuations in density of food resources.
A well-studied species is the red cushion star Oreaster reticulatus (Figure 2D), a species that
inhabits soft bottoms around reef areas such as sand, seagrass and cobble. Populations of this spe-
cies vary in their food selection, being either microphagous grazers or macrophagous predators,
and the proportion of the food sources utilized changes with respect to the availability of mac-
rofaunal prey (Scheibling 2013). Oreaster reticulatus forages on biofilms (microalgae and other
microscopic organisms), detritus, filamentous algae and seagrass in locations where animal prey
(sea urchins and sponges) is absent (Scheibling 1980d, 1982, Wulff 1995). Similarly, feeding by
18
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nail holes; also putty up any of the grain that may appear too open,
or else rub into the grain some lead mixed up very heavy.
The body, after having received three coats of lead, and been
puttied up, may now stand for two or three days. When it is again
taken in hand, sand-paper off any putty that may be above the level
of the surface; dust off, and brush on a level coat of lead, which must
dry hard and firm. Every coat of lead should be laid on as level as
possible, and made to fill up the grain of the wood as much as
possible. These coats are called “rough stuff.” The body may now
stand for three or four days, when it will be ready for the filling up.
There are two very important things to be studied in coach-
painting. First, to form a surface hard enough to hold out the varnish
and disguise the grain of the wood; and second, to have the first and
intermediate coats of paint sufficiently elastic to adhere and yield to
the natural action of the wood without cracking or flaking off. In
effecting one of these results we are apt to affect the other; and
nothing but the utmost care, on the part both of the manufacturer of
the essential ingredients and of the person who prepares them for
use, can insure durability.
The leather-covered portions are usually primed with two coats of
black Japan, reduced with a little turpentine.
A good stopping material for nail holes, &c., is made of dry lead
and Japan gold size. It is called “hard stopper.”
The rough coatings should dry firmly, possessing only sufficient
elasticity to bind them to the surface. The first coat will bear a trifle
more oil than the remaining ones, and should stand about four days
before the others are put on, which can be done every other day.
Five coats of filling up are next added, composed as follows:—
2 parts filling up stuff.
1 part tub lead.
2 parts turpentine.
1 part Japan gold size.
½ „ bottoms of wearing varnish.
The first coat should cover every portion of the lead surface, be
well brushed in, but not allowed to lie heavy at the corners. The
remaining coats may be applied reasonably heavy, but kept from
lapping over the edges or rounding the sharp corners, and thus
destroying the clean sharp lines of the body-maker.
Any defects noticed while filling in should be puttied or stopped,
ever bearing in mind that the perfection in finish aimed at is only
secured by care at every step taken.
The leather-covered parts generally have three additional coats of
filling in.
The time allowed for each coat to dry may be extended as far as
convenient, but there is nothing to be gained by allowing weeks to
intervene between the coatings. When a coat is hard it is ready for
another; and it is far better to have the body filled and set aside than
to divide the time between the coatings, and probably be compelled
to rub out the body before the last coat is firm. Of course, the time
occupied by the coats of filling in to dry varies according to its
composition. If much oil be used, it will take a longer time for each
coat to dry; but the above composition may be applied one coat
every other day.
The first coat may be applied rather thinner than the others, and is
improved by being mixed with a little more white-lead. It should also
be made more elastic than the succeeding ones, as it will then take a
firmer hold on the “dead” lead coat over which it is placed,
contributing a portion of its elasticity to that coat, and also cling more
firmly to the hard drying coats which follow.
The body having been filled in may be set aside to harden, or if the
smith is ready for it this is the best time for him to take it in hand, as
any dents or burns that he may cause can now easily be remedied
without spoiling the appearance of the vehicle. Later on, this is a
matter of great difficulty, if not impossibility. Any bruises should be
puttied; any parts which may happen to be burned must have the
paint scraped off bare to the wood. Prime the bare spots, and putty
and fill them to bring them forward the same as the general surface.
A material has recently been brought into the English market
called “permanent wood filling,” which is confidently recommended
as effecting a saving in time, expense, and labour, and at the same
time more effectually closing the pores of the wood than the ordinary
filling now in use. This invention is due to a Polish exile named
Piotrowski, who took refuge in America, and there introduced it about
1867, since which date it has found its way into the chief carriage
factories in the United States. It is applied to the bare wood, one coat
being given to bodies and two coats to carriage parts. This closes
the pores, holds the grain immovably in its place, and is so
permanent in its effect that neither exposure to dampness, nor
atmospheric changes, nor the vibrations to which a carriage is so
subject can affect the grain. The satisfaction which this material
appears to give to the Americans, who pride themselves on the
superiority of their carriage-painting, ought to induce our English
coach-builders to inquire after it; for if all that we hear of it be correct,
it must assuredly be a valuable acquisition to the paint shop.
In rubbing down use pumice-stone. It is best to begin on top and
follow on down, so that the filling water may not run down on to any
part that has been finished. Water should not stand for any length of
time on the inside of the body; and when the rubbing is completed
wash off clean outside and in, and dry with a chamois kept for the
purpose.
The body, when dry, receives a staining coat, and is to be carefully
sand-papered over, the corners cleaned out, and put on a coat or
two coats of dark lead colour, made of tub lead, lampblack, raw oil,
and a small quantity of sugar of lead, and reduced to a proper
consistency with Japan gold size and turpentine. When dry scratch
over the lead colour with fine sand-paper, which will make it appear
of a lighter colour; we shall then be able to detect any low or sunk
places by reason of the shadow. Putty up any imperfections with
putty made of lead and varnish, and when dry face down with lump
pumice and water. Follow with fine sand-paper, when the surface will
be in a condition to receive the colour coats. Sometimes, after
cleaning off, another coat of dark lead colour is laid on.
Analysing the foregoing, we find we have used—
1 priming coat of lead (or leather parts, 2 coats of
black varnish instead).
2 thin coats lead colour, and stopped up.
5 coats of filling up (8 coats on leather parts).
1 staining coat, rubbed down and cleaned off.
2 coats dark red colour, stopped up, and carefully rubbed down.
1 coat dark lead colour.
—
12 coats, and ready for colour.
So much for the body parts. To the carriage parts two coats of
priming are laid on, which are worked in the same way as those
applied to the body. All cavities are then stopped with hard stopper,
to which a little turpentine is added in order to make it sand-paper
easily. Two coats of quick-drying lead colour are then applied to the
wood parts. The whole is then well sand-papered down, and the
grain should be found well filled and smooth. A thin coat of oil lead
colour is then laid on, and when dry sand-papered down; any joints
or open places between the tire and felloes of the wheels are
carefully puttied up with oil putty. The carriage parts are then ready
for colour. This time we have applied—
2 coats of lead priming, stopped up.
2 coats of lead, thoroughly sand-papered.
1 coat (thin) of lead colour, sand-papered and puttied up.
—
5 coats, and ready for colour.
The colours are to be ground very fine, kept clean, and spread on
with the proper brushes. If the panels are to be painted different from
the other parts, lay on the black first, for if any black falls on the
panel colour it will occasion some trouble by destroying the purity of
a transparent colour. By repeatedly turning the brush over while
using it, there is less liability to accidents of this kind.
The colouring of the body is finished as follows:—For the upper
quarters and roof grind ivory black in raw oil to a stiff consistency,
add a little sugar of lead finely ground as a drier, and bring to the
required consistency with black Japan and turpentine. Lay on two
coats of this, and then two coats of black Japan, and rub down. Then
face off the moulding, and give a thin coat of dead black, after which
apply another coat of black Japan, and flat again. The whole should
then be varnished with hard drying varnish, flatted down, and
finished with a full coat of wearing body varnish. The varnish should
have at least three days to dry; five or six would be better. The first
coat of rubbing varnish may be applied thinner than the others, in
order to avoid staining the colours.
The pencils used on mouldings should be large enough to take in
the whole width at once, and let the colour run evenly along,
avoiding laps or stoppages, except at the corners, where it cannot be
helped. Avoid the use of turpentine in varnish if possible; but if the
varnish be dark and heavy, sufficient turpentine added to make it
flow evenly will not hurt it. The half elastic and fine bristle brushes
are better for working heavy varnish than the sable or badger.
In varnishing a body begin on the roof, bringing the varnish to
within 2 or 3 inches of the outer edges. Next, the inside of doors, &c.,
then the arch. When these are finished, start on the head rail on one
side, lay the varnish on heavy, and follow quickly to the quarter. The
edge on the roof, which was skipped before, is to be coated and
finished with the outside, thus preventing a heavy edge. Continue
round the body, finishing the boot last.
The frames and other loose pieces about a coach should be
brought forward along with the body, and not left as is often done.
The frames are most conveniently handled by a device similar to a
swinging dressing-glass; a base and two uprights stoutly framed
together, allowing space for the frame to swing. It is held in its
position by two pointed iron pins, one fixed and the other movable.
This is very convenient for varnishing, as the painter can examine
his work by tilting it to any angle, and thus detect any pieces of dirt,
&c.
If the body is to be lake in colour, the lake should be ground in raw
oil, stiff, and reduced with turpentine and hard drying varnish. The
same with dross black and Indian red. Over lakes and greens two
coats of hard drying varnish should be applied, and one coat of
finishing.
If the body is to be blue, mix ultramarine blue with one-half raw oil
and turpentine, and bring it to a workable consistency by thinning
with hard drying body varnish. Give the body two coats, and after
each a slight flatting; then give two more coats of the same with
varnish added.
When Prussian blue is used, two coats are applied, and white is
added, if necessary, to bring it to the required shade. The blues will
dry sufficiently well when merely ground in raw oil, stiff, and reduced
with turpentine, and it is better not to add a drier over blues; only one
coat of hard drying body varnish should be given, and one finishing
coat.
In no case should the painter allow his oil colours to dry with a
gloss. He must always flat them and give them the appearance of
dead colour. This is particularly important, in case rough stuff or
quick-drying colour is to be used over it.
The carriage parts are finished as follows:—Two coats of lead
colour are first laid on, composed in the same way as those for the
body before the colour is applied. Then stop all parts requiring it with
hard stopper, a little reduced with turpentine to sand-paper easily. To
the wood parts apply two coats of quick lead, composed of dry lead
and lampblack ground in gold size and thinned with turpentine.
Sand-paper down thoroughly, and the grain will be found smooth and
well filled up. A thin coat of oil lead colour is then applied, and sand-
papered down when dry; and at this stage any open parts between
the tire and felloe of the wheels, &c., should be again stopped up
with oil putty. A coat of colour varnish follows, then a second, with
more varnish added. The parts are then flatted and striped; another
light coat of clear varnish is given, and after being flatted down the
fine lines are added, and the whole is finished with a good coat of
wearing varnish.
The carriage parts are generally painted one or two tones lighter
than the colour of the panels of the body, except where the panel
colour is of a hue that will not admit of it. Certain shades of green,
blue, and red may be used on panels, but would not, when made a
tint or two lighter, be suitable for a carriage part. Dark brown, claret,
and purple lake would not be open to this objection, because, to the
majority of persons, they are colours which are pleasing to the eye,
both in their deep and medium tones.
When the panels are to be painted green, blue, or red, and the
painter wishes to carry these colours on to the carriage part, it is
better to use them for striping only, and let the ground colour be
black.
A carriage part painted black may be made to harmonise with any
colour used on the body, as the striping colours can be selected so
as to produce any desired effect. Brilliant striping can be brought out
on dark colours only, while, if the ground colour be light, recourse
must be had to dark striping colours to form a contrast. The carriage
part should not detract from the appearance of the body; that is,
there should be sufficient contrast between the two to bring out the
beauties of the body. A plainly finished body will appear to better
advantage on a showy carriage, and a richly painted body on one
that is not very ornate.
In striping the carriage parts, the bright colours should be used
sparingly. A fine line placed on the face of the spokes and naves,
and distributed over the inside carriage, would look far better than
when each side of the spokes, the faces, the naves, and felloes, &c.,
are striped on both sides.
The coatings of varnish contribute largely to the durability as well
as beauty of a carriage part. The ground and striping colours are
shown in their purity only after they are varnished and have a good
surface, and the test of wearing depends on the quantity and quality
of varnish applied.
Every carriage part should have at least two coats of clear varnish.
The first coat of varnish to be applied over the colour and varnish;
the second, a good finishing coat, possessing body, and good
wearing qualities. Ground pumice and water must be used to cut
down the varnish, otherwise the finishing coat will be robbed of its
beauty.
In laying on the finishing coat, avoid the extremes of putting it on
too thick or too thin. Lay on a medium coat. A thin one will appear
gritty and rough; and one too heavy will sink in and grow dim.
From the above description, it will be seen that painting a coach is
a tedious operation, and one which consumes a great deal of time in
its execution; but, if well done, the result will certainly be very
satisfactory. In no case should the painting be hurried, for by
allowing each coat of paint or varnish sufficient time to dry its
durability is insured.
A considerable amount of time is generally spent by the painter in
work which does not really belong to him—that of mixing and
grinding his colours. Where the muller and slab are used, they
occasion a great deal of labour, and the tones of the colours are
liable to be injured by the heat generated in the process; and even
where the hand mills are used, the process is by no means so
cleanly as it ought to be. And under the heading of waste, this must
always be a source of loss to the manufacturer, for the painter, for
fear of not mixing up sufficient colour for his use, generally prepares
too great a quantity, and as a rule, the surplus is waste, for it is no
use to employ stale colours in painting vehicles, however well it may
do in house painting.
What we want is to have the colours ready ground for the painter’s
hand, and against this has been urged the objection, that the
delicate colours would lose their purity, and all colours be more or
less affected by it. That this is utterly fallacious is seen by the fact
that paints and colours ready ground and prepared are the rule in
America. The invention of the machinery, &c., for this purpose, is due
to Mr. J. W. Masury, of New York, and he grinds pigments of the
hardest description to the most impalpable fineness without injuring
the tones of the most delicate; and by a process of his own
preserves them, so that the painter has nothing to do but to reduce
them to the consistency he may require for the work in hand. He
says they effect a saving of from 20 to 50 per cent. both of labour
and material. It is difficult to understand why so valuable an invention
is not more general in this country.
Irregularities in Varnish.
Varnish is subject to various changes after having been applied to

a body or carriage part. It crawls, runs, enamels, pits, blotches,
smokes or clouds over, and in the carriage parts gathers up and
hangs in heavy beads along the centre of the spokes, &c.
These irregularities will happen at times with the very best varnish
and the most skilled workmanship, and surrounded with everything
necessary to insure a perfect job.
The only reason that can be assigned for it is atmospheric
influence. These peculiarities have occupied a large portion of the
time of the trade, and no other solution has been arrived at than the
above.
The defects of varnishes should be divided into two classes: those
which take place while in the workshop and those which show
themselves after the vehicle has left the hands of the maker. The
defects which show themselves in the varnish room are those of
“spotting,” “blooming,” “pin-holing,” “going off silky,” “going in dead.”
Those which take place afterwards are “cracking,” “blooming,” “mud-
spotting,” and loss of surface, sometimes amounting to its almost
total destruction.
The two classes should be considered separately; and assuming
that the workmanship is of the best quality, the latter class of defects,
with the exception of blooming, are in no way attributable to the
varnish; and blooming is caused by the atmosphere being
overcharged with moisture, as would be the case before a storm,
and it is soon remedied. Cracking will arise from too great an
exposure to the sun, just as any other material will be damaged by
unfair treatment. Mud-spotting will arise from using the carriage in
muddy or slushy roads before the varnish is properly dry. The loss of
surface will depend largely on the coachman, who, from ignorance or
negligence, may rub down the panels of a carriage until its glossy
surface entirely disappears; and if the stable is contiguous to the
coach-house this destruction will be assisted by the ammoniacal
vapours arising from the manure, &c.
The other defects belong to the inherent nature of the varnish as
at present manufactured, and admitting the secondary cause to be
atmospheric influence, it is necessary to inquire why it is that varnish
should be subject to such influence. According to the usual way of
making varnishes, we know that various metallic salts and chemical
compounds are used to increase their drying properties. All these will
contain a certain definite amount of water, termed “water of
crystallization.” If deprived of this water they lose their crystalline
form, but they acquire a tendency of again assuming it by attracting
to themselves a proportionate amount of water when it is brought
within their power. Now the heat employed in making varnishes is
sufficient to expel this water; but the presence of the salts is sooner
or later detected, for when the varnish is applied to the work these
salts absorb moisture from the atmosphere, and by becoming
partially crystallized cause what is known as “blooming,” “spotting,”
and “pin-holing.” The tendency to bloom will always remain, even
after the varnish has hardened. But if any of these effects take place
in the varnishing room, while the varnish is drying, it will be fatal to
the appearance of the carriage.
To insure as near perfection as possible we want a substitute for
these objectionable driers, which will not be subject to atmospheric
influence.
CHAPTER XII.
ORNAMENTAL PAINTING.
monograms.
At the present time nearly all possessors of carriages have their
private marks painted on some part of the panels. These take the
form of monograms, initial letters, crests, and heraldic bearings or
coats of arms. The monogram is the commonest. For crests and
coats of arms a duty is levied, from which monograms are free.
A few examples are subjoined. They can be multiplied to any
extent; and designing monograms and initial letters would be
excellent practice for the apprentice.
Fig. 33.—Lay in C with dark blue, light blue,
and chrome yellow, No. 2; lighted with A to be
in Tuscan red, lighted with vermilion and
orange; V with olive green, lighted with a
bright tint of olive green and white. Separate
the letters with a wash of asphaltum.
Fig. 34.—Paint C a tan colour shaded with
burnt sienna, shaded with asphaltum to form
the darkest shades. Put in the high lights with
Fig. 33.—V. A. C.
white toned with burnt sienna. Colour I with
dark and light shades of purple, lighted with
pale orange; N to be lake colour lighted with vermilion. The above
may be varied by painting the upper half of the letters with the
colours named, and the lower portions in dark tints of the same
colour. When this is done, care must be taken
to blend the two shades, otherwise it will look
as if the letters are cut in two.
Fig. 35.—Paint the
upper half of O a light
olive green, and the
lower half a darker
tone of the same
colour; T to be lake,
lighted with vermilion
Fig. 34.—I. N. C.
above the division
made by the letter S,
no high lighting to be used on the bottom
portion of the stem; S to be painted red
Fig. 35.—O. T. S. brown, lighted with orange; or the colours
may be laid on in gold leaf, and the above
colours glazed over it.
Fig. 36.—This combination forms a
pleasing variety, and will afford good
practice in the use of the pencil. Lay in
the letters as indicated by the shading,
the letter V to be darker than A, and T
deeper in tone than either V or A. The
letters may all be laid on with gold leaf,
and afterwards glazed with colours to
suit the painter’s taste. The vine at the
base may be a delicate green tinged
with carmine.
Fig. 37.—This is of French design. Fig. 36.—V. A. T.
The letters furnish an odd yet attractive
style. It will be noticed that the stem of
the letter T covers the centre perpendicularly, and that the outer
lower portions of A and R are drawn to touch on the same line. The
main stems of these letters terminate in twin forms, arranged so as
to cross each other at the centre of the monogram and balance each
other on either side. In the matter of its colouring, it may be
mentioned that the letters in a monogram are
very often painted all in one colour, and
separated at the edges by a streak of white or
high light. Monograms painted in this manner
should be drawn so that the design will not be
confused by ornamentation; that is, the main
outlines of each letter should be distinctly
defined, and the spaces must be so arranged
as not to confuse the outlines. The pattern
here given may be coloured carmine, and the
Fig. 37.—A. R. T.
edges separated by straw colour or blue, and
the letters be defined by canary colour, or a
lighter tint of blue than the bodies of the letters are painted.
Fig. 38.—If the ground colour of the panels
is claret or purple the letters may be painted
with the same colour, lightened up with
vermilion and white, forming three distinct
tints; on brown, coat the letters with lighter
shades of brown; and so on with other
colours.
Fig. 38.—T. O. M.
Initial Letters.
A well painted initial letter is certainly quite equal to a monogram;
but then it must be well painted, because, as it stands alone, it has
only itself to rely upon for any effect, whereas, in a monogram, the
component letters mutually assist each other.
Fig. 39.—This letter possesses all the grace of outline that could
be desired in a single letter. Paint the letter in gold, shaded with
asphaltum and lighted with white. If a colour be used, have one that
agrees in tone with the striping on the carriage part; that is to say, if
blue be used in striping, then use the same kind of blue for the letter,
and so on with other colours.
We may here mention that all this kind of
painting is done on the last rubbing coat of
varnish, so that the letters receive a coat of
varnish when the finishing coat is given.
Fig. 40.—The
natural form of this
letter is graceful,
being composed of
curves bearing in
Fig. 39.—D. opposite directions,
and which blend into
each other, forming a continuous but varied
line. The ornamentation also falls into the
shape of the letter naturally. The upper and
lower ends of the letter terminate in three
stems, covered by three-lobed leafing, and
the main stem of the letter is preserved in Fig. 40.—S.
shape by appearing to grow out naturally from
its outer and inner edges.
Lay in the letter with gold, on which work out the design with
transparent colours. If colours only be employed the panel colour
may be taken as part of the colouring of the letter; for instance, if the
panel be dark brown, lake, blue, or green, mix up lighter tints of
whichever colour it may be, and considering the panel colour as the
darkest shade, lighten up from it.
Fig. 41.—This letter will please by the novelty of its ornamentation.
The body of the letter retains its natural outline almost wholly. From
the upper part of the thin stem springs a scroll, which curves
downward, reaching to the middle of the letter, and from this grows
out a second scroll, serving to ornament the lower portions.
Lay in the colour in harmony with the striping colour, deepening
the tone of the colour on the stem of the letter, as shown by the
shade lines. The leafing should be made out with light, medium, and
dark tints, blended into each other so as to avoid the scratchy
appearance which an opposite method produces.
Fig. 41.—V.
Crests and Heraldic Bearings.

It would be impossible to give anything like a comprehensive
series of these in this, or indeed in a very much larger work, as their
number and variety are so great. The examples subjoined are given
as exercises in colouring; and, if the student desires to extend his
studies in this direction, most stationers will supply him with sheets
of them at a trifling cost, and to them he may apply the principles
enumerated below.
Fig. 42.—This is a small ornament, but it
will disclose to the painter whether he has got
hold of the method of handling the “cutting-up
pencil.” If, in attempting the circular part, the
hand becomes inclined to be unsteady in its
motion, and create a lack of confidence, the
painter should practise until assured that the
hand will obey the will.
The ornamental part to be gold, shaded
Fig. 42.
with asphaltum, and high light with a delicate
pink, composed of flake white and light red.
The wreaths may be painted blue and white. Mix up three lines of
blue, placing the darkest at the bottom or lower part of each band
shown, as shaded in the figure. The white bands should not be of
pure white, but a light grey, made by mixing a little black with the
white colour. For the high light running along the centre of the
wreath, use white tinted with yellow. The space covered with
diagonal lines may either be left plain, showing the panel colour, or
barred across with grey lines made of flake white and black, tinged
with carmine.
Fig. 43.—This is the letter V combined
with a garter. Size in the entire pattern,
and lay the pattern in with gold, and glaze
over the inner part of the garter with a light
blue, the inner and outer edges to remain
gold. The flying ribbon to be pink,
composed of carmine and white, and the
shading to be clear carmine, with carmine
saddened with black for the deeper tones.
The stems of the letter V to be green,
shaded with a reddish brown, and the Fig. 43.
leafing to be the same colours.
Fig. 44.—Paint the cap crimson, the wreath green and grey,
lighted with a delicate pink. The circular part to be gold, shaded with
asphaltum tinted with carmine; the outside border of shield to be gold
also; the upper division of the shield to be red, deep and rich in tone.
The chevron, or white angular band across the shield, to be a grey,
lighted up with pure white. The lower division of the shield to be blue,
and the deep shades to be purple. Paint the leafing at the base with
a colour mixed of burnt umber, yellow, and lake; shade with
asphaltum tinted with carmine, and put in the high lights with orange
or vermilion.
Fig. 45.—This is from a design by Gustave Doré. It is an odd but
still pretty design. Lay in the whole of the pattern in gold; shade the
details with verdigris darkened with asphaltum; put in the high lights
with pink, composed of light red and white. The escutcheon may be
coloured with light brown, carmine, and dark brown. The edges of
the diagonal bar to be dotted minutely with vermilion.
Fig. 45.
Fig. 46.—Outline the garter

with gold; the buckle and slide to
be gold also. Fill in the garter
with light and dark tints of blue,
and put in the high lights with
canary colour. Paint the floral
Fig. 44. gorgons in brown shades, and
light with orange and clear
yellow. A small portion of lake added to
these browns will cause them to bear
out richly when varnished. Let the
medium lights and shades
predominate, and the high lights
added, first carefully considering their
true positions, and then touching them
in with sharp strokes of the pencil,
which will give life and “go” to the
details. The pendent stems with leaves
and berries may be coloured olive
green, and shaded with russet. When
Fig. 46. the painting of this ornament is dry it
will be considerably improved by
glazing.
Fig. 47.—The central pattern is Caduceus, a Roman emblem. On
the rod or centre staff the wings are represented “displayed,” and the
two serpents turning round it signify
power, the wings fleetness, and the
serpents wisdom.
This pattern would look well in
gold, with the dark parts shaded
with black to the depths shown on
the sketch; the lighter tones being
greys, warm in tone. The serpents
may be put in with carmine, as also
the wings and head, and the rod
carmine deepened with black.
Various treatments of colouring
may be applied to this pattern, and
thinking out some of these will be
very good exercise for the ingenuity
of the painter. Fig. 47.
Fig. 48.—
Put the pattern in in gold, separating the parts
where necessary with shadow lines, and
produce the effect of interlacing by a judicious
use of high light lines and deep black lines.
The best pencil suited to this class of
ornament is a “cutting-up” pencil an inch long.
Having traced the pattern on the panel,
commence by painting the crest, and next the
main upper left-hand division of the scroll
part, paying no attention to the leafing or
minor details. It will be noticed that the centre
Fig. 48. line of the heavy leafing is a part of the scroll
line, which passes from the wreath or ribbon
at the top, and is completed at the base; so
that to secure easy curves this line should be laid in through its
whole length, and the leafing or any minor dividing lines be governed
by it. Next lay in the other half of the pattern in the same manner,
and having secured these main curves the subordinate details may
be added.
Where two fine lines cross each other, the effect of one line
passing underneath the other may be produced by simply lighting
one of the lines across the intersection, which by contrast will make
the gold or colour of the other line appear darker, and as though the
lighted line passed over it and cast a shadow.
Paint the wreath blue and white, the crest to be merely lighted with
the colour used for high lighting the other parts.
Fig. 49.—This consists of a species of
dragon, having the head, neck, and wings of
a bird, and the body of a wild beast. He
supports a Norman shield, the “fess” or centre
part displaying a Maltese cross.
In painting this ornament, first get a correct
outline of the whole; then mix up two or three
tints of the colour you design painting it,
having a pencil for each, and a clean pencil Fig. 49.
for blending the edges, so that no hard lines
may appear at the junction of the different
colours. Lay on the shaded portions first, then the half lights, keeping
them subdued in tone, so as to allow for the finishing touches
showing clear and distinct.
On a claret-colour panel the whole may be painted in different
hues of purple and red. On a dark blue panel, varying shades of blue
lighter than the groundwork, and so with other colours. The shaded
portion must be distinct, and gradually connected with the lighter
portions by light tints of the shading colour.
Or the dragon may be painted grey, the high lights with the same
colour warmed up with yellow; the outline of the shield in gold; the
upper division, a light cobalt blue; the lower division, a pale orange;
the cross, brown, shaded with asphaltum; the wreath, blue and
white; and the flying ribbon and leafing in gold.
CHAPTER XIII.
LINING AND TRIMMING.

This is a department which requires great taste as well as skill. The
interior of a carriage should be lined with cloth and silk, or cloth and
morocco, with laces specially manufactured for the purpose. The
colours should correspond to or harmonise with the painting. Light
drab, or fawn colour, used to be a very general colour for the linings
of close carriages, such as broughams, because they at once
afforded relief to and harmonised with any dark colour that might
have been selected for the painting. But a severe simplicity of taste
has prevailed of late years in this country, and the linings of the
carriages have been made mostly dark in colour to correspond to the
colour of the painting. This is often carried to such an extreme as to
present an appearance of sameness and tastelessness. It is no
uncommon thing, for instance, to see a brougham painted dark
green, striped with black lines, and lined with dark green cloth and
morocco, with plain laces to correspond. This to us appears to be
only one degree removed from a mourning coach, and it will be a
great pity if such a taste prevails. On the other hand, violent
contrasts outrage all principles of good taste. Morocco and cloth, or
silk and cloth, of the same colour as the paint may be used for the
linings, but, as the painting should be relieved by lines that
harmonise with it, so should the linings be relieved by the laces and
tufts, which are intended to give life and character to it.
Landau Back, Quarter, and Fall.

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Ebffiledoc - 823download Textbook Oceanography and Marine Biology An Annual Review First Edition S J Hawkins Ebook All Chapter PDF

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Oceanography and Marine Biology: An

Annual Review First Edition S. J.

Oceanography and Marine Biology An Annual Review Volume

Marine Biology: - Function, Biodiversity, Ecology 1st

International Review of Cell and Molecular Biology

Developmental Biology of the Sea Urchin and Other

Biology and Ecology of Pharmaceutical Marine Tunicates

Marine Organisms as Model Systems in Biology and

Investigating Oceanography Raphael Kudela

Advances in Marine Biology 74 1st Edition Barbara E.

No claim to original U.S. Government works

Printed on acid-free paper

International Standard Book Number-13: 978-1-138-19786-2 (Hardback)

Introduction to the Special Issue ix

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

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

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

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

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

A Review of Herbivore Effects on Seaweed Invasions 421

Author Index 441

Systematic Index 471

Subject Index 475

1Ocean and Earth Science, National Oceanography Centre

Citadel Hill, Plymouth, PL1 2PB, UK

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

FILLING IN THE GRAZING PUZZLE: A SYNTHESIS

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

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

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

Data collection and definition of terminology

Facultative herbivores (ontogenetic changes in diet)

Facultative herbivores (spatial and temporal shifts in diet)

Ontogenetic changes in diet

Spatial and seasonal changes in diet

Varnish is subject to various changes after having been applied to

Crests and Heraldic Bearings.

Fig. 46.—Outline the garter

LINING AND TRIMMING.

Landau Back, Quarter, and Fall.

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