Seagrass and Reef Program for
Port Phillip Bay: Temperate
Reefs Literature Review
No. 11
May 2010
Seagrass and Reef Program for Port
Phillip Bay: Temperate Reefs Literature
Review
Neil Hutchinson, Taylor Hunt and Liz Morris
May 2010
Fisheries Victoria
Department of Primary Industries
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Neil Hutchinson
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Preferred way to cite:
Hutchinson, N., Hunt, T., and Morris, L. (2010).
Seagrass and Reef Program for Port Phillip Bay:
Temperate Reefs Literature Review. Fisheries
Victoria Technical Report No. 11, 61 pages.
Department of Primary Industries, Queenscliff,
Victoria, Australia.
ISSN
1835-4785
ISBN
978-1-74264-141-6 (print)
Temperate Reefs literature review
ii
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Executive Summary
The purpose of this document is to review the
existing scientific understanding, in an
international context, of the temperate reef
habitats in Victoria, particularly in Port Phillip
Bay. A variety of research has been conducted on
intertidal, subtidal, deep subtidal and canyon
reefs in Victoria both within Port Phillip Bay and
along the Victorian coast.
While some clear differences in species
distributions and assemblage structure have been
identified from eastern to western Victoria, the
majority of information on reefs in Victoria
provides only a “snapshot” at one particular
place or time of year. Examination of temporal
and spatial variation in the structure of
assemblages on different reef types, and more
basic information on which species occur and
their basic ecology and behaviour are lacking,
especially for deep and canyon reefs.
More information is required to enable us to
understand what is shaping many of Victoria’s
reef communities. There is a need to conduct
research that examines Victoria’s marine
ecosystems in a broader geographical context
across southern Australia, and also in the context
of coastal oceanography.
The amount of research and level of
understanding has been greatest for intertidal
reefs, is lower for subtidal reefs, and is very
limited for deep and canyon reefs. This is
probably a reflection of ease of accessibility and
logistical constraints.
Research on subtidal reefs in Port Phillip Bay has
been fragmentary, and there is a poor
understanding of the drivers influencing the reef
communities and how these differ from the open
coast. Further research on the physiological and
ecological drivers affecting subtidal reefs in Port
Phillip Bay is required.
Of special interest are the deep reef and canyon
habitats of Port Phillip Heads. This area has a
unique combination of deep water, strong
currents, and coastal sedimentary processes. The
sessile invertebrate communities are well
developed but their uniqueness is uncertain.
Detailed studies of taxonomy and assemblage
structure are required along with research into
the physical and ecological processes affecting
the system.
Temperate reefs literature review
iii
Table of Contents
Executive Summary............................................................................................. iii
Introduction ............................................................................................................ 1
Temperate reefs ......................................................................................................................................................... 1
Key reef types in Victoria ........................................................................................................................................ 1
1. Intertidal .......................................................................................................................................................... 1
2.
Subtidal ............................................................................................................................................................ 1
3.
Deep: ................................................................................................................................................................ 1
4.
Canyon: ............................................................................................................................................................ 1
Biogeographic patterns across Victoria ............................................................. 2
Oceanographic and productivity context of Victorian reefs......................... 5
Central Victoria ......................................................................................................................................................... 5
Western Victoria ........................................................................................................................................................ 5
Eastern Victoria ......................................................................................................................................................... 5
Upwelling ................................................................................................................................................................... 5
Current monitoring ............................................................................................... 7
Monitoring ................................................................................................................................................................. 7
Marine Habitat Classification ................................................................................................................................. 7
Ecosystem services ................................................................................................ 8
Physical habitats and biological communities ................................................ 9
Intertidal reefs ........................................................................................................................................................... 9
Subtidal reefs ........................................................................................................................................................... 11
Exposed subtidal reefs ......................................................................................................................................... 11
Sheltered subtidal reefs ....................................................................................................................................... 12
Deep reefs ................................................................................................................................................................. 14
Invertebrates ......................................................................................................................................................... 14
Fish ......................................................................................................................................................................... 15
Canyon reefs ............................................................................................................................................................ 17
Port Phillip Heads ................................................................................................................................................ 17
Other canyons ....................................................................................................................................................... 17
Temperate reefs literature review
iv
Ecological and physical environmental drivers ............................................ 19
Intertidal reefs .......................................................................................................................................................... 19
Top-down / disturbance processes ..................................................................................................................... 20
Bottom-up / supply side processes ..................................................................................................................... 22
Competition ........................................................................................................................................................... 24
Subtidal reef ............................................................................................................................................................. 25
Physical drivers ..................................................................................................................................................... 25
Ecological drivers .................................................................................................................................................. 27
Major commercial fisheries.................................................................................................................................. 30
Deep reefs and canyons .......................................................................................................................................... 32
Physical drivers ..................................................................................................................................................... 32
Ecological drivers .................................................................................................................................................. 33
Knowledge gaps ....................................................................................................................................................... 34
Intertidal reefs ....................................................................................................................................................... 34
Subtidal reefs ......................................................................................................................................................... 34
Deep reefs and canyons ....................................................................................................................................... 35
References ............................................................................................................. 37
Appendix 1 Tables............................................................................................... 51
Appendix 2 Conceptual Models ....................................................................... 56
Temperate reefs literature review
v
List of Figures
Figure 1. Key relationships and drivers on intertidal reefs in Victoria............................................................. 57
Figure 2. Key relationships and drivers on subtidal reefs in Victoria............................................................... 58
Figure 3. Key relationships between invertebrates on subtidal reefs in eastern and western Victoria ........ 59
Figure 4. Key relationships and drivers on deep reefs in Victoria .................................................................... 60
Figure 5. Key relationships and drivers on canyon reefs in Victoria ................................................................ 61
List of Tables
Table 1. Six bioregions relevant to Victoria as presented in Interim Marine Coastal Regionalisation of
Australia Technical Group, IMCRA (2006) ................................................................................................... 51
Table 2. Interim intertidal marine habitat (MHC) categories for Victoria, Ferns et al. (2000). ....................... 52
Table 3. Interim shallow subtidal (0 – 2.5 m) marine habitat (MHC) categories for Victoria, Ferns et al.
(2000). ................................................................................................................................................................. 52
Table 4. Primary shallow habitat classification scheme as presented in Ball et al. (2000). ............................. 53
Table 5. Potential physical drivers responsible for shaping deep reef assemblages in Victoria. Summarised
from Edmunds et al. (2006b) ........................................................................................................................... 54
Table 6. Potential ecological drivers responsible for shaping deep reef assemblages in Victoria.
Summarised from Edmunds et al. (2006b) .................................................................................................... 55
Temperate reefs literature review
vi
Introduction
Temperate reefs
Key reef types in Victoria
Temperate reefs are defined here as hard
substrata in marine waters averaging
temperatures below 18C (The University of
Queensland 2001; Port of Melbourne
Corporation 2007c). They are widely distributed
in Australia, and extend around southern
Australia from Kalbarri in Western Australia to
Noosa in Queensland.
For the purposes of this literature review we
have grouped reefs in Victoria into four broad
categories:
In Victoria, temperate reef habitats cover
extensive areas of the coastline and are known
for their high biological complexity, species
diversity, species richness, level of endemism
and productivity (Keough and Butler 1996;
Edmunds et al. 2003).
These habitats are of social and cultural value to
people due to a variety of indigenous, aesthetic,
recreational and historical aspects (Keough et al.
1990; Edmunds et al. 2006a). They also hold
significant economic value by supporting two of
the most valuable commercial fisheries in
Australia, in abalone and rock lobster, plus other
valuable activities such as recreational fishing,
diving and a range of other tourism activities
(Environment Conservation Council 2000;
Edmunds et al. 2003; Department of Primary
Industries 2009b). But, despite the great
importance of temperate reefs, they are not well
known scientifically (Keough and Butler 1996).
The purpose of this literature review is to
critically evaluate the current understanding of
Victoria’s temperate reefs, with particular
reference to reef communities, ecological and
physical environmental drivers, dynamics, key
structuring and ecosystem services, key
relationships, and any key current relevant
research/monitoring programs. The review will
also construct conceptual models, summarizing
key relationships and provide an indication of
scientific uncertainty.
1. Intertidal
Reef that is periodically exposed to air at low
tide but is submerged or directly influenced by
sea water at high tide. Intertidal reefs are also
known as rocky shores.
2. Subtidal
Reef that is never exposed to the air from tidal
influences, generally covering depths of 2.5-20
m. The review will distinguish and discuss
differences in communities and processes on
subtidal reefs split into two categories based on
wave exposure and currents:
 Exposed subtidal reefs found on the
open coast and Port Phillip Heads
 Sheltered subtidal reefs found within
Port Phillip Bay.
3. Deep: Reef that is located 20 m or deeper.
4. Canyon: Deep reef located in canyons
(geomorphological formations). Particular
attention will be given to the Port Phillip
Heads deep reef canyon.
These categories were selected as they represent
reef types across Victoria. They distinguish reef
types based on their exposure to key physical
modifiers including depth, tidal range, energy
(wave and currents) and photic influences.
These factors have been shown to be
fundamental drivers of the distribution patterns
of benthic marine communities (O'Hara et al.
1999; Ferns et al. 2000; Ball et al. 2006; Coleman et
al. 2007).
Temperate reefs literature review
1
Biogeographic patterns across Victoria
Scientific research on temperate reef
communities in Victoria has, to some extent,
focused on the classification of marine waters
into bioregions. By examining qualitative
measures of biological or physical variables at
intertidal and subtidal sites across Victoria,
broad-scale biogeographic patterns have been
identified and subsequently supported in later
studies.
Early studies of intertidal and subtidal
temperate reef communities along the coast of
southern Australia categorised communities into
three distinct marine biogeographic provinces
(Bennett and Pope 1952; Dartnall 1974; Edgar
1984; Womersley and King 1990).
These provinces were:

The western, warm temperate, “Flindersian
Province”

The eastern, warm-temperate, “Peronian
Province”

The southern, cool-temperate, “Maugean
Province”
Edmunds et al. (2000) briefly reviewed these
studies and surmised that the coast of Victoria is
situated at the confluence of these provinces and
that the existing biological communities there
include a mixture of typically western, eastern
and southern species.
Typical western, Flindersian, species on
Victorian reefs include: the algae Caulerpa
brownii, Zonaria turneriana, Seirococcus axillaris,
Carpoglossum confluens, Cystophora monilifera,
Sargassum decipiens, Sargassum varians,
Sonderopelta coriacea and Melanthalia obtusata; sea
urchins including Holopneustes porossisimus and
Holopneustes inflatus; the abalone Haliotis
laevigata and Haliotis scalaris; and fish such as the
horse-shoe leatherjacket Meuschenia hippocrepis,
yellow-striped leatherjacket Meuschenia
flavolineata and Victorian scalyfin Parma victoriae.
Typical eastern, Peronian, species on Victorian
reefs include: the algae Phyllospora comosa; the
sea urchins Centrostephanus rodgersii and
Holopneustes purpurascens; the sea squirt Pyura
stolonifera; and fish including eastern hulafish
Trachinops taeniolatus, silver sweep Scorpis
lineolatus, black drummer Girella elevate, white
Temperate reefs literature review
2
ear Parma microlepis and mado Atypichthys
strigatus.
Prominent southern, Maugean, species on
Victorian reefs include: algae such as the string
kelp Macrocystis angustifolia, bull kelp Durvillaea
potatorum, Splachnidium rugosum, Zonaria
angustata and Cystophora torulosa; the sea stars
Patiriella brevispina, Nectria ocellata and Fromia
polypora; and fish including southern hulafish
Trachinops caudimaculatus and southern sea carp
Aplodactylus arctidens.
In addition to provincial influences, Victorian
communities are also composed of species
distributed throughout all of southern Australia
including algae such as common kelp Ecklonia
radiata, Pterocladia capillacea, Phacelocarpus
peperocarpus, Haliptilon roseum, Sargassum sp. and
Cystophora sp.; the common sea urchin
Heliocidaris erythrogramma; and the eleven armed
sea star Coscinasterias muricata.
The high turnover of species in the central
Victorian area was reaffirmed in a study based
on examining echinoderm and crustacean
species distributions along the coast of southern
Australia by O'Hara (2000a). O'Hara (2000a)
analysed museum collections sampled from
within 62 spatial units between Western
Australia and northern New South Wales. The
study determined that 49% of the total species in
southern Australia (739) occur in Victoria (362)
and the species distributions and turnover were
consistent with previous biogeographical
classifications (Bennett and Pope 1952; Edgar
1984; Womersley and King 1990).
Given the fact that these different regions meet
in Victoria, allowing comparative work to be
carried out at similar latitudes within a
relatively small geographic area, surprisingly
little work has been conducted on the ecology of
rocky reefs in the State.
Waters et al. (in press) recently provided an a
priori test for the existence of Maugean,
Flindersian and Peronian biogeographical
provinces across southern Australia. The study
quantitatively analysed distributional data of
1,500 algal species and identified the three
distinct biogeographical assemblages, consistent
with previous biogeographical classifications
(Bennett and Pope 1952; Edgar 1984; Womersley
and King 1990). The authors recommended that
broad provinces be applied as a regional
framework for understanding and managing
Australia’s marine biodiversity, particularly for
integrating the ongoing discovery of biological
variation at finer scales. The authors support
their recommendation through (1) arguing
against the relevance of a biogeographical
classification of Australia’s coastline based on
physical variables (IMCRA 1998) and (2)
documented evidence for ecological variation
(aside from species composition) across
southern Australia.
The Interim Marine Coastal Regionalisation of
Australia Technical Group, IMCRA (1998), used
measurements of bathymetry, coastal
geomorphology, sediments, currents, tides,
water chemistry and water temperature to
classify Victoria into six bioregions (Table 1).
Waters et al. (in press) argue that the broad
biological relevance of these bioregions remain
unclear because they are derived from gross
biophysical features rather than based on large
numbers of taxa. The authors go on to question
the effectiveness of IMCRA (1998) studies in
defining regional biodiversity and shaping
conservation policy.
Evidence exists for ecological variation across
southern Australia aside from species
composition (Connell and Irving 2009). Connell
and Irving (2008) sought to provide a
biogeographical framework for the
interpretation of variation in the ecology of, and
threats to, subtidal rocky landscapes. The study
quantified the frequency and size of patches of
major benthic habitats across the southern coast
of Australia to establish biogeographical
patterns that may result from contrasting
regional-scale processes. Key findings from the
study include distinct biogeographical
patterning in the extent of kelp forests as related
to the Flindersian and Peronian provinces.
Flindersian coasts have higher kelp forest cover
and higher kelp forest heterogeneity relative to
Peronian coasts. The patchwork of forests in the
Peronian province is created by the foraging
activities of the black urchin Centrostephanus
rogersii, a species that is not recorded from the
Flindersian province. Distinct differences in
concentrations of nutrients, chlorophyll a, kelp
forest canopy-morphology, kelp forest
understorey associations and broad degree of
wave exposure between Flindersian and
Peronian coasts also support the finding of
ecological variation, aside from species
composition, across southern Australia (Connell
2007; Connell and Irving 2008; 2009; Waters et al.
in press).
Finer-scale bioregions in Victoria have also been
described in order to better understand marine
biodiversity at the ‘habitat’ and ‘community’
levels. Ferns et al. (2000) provided a review of
classifying Victoria’s marine ecosystems,
recognising the importance of both physical and
biological factors and the use of qualitative and
quantitative attributes. The authors
acknowledge previous classifications at the
broad-scale bioregion level, and then describe
two approaches in assessing and understanding
marine biological communities at the ‘habitat’
and ‘community’ levels.
The first approach is to classify intertidal and
subtidal marine ‘habitats’ using qualitative
attributes collected from remote sensing and
field survey techniques. Qualitative descriptions
of seafloor substratum and dominant biota are
generally arranged in a hierarchical system to
create Marine Habitat Classes (MHCs). MHCs are
applied over scales of 1 m – 100 km that have
distinct physical and/or biological ‘habitat’
attributes. MHC attributes were identified for
intertidal and subtidal areas by Ferns et al. (2000)
(Tables 2 and 3).
Edmunds et al. (2000) also provided a
quantitative classification of the biological
communities in Victoria. The classification was
derived from quantitative sampling of biological
communities of macrophytes,
macroinvertebrates and fish associated with
rocky reefs using visual census techniques.
These biological data provided attributes for
specifically defined Marine Ecological
Communities (MECs), which were applied to
areas at scales from 1 m – 1 km and consisted of
distinct biological communities. The results of
the study indicated that there was a
considerable overlap of the provincial
influences, particularly in the area of central
Victoria where the Flindersian components
extend as far east as Cape Liptrap, and the
Peronian components extend as far west as Cape
Otway. Ferns et al. (2000) state that MECs
potentially serve as useful indicators for
reporting on the long-term status of marine
communities.
Presently, MECs for macrophytes, invertebrates
and fish have been delineated for a selection of
subtidal reefs across four areas of coastline in
Victoria including Port Phillip Heads, Phillip
Island, Bunurong and Wilsons Promontory
(Ferns et al. 2000). But due to the lack of similar
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3
studies determining MECs along the remainder
of the Victorian coast, local temperate reef
community information is scarce. This is,
however, gradually being addressed.
In recent years, a number of studies have
examined Victoria’s subtidal reefs using
combinations of georeferenced aerial and
satellite images, echo-sounders, GIS and
ground-truthed video drops/transects and diver
surveys. For example, the “Victorian Marine
Temperate reefs literature review
4
Habitat Mapping Project” and other projects are
covering reefs along the majority of Victoria’s
coastline (Bax and Williams 2000; Bax and
Williams 2001; Beaman et al. 2005; Ball and Blake
2007a; b; Holmes et al. 2007; Monk et al. 2008;
Blake et al. 2009a; b; Rattray et al. 2009). These
surveys have described a wide range of reefs
along the Victorian coast at the habitat level and
broadly describe the make-up and distribution
of biota in relation to substrate type at the
community level.
Oceanographic and productivity context
of Victorian reefs
Central Victoria
The oceanography of the central area of Victoria
is dominated by the relatively shallow waters of
Bass Strait. The Strait consists of a shallow
platform, mostly about 70 m below sea level,
flanked by 4-5 km deep ocean to the east and
west and by land to the north and south.
Although tidal currents can be strong in certain
areas, such as the eastern and western entrances
to Bass Strait (Black 1992), the net tidal
circulation tends to be small. In central Victoria,
the local tidal dynamics are strongly influenced
by the two large bays, Port Phillip Bay and
Western Port. The primary determinants of net
water movement in the region are wind-driven
currents and coastal trapped waves (Middleton
and Black 1994). The density structure of Bass
Strait ranges from well mixed in winter to
strongly stratified in central areas in summer
(Baines and Fandry 1983). Bass Strait is generally
considered to have a low nutrient status because
nutrient rich water from the deep ocean is
generally absent or diluted (ASR 2008).
Port Phillip Bay is a large (2000 km2), semienclosed, predominantly tidal embayment
linked to the ocean of Bass Strait by a narrow
entrance. The hydrodynamics are characterised
by an entrance region where fast (3 m s-1) ebb
and flood jets dominate the circulation, a large
flood-tidal delta known as the Sands region,
where strong currents occur in the major
channels, and an ‘inner’ zone where tidal flows
are weak and circulation is predominantly by
wind-driven currents (Black et al. 1993). Tides
are semidiurnal and the amplitude inside the
bay is less than 1 m. Salinity in Port Phillip Bay
is essentially marine and recently has become
hypersaline in comparison with Bass Strait
(Nicholson and Longmore 2009). Although there
is significant nutrient input to the Bay from
sewage treatment, effects of eutrophication are
presently only seen in localised areas.
Western Victoria
The oceanography of western Victoria is
influenced by the major western boundary
current: the Leeuwin current. The winter
circulation in Victoria is characterised by a west
to east coastal current (Cirano and Middleton
2004). The eastward coastal current is partly
driven by the Leeuwin current to the west
(Cirano and Middleton 2004). The Leeuwin
current is a warm water current, and the
significant inter-annual variation that has been
recorded in the eastern coastal current (Cirano
and Middleton 2004) would be likely to lead to
inter-annual variation in water temperature to
the west of Bass Strait. The Leeuwin current,
and by extension the eastern coastal current, is a
warm, low-salinity current that is considered to
be nutrient poor (Connell 2007; Suthers and
Waite 2007).
Eastern Victoria
Eastern Victoria is influenced by the major
eastern boundary current: the East Australian
Current (EAC) (Suthers and Waite 2007; ASR
2008). The EAC often generates warm-core
eddies, or circular flows, up to several hundred
kilometres across. These eddies drift south and
can be found off the central east coast of
Tasmania (Suthers and Waite 2008; ASR 2008).
Although most of the water from the EAC is
directed into the Tasman Sea, some EAC water
also enters eastern Victoria, especially over
summer. The EAC is characterised by upwelling
in a number of areas (Suthers and Waite 2007)
and is generally considered more nutrient rich
and productive than the Leeuwin current
(Connell 2007).
Upwelling
As well as the influence of broad-scale currents
and their associated nutrient regimes, local-scale
areas of upwelling, with associated nutrient and
productivity increases, can occur though-out
Victoria when wind and oceanographic
conditions are suitable. Upwelling is an
oceanographic phenomenon where surface
winds along coastal areas drive dense, cool, and
nutrient-rich water towards the surface.
Upwelling has been recorded in eastern Victoria
(Gibbs et al. 1986; Rochford 1977; Butler et al.
2002b) and to a lesser extent in central areas of
Bass Strait (ASR 2008). By far the strongest area
Temperate reefs literature review
5
of upwelling and nutrient enhancement in
Victoria occurs in the west. The Bonney
upwelling is the most prominent upwelling in
southeast Australia and is driven by the
prevailing south-easterly winds in the region.
Between November/December and March/April,
upwelling plumes are regularly observed along
the Bonney Coast from Robe, SA to Portland,
Vic which make the area highly productive
(Butler et al. 2002b), and all reefs in the area may
be influenced to some degree by this
phenomenon.
The Bonney Coast is within the Otway region as
classified by the IMCRA (Butler et al. 2002b) and
was described by Womersley (1984) as being
part of the Maugean province, of which it forms
the western extension (Butler et al. 2002b).
Temperate reefs literature review
6
Current monitoring
Monitoring
The Victorian Subtidal Reef Monitoring
Program (SRMP) and Intertidal Reef Monitoring
Program (IRMP) were established by the
Victorian Government to provide information
on the status of Victorian reef flora and fauna
(focussing on macroalgae, macroinvertebrates
and fish). The SRMP uses standardised
underwater visual census methods based on an
approach developed and applied in Tasmania
by Edgar and Barrett (1997). This includes
monitoring the nature and magnitude of trends
in species abundances, species diversity and
community structure through time, with
particular emphasis on Marine National Parks
and Sanctuaries in Victoria (Edmunds et al.
2003).
The SRMP was initiated in May 1998 with 15
sites established on subtidal reef habitats in the
vicinity of Port Phillip Heads Marine National
Park. Since then, the SRMP has been expanded
to reefs in the vicinity of the Bunurong Marine
National Park (12 sites), Phillip Island (6 sites),
and Wilsons Promontory Marine National Park
(20 sites) and a further twelve Marine National
Parks and Marine Sanctuaries including: Point
Cooke, Jawbone, Ricketts Point, Merri, Marengo
Reef, Eagle Rock, Beware Reef and The Arches
Marine Sanctuaries and Point Addis, Cape
Howe, Point Hicks and Twelve Apostles Marine
National Parks. This program is currently
implemented by Parks Victoria, in association
with the Department of Sustainability and
Environment (Edmunds et al. 2006a). The data
include quantitative estimates of large mobile
fishes, cryptic fishes, megafaunal invertebrates
and macroalgal species.
The IRMP was initiated in April 2003 with 14
sites established on intertidal reef habitats
within, and in the vicinity of, the following
marine protected areas: Point Addis Marine
National Park and Point Danger, Barwon Heads,
Point Cooke, Jawbone, Ricketts Point and
Mushroom Reef Marine Sanctuaries.
Although the monitoring program will begin to
adequately reflect average trends and patterns
as the surveys continue over longer periods such
as multiple years to decades (Edmunds et al.
2003), Ball et al. (2006) state that macrophyte
communities identified in the study have been
generally compatible with dominant reef biota
MHC categories.
The Abalone Assessment Monitoring program
run by Department of Primary Industries (DPI)
is an annual fishery independent survey across
250 subtidal reef sites in Victoria that
commenced in 2002. The data are collected by
underwater visual census and includes
quantitative estimates of abundance of abalone
and prevalent organisms other than abalone
such as macroalgae, predators and competitors
and abiotic features.
Marine Habitat Classification
Since the Victorian interim marine habitat
classification scheme presented in Ferns and
Hough (2000) there has been several steps
forward in improving marine habitat
classification not only on a temperate reef level
but on a nationwide marine habitat level. These
advancements will allow future studies to
further our understanding of the distribution
and abundance of different temperate reef
habitats in Victoria.
Ball et al. (2006) reviewed and documented the
relationship between relevant Australian and
international local-level marine habitat
classification systems and the Victorian interim
marine habitat classification scheme presented
in Ferns and Hough (2000). The authors then
presented a revised marine habitat classification
system to support mapping of shallow (assumed
0 - 20 m depth) habitats at Victoria’s Marine
National Parks and Sanctuaries. The primary
habitat classification scheme divides
classification into five levels of modifiers to
classify habitats mapped from underwater
(Table 3). The purpose of this habitat mapping
classification is to assist in the selection and
evaluation of candidate representative marine
protected area locations, to allow more accurate
description of the spatial extent and distribution
of shallow seabed habitats, and to provide more
detailed biological information on marine
protected areas to support assessment of
management performance.
Temperate reefs literature review
7
Ecosystem services
Ecosystem services are the direct and indirect
benefits supplied to human societies by natural
ecosystems (Daily et al. 1997) including, for
example, key fishery species in the reef
community, or productivity values associated
with the reef.
Temperate reefs are known for their high
biological complexity, species diversity, species
richness, level of endemism and productivity
(Keough and Butler 1996; Edmunds et al. 2003).
Temperate reef habitats have social and cultural
values including indigenous, aesthetic,
recreational and historical aspects (Keough et al.
1990; Edmunds et al. 2006a). They also hold
significant economic value by supporting two of
the most valuable commercial fisheries in
Australia, in abalone and rock lobster, plus other
valuable activities such as recreational fishing,
diving and other tourism activities
(Environment Conservation Council 2000;
Edmunds et al. 2003; Department of Primary
Industries 2009b). Keough and Butler (1996) add
that the temperate reefs in Australia are
important for commercial and recreational
fisheries and for recreational diving, and they
are potential sources of natural products for the
pharmaceutical industry
The subtidal and deep reefs (including the
canyon) throughout Port Phillip Bay are
described as ecological assets of Port Phillip Bay
Temperate reefs literature review
8
(Port of Melbourne Corporation 2007a; b;
Connell and Irving 2009). The south of the Bay,
together with the Entrance, is popular for
recreational diving because the water is
generally clear and the area contains high
quality diving sites associated with the deep
canyon in the Entrance, marine national parks
and shipwrecks. Key dive sites across the south
of the Bay include Lonsdale Wall, which extends
to depths of 45 m, and Popes Eye, which is a
renowned location for trainee divers, snorkellers
and for bird watching and general sightseeing
(Port of Melbourne Corporation 2007a).
An example of the importance of reefs in Port
Phillip Bay in terms of ecosystem services is the
recent State Government initiative to create
artificial reefs for recreational angling in the Bay.
This indicates that natural reefs are considered
to be an important and limited resource (P.
Hamer, pers comm.)
Other examples of the ecosystem services
provided by reefs in Port Phillip Bay and
elsewhere include protection from beach erosion
by acting as a wave break, and also associated
algae act as a nutrient sink and site of detritus
production that underpins the detrital food
chain in soft bottom habitats.
Physical habitats and biological
communities
Intertidal reefs
Intertidal reefs vary in structure from steep
sloping rock faces to relatively flat or gently
sloping boulder fields and rock platforms,
with a variety of features including cobble
fields, vertical steps, undulations in the reef,
crevices, patches of sand and rock pools
caused by a variety of processes such as
weathering (Edmunds et al. 2004).
Bennett and Pope (1952) surveyed 16 sites on
exposed intertidal reefs along the coast of
Victoria and in spite of differences in localities,
five major zones and subsequent community
assemblages were identified:

Melaraphe-lichen zone: species present
include herbivorous molluscs (Nodilittorina
spp., previously called Melaraphe spp.),
lichens, crustaceans such as small crabs,
amphipods and isopods, and occasionally
predatory gastropods.

Barnacle-mussel zone: species present
include barnacles and mussels.

The mixed algal turf or Galeolari zone:
species present include mixed algae in
exposed areas with the serpulid Galeolaria
caespitosa in sheltered areas.

The Poneroplax or “Bare” zone: species
vary along the coast and with wave action.
Chitons of the genus Poneroplax are
always present, and on many shores east
of Cape Otway ascidians are found.

The large brown algae zone: species
present include large fucoid kelp algae.
Such patterns of vertical distributions in
organisms are well chronicled on rocky shores
internationally, for example bare zones at the
mid shore level (Mettam 1994; Kaehler and
Williams 1997).
Zonation of algae in Port Phillip Bay was
briefly reviewed by Spencer (1970), and has
been examined on basalt reefs (King et al. 1971;
O'Brien 1975; Brown et al. 1980). Clear vertical
zones were apparent on reefs in the Bay and
while species present varied to some extent
between sites, some of them were commonly
found. For example:

Upper eulittoral zone: species include
Enteromorpha spp.

Mid eulittoral zone: species include
Porphyra spp, Ulva spp, Gelidium pusillum

Lower eulittoral zone: species include
Codium sp., Caulerpa sp., Polysiphonia sp.
More recent studies (O'Hara et al. in press)
have collected data on 65 intertidal reefs across
Victoria as part of an effort to detect impacts
on assemblages, and there are ongoing
qualitative surveys being conducted at Barwon
Heads by local school children.
Several recent monitoring studies of Victoria’s
Marine Protected Areas have described the
dominant biota present on Victorian temperate
reefs. One of the most common algal species
on intertidal reefs is the perennial fucoid alga
Neptune’s necklace, Hormosira banksii, which
can form extensive monotypic stands at midtidal levels of rock platforms (Keough and
Quinn 1998). Other common algae species that
can form mats with or without the presence of
H. banksii include the green algae Ulva spp.
and Enteromorpha spp, coralline algae and
filamentous brown algal turfs (Edmunds et al.
2004). Less conspicuous is a thin layer of
microscopic algae growing directly on the
surface of the reef, which is an important food
source for species of grazing molluscs
(Edmunds et al. 2004; Gilmour and Edmunds
2007; Stewart et al. 2007).
The invertebrate fauna on intertidal reefs,
which tend to exhibit patterns of zonation in
the same way as shown by algae (King et al.
1971), are dominated by herbivorous and
predatory molluscs. Common herbivorous
species on Victoria’s shores include the top
shell Austrocochlea porcata, the variegated
limpet Cellana tramoserica and conniwinks
Bembicium spp. Less common species include
the warrener Turbo undulatus, the black nerite
Nerita atramentosa and the predatory
Temperate reefs literature review
9
gastropods Cominella lineolata and Lepsiella
vinosa (Edmunds et al. 2004; Gilmour and
Edmunds 2007).
Sessile invertebrate species create encrusting
mats on the surface of the reef such as the
mussels Xenostrobus pulex and Brachidontes
rostratus and tubeworms Galeolaria caespitose,
whilst there are a variety of other mobile
invertebrates such as small crabs and fish
species that are either present in the intertidal
zone throughout the tidal cycle and refuge in
rockpools, or move into the area at high tide
(Edmunds et al. 2004; Gilmour and Edmunds
2007; Stewart et al. 2007).
As is common on shores elsewhere, both in
Australia and worldwide (Little et al. 2009),
differences exist between biota present on
exposed intertidal reefs along the Victorian
coastline and sheltered intertidal reefs within
Port Phillip Bay. Gilmour and Edmunds (2007)
surveyed intertidal reef biota of Central
Victoria’s Marine Protected Areas and found
species richness of algae and invertebrates, and
algal cover, to be higher on the exposed
intertidal reefs along the coastline than on the
sheltered intertidal reefs inside Port Phillip
Bay. Stewart et al. (2007) reaffirmed their
findings and also found that mat forming
mussels were components of more intertidal
reef communities outside Port Phillip Bay than
inside the Bay.
Intertidal reefs in Victoria are known to show
the same general patterns as those in eastern
Australia (Bennett and Pope 1952).
Underwood and Kennelly (1990) stated that
“kelps are represented by Durvillaea potatorum,
Ecklonia radiata and Phyllospora comosa;
Macrocystis angustifolium is present in some
places. Cystophora intermedia are found in
eastern Victoria and reappear in South
Australia. Tunicates, such as the cunjevoi
Pyura stolonifera, are only found in some places
in Victoria and low-shore regions are occupied
by large chitons and encrusting calcareous
algae. Above the regions dominated by algae
are areas occupied by barnacles. Conspicuous
amongst these barnacles are bands of foliose
algae (e.g. Splachnidium rugosum and Ileafascia
in summer).”
Temperate reefs literature review
10
Several authors have investigated specific
groups of species in Victoria. For example, a
review by Sanderson (1997) described algal
communities in more depth and Light (1992)
reviewed literature on the benthic flora of Port
Phillip Bay, which included information from
both soft and hard substrates.
Surveys of Port Phillip Bay in the 1950-60s
described 173 species of subtidal algae, which
included numerous species attached to hard
reefs, particularly in the Port Phillip Heads
region (Womersley 1966). Womersley (1966)
noted that species richness was lower inside
than outside the Bay and suggested that this
may be due to a variety of factors including
differences in water depth and temperature.
He also mentioned the comparable lack of
rocky reefs within Port Phillip Bay. Hope Black
(1971) used this data to examine distributions
with substrate, in particular concentrating on
the importance of reefs in Port Phillip Bay as
habitat and summarised the distribution of
reefs in Port Phillip Bay as 4 main groups in
respect to rock type:
1.
Dune limestone: in and adjacent to Port
Phillip Heads, including Popes Eye
(artificial basalt structure)
2.
Oligocene basalt: from Corio Bay to
Williamstown
3.
Tertiary ironstone of the Miocene clays
and sandstones: north and east shores
4.
Granites: extending from Martha Pont
seawards. Form off-shore reefs.
Light and Woelkerling (1992) summarised
descriptions previously made of algae in the
intertidal and subtidal zones of Corio Bay
(Womersley 1966; Hope Black 1971), the
Werribee region (Womersley 1966; Spencer
1970; Hope Black 1971; Brown et al. 1980),
Altona Bay – Hobsons Bay (Spencer 1970),
Carrum and Portsea, and discussed
unpublished data for Werribee (Brown et al.
1980), Gloucester Reserve (O'Brien 1975),
Williamstown and Portarlington (Lewis 1975).
Subtidal reefs
Subtidal reefs are defined here as reef that is
never exposed to the air from tidal influences
and generally covering depths of 2.5-20 metres.
They can be split into two categories:
1.
Exposed subtidal reefs
2.
Sheltered subtidal reefs
Exposed subtidal reefs are physically
characterised by a high degree of exposure to
wave and current energy.
Exposed subtidal reefs exist along the majority
of the Victorian coastline and include the reefs at
the entrance of Port Phillip Bay, due to their
high degree of exposure to ocean swells and
tidal energy (McShane et al. 1986; Port of
Melbourne Corporation 2007a). Certain taxa
have been described in some detail off the coast
of Victoria yet gaps still remain in our
knowledge. For example, a review of
opisthobranch molluscs (Burn 2006) in Victoria
described the range of species that occur in
habitats such as the rocky intertidal zone and
subtidal rocky reefs from Cape Otway to
Wilsons Promontory, but noted that little is
known of species in shallow waters off eastern
Victoria (Wilsons Promontory to Cape Howe).
Algae
In Victoria, subtidal reef communities are
typified by their prominent biota of algae
(Keough and Butler 1996; Edmunds et al. 2006a;
Connell 2007). Sanderson (1997) described how
reefs show broad patterns of zonation along the
depth gradient: Hormosira banksii is the
dominant species observed on intertidal rock
platforms; Durvillaea potatorum with mixed
brown and green algae inhabits the seaward
edge of intertidal rock platforms; Phyllospora
comosa occurs in large beds at depths of ~3–5 m;
at exposed sites, Ecklonia radiata is typically
observed at depths >7 m and often growing with
P. comosa; and giant kelp Macrocystis angustifolia
occurs at Port Phillip Heads MNP - Point
Lonsdale and at Point Nepean.
Other research on subtidal algae on exposed
subtidal reefs in Victoria is thoroughly reviewed
by Light and Woelkerling (1992) and Sanderson
(1997). These also include reports on subtidal
reef algae at Black Rock between Ocean Grove
and Torquay (Rollings et al. 1993; Chidgey and
Marshall 1994) and other areas throughout
Victoria and the region (Cheshire and Hallum
1989; Land Conservation Council 1993).
On exposed subtidal reefs at the entrance to Port
Phillip Bay, high vertically-structured brown
algae (kelps) such as Macrocystis angustifolia,
Ecklonia radiata and Phyllospora comosa are often
dominant. The green alga Caulerpa is also
usually present plus a wide variety of low
structured, understorey red algae species
including Plocamium, Pterocladia, Melanthalia and
Coralina (Port of Melbourne Corporation 2007b).
Vegetation has been surveyed at a number of
subtidal reef sites along the coast of Victoria
resulting in the identification of two distinct
vegetation groups together with the following
observations recorded by O'Hara (2000b; 2001):

Ecklonia/Phyllospora: Form dominant
canopies in many exposed open-coast
localities across Victoria. Phyllospora comosa
forms a dense canopy on high relief, shallow
(3 to 8 m), exposed or semi-exposed reefs
but is often absent from low-relief reef.
Ecklonia radiata often occurs in combination
with Phyllospora or in combination with
Macrocystis, Cystophora and other brown
algae when Phyllospora is absent and is the
dominant species on deeper areas of reef
(Ball et al. In prep).

Cystophora/Sargassum: A diverse mixed-algal
assemblage occurs in some Victorian
subtidal reefs where the usual
Phyllospora/Ecklonia canopy is absent. These
sites are visually characterised by the fucoid
algae Cystophora, Sargassum, Seirococcus and
Acrocarpia, with many other brown, red and
green algae interspersed (including Ecklonia
and Macrocystis).
Invertebrates
Exposed subtidal reefs in Victoria typically have
high abundances of large-mobile predatory and
grazing invertebrates (Edmunds et al. 2006a).
Predatory mobile invertebrates include: rock
lobster Jasus edwardsii, red bait crab Plagusia
chabrus, dogwhelk Dicanthais orbita, octopus
Octopus maorum and seastars Coscinasterias
muricata. Grazing mobile invertebrates include:
abalone Haliotis rubra and H. laevigata, warrener
Turbo undulatus and sea urchins Centrostephanus
rogersii, Heliocidaris erythrogramma, Holopneustes
and Amblypneustes (Edmunds et al. 2000;
Australian Government 2005; Port of Melbourne
Corporation 2007b; c).
The low abundance of the grazing urchin
Centrostephanus rodgersii on exposed reefs
Temperate reefs literature review
11
throughout the majority of Victoria results in
urchin barren habitat being restricted to the fareast where it is present in high abundance
(Keough and Butler 1996; O'Hara 2000b; Ferns
2003; Department of the Environment and
Heritage 2005). Several studies have been
conducted on the creation and maintenance of
urchin barrens elsewhere in southern Australia
and New Zealand (Andrew and MacDiarmid
1991; Andrew 1993; Andrew and O'Neill 2000).
In Victoria, urchin barrens have been described
at Cape Howe Marine National Park (Ball and
Blake 2007b). There are also various small
species of crustaceans and molluscs that occupy
niches as grazers, predators or foragers
(Edmunds et al. 2006a).
Sessile invertebrates present on exposed
subtidal reefs can be found in several body
forms:

Prominent colonial or modular sessile fauna
such as sponges, ascidians, soft corals,
hydroids and bryozoans are abundant on
subtidal reefs, usually in high abundances
within well-shaded rock faces, overhangs
and caves, and on the underside of boulders
(Keough and Butler 1996; Keough 1999)

Solitary or unitary forms of sessile fauna
include barnacles, some ascidians (eg.
Pyura), bivalve molluscs and polychaetes
(Keough and Butler 1996).
Fish
Fish are a dominant biotic component of
exposed subtidal reef ecosystems in Victoria
with wrasse (Labridae), morwong
(Cheilodactylidae) and leatherjacket
(Monacanthidae) making up the majority of
species (Edmunds et al. 2000; Edmunds et al.
2006a). Wrasse are a roaming predatory species
that feed on small molluscs and crustaceans,
while morwongs are herbivorous or omnivorous
and leather jackets are generally omnivorous or
carnivorous picker-type species (Edmunds et al.
2000). Exposed subtidal reef fish species also
commonly include herring cale Odax cyanomelas,
sea sweep and scaly fin (Port of Melbourne
Corporation 2007c).
Reef fish tend to include algavores, carnivores
and omnivorous species. Russell (1983) noted
that “most rocky reef fishes have broadly
generalised feeding habits, and foods taken
mainly reflect those organisms of suitable size
that are abundant and most readily available”.
Gillanders (1995) has shown that the diet of
rocky reef fish of different size classes can vary
Temperate reefs literature review
12
with depth and habitat, with juveniles feeding in
shallow areas of fringing habitat and adults in
deeper turf and barrens habitats, reflecting the
distribution of the species with depth.
Some reef-related fish species show regular
patterns of seasonal movement of adults on and
off reefs, such as Hypoplectrodes maccullochi at
Cape Banks NSW, possibly due to changes in
prey abundance or because adults move to other
areas to spawn (Webb and Kingsford 1992;
Holbrook et al. 1994)
Sheltered subtidal reefs are physically
characterised by a low degree of exposure to
wave and current energy.
Sheltered subtidal reefs occupy a relatively
small part of Port Phillip Bay, about 8 km2, or
less than 0.5%, of the seafloor (McShane et al.
1986; Port of Melbourne Corporation 2007c).
These reefs are generally within 1 km of shore in
shallow water depths less than 4 m (Port of
Melbourne Corporation 2007c). Port of
Melbourne Corporation (2007c) described the
distribution and lithology of Port Phillip Bay
reefs. Basalt reefs exist along the northern
shoreline at Williamstown extending west to
Point Lillias and sandstone reefs exist along the
north-eastern shoreline from Ricketts Point to
Point Ormond. There are also patches of reefs on
the south-eastern shoreline from Dromana to
Frankston and on the Bellarine Peninsula from
Clifton Springs to St Leonards (Port of
Melbourne Corporation 2007a).
Several artificial reefs exist within Port Phillip
Bay, including newly deployed reefs for
recreational fishing in the north of the Bay.
Previous studies such that carried out by Lewis
(1983) have noted that algae on artificial
structures in Port Phillip Bay usually includes
several introduced species.
Algae
Research on algae in Port Phillip Bay is
reviewed in Light and Woelkerling (1992) and
Sanderson (1997), and includes reports on
subtidal reef algae in: the Werribee region
(Brown et al. 1980); Williamstown, St Leonards,
Corio Bay (King et al. 1971; Lewis 1975); Altona
and Portarlington (Spencer 1972); and Hobsons
Bay (O'Brien 1975).
The surfaces of sheltered subtidal reefs in Port
Phillip Bay are typically patchier near the
exposed entrance, with various red and brown
algae and the green algae species Caulerpa and
Ulva, while other areas are characterised by the
kelp Ecklonia radiata or the introduced species
Undaria pinnatifida (Campbell 1999; Port of
Melbourne Corporation 2007c).
macroalgal cover, and possibly the effects of
spear fishing (Jenkins et al. 1996)
Invertebrates
Research on invertebrates on subtidal reefs
within Port Phillip Bay is sparse, although some
research is available that has examined
relationships between invertebrates in the bay,
such as studies by Day et al. (1995) and McShane
and Smith (1986) on starfish predation.
Fish often display complex relationships with
their surroundings. A whole suite of factors are
known to affect temperate reef fish assemblage
structure and populations. For example, habitat
type and movement patterns can influence the
size range and age classes found at different
sites (Wheatley 2000).
Fish
Sheltered reef fish species commonly include
southern hulafish Trachinops caudimaculatus,
weed fish Heteroclinus perpicillatus, old wives
Enoplosus armatus, toadfish (Aracanidae),
leatherjackets (Monacanthidae), globe fish
(Diodontidae), zebra fish Girella zebra, blennies
(Blennidae) and seahorses Hippocampus sp..
Wheatley (2000) examined fish assemblage
structure, species richness and density on
sheltered reefs in Port Phillip Bay in relation to
macro-algae, depth, substrate type etc., as well
as how habitat mediates recruitment,
competition and predation, with an emphasis on
the leatherjackets Meiischenia freycineti and
Meiischenia hippocrepis. This study found that
fish assemblages were most similar on reefs that
were closer together than distant reefs, and
suggested that these patterns were due to
similarities in larval supply, wave action and
tidal movement as well as to differences in
macroalgal assemblage structure. M. hippocrepis
were only recorded on reefs, while M. freycineti
were mainly found on reefs as adults following
recruitment to seagrass beds. Mean growth rates
differed between adults of the two species on
different reefs, and adults appeared to be
permanent residents on individual reefs, with
no evidence of movement between reefs being
found. While M. freycineti juveniles and subadults moved between coastal seagrass beds and
offshore reefs over sand patches, the same sand
patches appeared to restrict movement of adults,
a pattern of movement also described for this
and other Monacanthid species in NSW (Bell et
al. 1978).
Coleman (1972) collected fish from sheltered
rocky reefs at Williamstown, Sandringham,
Rickett’s Point, Mornington, Mt. Martha, Safety
Beach and Sorrento Bay. Fish from 31 families
were recorded and observations of basic ecology
and biology were made including habitat use
and feeding behaviour. This study briefly
discussed how food availability, substrate type
and exposure may influence the distribution of
these species.
Sheltered subtidal reefs are also important to
snapper Pagrus auratus and King George whiting
Sillaginodes punctata at different stages of their
life cycles (Port of Melbourne Corporation
2007c).
The structure of fish communities inhabiting
reef-algal and unvegetated sandy areas at six
sites has been studied by underwater visual
census over 18 months in Port Phillip Bay
(Jenkins et al. 1996). A total of 75 species
representing 36 families were recorded. Species
richness and total abundance varied both
spatially and temporally but were always
greater on the reefs (Jenkins et al. 1996). In
general, the northern bay sites were
characterised by small cryptic species with an
increase in conspicuous water column species in
the south of the Bay (Jenkins et al. 1996).
Differences in community structure between
these sites may be related to reef topography,
Such patterns of ontogenetic movement between
habitats as individuals grow has been reported
for other species in the Bay such as Sillaginodes
punctatus (that moves between seagrass
beds/reef-algae and sand areas ) (Jenkins et al.
1996; Jenkins and Wheatley 1998), as well as
elsewhere in the region, e.g., Pseudolabrus
celidotus (Jones 1984) in New Zealand and
Achoerodus viridis (Gillanders 1997b; Gillanders
1997a) in NSW.
Temperate reefs literature review
13
Deep reefs
Deep reef habitat is defined here as rocky
habitat at depths greater than 20 m (Ball and
Blake 2007a; Port of Melbourne Corporation
2007b). In western Victoria, recognised deep reef
sites include Discovery Bay Deep, offshore Cape
Bridgewater and Cape Nelson, Port Fairy Deep,
offshore Moonlight Head, Cape Otway, Apollo
Bay and Nine Mile Reef (Edmunds et al. 2006b;
Ball and Blake 2007a; Ball et al. in prep). Other
features include a submarine scarp between
Barwon Heads and Cape Otway on the Otway
Shelf (Jennings 1958) and a cliff at ~45 m depth
that extends for ~20 km between Point
Roadnight and Sugarloaf Creek (Gill et al. 1980).
In central Victoria, deep reef sites include Port
Phillip Heads, including The Canyon, Point
Addis and Wilson’s Promontory (Edmunds et al.
2006b; Ball and Blake 2007a). Deep reefs habitats
are also located in Port Phillip Bay itself at
Schnapper Deep, Portsea Hole, Spectacular Reef
and Far Side Reef, and The Rip (Edmunds et al.
2006b; Port of Melbourne Corporation 2007a). In
eastern Victoria, smaller areas of deep reef exist
near Point Hicks and Cape Howe (Edmunds et
al. 2006b). Mapping has been performed using
video transects and side-scan sonar to provide
quantitative descriptions of reef biota on sites
including: granite reefs such as “Point Hicks
Reef”and“New Zealand Star Bank”; low relief
sandstone/limestone reefs such as “Broken
Reef”; and high/low relief limestone reefs such
as the ”Howe/Gabo complex” (Bax and Williams
2001). Broader-scale surveys have also been
conducted of substrate type and benthic
organisms throughout the Bass Strait which
have included reefs such as “New Zealand Star
Bank” (O'Hara 2002; Passlow et al. 2006).
At New Zealand Bank, Beaman et al. (2005)
described the biota that occurred on broadly
categorised habitats. In this particular study,
“high relief granite” outcrops at ~30-45 m were
described as closely resembling those of similar
habitats along the coast of NSW, where
communities include large sponges, ascidians,
cnidarians, bryozoans and reduced algal cover
(Underwood et al. 1991; Andrew 1999; Beaman et
al. 2005).
“Deep reef/urchin barrens” described at this site
at ~40-50 m (Beaman et al. 2005) closely resemble
those described in NSW and elsewhere
(Underwood et al. 1991), with large concentrated
patches of the urchin Centrostephanus rodgersii on
deep reef, inhabiting areas consisting
Temperate reefs literature review
14
predominantly of encrusting algae and, where
urchins are low in number, many sessile
invertebrates such as sponges and seawhips.
Edmunds et al. (2006b) states that few studies
have been done on deep reef biology due to the
logistical difficulties associated with working at
these depths. While some deep reef invertebrate
species have been identified, most remain
unidentified due to the paucity of collected
specimens. Consequently, our ecological
understanding of deep reef assemblages is
limited, especially with respect to patterns and
factors that influence the diversity and
abundance of deep reef species.
In general, worldwide research on deep reefs
remains sparse in comparison with that on
shallow reefs.
Invertebrates
Deep reef biota is typified by invertebrate
animals rather than algae, usually in the form of
sessile, filter feeding fauna (Tissot et al. 2006;
Sanchez et al. 2009). Organisms such as sponges,
octocorals, bryozoans and ascidians usually
dominate rock faces on deep reefs (Keough and
Butler 1996; O'Hara et al. 1999; Edmunds et al.
2006b). For example, the rocky wall within
Portsea Hole is characterised by an abundance
of sponges, ascidians and bryozoans (Elias et al.
2004; Edmunds et al. 2006b). This is partly due to
the ability of species such as sponges to survive
in low light conditions that algae is unable to
survive in (Sorokin et al. 2008).
Communities of sessile invertebrate animals are
sometimes collectively known as a ‘sponge bed’
(Butler et al. 2002a). Sponge bed species can be
encrusting or grow in a variety of forms whilst
attached to the substratum (Edmunds et al.
2006b). The most common algae present on deep
reefs are encrusting coralline red algae which is
able to tolerate low levels of penetrating light
(Edmunds et al. 2006b). Sponge bed
communities are the typical deep reef biota at
sites across Victoria but there are differences in
assemblages between deep reef sites (Edmunds
et al. 2006b; Ball and Blake 2007a; Ball et al. in
prep). Butler et al. (2002a) have surveyed
sponges throughout the Bass Strait.
Recent surveys have concentrated on the area of
reef known as The Rip, in Port Phillip Heads,
which is more complex than other reefs within
the Bay and includes “a variety of substrate
structures including vertical walls, caves, ledges,
reef slopes, reef flats and steep-sided chasms”
(Edmunds et al. 2006b). These diverse
microhabitat types may result in a high diversity
of species and (Edmunds et al. 2006b).
Edmunds et al. (2006b) found assemblages of
benthic fauna were different between all deep
reefs within Central Victoria and Port Phillip
Bay collectively. The Port Phillip Bay deep reef
assemblages were dominated by sponges,
occupying 70 to 90% of the rocky substratum.
The Point Addis assemblage was dominated by
upright sponges (arborescent, massive and
flabellate growth forms), but cnidarians
including hydroids were entirely absent.
Wilson’s Promontory had a low coverage of
encrusting sponges and hydroids, with high
abundances of red and brown algae and the
gorgonian fan Pteronisis sp.. The Port Phillip
Heads assemblage was dominated by encrusting
sponges, hydroids, ascidians and bryozoans,
many of which are considered rare, and,
according to Edmunds et al. (2006c) there are
many identifiably different assemblage types
present.
Within Port Phillip Heads, further differences in
assemblage structure were apparent between
areas (Edmunds et al. 2006b):

Catacombs Ridge, the Plateau and Nepean
Bank contained a high abundance of
encrusting sponges and hydroids

Deep cobble habitats, such as Uelmans Deep
were characterised by low abundances of
sessile organisms

Reef walls at ~25 m were dominated by
encrusting osculated sponges and other
sessile species, e.g. Portsea Hole

Reef slopes in the area were characterised by
dominant assemblages of “massive ruffled
sponges, rambling grey encrusting sponge
and other encrusting sponges”.
Other sessile species that occur in the reef
assemblages of Port Phillip Heads include
hydroid fans, soft corals, gorgonian corals,
crustose and arborescent bryozoans, colonial
ascidians, and solitary ascidians (Edmunds et al.
2006b). It has been reported that over 271
validated species of sponges have been collected
from Port Phillip Heads, which is a large
proportion of the 523 species identified to date
from Victoria, and the 1416 identified to date
throughout Australia (Flora and fauna
guarantee - scientific advisory committee 2009).
Notably, 115 of the species recorded in Port
Phillip Heads are unknown from any other area
(Flora and fauna guarantee - scientific advisory
committee 2009). However, detailed taxonomic
studies are still incomplete on these species in
Port Phillip Heads and as sponges are known to
exhibit variable growth patterns in relation to
currents, wave action, predation, etc. (Palumbi
1986; Kaandorp 1999; Bell and Barnes 2000; Hill
and Hill 2002), and species can adapt their gross
morphology to changes in their environment
within weeks (Palumbi 1984; Kaandorp and
Kluijver 1992), more in depth taxonomic studies
are required to confirm these records.
Similarly, this area also exhibits a high diversity
of Bryozoans compared to other areas (Ponder et
al. 2002), has been described as an area
containing highly species rich hydroid fauna
(Watson 1982), and contains at least one ascidian
species known only from Port Phillip Heads
(Flora and fauna guarantee - scientific advisory
committee 2009).
While studies of organisms on deep reefs in Port
Phillip Heads remain sparse, work has been
conducted elsewhere on species such as sponges
that are predominant on this type of reef
(Roberts and Davis 1996; Roberts 1996).
While recent surveys of Port Phillip Heads have
begun to identify those species that make up
sessile communities, little is known about the
mobile invertebrates that occur on deep reefs in
Port Phillip Bay. This is partly due to sampling
strategies that have been used, with video
transects unable to record species that are often
cryptic in nature and shelter in cracks and
crevices in the substrate as well as within cryptic
microhabitats created by sessile organisms. In
addition, technical difficulties involved in
surveying deep reefs also make this problematic
(Edmunds et al. 2006b). Edmunds et al. (2006b)
suggests that these organisms play an
ecologically significant role, providing food for
carnivorous fishes on deep reefs in Port Phillip
Bay, and are likely to include a variety of
crustaceans and molluscs.
Fish
Deep reef fish fauna include common subtidal
reef fish species, but other species that are rare
on shallow reef may also be present including
boarfishes Pentacerotidae), splendid perch
Callanthias australis, butterfly perch Caesioperca
Lepidoptera and banded seaperch Hypoplectrodes
nigroruber (Edmunds et al. 2006b). Fish
assemblages typically begin to change at depths
greater than 20 m, with the loss of the kelp-
Temperate reefs literature review
15
associated wrasses and leatherjackets, and the
appearance of deeper water species (O'Hara et
al. 1999). On deep reefs within Port Phillip Bay,
fish assemblages are reasonably diverse and
include a variety of planktivores, roaming
carnivores and omnivore/herbivores that occur
in moderately high abundances (Edmunds et al.
2006b).
The relationships between fish distributions and
reef habitat in Victoria have not been examined
to a large extent, apart from recent studies on
reef down to ~60 m at Cape Howe Marine
National Park (Moore 2008; Moore et al. 2009)
using stereo-Baited Remote Underwater Video
Systems (stereo-BRUVS). This study used
habitat suitability modelling to successfully link
the distribution of fish species with
environmental characteristics including depth,
substrate type and reef complexity (Moore et al.
2009). This allowed an examination of subtle
differences in biological and topographic
complexity and their importance in structuring
species distributions. For example:

The eastern blue grouper Achoerodus viridis,
a carnivore which feeds on a variety of prey,
was distributed in relation to the presence of
solid reef and boulders

The green moray Gymnothorax prasinus was
linked positively to reef and broad-scale
topographic complexity

The eastern blue-spotted flathead
Platycephalus caeruleopunctatus, was closely
affiliated with sand (Hutchins and
Swainston 2002) and was present in depths
Temperate reefs literature review
16
of > 60 m over sediment areas and some
reef-and-sediment areas, and in relation to
algae and invertebrates

The draughtboard shark Cephaloscyllium
laticeps was present across all substrata in
depths of ~ 60 m, and was predicted to be
present out to the deepest regions of the
park at ~110 m
Surveys of habitat structure and fish community
structure on deep reefs from 25-200 m from
Gippsland Lakes – Cape Howe and southern
NSW found a correlation between communities
and depth, latitude and seabed type (Williams
and Bax 2001). For example, 61% of fish species
surveyed were associated with either reef or soft
substrates, and the remainder were associated
with both. Species that were associated only
with reefs in the study included: butterfly perch
Caesioperca lepidoptera, eastern orange perch
Lepidoperca pulchella, bearded rock cod
Pseudophycis barbata, chinaman leatherjacket
Nelusetta ayraudi, barber perch Caesioperca razor,
splendid perch Callanthias australis, striped
trumpeter Latris lineate, mado Atypichthys
strigatus, bastard trumpeter Latridopsis forsteri,
common bullseye Pempheris multiradiata, maori
wrasse Ophthalmolepis lineolata, longfin pike
Dinolestes lewini, largetooth beardie Lotella
rhacinus, bluethroat wrasse Notolabrus tetricus,
southern conger Conger verreauxi, sergeant baker
Aulopus purpurissatus, swallowtail Centroberyx
lineatus, pigfish Bodianus sp., silver sweep Scorpis
lineolata, and thresher shark Alopias vulpinus.
Canyon reefs
Assemblages within canyons vary with
substrate depth, size, and complexity (Harris
2007; Williams et al. 2009b) and these habitats
may be particularly sensitive to anthropogenic
disturbance in the form of commercial fishing
which is concentrated at ~50-1300 m depth
(Larcombe et al. 2002; Williams et al. 2009b).
anemones and ascidians (Port of Melbourne
Corporation 2007d). Sponges make up 67% of
the biological groups in the canyon, hydroids
14% and all other groups less than 2% each (Port
of Melbourne Corporation 2007d). Edmunds et
al. (2006b) also found encrusting sponges and
hydroids to dominate these assemblages.
Port Phillip Heads
Other canyons
The entrance of Port Phillip Bay contains a key
geomorphological feature known as “The
Canyon” which meanders more than 3 km from
1 km south‐west of Point Nepean through the
Entrance and 1 km north‐west of Point Nepean
inside the Entrance. It covers an area of ~100-120
ha, with a depth range to ~100 m, but also
includes two shallow banks, Nepean Bank and
Rip Bank, which lie in the shipping channel
through the Entrance (Port of Melbourne
Corporation 2007d). The canyon contains a
distinctive ecological environment unlike other
environments in the Bay or elsewhere in
Victorian waters. Reef communities within the
Canyon are present on mixed substrates of
Aeolian calcarenite and basaltic rock and
experience large volumes of water movement,
with the tidal flow between the heads reaching
speeds of up to 5 ms-1 (Edmunds et al. 2006b).
The water passing through Port Phillip Heads
carries high volumes of suspended particles
such as sand on the flood tide and sediments
from the Bay on the ebbing tide which reduce
light levels, but increase nutrient levels
(Edmunds et al. 2006b).
Little is known about abyssal reefs found deep
on the continental slope (>200-400 m), such as
the “Cascade” off east Gippsland, which is
thought to be dominated by cold water corals
that form a habitat for a variety of invertebrate
species (O'Hara et al. 1999).
Invertebrate communities are common deep reef
biota, but are difficult to study because of the
strong tidal currents, ocean waves and seabed
depths that are beyond the normal range of
depth for scuba divers. Consequently, the
Canyon’s habitats and associated biota have
only been examined in detail by using remotely
operated vehicles (ROVs) and, to a lesser extent,
divers using underwater video and still
photography (Port of Melbourne Corporation
2007d). These ecological studies confirmed that
much of the rocky seabed beyond 20 m in the
canyon consists of highly sculptured calcarenite
reef (including vertical walls, pinnacles, crevices
etc) with sandy and rubble patches on the flatter
regions and in the deep basins. The permanent
rocky habitat is totally covered with a range of
sessile invertebrates, particularly encrusting
sponge communities, with a variety of
component biota including hydroids, bryozoans,
Recent mapping of geomorphic features and
assemblages from 200-2000 m has, however,
been conducted in south-east Australia (Harris
et al. 2005) and has been used in the design of
deep water Marine Protected Areas (Harris
2007), although mapping is still incomplete at
the current time (Williams et al. 2009b). As such,
relatively little is known about organisms that
inhabit deep reefs and rocky canyons in the
region, with surveys of similar habitats
elsewhere in Australia often finding a high
percentage of un-described species (Poore et al.
2008). For example, recent surveys in 2009 are
the first to collect samples of invertebrate
megafauna from deeper than 1800 m around
Australia (Williams et al. 2009a)).
Some work has been conducted off southeast
Australia. Surveys were recently undertaken in
deep-water marine reserves within the “Southeast Commonwealth Marine Reserve Network”
(Department of the Environment 2008) that were
gazetted in 2007 as part of the “National
Representative System of Marine Protected
Areas” (Williams et al. 2009b). Off East
Gippsland, areas of rocky reef are exposed from
muddy sediments on the steep slopes of “Big
Horseshoe Canyon”, part of the Bass Canyon
system. The area is the largest canyon that has
been sampled in the south-east for benthic
biodiversity (Williams et al. 2009b). This canyon
system, at a depth of ~1500 m, has a total area of
~319 km2 (Williams et al. 2009b), and the
majority of the diverse, abundant, sessile
megafauna in the canyon is associated with
these exposed hard surfaces (Kloser et al. 2001).
Percentage cover of “rich fauna” as determined
from underwater video peaked at 200-300 m
(Williams et al. 2009b). Organisms include
diverse, abundant, filter-feeding species, such as
Temperate reefs literature review
17
dense beds of large sponges and the stalked
crinoid Metacrinus cyaneus, and numerous
species of octocoral (especially gold corals). This
site is the type locality for M. cyaneus and it is
the only known location of the species off southeastern Australia (Kloser et al. 2001; Williams et
al. 2009b). Above 600 m fisheries are important
in this area (Harris 2007).
In contrast, the Zeehan canyon complex to the
south-west of Victoria consists of mostly soft
sediment substrates, with few exposed rock
surfaces (Williams et al. 2007). Canyons in this
area exist every ~7 km and include fisheries for
several species including blue grenadier, crab,
ling and rock lobster (Harris 2007). According to
Harris (2007) “conservation values include links
between canyons, ocean currents, upwelling
processes and flows through Bass Strait”.
Temperate reefs literature review
18
Several canyons in south-east Australia are the
locations of the largest known aggregations of
feeding and spawning fishes in the South-East
Fishery region (Williams and Kloser 2004). For
example, the Zeehan Canyon complex provides
“areas for high-order predator foraging and blue
grenadier spawning” (Harris 2007). Elsewhere,
plankton and nekton have been found to be
elevated in and around canyons, in addition to
high densities of megafauna (Beaman et al.
2005).
The use of abiotic classification of habitats to
infer levels of biodiversity in these deep systems
is a contentious issue (Harris et al. 2009; Williams
et al. 2009a; Williams et al. 2009b), which
indicates how little is currently understood
about assemblages in these systems.
Ecological and physical environmental
drivers
Major environmental factors influence reef
communities in Victoria. Womersley and King
(1990) split these factors into four key groups:
roughness and complexity, and shading
(Womersley and King 1990; Underwood and
Chapman 1995; Little et al. 2009).
Dynamic factors:
Three features of the tides affect intertidal reef
communities: the rise and fall of the tide,
variation between neap and spring tides and
daily progression of the tides (Underwood and
Chapman 1995; Little et al. 2009).

Tide - period and amplitude, cycles

Water movement – currents, upwelling, surf
and wave action, storms

Wind – effect on tide, water movement, and
humidity
Physical factors

Light – quality, quantity, periodicity

Sea temperature
stratification

Air temperature (in conjunction
emersion)– variation, range

Relative humidity

Rainfall

Substratum
–
variation,
range,
with
Chemical factors

Salinity

Nutrients

Availability of gases (O2 and CO2)

pH

Pollution
Biotic factors

Competition for space, light, nutrients etc

Grazing/predation

Epiphytism

Symbiotic relationships
Intertidal reefs
Tides are, perhaps, the most important
environmental factor causing gradients across
intertidal reefs. The direct effects will be
modified by factors such as wave action, air
temperature, humidity, freshwater run-off,
desiccation stress, rock type, aspect, surface
Waves cause major variation in assemblage
structure of intertidal reef communities from
exposed to sheltered areas (Underwood and
Chapman 1995). The importance of the
magnitude of wave exposure in shaping
community structure has long been recognised
(Lewis 1964; Stephenson and Stephenson 1972),
with comparisons and contrasts made between
exposed and sheltered intertidal reefs (e.g.
Seapy and Litter 1978).
A diverse range of other factors and processes
influence assemblage structure and a variety of
work has been carried out worldwide that
quantifies how these interact with each other.
For example, while molluscan grazers may
impact algal cover and shape sessile assemblage
structure at local scales, the availability and
supply of algal propagules is also important and
may in turn be partially driven by processes at
broader spatial scales such as benthic-pelagic
coupling and oceanic currents (Schiel 2004;
Underwood and Chapman 2007; Little et al.
2009).
At local scales, gradients can be seen in the
relative importance of physical and biological
factors on intertidal reefs. Physical factors, such
as desiccation stress, generally increase in
importance up the shore, while biological factors
such as competition and predation increase in
importance down the shore. Towards the top of
the shore where physical stress is high, algal and
sessile invertebrate cover is low, while towards
the bottom of the shore where physical stress is
low, percentage cover of algae and sessile
invertebrates is high. In the mid shore, where
physical and biological stress levels may be of
equal importance, a bare zone is often found. In
this zone, there is typically a low cover of algae
due to a mixture of desiccation stress and
Temperate reefs literature review
19
grazing; competition for free space is therefore
usually low (Connell and Gillanders 2007; Little
et al. 2009).
There is a large body of research that focuses on
the importance of disturbance in the structuring
of intertidal reef assemblages. Disturbance is
important, as free space is a limiting factor at
some tidal heights on intertidal reefs and the
creation of free space can lead to changes in the
structure of assemblages (Little et al. 2009).
Initially this research concentrated on predation
as a key driver (Paine 1966) but this was
followed by a general recognition that any
disturbance that created space was fulfilling the
same function (Connell and Gillanders 2007;
Little et al. 2009).
The magnitude and timing of disturbance events
such as storms, ice scour and boulders
overturning can create patches of space that
allow new individuals to recruit and may lead to
a complex mosaic of patches at various states of
succession (Connell and Gillanders 2007; Little et
al. 2009). On some shores, the impact of
disturbance events may be relatively
predictable, for example low on shores that have
high algal cover (Connell and Gillanders 2007).
Another key factor that affects the ecology of
intertidal reefs is the arrival of propagules or
“supply side ecology” (Underwood and
Chapman 2007). Planktonic dispersal and
survival of propagules is highly variable, and
consequently, spatial and temporal variability in
the number of propagules arriving is also highly
variable. The interactions between these new
recruits and the existing assemblage will vary
greatly depending on this “patchiness” in larval
supply (Underwood and Chapman 2007). Apart
from supply of propagules, other bottom up
processes relate to the interaction of coastal
oceanography with intertidal communities, for
example the effect of coastal upwelling on the
supply of phytoplankton, particulate matter and
nutrients to intertidal assemblages (Menge
2000). Top down and bottom up processes can
interact to influence the dynamics of
communities on intertidal reefs (Menge 2000).
The following sections examine research that
has been conducted locally.
Top-down / disturbance processes
In Victoria, several studies have been conducted
that examine natural disturbance on intertidal
reef communities. For example, Bennett (1990)
examined disturbance within beds of the
commonly occurring alga Hormosira banksii, and
Temperate reefs literature review
20
several other authors describe the effects of very
hot days combined with afternoon low tides on
this species (Keough and Quinn 2000; Bellgrove
et al. 2004) and mobile gastropods (Parry 1982).
Parry (1982) also described predation on
intertidal limpets by oystercatchers and wrasse.
The majority of disturbance studies in Victoria
have, however, been focused on anthropogenic
disturbance events, such as the recreational use
of reefs and the effects of trampling and the
harvesting of invertebrates for food or bait
(Addison et al. 2008) or the effects of pollution
(Bellgrove et al. 1997; Hindell and Quinn 2000).
In a study that has collected data from 65
intertidal reefs across Victoria, a number of
bioassessment methods have been tested to
determine their ability to detect impacts on reef
assemblages (O'Hara et al. in press).
Human impacts on rocky shore communities
have been chronicled in Victoria, and appear to
take two main forms: the collection of seafood
from the intertidal zone; and trampling of
organisms by visitors walking on the shore
(Povey and Keough 1991; King 1992; Keough et
al. 1993; Keough and Quinn 1998; Keough and
Quinn 2000; Addison et al. 2008). Such impacts
are well known and have been shown elsewhere
in Australia and internationally (Underwood
1993; Fletcher and Frid 1996).
Human predation (collecting)
Local studies within Port Phillip Bay, focusing
on the collection of gastropods from the
intertidal zone, have demonstrated that the
mean size and abundance (one species only) of
target species were reduced at an impacted site
compared to protected sites nearby (Keough et
al. 1993; Keough and Quinn 2000). The indirect
effects of this type of human predation have
been found to be subtle and related mainly to
the abundance of microalgae. For example,
frequent removal of cellanid limpets can result
in an increase in microalgal densities while
removal of nerites results in a decrease in
microalgae (Sharpe and Keough 1998).
Such removals may have more complex effects
than currently appreciated, and a more in depth
examination is required. For example, it has
been suggested that removing the largest
individuals from a population of the limpet
Cellana tramoserica may eliminate the bulk of the
reproductive effort of this species (Parry 1982),
which may in turn impact community structure
on the targeted shore as well as nearby areas.
There is almost no ecological information
available about another main target species in
Victoria, Turbo undulatus, and so it is impossible
to speculate what the effect of its removal might
be (Keough and Quinn 2000).
Other studies undertaken on the Victorian coast
at shores visited less frequently have found little
evidence to suggest there is an impact of
harvesting (Keough and King 1991; King 1992),
although Addison et al. (2008) suggest that
shellfish collections at Sorrento may have a
broader impact on intertidal assemblages.
There have been no studies on the harvesting of
other taxa for bait in Victoria, although in New
South Wales there is some evidence that the
removal of other taxa such as crabs and the
tunicate Pyura stolonifera is unsustainable
(Fairweather 1991; Underwood 1993). This may
also hold true in Victoria, where fertilisation
success in isolated spawning individuals (>2 m
apart) of P. stolonifera at Barwon Heads has been
shown to be limited by a scarcity of sperm
(Marshall 2002). Removal of individuals may
therefore directly reduce reproductive capacity
within local populations.
Fisheries Victoria is currently liaising with local
communities to determine which species they
preferentially target on intertidal reefs; this
information could be used to direct future
research examining this type of disturbance.
Trampling
The other main disturbance caused by the
recreational use of reefs is trampling. This has
been recognised as an issue of concern in
Victoria (Carey et al. 2007). There is evidence
that trampling can impact intertidal reef
assemblages both in Victoria (Povey and
Keough 1991; Keough and Quinn 1998) and in
other parts of the world (Schiel and Taylor 1999;
Benedetti-Cecchi et al. 2001). Studies undertaken
within Port Phillip Heads Marine National Park
have found that trampling produced marked
effects in two major habitats. In beds of coralline
algae, the effects were short lived, but in beds of
Hormosira, algal cover was greatly reduced and
the number of grazing molluscs was increased
in trampled areas and recovery was still not
complete >400 days later (Povey and Keough
1991). This pattern was also evident in a
separate, longer-term study, where two of three
sites were able to recover between trampling
events whilst a third site did not recover
(Keough and Quinn 1998). Keough and Quinn
(1998) propose a conceptual model where
Hormosira can be likened to a keystone species
(Paine 1995) whereby they provide damp, shady
refuges for some species but also reduce the
cover of microalgae - the main food source for a
number of gastropod grazers. The mobile
herbivores emigrate in response to increasing
cover of Hormosira and when Hormosira cover is
reduced, microalgal abundances increase
resulting in increased densities of grazers. The
authors of this study were unable to determine
why one site showed no recovery over the sixyear study but they discuss the potential
ramifications of sites having different levels of
resilience and the management options
associated with this. A management program
within the Marine National Park system will be
targeting trampling as a hazard and this may be
able to provide data on which shores are more,
or less, resilient to trampling (Carey et al. 2007).
Pollution
The release of pollutants is another major type of
disturbance event that impacts assemblages on
intertidal reefs in Victoria. Reefs within Port
Phillip Bay are more likely to be impacted by a
range of toxicants due to their proximity to
urban and industrial areas, harbours, ports and
shipping channels. There have been numerous
studies documenting the superimposition of
male sexual characteristics on female gastropods
following exposure to TBT (tributyltin) -based
antifouling paint since this was first
demonstrated (Bryan et al. 1987; Gibbs et al.
1987). Foale (1993) described the incidence of
imposex in the whelk Thais orbita at sites around
Port Phillip Bay and found that the incidence of
imposex corresponded strongly to the proximity
to harbours or marinas although mean body
burdens of TBT were not particularly high. The
reduction in reproductive potential of intertidal
predators such as whelks has the potential to
impact the structure of intertidal reef
assemblages (Spence et al. 1990).
Studies of intertidal reef assemblages affected by
TBT have not been undertaken in Port Phillip
Bay, although Parry (1982) considered predation
by whelks to be a minor cause of mortality in the
four species of limpet he studied at a San Remo
shore. A recent study (Rees et al. 2001) found
that the severity and extent of imposex has been
reduced except at locations that were adjacent to
major ports or certain harbours. It is possible
that in some areas sediments are providing a
source of TBT although no data exists on this in
Port Phillip Bay.
Other toxicants, such as copper, have also been
shown to cause imposex in certain species (Nias
et al. 1993). No specific research on the effects of
other toxicants on intertidal reef assemblages
Temperate reefs literature review
21
within Port Phillip Bay or coastal Victoria have
been conducted to our knowledge.
Bottom-up / supply side processes
Pollution
A form of pollution that primarily has a bottomup effect through the elevation of nutrients is
the discharge of sewage effluent. The majority of
studies considering the effects of sewage
effluent on intertidal reef assemblages in
Victoria have been undertaken around Boags
Rock. However the Western Treatment Plant
also discharges secondarily treated sewage
effluent from Melbourne into Port Phillip Bay.
Most of the studies considering the impacts of
this discharge have been focused on the
predominantly soft sediment environment in the
vicinity of these discharges. Axelrad et al. (1981)
sampled the macrophyte assemblages at three
basalt reefs within the Bay and found that the
intertidal zone at Werribee was characterized by
Centroceras clavulatum throughout the year and
Ulva lactuca and Enteromorpha compressa in
summer / autumn. Blooms of the blue green alga
Cladophora sp. were also observed. We are not
aware of more recent studies specifically aimed
at the impacts of sewage effluents within Port
Phillip Bay.
Long-term monitoring of the macroalgal
assemblage around the Boags Rock discharge
found that fewer species were recorded at sites
closer to the outfall, as well as a loss or
reduction of Hormosira coupled with an increase
in algal turfs and the polychaete Boccardia
proboscidea adjacent to the discharge (Brown et al.
1990). These patterns are considered typical of
those that have occurred around sewage
discharges throughout the world (Littler and
Murray 1975; Fairweather 1990). There is no
clear consensus to explain the loss of H. Banksii
(or other dominant brown algae around the
world) following exposure to sewage effluent
(Bellgrove et al. 1997). At Boags Rock there was
no evidence that the sewage effluent
detrimentally affected the availability of
propagules or macroalgal recruitment (Bellgrove
et al. 1997). There were, however, very high
densities of propagules of opportunistic taxa
such as Ulva and Enteromorpha and high
recruitment of these taxa at polluted sites. Other
studies have found that high concentrations of
treated sewage effluent inhibit zygote
germination and embryo growth of Hormosira
(Doblin and Clayton 1995) and so propagule
supply and recruitment may still be important
in the loss of Hormosira in sewage affected areas.
Temperate reefs literature review
22
It is likely that the small dispersal shadow of
Hormosira, combined with potential impacts on
establishing zygotes, will limit the reestablishment of this species in areas adjacent to
sewage discharges (Bellgrove et al. 1997)
Another study undertaken at Boags Rock has
found that the recruitment of the intertidal
mussel Brachiodontes rostratus was facilitated by
the secondarily-treated effluent while shell
growth was reduced and mortality of larger
individuals was increased (Hindell and Quinn
2000). This study also discussed the implications
for intertidal fauna that use the habitat created
by mussel clumps.
Larval supply / recruitment
It is well recognised that a vital consideration in
rocky shore ecology is variability in recruitment
in both space and time. Recruitment depends on
larval supply, local hydrodynamics, larval
behaviour, predation / herbivory and the
availability of adequate resources to allow
settlement and survival and so recruitment to a
population. Quinn (1988) undertook an
experimental study to investigate the
importance of conspecific adults, macroalgae
and height on the shore to recruitment of the
intertidal limpet, Siphonaria diemensis. In this
study there was no effect of the density of adults
on the density of recruits or height on the shore
but there was an effect of macroalgal cover; with
significantly greater recruitment to areas with
encrusting macroalgae.
Studies of intertidal macroalgal species in
Victoria have concluded that while recruitment
cannot be predicted directly from the supply of
propagules, the two processes are linked
(Bellgrove et al. 2004). Results indicate that while
pre- and post-settlement processes are likely to
influence macroalgal distribution and
abundance, the temporal and spatial variability
in the supply and recruitment of propagules can
explain much of the patchiness in macroalgal
assemblages. A study on the effects of the
grazing gastropod Bembicium nanum on the
recolonization of algae at Aireys Inlet found that
B. nanum significantly reduced recolonization of
the ephemeral brown alga Scytisiphon lomentaria
but had no effect on recolonization of Hormosira
or the green alga Enteromorpha intestinalis (Braley
et al. 1991). A separate study has found that
micrograzing copepods have the potential to
greatly influence the density of various
macroalgal taxa, while a littorinid mesograzer
may reduce the both the density and number of
macroalgal taxa (Bellgrove 1998). This study also
found that two limpets studied (Patelloida
latistrigata and Cellana tramoserica) did not have
any significant effects on the densities of
macroalgal taxa recruiting (Bellgrove 1998). This
contrasts with studies at other locations that
have shown rapid colonisation of foliose algae
following the exclusion of patellid limpets
(Hawkins 1981; Underwood and Jernakoff 1981).
There have also been a number of studies
specifically focused on Hormosira. While this
species has a broad distribution throughout
south-eastern Australia, dispersal of propagules
is thought to be limited (Bellgrove et al. 1997;
2004). However, investigations into the
occurrence of ‘outbreeding depression’ at a
regional scale led researchers to speculate that
Hormosira may be capable of longer distance
dispersal than previously thought (McKenzie
and Bellgrove 2006). Detached fronds of
Hormosira have been shown to be capable of
releasing gametes for up to 8 weeks following
detachment and floating fertile fronds may be
an important mechanism for facilitating longdistance dispersal in this species (McKenzie and
Bellgrove 2008). Further studies investigating
genetic diversity over the broad-scale
distribution of Hormosira using genetic markers
may give valuable insight into gene flow and
long distance dispersal and are already
underway.
In a study that quantified species richness of
invertebrates on 11 rocky intertidal shores
separated by a biogeographical barrier (Ninety
Mile Beach), it was revealed that species
richness and species composition did differ on
each side of the barrier and that the distribution
of species was not related to their potential for
dispersal (Hidas et al. 2007). The three species
studied with direct development were found on
both sides of the barrier, while seven of the eight
species restricted to one or other side of the
barrier had planktonic larvae. All species
present on Red Bluff, an isolated rock platform,
did, however, have planktonic larval stages. In
contrast, a study looking at the mitochondrial
DNA of the highly-dispersive intertidal
gastropod Nerita atramensota found a
biogeographical disjunction that was unrelated
to either the Coorong or Ninety Mile Beach;
instead, Wilsons Promontory was the point of
disjunction (Waters 2005). The authors
concluded that this sharp biogeographical
disjunction was in marked contrast to the
species’ high dispersal abilities and supports
other studies that have noted that Wilsons
Promontory may have considerable
biogeographic significance for Australia’s
southern temperate region (O'Hara and Poore
2000).
An ongoing study looking into the connectivity
of intertidal gastropod molluscs with varying
dispersal potentials within marine national
parks is currently comparing recruitment
patterns of gastropods in Marine Protected
Areas (MPA) and non MPA sites in Port Phillip
Bay and the central coast of Victoria (Bathgate
pers. comm.)
Nutrients
The supply of nutrients is considered another
important bottom-up process and to date there
has been very little attention in Victoria to the
impacts of nutrients on rocky shore ecology.
There have been studies that consider the
impacts of sewage discharges on intertidal reef
assemblages (see discussion above) but it is not
always easy to separate the effects of increased
nutrients from other aspects of the discharge
(freshwater, suspended solids, toxicants etc.). A
manipulative study has been undertaken at
Point Nepean and found no effect of adding
nutrients or removing grazers (separately or
together) on the algal assemblage monitored
(Baker 2006). This finding is in stark contrast to
work in other parts of the world (Thompson et
al. 2000; Bokn et al. 2003) and other parts of
Australia (Underwood 1984; Gorgula 2004) and
may relate to the time of year the experiments
were undertaken (summer) when propagule
supply was likely to be low (Clayton 1990;
Bellgrove et al. 2004) and physical stress (heat)
was high. A manipulative study undertaken in
the shallow subtidal in south Australia found
that increasing nutrients had no effect on algal
assemblages in the presence of canopies and, in
the absence of canopies, only had an effect on
algal assemblages in the presence of grazers
(Russell and Connell 2005). They attributed
these complex interactions to the fact that the
area of coast studied had few grazers (weak topdown control by grazers) and was relatively
oligotrophic. There is some evidence that the
coastal regions of Victoria are also under weak
consumer control compared to eastern Australia
(Beovich and Quinn 1992; Bellgrove 1998;
Sharpe and Keough 1998; Burton 1999) and this
may affect the response of assemblages to
elevated nutrients. Ambient nutrient levels
measured at Cheviot Beach (Parry and Restall
2007) were very similar to those cited in the
South Australian study and considerably less
than nutrient concentrations in New South
Wales (Russell and Connell 2005). Further
Temperate reefs literature review
23
studies are required to investigate how nutrient
supply might affect recruitment of Hormosira
and other algal species, grazer control,
competitive interactions and responses to
anthropogenic disturbances such as trampling.
Competition
Studies of competition on intertidal reefs in
Victoria have primarily focused on exploitative
intra and interspecific competition of
herbivorous gastropods. Studies undertaken at
San Remo have found that densities of
Austrocochlea constricta and Bembicium nanum
were not restricted by interspecific competition
for food, although intraspecific effects did occur
in summer and autumn due to a reduction in
food supply (Quinn and Ryan 1989). Quinn and
Ryan (1989) also found that growth was not a
fixed characteristic in Siphonaria diemensis and
can change rapidly in response to variations in
food supply; growth decreased with increased
densities of conspecifics, and shell growth and
tissue weight were reduced in an experimental
treatment with reduced food compared to a
normal food treatment. Beovich and Quinn
(1992) found no evidence that Siphonaria
lomentaria were food limited by Cellana
tramoserica in winter and spring at this site.
Studies on basalt rock in Port Phillip Bay have
found that Cellana tramoserica individuals are
strong exploitative competitors for food
resources and when exposed to high densities of
conspecifics experience reduced growth and
increased mortality (Wright 1989; Marshall and
Keough 1994; Merory 1997). These results are
consistent with studies from other parts of
Victoria (Parry 1982; Burton 1999) and New
South Wales (Creese and Underwood 1982).
Marshall and Keough (1994) also found an
asymmetry in the competitive ability of cellanid
limpets, with smaller individuals able to out
compete larger individuals. They speculated
that smaller limpets were able to access a food
supply within the rough rock surface that larger
limpets with their larger radulae were not able
to access. This hypothesis was later tested by
repeating the experiment on basalt rock at
Williamstown and limestone rock on the
exposed Point Nepean shore (Keough et al.
1997). This study confirmed predictions that
asymmetry in competitive abilities of limpets
would not be apparent on a smoother
substratum and was further supported from
results from another study undertaken at Point
Nepean (Burton 1999). Cellanid limpets were
also shown to be competitively superior to a
Temperate reefs literature review
24
pulmonate, spihonariad limpet at the exposed
rocky shore of Point Nepean (Burton 1999), in
contrast to a previous study undertaken at San
Remo (Beovich and Quinn 1992). In the Burton
(1999) study and that of Keough et al. (1997) it
was shown that while patterns of cellanid
growth varied in response to competitive
pressures, microalgal abundance did not. This is
in contrast to findings from other parts of
Australia (Underwood 1984) and other parts of
the world (Little et al. 2009). Further
investigation concluded that feeding behaviour
was not important to limpet growth and
mortality, and that the reasons for the observed
effect of increased densities on growth and
mortality at Cheviot Beach (Point Nepean)
remain unresolved (Burton 1999).
There is evidence that the outcomes of
exploitative competition in Victoria can depend
on rock type and seasonal effects of food supply,
and appear to differ in some respects from
similar studies undertaken in New South Wales.
While there is evidence of competition for food
in Victorian populations, it does not appear to
be driving distributions as in New South Wales.
While in some instances removal or reduction in
grazer densities on Victorian shores did cause an
increase in algal growth, the magnitude of
effects was much smaller than observed in New
South Wales and other parts of the world.
Subtidal reef
A wide range of factors and processes are
responsible for the distribution and structure of
assemblages on subtidal reefs. These have
recently been reviewed in some depth (Connell
2007; Connell and Irving 2009; Wahl 2009), and
the descriptions that follow below cover work
that has been conducted in Victoria, with
relevant examples from elsewhere also included
to provide context.
Physical drivers
Biogeographical provinces or regions have been
demonstrated to influence the distribution and
composition of subtidal reef communities in
Victoria. Waters et al. (in press) recently
provided an a priori test for the existence of
Bennett and Pope’s (1952) Maugean (southeastern), Flindersian (western) and Peronian
(eastern) biogeographical provinces across
southern Australia. The study quantitatively
analysed distributional data from 1,500 algal
species and identified the three distinct
biogeographical assemblages, consistent with
traditional qualitative provinces. Connell and
Irving (2008) quantified the frequency and size
of patches of major benthic habitat on across the
southern coast of Australia and went on to show
that biogeography had a fundamental influence
on the patterns of abundance and composition
of subtidal assemblages across regional scales.
The most fundamental patterns related to (1) the
proportion of rock covered by kelp forests, as
related to particular functional groups of
herbivores, and (2) the small-scale heterogeneity
that characterises these forests.
Prior to this, O'Hara (2001) tested the
assumption that habitat defined by various
biological and environmental variables are
surrogates for biodiversity. The study collected
and analysed benthic floral and faunal samples
from subtidal reefs at 23 sites along the coast of
Victoria, and determined that habitats defined
by dominant vegetation, and to a lesser extent
region, supported consistent floral and faunal
assemblages. For each sample, habitat
parameters were recorded including dominant
vegetation, geographic region, depth, rock type
and exposure. Although physical variables were
determined to have little emphasis as drivers,
the author acknowledges that the experimental
design was probably not an appropriate test for
the effect of depth range, rock type and
exposure on biodiversity.
In contrast, Edmunds et al. (2000) specifically set
out to establish which physical variables
influence the distribution and composition of
reef communities. The study involved surveying
seven subtidal (8 – 18 m) sites between Cape
Schank and Wilsons Promontory to provide a
variety of reef substratum types, recording
biological (density of macroinvertebrates,
canopy forming plants, cover and biomass of
understorey macroalgae) and physical
(substratum lithology, relief, interstitial space,
and complexity, depth, exposure, aspect,
distance from shore, slope, longitude and
sediment cover) variables. Results of the study
found that many reef inhabiting biota were
closely associated with particular physical
variables. Invertebrates were closely associated
with combinations of substratum structure (i.e.
interstitial space, complexity, slope and relief),
depth and longitude whilst aspect and exposure
were also of some importance.
The slope of rock has been described as the most
striking feature influencing the composition of
assemblages of rocky subtidal systems by
Connell (2007). The position of rock affects key
physical processes such as light, flow and
sedimentation, as well as biological processes
including consumer foraging, competition,
facilitation and recruitment. Edmunds et al.
(2000) found algae had less correspondence with
physical data, but exposure, depth and reef
slope were reasonably good explanatory
variables. Substratum type lithology as a single
variable was not found to be a major
determinant, although some species occurred
predominantly on one substratum type. The
authors go on to state that the relationships
between variables measured do not necessarily
imply causality, but the physical-biological
relationships described are similar to those
found in studies of temperate reef assemblages
in New Zealand (Choat and Schiel 1982; Choat
and Ayling 1987).
In New Zealand, depth and exposure have been
shown to play an important role in algal and
echinoid distribution (Choat and Schiel 1982),
and echinoid and algal habitat have been shown
to play an important role in determining
associated fish fauna (Choat and Ayling 1987).
These findings have been reiterated in
Australian studies with depth and wave
exposure shown to influence the distribution
and composition of subtidal temperate reef
communities in southern Australia. Edgar (1984)
Temperate reefs literature review
25
described benthic floral and faunal assemblages
along the eastern, southern and western
Tasmanian coasts and related them to a subtidal
zonation pattern of wave exposure and depth.
Results of the study found that major
assemblages of benthic organisms within the
same cool-temperate biogeographical provinces
(Maugean and Flindersian), can be predicted
reasonably accurately by reference to wave
exposure and depth. Coleman et al. (2007) found
depth to be a strong driver of patterns of mobile
invertebrates in kelp forests on the temperate
coast of Western Australia. Their study found a
greater abundance and richness of common taxa
in holdfasts from shallow relative to deep
waters.
Such differences in the distribution of species
with depth can also be seen for a variety of reef
fish. For example, the reef fish Achoerodus viridis
has been shown to decrease from shallow to
deep areas of reef and from inner to outer areas
of estuaries, although large individuals
increased in numbers on more exposed coastal
reefs (Gillanders 1997a; Gillanders 1997b). Fish
found on rocky reefs in New South Wales, in
particular labrids, have been found to show
distinct patterns in diversity and abundance in
relation to wave exposure and depth (Fulton
and Bellwood 2004). Few species were found to
be abundant in exposed shallow habitats, while
several species were restricted to sheltered
habitats (Fulton and Bellwood 2004). Seasonal
fluctuations in water temperature may also
result in changes in the abundance of fishes on
temperate reefs (Parker Jr 1990).
Studies describing marine biodiversity in
Victoria have found depth and geography to
have the strongest influence on the composition
of communities. Ferns et al. (2000) described
temperate reef biodiversity through mapping
and analysis of biological data and concluded
that depth is an important factor in the
distribution of dominant biota because of
localized conditions such as substratum
topography/complexity and wave exposure.
Victorian temperate reef community Marine
Habiat Class (MHC) attributes generally differ
between shallow (0 - 2.5 m), moderate (2.5 – 20
m) and deeper (>20 m) subtidal depths. Broad
patterns of algal species zonation along the
depth gradient have also been shown in Victoria
(Sanderson 1997): Hormosira banksii is the
dominant species observed on intertidal rock
platforms; Durvillaea potatorum with mixed
brown and green algae inhabits the seaward
edge of intertidal rock platforms; Phyllospora
Temperate reefs literature review
26
comosa occurs in large beds at depths of ~3–5 m;
at exposed sites, Ecklonia radiata is typically
observed at depths >7 m and often growing with
P. comosa; and, giant kelp Macrocystis angustifolia
(an indicator species for the Maugean
biogeographical province) occurs at Port Phillip
Heads MNP - Point Lonsdale and at Point
Nepean.
In South Australia, Connell (2003a; b; 2005) has
investigated the importance of physical
processes such as light penetration and
sedimentation in structuring benthic
communities, and a number of studies
elsewhere have also shown that subtidal
assemblage distributions vary due to a variety of
physical factors including light availability,
wave action and storm disturbance (Dayton et al.
1984; Schiel and Foster 1986). Connell (2003a)
tested the hypothesis that different temperate
reef-algal habitat types will converge to become
like those under Ecklonia radiata if subjected to
the same low light and accumulation of
sediment without the presence of E. radiata. The
study found habitats did not converge under
treatments. The author concludes that canopies
place strong constraints on the presence and
abundance of many taxa, but not encrustingalgal habitats which beneficially coexist as
understorey. Connell (2003b) then went on to
investigate why many sessile invertebrates are
anomalously absent from understorey
communities. A series of experiments were
applied to partition both positive and negative
habitat modifying effects by kelp for sessile
invertebrate recruitment. The key finding of the
study was that a reduction of light intensity and
removal of sediment by canopies acted to
facilitate recruitment, but physical abrasion by
the canopy acted as a negative force to
overpower these positive effects for recruitment
of sessile invertebrates. Connell (2005)
experimentally separated the positive and
negative effect of light penetration and
sedimentation on the assembly and maintenance
of three subtidal habitats that characterise much
of the world’s temperate coastline: encrusting
coralline algae, articulate coralline algae and
filamentous turf-forming algae. The study found
shade to cause a positive effect for encrusting
coralline algae by facilitating the retention of
space without overgrowth; however, it caused a
negative effect for articulate coralline and turfforming algae by restricting growth and space
persistence. Sediment deposition caused a
negative effect on encrusting coralline algae, a
positive effect on articulated coralline algae and
a neutral effect on turf-forming algae. This
finding explains the presence of articulated
coralline and turf-forming algae on humandominated coasts following the loss of canopy
forming algae on reefs with heavy
sedimentation. The studies demonstrate the key
role of physical factors associated with habitatforming canopy algae, particularly in relation to
human-dominated coasts with heavy
sedimentation.
Another physical driver of subtidal reef
communities is the presence of oceanographic
features such as upwelling that can alter regimes
of variables such as nutrients and temperature.
For example, Butler et al. (2002b) states that the
Bonney upwelling area contains “distinct colder
water flora; rich assemblages of sessile filter
feeders such as sponges, bryozoans and corals;
feeding grounds for seabirds, fishes, whales and
other higher order predators such as fur seals
and penguins; and a productive fishing ground
for rock lobster, sustaining a relatively large
fishing industry”. Butler et al. (2002b) described
how limestone reefs in the area are often
covered by dense assemblages of molluscs,
sponges, bryozoans and red algae, with
upwelling related assemblages of bryozoans,
sponges and axoozanthellate coral on the shelf
edge and upper slope, and off Portland, diverse
cyclostome bryozoans are found on the deep
shelf.
Ecological drivers
Habitat structure
Ecological components of temperate reefs have
been shown to influence the distribution and
composition of subtidal reef communities in
Victoria. Algae, in particular canopy forming
species such as kelps, act as foundations and
strong ecological drivers to entire subtidal
systems because they provide important food
and habitat structure for other organisms on the
reef (Duggins 1989; Keough and Butler 1996).
Variation in the configuration of subtidal algae
in southern Australia and northern New
Zealand (e.g. monotypic or mixed stands of
algae) may influence the composition and
abundance of associated organisms (Goodsell et
al. 2004) such as understorey algae and sessile
invertebrates (Irving et al. 2004).
O'Hara (2000b; 2001) used multivariate analysis
and determined that dominant vegetation is a
primary ecological driver of subtidal reef
habitats. The study concluded that larger kelps
(e.g. Phyllospora, Ecklonia, and Macrocystis)
supported a relatively species-poor epiphytic
community compared with Cystophora and
Sargassum species. However, Ecklonia and
Macrocystis had large branching holdfasts that
provided shelter to many larger cryptic animals,
and provided stable substratum for many sessile
invertebrates and smaller algae.
Algae can influence the presence of other marine
biota such as fish and invertebrates (Andrew
1993; Edmunds et al. 2006a). Variation in algal
cover and other biotic factors can show
considerable variation spatially and temporally
(Dayton et al. 1984; Schiel and Foster 1986), and
this can subsequently be seen in the distribution
and abundance of associated invertebrate and
fish species (Russell 1977; Bodkin 1988;
Holbrook et al. 1990a, b; Schmitt and Holbrook
1990a; Anderson 1994).
A large body of work has been conducted in
California, USA on temperate reef systems.
Canopy-forming kelps such as Macrocystis
pyrifera can directly affect the densities of fish
species that use the kelp as a nursery area
and/or adult habitat (Ebeling and Laur 1985;
Bodkin 1988; Holbrook et al. 1990a; Carr
1994).The effects of increases or decreases in
different types of algal community have been
found to be strongly related to the resources
required by different life history stages of fishes
(Holbrook et al. 1990a), as well as to the differing
needs of individual species of fish (Holbrook et
al. 1990b). For example, positive effects have
been chronicled for species that use kelp as a
nursery ground or adult habitat; whereas
negative effects can be seen for fish that rely on
understorey species that are impacted by
shading effects and which may serve as
important sources of food (Holbrook et al. 1990a;
Schmitt and Holbrook 1990a; b).
While work on temperate reefs in California,
suggest that the distribution and abundance of
canopy and understorey algae may partially
explain spatial variation in the species
composition of fish recruitment among
temperate rocky reefs at the local level (Bodkin
1988; Carr 1989), heterogeneity of other habitat
features, e.g. the presence of sand patches, is
also likely to drive spatial variability in the
distribution of reef fishes, particularly at small
spatial scales (Garcı´a-Charton and Pe´rezRuzafa 2001). Macroalgal structure cannot,
therefore, necessarily be used as an indicator of
recruitment strength for algae associated fish on
reefs (Anderson 1994).
Effects of habitat forming biota are not just
limited to plants, e.g. sponge assemblages, have
Temperate reefs literature review
27
also been shown to directly influence the
distribution of reef fish in New South Wales
(Curley et al. 2002) as well as other areas, such as
the Mediterranean (Sanchez-Jerez et al. 2002).
Herbivory and Grazing
Herbivorous fish can make up to 47% of the fish
species present on some Victorian subtidal reefs
(Jones and Andrew 1990), and these species are
often associated with their preferred algal food
(Port of Melbourne Corporation 2007c). For
example, fish species such as herring cale Odax
cyanomelas feed almost exclusively on patches of
Ecklonia radiata and are more abundant on the
southern shallow reefs where these species are
more dominant (Jones and Andrew 1990). The
Victorian scalyfin Parma victoriae feeds in areas
where kelp is absent and red algal turfs are
present (Jones and Andrew 1990). Territorial
herbivorous fish such as P. victoriae and O.
cyanomelas may affect algal distribution and
abundance by (1) direct feeding effect due to
consumption of algae, (2) weeding out of algae
to promote growth of preferred algae and (3)
effects due to exclusion of other herbivorous
fish.
Herbivorous fish are generally found to have a
localised impact on algae where as grazing
urchins like Centrostephanus rodgersii appear to
have a major effect on spatial patterns in the
distribution of macroalgae (Jones and Andrew
1990).
Sea urchins are described throughout the
literature as key components of temperate reef
ecosystems as they graze on algae and modify
habitat, influencing the distribution and
composition of subtidal reef communities
(Australian Government 2005). Several species
of urchins exist in Victoria but only the black
urchin Centrostephanus rogersii causes ‘urchin
barren’ or ‘white rock’ habitat which is devoid
of macroalgae (Australian Government 2005).
Urchin barrens are restricted to the far-east of
Victoria where C. rogersii is present in high
abundance (Keough and Butler 1996; O'Hara
2000b; Ferns 2003; Department of the
Environment and Heritage 2005). Several
studies have been conducted on the creation,
maintenance and effect of urchin barrens
elsewhere in southern Australia and New
Zealand (Andrew and MacDiarmid 1991;
Andrew 1993; Andrew and O'Neill 2000). In
Victoria, urchin barrens have been described at
Cape Howe MNP (Ball and Blake 2007b). The
presence of C. rogersii barrens represents one of
the major differences between the Perionian and
Temperate reefs literature review
28
Flindersarian biogeographical provinces
(Connell 2007)
C. rodgersii and another barren forming urchin
Evenchinus chloroticus appear to have a major
impact on habitats, and patterns in the
abundance and foraging of other species.
Herbivorous fish species such as Odax and
Parma victoriae were found to be influenced by
the grazing of both of these urchin species, but
the authors conclude that it is unknown if this
finding is the representative of all sea urchinfish species interactions (Jones and Andrew
1990). Andrew and Stocker (1986) studied the
microhabitat, occupancy, dispersion and
movement of E. chloroticus. The study found sea
urchins were positively associated with
encrusting coralline algae but movement with
regards to algae needs to be further investigated.
Andrew (1993) investigated the effect of
availability of shelter on the foraging behaviour
of C. rogersii, and on the abundance of
invertebrates and algae. The study found that
urchin recruitment was not restricted to high
shelter areas, but barren creation was heavily
reliant on shelter and urchin grazing caused
reductions in the density of foliose algae and
limpets. Andrew and O'Neill (2000) mapped
shallow subtidal habitats in New South Wales
and estimated the coverage of C. rogersii barren
habitat. The study found that the extensive
coverage of the barren habitat in New South
Wales is likely to limit the productivity of the
abalone industry and the development of a sea
urchin fishery may have large impacts on
habitat representation on near shore reefs. A
commercial fishery is currently developing for
sea urchins in Victoria with harvesting of the
white sea urchin Heliocidaris and C. rogersii
recording a catch valued at $191,199 in 2003/04
(Australian Government 2005).
In New South Wales, Gillanders and Kingsford
(1998) found small Achoerodus viridis in greater
numbers in kelp beds than in adjacent urchin
barrens, although the species appeared to be
flexible in its use of habitats on reefs, suggesting
that populations would survive readily if more
urchin barrens were created in an area . In
north-eastern New Zealand, some species of reef
fish have shown similar patterns, while others
were more closely linked to kelp or urchin
barrens (Anderson and Millar 2004). Parma
microlepsis in NSW have been shown to exhibit
exactly the opposite pattern (Holbrook et al.
1994).
Larval supply/recruitment
Like intertidal habitats, larval supply,
connectivity and recruitment are key processes
structuring communities on sub-tidal reefs
(Keough and Swearer 2007).
Recruitment processes in blacklip abalone,
Haliotis rubra, were related to regional
hydrodynamics in eastern Victoria (McShane et
al. 1988). The study suggested that dispersal of
abalone larvae is primarily local. Results
indicated that local reef topography could
sufficiently attenuate currents for larvae to stay
on the parent reef for the full 3 -7 day pelagic
period.
Hunt (2007) investigated the ecological
determinants for recruitment in a reef-associated
planktivorous fish in Port Phillip Bay, the
southern hulafish Trachinops caudimaculatus. The
study surveyed the density of fish populations
and various microhabitat characteristics and
analysed them together to detect relationships.
Microhabitat characteristics explained 85% of
the spatial variation in recruit density, with food
accessibility characteristics in the form of high
plankton supply and low suspended sediment
being the most important for recruitment.
Jenkins and Wheatley (1998) studied the
recruitment of fish in shallow sub-tidal habitats
in Port Phillip Bay and found that while most
emphasis in the past has been placed on
seagrass beds as nursery areas for juvenile fish,
reef-algal habitat showed similar levels of
recruitment to seagrass habitats for a number of
species such as King George whiting, Sillaginodes
punctata. Juveniles of most species commonly
thought to use seagrass as a nursery area were
also found to recruit to shallow subtidal reef.
Exceptions were relatively specialised groups
such as pipefish.
Although the number of studies to date in
Victoria on recruitment to subtidal reefs is
limited, the are currently a number of studies
underway addressing issues of larval supply,
connectivity and recruitment of fish to subtidal
reefs in Port Port Phillip Bay (S.E. Swearer, P.A.
Hamer, pers. comm.).
Competition
In contrast to intertidal reefs, little research has
been conducted on competition on subtidal reefs
in Victoria. Competitive relationships for food
and space are likely to exist between abalone
and sea urchins as both co-occur in reef
ecosystems and feed on drift algae (Jenkins
2004). Urchins may be better competitors where
food is limiting as they can graze directly on
macroalgae and create barrens where abalone do
not occur (Andrew and Underwood 1992).
Jenkins (2004) states that most work would
suggest abalone fluctuations may be affected by
urchin abundances and the reverse is less likely.
However, there are suggestions that when food
is not limiting abalone are superior competitors
for space, such as in crevices.
At Popes Eye in Port Phillip Bay, research on
Parma victoriae found that territory size was
related to intraspecific interactions, with the size
of territories increasing considerably when
neighbours were removed. No correlation
existed between territory size and fish body size
or age, and no relationship was found between
territory size and food abundance following
experimental manipulations of algal food
abundance (Norman and Jones 1984). P. victoriae
with larger territories do, however, have a wider
variety and abundance of food types as larger
territories typically include a wider range of
microhabitats (Jones and Norman 1986).
Predation
Predation is a top-down process that has been
linked to trophic cascades in subtidal
ecosystems. This occurs where removal of large
predators from the reef ecosystem leads to an
increase in lower trophic levels that can have
profound effects on the ecosystem. An example
of this occurs in New Zealand where removal of
snapper (Pagrus auratus) and rock lobster (Jasus
edwardsii) leads to a proliferation of urchins and
subsequent barren formation (Shears and
Babcock 2002). When marine protected areas
were declared, there was a subsequent increase
in lobsters and snapper, and a corresponding
growth of kelp forests (Shears and Babcock
2002).
Rock lobster are considered to be a keystone
species on temperate reefs in Victoria because
their feeding activities can have a significant
effect on the structure of reef ecosystems
(Jenkins et al. 2005b). Adult rock lobsters are
carnivorous and feed mostly at night on a
variety of bottom dwelling invertebrates such as
molluscs, crustaceans and echinoderms
(Department of Primary Industries 2009b). In
Tasmania, the spread of Centrostephanus rodgersii
barrens may be related both to climate change
(facilitating a range extension) and heavy fishing
of rock lobsters allowing urchin populations to
increase and lead to barren formation (Connell
2007).
Temperate reefs literature review
29
The few studies of predation on subtidal reefs in
Victoria have focussed on the 11 arm sea stars
Coscinasterias muricata. Day et al. (1995) found
that C. muricata fed predominantly on mussels
and to a lesser extent other molluscs and
invertebrates. Abalone were rarely eaten except
where aggregations were found eating abalone
when other prey were scarce. Abalone were
found to have a number of behavioural
responses that increased the likelihood of
escaping predation. McShane and Smith (1986)
recorded very high mortality of tagged abalone
as a result of predation by a dense aggregation
of C. muricata on a subtidal reef in Port Phillip
Bay.
Major commercial fisheries
Subtidal reefs support the two most valuable
commercial fisheries in Victoria: abalone and
rock lobster. The Victorian abalone fishery
commenced in 1962 and includes blacklip,
Haliotis rubra and greenlip, H. laevigata species.
Abalone are the most valuable commercial
fishery in Victoria, worth currently about $28.5
million (Department of Primary Industries
2009a). The rock lobster, J. edwardii fishery is the
second most valuable commercial fishery in
Victoria after abalone. The 2008/09 catch was
valued at about $14.3 million (Department of
Primary Industries 2009b).
After a planktonic larval stage, both abalone and
rock lobster settle to a benthic rocky reef
existence on coastal reefs throughout Victoria
(McShane 1999; Department of Primary
Industries 2009b). Abalone are found on
Victorian reefs to depths of 60 m but they are
most commonly found in shallow water less
than 10 m deep (Harry Gorfine pers. comm,
McShane 1999). Greenlip abalone are patchily
distributed west of Wilsons Promontory whilst
blacklip abalone have a relatively consistent
distribution,, extending throughout Victoria
(Harry Gorfine Pers. comm.). Southern rock
lobster are found to depths of 150 metres, with
most of the commercial catch coming from
inshore water less than 100 metres deep
(Department of Primary Industries 2009b). The
abundance of rock lobster decreases from west
to east in Victoria reflecting a decreasing area of
suitable rocky reef habitat (Department of
Primary Industries 2009b). The ecology of
abalone and rock lobster has been investigated
in several studies revealing relationships with
biotic and abiotic features of Victorian temperate
reef habitats.
Temperate reefs literature review
30
Abalone
A number of studies have been conducted in
Victoria on abalone movement and dispersal,
particularly in terms of implications for
abundance and mortality estimates for stock
assessment purposes. Abalone were found to
move and re-aggregate after removal by
experimental fishing, potentially affecting
estimates of abundance (Officer et al. 2001).
Another study followed dispersal of tagged
abalone from experimental plots (Dixon et al.
1998). Dispersal contributed 40-60% of tag
disappearance and potentially has a significant
affect on mortality estimates. The magnitude of
movements varied with habitat quality and is
potentially affected by fishing (Dixon et al. 1998).
Jenkins (2004) summarised the ecosystem effects
of abalone fishing by reviewing papers that
investigated interactions between abalone and
the Victorian temperate reef ecosystem
including feeding, competition, commensalism,
predation and parasitism. Abalone appear not to
have a structurally important role in the reef
ecosystem due to their passive feeding on drift
macroalgae (Shepherd and Clarkson 2001).
Jenkins (2004) concluded that the literature
suggests a low effect of abalone fishing on the
ecosystem in comparison to other fishing
activities, such as trawling and dredging.
However, this should not circumvent rigorous
experiments into the impacts of the fishery and
investigation into ecological interactions.
Following this conclusion, Jenkins et al. (2005a)
analysed abalone/ecosystem monitoring datasets from the Abalone Assessment Monitoring
program conducted by DPI and the Subtidal
Reef Monitoring Program conducted by Parks
Victoria/DSE to identify ecological interactions
and potential indicator species for abalone. The
taxon that showed positive correlation over the
broadest geographical area was encrusting algae
or “pink rock”, which abalone larvae are
dependent on for settlement (Daume et al. 1999).
Considering abalone undergo limited dispersal
(Prince 2003), Jenkins et al. (2005a) described the
correlation of abalone and crustose coralline
algae as a reasonable expectation.
Rock lobster
A recent project by Ball et al. (in prep) used
underwater video techniques to (1) describe
critical habitat of rock lobsters and (2) identify
potential interactions between rock lobster and
other species. A total of twelve fixed sites were
surveyed using underwater video in the western
and eastern zones of the Victorian rock lobster
fishery. The video habitat data was analysed
with rock lobster catch data to assess
interactions between rock lobsters, habitat and
other species. Results for the analyses
determined that distinct habitat assemblages
could be linked with depth, and a number of
habitat parameters were identified as being
important in determining rock lobster
distributions. These variables were depth,
proportion of continuous rocky reef, percentage
cover of P. comosa, percentage cover of
understorey red algae and percentage cover of
sessile invertebrates. A positive relationship was
found between rock lobster and patchy reef,
which may provide rock lobster with greater
foraging opportunities and protection compared
with continuous reef. The study concludes by
stating that it only investigated the relationship
between physical and biological habitat and
rock lobster catches, but oceanographic
influences such as wave energy and nutrient
rich upwellings may also be linked to rock
lobster distribution and abundance.
Temperate reefs literature review
31
Deep reefs and canyons
The Port Phillip Heads region contains deep reef
habitat and species that are unique within Port
Phillip Bay (Edmunds et al. 2006b). These
habitats and the biota resident there were
surveyed prior to recent channel deepening
activity, primarily using video from ROVs, as
were other deep reef habitats within Port Phillip
Bay which included the Rip (between Point
Lonsdale and Point Nepean), Schnapper Deep,
Portsea Hole, Spectacular Reef and Far Side Reef
(diver video), and at Wilson’s Promontory (drop
video) and Point Addis (towed video) on the
open coast (Edmunds et al. 2006b). This study
described a variety of substrate types in the Rip
and on other deep reefs within Port Phillip Bay,
with deep areas of The Rip having more
complex surface complexity than shallow areas
(Chidgey 2006). Deep reefs in Port Phillip Heads
contain a range of assemblage types, some of
which are different to all other reefs surveyed
within and outside Port Phillip Bay, and are
mainly dominated by encrusting sponges and
hydroids (Edmunds et al. 2006b).
Many of the factors and processes responsible
for shaping assemblages on deep reefs are the
same as those that occur on shallow subtidal
and intertidal reefs (see previous discussion).
For example, substrate angle can effect the
distribution of sponges on deep reefs, and
sponge diversity also varies with depth
(Maldonado and Young 1996).
Edmunds et al. (2006b) discussed the biology of
organisms on deep reefs in Victoria, with the
assumption that this was broadly similar to
species studied on shallow reefs, and described
a variety of physical and ecological drivers
related to the ecology of deep reefs in Victoria,
particularly those in and around Port Phillip
Heads. These processes often interact with each
other and can therefore exert a variable
influence on species found on and around deep
reefs. Very little, if any, research has been
conducted in Victoria on these patterns and
processes and that done elsewhere in deep
water has tended to concentrate on more widely
distributed soft sediments (Levin et al. 2001) This
is due in part to the perceived difficulties
associated with sampling and conducting
manipulative experiments on deep reefs.
A summary of the likely factors involved in
shaping the structure of assemblages are laid out
in Tables 5-7 following a recent review by
Temperate reefs literature review
32
Edmunds et al. (2006b) and several of these
drivers are discussed below.
Physical drivers
As light intensity diminishes with depth,
animals replace plants as the dominant life form
(Keough and Butler 1996; O'Hara et al. 1999).
Algae, including encrusting red coralline algae
that are often found on deep reefs, is typically
replaced by attached invertebrates such as
sponges, bryozoans, corals and ascidians
(Edmunds et al. 2006b).
Recent surveys carried out on a number of deep
reefs in Victoria have described how the
dominant biota shows zonation with depth (Ball
et al. in prep). Phyllospora comosa (crayweed) was
observed to depths of ~29 m, and typically grew
with the kelp Ecklonia radiate which was the
dominant species observed to depths of ~ 48 m.
E. radiata was the dominant species at the
majority sites, while P. comosa alone or with E.
radiata was an important component of the biota
at a range of sites at Portland, Warrnambool,
Warrnambool West, Port Fairy West and from
Ocean Grove to Torquay. At 60–80 m, sessile
invertebrates began to replace these dominant
large brown macroalgae.
Ryan et al. (2005) described how submarine
canyon and shelf topography can influence
physical–biological coupling. Considerable
amounts of water circulate through deep water
canyons due to a range of factors including
temperature gradients, eddies, oceanic currents
and upwelling (Breaker and Broenkow 1994).
Such movements of large bodies of water have
been shown to drive the movement of relatively
large quantities of sediments and sand (Hume et
al. 2000), which has ramifications for organisms
that live in these environments. For example,
high levels of organic matter in sediments may
influence species richness (Curdia et al. 2004),
excess sediments may cover and destroy
assemblages (Okey 2003), or nutrient transport
may be affected as in Perth Canyon off Western
Australia where interactions occur between the
Leeuwin Current and upwelling (Hume et al.
2000; Rennie et al. 2009).
Interactions between topography and currents
can also influence the distribution of deep reef
fishes. On deep shelf areas where cold water is
uplifted from slopes, areas can exist where
waters are nutrient rich and key prey are
abundant (Bax and Williams 2000), for example,
at the head of Horseshoe Canyon where
productive fishing grounds are found (Bax and
Williams 2001). Strong currents may effect
certain species due to their physical structure,
and at a smaller scale, cracks, crevices and holes
of varying sizes can provide microhabitat and
influence the distribution of species in relation
to water movement (Beaman et al. 2005). For
example, currents in Port Phillip Heads are
likely to have a greater effect on erect sponge
species than encrusting species, as has been seen
elsewhere (Bell and Barnes 2000; Bell et al. 2002),
and may result in certain sponge species being
found in encrusting rather than erect forms
(Abdo et al. 2008).
(Hypoplectrodes nigroruber). Schools of barber
perch (Caesioperca razor) are replaced by the
related butterfly perch (Caesioperca Lepidoptera)
(O'Hara et al. 1999). While fish present on
shallow subtidal reefs include algavores,
omnivores and carnivores, those on deep reefs
are typically carnivorous as algae are typically
not present at depth. Although common on
rocky reefs, sponges, hydrozoans, anthozoans,
bryozoans, and ascidians are thought to be
largely unpalatable to reef fish (Russell 1983); it
is therefore likely that fish at these depths are
feeding on associated mobile invertebrate fauna.
Sponge assemblages are more stable on solid
reef rather than boulder substrates, but are more
diverse on boulders (Carballo and Nava 2007).
This may be due in part to the fact that sponges
on boulders can occur on surfaces at a variety of
angles which produces microhabitats with
varying degrees of shading and water
movement. Boulders may also be more prone to
disturbance, and intermediate levels of
disturbance are generally thought to increase
diversity (Sousa 1979).
Competition
Sponges in deep water have been shown to be
important competitors for space (Suchanek et al.
1983) and may live for long periods (Ayling
1981), which is likely to effect the distribution of
other sessile species on deep reefs/canyons such
as hydroids and bryozoans. The competitive
ability of sponges is also increased by their
ability to regrow swiftly after injury (Duckworth
2003), and the ability of some encrusting species
to grow quickly when free space becomes
available yet grow slowly when it is limited and
they are undisturbed (Roberts and Davis 1996).
Ecological drivers
Herbivory and grazing
Verges et al. (2009) found that herbivory differs
with depth from shallow to deep reefs and has
an important effect of the vertical distribution of
algae to ~40 m.
In deeper areas of reefs, sessile species such as
sponges provide structure and diversity, and
may provide either direct or indirect resources
for fish (Tissot et al. 2006). Species richness and
fish densities vary from shallow to deep reefs
(Love et al. 2009), and heterogeneous habitats,
substrate type and complexity have been shown
to influence the distribution of fish assemblages
on deep reefs (Yoklavich et al. 2000; Anderson et
al. 2009).
Habitat structure
The distribution of fish fauna is governed by
biologically formed habitat structure as well as
by food, and begins to change with the loss of
the kelp-associated wrasses and leatherjackets,
and the appearance of fishes such as boarfishes
(family Pentacerotidae), splendid perch
(Callanthias australis) and banded seaperch
While sessile species may compete for free
space, they can also have positive interactions
on each other. For example, Abdo et al. (2008)
found that neighbouring sponge species
protected each other from environmental
impacts.
Larval supply/recruitment
Sponges can reproduce asexually and spread
through the water column or across substrates
or neighbouring individuals. They can also
reproduce sexually, releasing larvae, propagules
or gametes into the water column. While asexual
reproduction and growth may be of primary
importance to sponges (Jackson 1986), dispersal
and recruitment are also important in sustaining
populations and shaping assemblage structure
(Keough 1988; Underwood and Keough 2001).
The presence of large sessile species may effect
larval deposition and food availability of
adjacent species by altering fluid dynamics in
the immediate area (Nowell and Jumars 1984).
Temperate reefs literature review
33
Knowledge gaps
Recent and current efforts to map reef biota in
relation to substrate and habitat type are
commendable and are providing a wealth of
information on local systems. However, the
majority of information on reefs in Victoria
provides only a “snapshot” of assemblage
structure and the distribution of species at one
particular time of year or sampling site. What is
lacking are examinations of temporal and spatial
variation in the structure of assemblages on
different reef types, and in many cases more
basic information on which species occur and
their basic ecology and behaviour. While general
patterns that shape reef communities have been
examined world wide, and to some extent
locally, knowledge of processes and species
interactions effecting local species are less well
understood. This is problematic as this
information is likely to be important in helping
us to understand what is shaping many of these
communities.
the physical and ecological processes affecting
the system.
While understanding what species are in
assemblages and how and why they vary is
important, we also need to conduct research that
examines marine ecosystems in a broader
geographical context across southern Australia.
Victoria’s placement at the convergence of
several biogeographical provinces (i.e.
Flindersian, Peronian and Maugean) make it the
ideal place to conduct studies on a variety of
questions at these broader scales.
The amount of research and level of
understanding has been greatest for intertidal
reefs, is lower for subtidal reefs, and is very
limited for deep and canyon reefs. This is
probably a reflection of ease of accessibility and
logistical constraints.
Research on subtidal reefs in Port Phillip Bay has
been fragmentary, and there is a poor
understanding of the drivers influencing the reef
communities and how these differ from the open
coast. Further research on the physiological and
ecological drivers affecting subtidal reefs in Port
Phillip Bay is required.
Of special interest are the deep reef and canyon
habitats of Port Phillip Heads. This area has a
unique combination of deep water, strong
currents, and coastal sedimentary processes. The
sessile invertebrate communities are well
developed but their uniqueness is uncertain.
Detailed studies of taxonomy and assemblage
structure are required along with research into
Temperate reefs literature review
34
In the following we outline knowledge gaps that
exist for each of the reef systems that we have
discussed in this review.
Intertidal reefs
Much of the work on reefs in Victoria that has
examined patterns and processes such as
competition and settlement has been conducted
on intertidal reefs.
There are, however, still a number of knowledge
gaps that include:

Basic information on the biology and ecology
of a range of species

Quantitative examinations of spatial and
temporal variation of assemblages on many
shores, both in regional Victoria and within
Port Phillip Bay

Potential impacts of sedimentation, pollution
etc on assemblages

Linkages with adjacent assemblages from
different habitats
-
e.g. how highly mobile predators such
as fish and birds utilise intertidal rocky
reef habitats

The ecology and importance of species that
are becoming targets of food and bait
collection, but which have not traditionally
been collected in Victoria

Interactions between coastal oceanography
and intertidal communities, i.e. effects of
upwelling such as on the Bonney coast

How water level rises will affect reef
assemblages
-
e.g. are there low lying reefs that will be
completely covered?
Subtidal reefs
Plummer et al. (2003) summarised the habitat
types and species found on shallow reefs within
marine national parks and sanctuaries in
Victoria. For many sites, they noted that while
anecdotal or qualitative data existed, quantitative
data were often lacking.
It has been suggested that on shallow reefs in
Victoria, surrogates such as dominant vegetation
and region may be more accurate for predicting
assemblage structure, at least for algae and
invertebrates (O'Hara 2001), than substrate alone,
and indeed recent predictive models appear to be
satisfactory for linking substrate and depth with
macroalgae (Holmes et al. 2008).

Determining sources of larval recruitment
and causes of variability in recruitment

Standardising commercial catch rates and
determining stock status
Gaps in our knowledge about shallow reefs,
including those in Port Phillip Bay and along
Victoria’s open coastline include:

Investigating the effect of seawater
temperature of rock lobster ecology
including the effect of the Bonny upwelling
nutrient rich intrusion into western Victoria.

Basic information on what species are found
and assemblage structure
-
Algae
-
Sessile invertebrates
-
Mobile invertebrates
-
Fish

Quantitative data examining patterns of
spatial and temporal variation

Relationships with assemblages from
adjacent habitats, e.g. sand and seagrass
patches

The impact of introduced species

Relationship with coastal oceanography such
as the Bonney upwelling

Processes driving the ecology of sheltered
subtidal reefs in Port Phillip Bay compared to
exposed reefs on the open coast

Very little research has been carried out on
invertebrates on subtidal reefs of Port Phillip
Bay

Processes relating to the export of drift algae
and detritus from subtidal reefs and the
subsidy to other habitats (Vanderklift and
Wernberg 2008; Crawley et al. 2009).
Key knowledge gaps for commercial fisheries of
abalone and rock lobster exist and have been
identified for the purposes of this literature
review.
Abalone research is required (Harry Gorfine
pers. comm.) into:

The quantification of stock-recruitment
dynamics of the fishery

Age validation of individual abalone
specimens and

Modelling the fishery as an entire system
including management performance
measures.
Rock lobster research is required (David Reilly
pers. comm.) into:
Deep reefs and canyons
Little is known about deep reefs in Victoria, or
the biology and ecology of organisms that live on
them, due in part to difficulties associated with
conducting observational work or manipulative
experiments in situ. Almost nothing is therefore
known about how various biotic factors influence
the distribution, survival, etc. of species found on
deep reefs, except for those species such as rock
lobster and abalone, where some information is
available due to directed research on these
valuable fisheries species (see previous
discussion of these fisheries).
Without basic information on the life history of
organisms and details of how they interact with
each other and the environment, it may not be
possible to determine to what degree potential
disturbance events have an impact on individual
species or assemblages of species.
Gaps in our knowledge about deep reefs,
including those in Port Phillip Heads include:

Basic information on what species are found
and assemblage structure; this is true for
both sessile species and associated mobile
invertebrates

Spatial and temporal patterns of distribution
of species on deep reefs through Victoria

Which species are ecologically important

Rarity of species and assemblages in a wider
geographical sense

Processes determining the structure of
communities in deep reefs and canyons,
especially in Port Phillip Heads (current,
sediment
loads,
plankton/seston
concentration)

Identification of species and assemblages that
are vulnerable to natural and human
disturbance/impacts

The uniqueness of sessile invertebrate species
in Port Phillip Heads deep reef and Canyon
compared to other deep reef and canyon
habitats in Victoria
Temperate reefs literature review
35

Trophic linkages between deep reefs and
canyon communities in Victoria and other
marine and terrestrial communities.
Due to this knowledge gap, key questions that
we are currently unable to answer with
confidence include:

How the removal, increase or introduction of
species may effect assemblage structure
Temperate reefs literature review
36

How communities change on a temporal
basis, whether that be seasonally or between
years

How assemblages may recover from pulse or
press disturbance events

How climatic or oceanographic changes may
influence the distribution and health of
assemblages.
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Appendix 1 Tables
Table 1. Six bioregions relevant to Victoria as presented in Interim Marine Coastal Regionalisation of
Australia Technical Group, IMCRA (2006)
 Otway
Location: Cape Jaffa to slightly north of Apollo Bay and including King Island
Very steep to moderate offshore gradients. High wave energy. Currents generally slow, but moderately
strong through entrance to Bass Strait. Cold temperate waters subject to nutrient rich upwellings.
 Central Victoria
Location: Cape Otway to west of Wilsons Promontory to Flinders Is.
Very steep to steep offshore gradients dominated by cliffed shorelines. Sea-surface temperature is
representative of Bass Strait waters. Moderate wave energy.
 Flinders
Location: Eastern Entrance to Bass Strait and including Wilsons Promontory,
Rapid changes in offshore gradient. Granitic coastline exposed to submaximal swells on east-facing
shores of Flinders Island and moderate to low swells elsewhere. Sandy beaches of moderate length with
seagrass beds prevalent in shallow water. High tidal range » 3 m and strong tidal currents. Sea-surface
temperature is representative of Bass Strait waters. Waves highly variable.
 Twofold Shelf
Location: East of Wilsons Promontory and north to Tathra (36°48’S),
Submaximally exposed coastline with long sandy beaches broken by rocky headlands and numerous
coastal lagoons. Moderate tidal range ~ 2 m. Mean annual sea-surface temperature reflects the influence
of warmer waters brought into Bass Strait by the East Australian Current. Variable wave energy.
 Victorian Embayments
Location: Victorian bays, inlets and estuaries e.g., Port Phillip Bay
Confined bodies of water that total in excess of 3000 km 2 and individually vary in size from 1950 km2 to
less than 1 km2. They are generally basin-shaped, less than 25 m deep, have limited fetch and are
dominated by depositional environments.
 Central Bass Strait
Location: Central offshore region of Bass Strait
The region is about 60,000 km2 in size and lies in the central area of Bass Strait. The sea floor is shaped
like an irregular saucer with water depth varying from about 80 m at its centre to 50 m around the
margins. The substrate of central area is mainly mud. Tidal velocities vary from <0.05 ms–1 in the central
area to as high as 0.5 ms–1 at the margins where the islands and promontories form the western and
eastern entrances to Bass Strait. Water mass characteristics are complex and vary seasonally representing
the mixing of the different water masses present on western and eastern side of the Strait.
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Table 2. Interim intertidal marine habitat (MHC) categories for Victoria, Ferns et al. (2000).
Table 3. Interim shallow subtidal (0 – 2.5 m) marine habitat (MHC) categories for Victoria, Ferns et al.
(2000).
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Table 4. Primary shallow habitat classification scheme as presented in Ball et al. (2000).
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Table 5. Potential physical drivers responsible for shaping deep reef assemblages in Victoria.
Summarised from Edmunds et al. (2006b)
o
o
o
o
o
Sedimentation
 Sediments may naturally settle and cover rock
 Strong wave action and currents may expose rocky reefs and keep them free of sediments
 With increasing depth, wave action decreases and sedimentation may increase
 Sediment transport from adjacent habitats may be such that reefs are heavily sediment
effected even in the presence of strong currents
 Reef steepness and relief can influence sediment build-up, i.e., more vertical surfaces are less
likely to accumulate sediment
Geological structure
 The type of rock and geological history can influence the size and structure of reefs
 Surface roughness can effect settlement, increase surface area and influence water movement
over the rock surface and therefore the transport of food, oxygen, nutrients etc (Thomas and
Atkinson 1997)
 Reef structure may influence how sediments and loose rocks smother or scour reef surface
and sessile organisms
Water movement, including waves and currents
 May erode reefs
 Can impact sediment deposition and transport
 Influence the type and distribution of organisms on the reef
Light climate
 High levels of sedimentation and increasing depth can influence light levels
 Reef aspect and shading at a variety of scales influence light availability
 Where light levels are sufficient, algae out compete sessile invertebrates on rocky reefs, e.g.,
sponges and corals
Depth
 Interacts with factors listed above
 Deep reefs tend to be dominated by sessile invertebrate assemblages, e.g., sponge gardens
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Table 6. Potential ecological drivers responsible for shaping deep reef assemblages in Victoria.
Summarised from Edmunds et al. (2006b)




Sessile invertebrates spend their adult life attached to the substratum
o Species such as sponges, may exhibit different physical characteristics depending on
environmental conditions
o Sponges require suitable seabed for them to attach and resist the actions of currents and
waves and are often found in underwater tunnels and notches that are surrounded by
kelps (Keough 1999)
o Sponges, cnidarians, bryozoans, and colonial ascidians dominate deep reef assemblages
o The age of individuals and colonies on deep reefs in Victoria cannot be assumed to be
correlated with size
o There may be complex interactions which will not be known without study of the basic
ecology of species,
o Sessile clonal species such as sponges often release actively swimming larvae which
travel relatively short distances before recruitment, even in high currents , while some
species produce crawling larvae
o Larvae can react to a range of stimuli such as light levels and gravity (Keough et al. 1996)
Food availability effects the distribution of species, and while mobile invertebrates and fish can
move to areas where food is available, sessile species are predominantly suspension feeders that
require food in the form of plankton that is available in the water column.
Little is known about the small crustaceans, molluscs etc that are associated with sessile
communities on Victoria’s deep rocky reefs.
o Therefore their role in shaping assemblage structure is not understood.
Little is known about the behaviour and diet of predatory fish on Victoria’s deep reefs, or their
impact on assemblage structure
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Appendix 2 Conceptual Models
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Figure 1. Key relationships and drivers on intertidal reefs in Victoria
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Figure 2. Key relationships and drivers on subtidal reefs in Victoria
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Figure 3. Key relationships between invertebrates on subtidal reefs in eastern and western Victoria
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Figure 4. Key relationships and drivers on deep reefs in Victoria
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Figure 5. Key relationships and drivers on canyon reefs in Victoria
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