This is the published version:  
 
Hirst, Alistair J., White, Camille A., Green, Corey, Werner, Guy F., Heislers, Simon and Spooner,
Daniel 2011, Baywide anchovy study sub‐program : milestone report no. 4 (2011) Department of
Primary Industries, Queenscliff, Vic.


Available from Deakin Research Online: 
 
http://hdl.handle.net/10536/DRO/DU:30059164



Reproduced with the kind permission of the copyright owner.

Copyright : 2011, Department of Primary Industries

Baywide Anchovy Study Sub-
Program

Milestone Report No. 4 (2011)

No. 150
October 2011
Baywide Anchovy Study Sub-Program

Milestone Report No. 4 (2011)

Alastair J. Hirst, Camille A. White, Corey Green, Guy F. Werner,
Simon Heislers, and Daniel Spooner

October 2011

Fisheries Victoria
Technical Report Series No. 150
If you would like to receive this
information/publication in an
accessible format (such as large
print or audio) please call the
Customer Service Centre on:
136 186, TTY: 1800 122 969,
or email
customer.service@dpi.vic.gov.au
Copyright  The State of Victoria, Department of
Primary Industries, 2011.
This publication is copyright. No part may be
reproduced by any process except in accordance
with the provisions of the Copyright Act 1968.
Authorised by the Victorian Government, GPO
Box 4440, Victoria 3001.
Printed by Fisheries Victoria, Queenscliff,
Victoria

Published:
Preferred way to cite:
Fisheries Victoria Department of Primary
Hirst, A.J., White, C.A., Green, C., Werner, G.F.,
Industries, Queenscliff Centre PO Box 114,
Heislers, S., and Spooner, D. (2011). Baywide
Queenscliff, Victoria 3225 Australia.
Anchovy Study Sub-Program. Milestone Report
No. 4 (2011). Technical Report No. 150.
Department of Primary Industries, Queenscliff,
General disclaimer Victoria, Australia. 36 pp.
This publication may be of assistance to you but
the State of Victoria and its employees do not
guarantee that the publication is without flaw of
ISSN 1835-4785
any kind or is wholly appropriate for your
particular purposes and therefore disclaims all ISBN
liability for any error, loss or other consequence
which may arise from you relying on any
information in this publication.

ii
Executive Summary
The abundance of the anchovy, Engraulis
australis, in Port Phillip Bay (PPB) during winter
2011 was determined using a demersal trawl net
and sonar at 67 sites.
A total of 231,711 anchovies weighing 795 kg
were caught during the survey. Anchovies were
primarily recorded throughout the central and
eastern part of PPB in an area covering
approximately 762 km2. The total biomass of
anchovies in PPB during May–June 2011 was
estimated to be as 523.8 ± 27.6 (se) tonnes.
Anchovy aggregations were patchily distributed
throughout PPB in May–June 2011. Dense
patches of anchovies were detected offshore at
Sandringham, Aspendale, Frankston and
Portarlington, and throughout the deeper,
central regions of PPB. Anchovy populations
have displayed a consistent spatial distribution
in PPB over the duration of the surveys from
2008 to 2011. This spatial pattern probably
reflects variation in zooplankton availability, as
a food source, at a large spatial scale in PPB.
The diet of juvenile anchovy were analysed by
examining the stomach contents of fish
collected in the 2011 survey. Stomach contents
were dominated by digested material that was
largely unidentifiable, although the identifiable
component of the stomach contents was
dominated by calanoid copepods.
Nocturnal feeding activity was analysed by
examining trends in stomach fullness over the
night surveys. Fish stomachs tended to be fuller
earlier in the night following sunset, compared
with fish collected during the early morning
prior to sunrise, suggesting that feeding activity
during the night is low.
Most anchovies in PPB during May–June 2011
belonged to the 0+ and 1+ age-cohorts, and few
anchovy older than 1+ were recorded. Adult fish
(>1+) are believed to move out of PPB into Bass
Strait in late summer/autumn, returning in
spring to spawn.
The spatial distribution of anchovy age classes
in PPB varied across all annual survey years
(2008–2011), with no consistent pattern.
Spawning dates calculated from otolith daily
growth increments ranged from September to
April, with little variation between years. Peak
spawning period, based on the modal length of
the 0+ age-cohort in each year, was estimated to
occur from late November to early January.
Conclusions
The distribution and age-class structure of
anchovy in PPB during May–June 2011 were
consistent with the results of previous annual
surveys (2008–2010) and the known attributes of
anchovy populations in other parts of Australia.
The results of the 2011 survey are also consistent
with the conceptual ‘population and life history’
model developed for anchovy in PPB (Hirst et al.
2010). This model is further complemented by
additional information on diet, feeding activity
and spawning season generated during 2011.
The model features the following characteristics:
• PPB is an important spawning and nursery
area for anchovy within its Victorian
distribution
• Anchovy population in PPB is dominated by
juvenile (0+ and 1+ ) anchovy during May-
July, with few adult (>1+) fish present
• It is believed that adult fish move out of PPB
following spawning to occupy offshore
waters during winter months
• Anchovy in PPB exhibit an elongated
spawning season, extending from September
to April, with peak spawning estimated to
occur from late November to early January
• Calanoid copepods were the major
component of the diet of juvenile anchovy
during 2011
• Anchovy are an important food source for
many predatory fish and seabirds, and
provide an important trophic link in PPB
between zooplankton productivity and
higher trophic levels.
iii
Table of Contents

Executive Summary............................................................................................. iii

Conclusions...............................................................................................................................................................iii

Introduction ............................................................................................................ 1
This report ...................................................................................................................................................................1

Project Design and Methods ............................................................................... 2

Sampling .....................................................................................................................................................................2
Anchovy diet...............................................................................................................................................................2
Exceptions ...................................................................................................................................................................2

Results...................................................................................................................... 5
Abundance and biomass ..........................................................................................................................................5
Population structure..................................................................................................................................................5
Distribution ................................................................................................................................................................5
Diet ...............................................................................................................................................................................5
Other pelagic species ................................................................................................................................................6
Abundance and Biomass .......................................................................................................................................6
Distribution..............................................................................................................................................................6

Discussion............................................................................................................... 7
Population structure..................................................................................................................................................7
Diet ...............................................................................................................................................................................8
Distribution ................................................................................................................................................................8
Abundance ..................................................................................................................................................................9
Interactions with other species................................................................................................................................9
Conclusion ................................................................................................................................................................10

Acknowledgements............................................................................................. 13

References ............................................................................................................. 14

Appendix 1. Materials and Methods ............................................................... 17

Abundance and biomass ........................................................................................................................................17
Distribution ..............................................................................................................................................................17
Age..............................................................................................................................................................................17

v
 Annual ................................................................................................................................................................... 17
Daily increment .................................................................................................................................................... 17
Anchovy diets .......................................................................................................................................................... 17

Appendix 2 Results ............................................................................................. 20

Quality Control........................................................................................................................................................ 20
Anchovies ................................................................................................................................................................. 20
Abundance and biomass ..................................................................................................................................... 20
Distribution ........................................................................................................................................................... 20
Population structure ............................................................................................................................................ 20
Diet ......................................................................................................................................................................... 21
Other pelagic species.............................................................................................................................................. 21
Abundance and Biomass..................................................................................................................................... 21
Distribution ........................................................................................................................................................... 22

Appendix 3 Data Files ........................................................................................ 36

vi
List of Tables
Table 1. Summary of sampling design undertaken in 2011 survey .................................................................... 4
Table 2. Conceptual understanding of anchovy population in PPB gained from CDBMP surveys 2008–
2011 and the literature..................................................................................................................................... 12
Table 3. Number of anchovies aged at each site for annual PPB surveys in 2008–2011. ............................... 19
Table 4. Parameters for linear regressions between length (mm) and daily age (days) for anchovy aged
using daily growth increments in 2008, 2009, 2010 and 2011 (age (days) = slope*length + constant). . 19
Table 5. Vertical distribution of clupeoid fish within 5 m of the seabed: cumulative % of fish echoes at 1 m
intervals. *n=1014 images............................................................................................................................... 20
Table 6. Mean biomass of anchovies /trawl tow (+ se) for the region identified in the 2011 survey (Table 2,
Figure 2), approximate area occupied by anchovies in PPB, and estimated biomass of anchovies in
PPB (+ se tonnes) in the 2011 survey. ............................................................................................................ 23
Table 7. Mean length (mm) and standard error of anchovies of different ages in PPB in the period June–
July 2008, 2009, 2010 and 2011. N: number of anchovies aged for each age cohort.* .............. 28
Table 8. Peak spawning period estimated from age (days) of 0+ anchovy at the modal length (mm) in
2008, 2009, 2010 and 2011................................................................................................................................ 31
Table 9. Mean length (mm), stomach fullness (%) and number of prey items recorded for fish examined
at 8 sites in May-June 2011.............................................................................................................................. 32
Table 10. Summary of anchovy stomach content analysis undertaken in 2011. ............................................. 32
Table 11. Biomass and abundance of all pelagic species caught in the 2011 survey. ..................................... 34

List of Figures
Figure 1. Fixed sampling sites surveyed using either trawl or sonar in Port Phillip Bay during May–June
2011. ..................................................................................................................................................................... 3
Figure 2. Fixed sites sampled for anchovies during the 2011 annual survey. Map displays fixed sites at
which sonar indicated anchovies were: present and trawled, present and not trawled, or absent
(Hirst et al. 2011)................................................................................................................................................. 4
Figure 3. Total catch (bars), A) catch rate (kg/day), and B) effort (days) for the commercial anchovy
fishery in PPB 1978/79 to 2010/11................................................................................................................... 11
Figure 4. Distribution of anchovy biomass (kg/10 minute tow) in the 2011 annual survey in PPB. Polygon
indicates the approximate area in which anchovies were recorded by trawl or sonar at fixed sites
during 2011. ...................................................................................................................................................... 23
Figure 5. Polygons showing the approximate distribution of anchovies in PPB during annual surveys
conducted in 2008–2011. ................................................................................................................................. 24
Figure 6. Acoustic survey 2011: percentage area of fish echoes within 2 m of seabed for underway
sampling between fixed sites. The polygon shows the primary region of anchovy distribution
inferred from fixed sampling conducted in May–June 2011...................................................................... 25
Figure 7. Age-length frequency distributions of anchovies from annual surveys in PPB during June-July
2008, June 2009, June 2010 and May-June 2011............................................................................................ 26
Figure 8. Length-frequency distributions of anchovies from annual surveys in Port Phillip Bay 2008–2011.
Number is the projected number of anchovies in each size category (1 mm increments) based on the
number of fish measured as a proportion of the total number of anchovy caught at each site (see
Parry et al. 2009). Note, y axes differ between years. The arrow indicates the modal length of the 0+
age-cohort in each year used in calculations of peak spawning period (Table 8)................................... 27
Figure 9. Distribution of age cohorts surveyed in May-June 2011; % 0+, 1+, and 2+ fish aged at each site. 29

vii
Figure 10. Estimated growth based on daily growth increments of anchovy in PPB for 0+ age-cohort fish
collected in 2008, 2009, 2010 and 2011 surveys. Lines = linear regressions between length and age in
each year. Analysis shown in Parry et al. (2009) only includes fish <90 mm length for the purpose of
validating the 0+ age-cohort. .......................................................................................................................... 30
Figure 11. Range of spawning dates (months) encountered for 0+ age-cohort fish aged using daily growth
increments in 2008, 2009, 2010 and 2011. ...................................................................................................... 31
Figure 12. Relationship between mean (± se) anchovy stomach fullness (%) and nominal sampling time
(24 hour scale) for anchovies collected from 31 May–4 June 2011. A quadratic function has been fitted
to the data following Tudela and Palomera (1997). Vertical lines indicate position of sunrise (approx.
7.25 am) and sunset (5.10 pm). ....................................................................................................................... 33
Figure 13. Non-metric multidimensional scaling (nMDS) plot displaying the compositional similarity of
anchovy stomach contents of fish collected at eight sites in 2011. n = 10 fish stomachs site-1 ............... 33
Figure 14. Distribution and biomass (kg/10 min tow) of the three most abundant pelagic species (other
than anchovy) caught during the anchovy survey in 2011. ....................................................................... 35

viii
Introduction
Anchovy, Engraulis australis, are an important
prey item of numerous fish species and of little
penguins in Port Phillip Bay (PPB).
The Baywide Anchovy Study Sub-Program is
described in the Channel Deepening Baywide
Monitoring Programs (CDBMP), Anchovy Study
Detailed Design - CDP_ENV_MD_016 Rev 3
(PoMC 2010). The objective of this sub-program
is to collect data on anchovy abundance,
distribution and population structure that will
fill knowledge gaps and assist in the assessment
of changes observed in other Baywide programs
(PoMC 2010).
This report
This report summarises results for the fourth
annual survey in PPB, examining anchovy
abundance, distribution and population
structure during May–June 2011.
This report contains:
• Estimates of anchovy abundance and
biomass for May–June 2011
• Distribution of anchovies for May–June 2011
• Population structure (length frequency and
age) of anchovies in PPB for May–June 2011
• Distribution of other pelagic species with
relatively high biomass in PPB for May–June
2011
• Discussion of the abundance, distribution
and population structure of anchovy in PPB
for May–June 2011, including comparison
with June–July 2008, June 2009 and June
2010 survey results
• A conceptual ‘population and life-history’
model summarizing the status of current
knowledge on the population structure,
distribution, abundance and ecological
significance of anchovy in PPB.
1
Project Design and Methods
The methods used in the present study are
described below, and in PoMC (2010), Parry and Exceptions
Stokie (2009), Parry et al. (2009), Hirst et al. (2010, An exception to the Detailed Design -
2011) and Appendix 1. CDP_ENV_MD_016 Rev 3 (PoMC 2010).for this
study period is documented in Exception Report
Sampling ER2011–91, and summarised as follows:
Sampling was undertaken at night from 31 May • Only one trawl net was used (instead of
to 4 June 2011 during a new moon so that all two) throughout the 2011 annual anchovy
sampling occurred in near maximal darkness to survey.
minimise possible aggregation of anchovies. This exception has not changed the conclusions
All trawling was undertaken by the “Western reached in this report.
Alliance”, a 22 m commercial fishing trawler.
A fixed-site sampling design was used (Parry et
al. 2009, PoMC 2010), and the location of all sites
is shown in Figure 1. Fixed sites were sampled
using both trawl net and sonar, and sonar
sampling was also undertaken whilst travelling
between fixed sites. The fixed-site sampling
methods employed in this study are dictated by
decision rules summarised in Hirst et al. (2011).
A total of 67 fixed sites were sampled using
trawl and sonar, or sonar only, in 2011 (Table 1,
Figure 2):
• 32 sites using trawl and sonar (including
two zero calibration sites where sonar
indicated anchovy were absent)
• 35 sites using sonar only (including four
sites where sonar indicated anchovy were
present, but which could not be trawled
(Hirst et al. 2011).
The distribution of aggregations of clupeoid fish
(anchovy, sandy sprat, pilchard) within 5 m of
the benthos was measured using sonar,
according to the method described by Parry et al.
(2010).

Anchovy diet
The contents of anchovy stomachs collected at
eight sites in 2011 were examined and analysed
for spatial differences using multivariate
statistics.

2
Figure 1. Fixed sampling sites surveyed using either trawl or sonar in Port Phillip Bay during May–June 2011.

3
Figure 2. Fixed sites sampled for anchovies during the 2011 annual survey. Map displays fixed sites at
which sonar indicated anchovies were: present and trawled, present and not trawled, or absent (Hirst
et al. 2011).

Table 1. Summary of sampling design undertaken in 2011 survey

Sampling method No No of sites No of sites
of (subset of (subset)
sites total)
Sampled all methods 67
Trawl and Sounder 32
Trawled where sounder indicated anchovies 30
Trawled where sounder indicated no anchovies 2
Sounder only 35
Sounder indicated absence of anchovies 31
Sounder indicated anchovies present, but not trawled 4

4
Results
Results for the 2008, 2009 and 2010 annual
surveys were presented in Parry and Stokie
(2009), Parry et al. (2009) and Hirst et al. (2010).
Preliminary results for the 2011 annual survey
were presented in Hirst et al. (2011). Additional
results for the 2011 annual survey are provided
in Appendix 2, and summarised below.
Abundance and biomass
A total of 231,711 anchovies weighing 795 kg
were caught during the 2011 survey.
Anchovies occurred in an area of PPB covering
762 km2 in 2011. The total biomass of anchovies
in PPB during May-June 2011 was estimated as
523.8 ± 27.6 tonnes.
Population structure
Anchovy populations in PPB comprised 0+, 1+,
2+ and 3+ age-cohorts in 2008, 2009 and 2010,
and 0+, 1+ and 2+ age-cohorts in 2011. The
dominant age cohorts in all years were 0+ and
1+. The mean lengths of anchovy in the 1+ and
2+ age-cohorts were slightly lower in 2011
compared with the same cohorts in 2008, 2009
and 2010. The mean lengths of anchovy in the
0+ age-cohort were similar in 2008–2011.
Timing of spawning estimated from the 0+ age-
cohort ranged from:
• September 2007 to February 2008, with peak
estimated late November 2007 (modal length
75 mm), for fish collected in June–July 2008
• September 2008 to April 2009, with peak
estimated early January 2009 (modal length
65 mm), for fish collected in June 2009
• September 2009 to March 2010, with peak
estimated December 2009 (modal length 75
mm), for fish collected in June 2010
• October 2010 to March 2011, with peak
estimated December 2010 (modal length 70
mm), for fish collected in May–June 2011.
Distribution
The distribution of anchovies in PPB during the
May–July survey period was similar in 2008,
2009, 2010 and 2011. Anchovies were found
further north in 2009 than 2008, 2010 or 2011;
and further west in 2010 and 2011 compared
with anchovy distribution in 2008 and 2009.
Anchovy were patchily distributed within the
region in which they occurred in 2011. Dense
patches of anchovies were common throughout
the deeper, central region of PPB and offshore at
Sandringham, Aspendale, Frankston and
Portarlington.
In 2011, there were differences in the spatial
distribution of anchovies of different ages within
PPB:
• 0+ anchovy were distributed throughout
PPB but were more prevalent in the central
part of PPB
• 1+ anchovy were distributed throughout
PPB but were more prevalent around the
periphery of PPB
• 2+ anchovy distribution displayed no
spatial pattern, but comprised only a small
proportion of fish aged at all sites.
The spatial distribution of anchovy age classes
in PPB varied across all years of the survey,
with no pattern evident.
Diet
Stomach contents were analysed for 79
anchovies collected from eight sites across PPB
in May-June 2011. The relationship between %
stomach fullness and nominal time (24 hour)
indicated that stomachs tended to be fuller,
earlier in the night following sunset than during
the early morning prior to sunrise.
The majority of the material found in the
stomachs of anchovies collected was heavily
digested unidentifiable material. Where stomach
contents could be identified, the diet of
anchovies was dominated by calanoid
copepods.
There was a high degree of overlap between the
diets of fish collected from different sites, with
the exception of Dromana 7 m. Stomachs of fish
collected from Dromana 7 m contained the
smallest volume of material examined, and
were dominated by benthic prey including
gammarid and caprellid amphipods.
5

Other pelagic species
Abundance and Biomass
The most abundant pelagic species caught in
2011 were anchovies, with 795 kg caught. Sandy
sprat, yellowtail scad and pilchards were the
next most abundant pelagic species caught.
Only 70 kg of blue jellyfish (Catostylus mosaicus),
the most abundant pelagic species caught in
previous annual surveys, were recorded in 2011.
6
Distribution
Sandy sprat, yellow-tailed scad and pilchards
were found throughout PPB. Sandy sprat were
more abundant in the north-east region of the
Bay. Yellow-tailed scad and pilchards were
patchily distributed with the majority of the
total catch for these two species caught at only
two sites in 2011.
Discussion
Australian anchovy (Engraulis australis) are
small, schooling planktivorous fish that occupy
pelagic waters in habitats ranging from estuaries
to the continental shelf, from Cape Capricorn in
Queensland to Shark Bay in Western Australia
and south to Tasmania (Ward et al. 2003,
Dimmlich et al. 2004, Gomon et al. 2008). In
Victoria, spawning generally occurs in estuaries,
bays and inlets and extends from approximately
October to March (Arnott and McKinnon 1985,
Jenkins 1986), with peak activity typically
occurring in mid-summer, possibly associated
with peaks in food abundance (Hoedt and
Dimmlich 1995, Ward et al. 2006). Large
numbers of juvenile anchovy (0+ and 1+ year-
class fish) can be found in estuaries and bays
from late summer through to winter, and
constitute the majority of the resident
population biomass. Adult anchovy (2–5+ year-
class fish) tend to be temporary inhabitants of
estuaries and bays, migrating into the open sea
between February and May and returning
between October and December to spawn
(Blackburn 1950, Hoedt et al. 1995).
Little was known about the population
structure, distribution and abundance of
anchovy in PPB prior to 2008 when the present
study began. This report summarizes work
undertaken in the final year of the present study
(May-June 2011), and builds on a conceptual
‘population and life-history’ model for anchovy
in PPB described by Hirst et al. (2010) using
information collected over the full term of this
study (2008–2011) (Table 2).
Population structure
Anchovy population structure in PPB during
winter sampling (May-July) in 2008–2011 was
consistent with that reported for anchovy
populations in bays and estuaries elsewhere
(Scharf et al. 2002, Dimmlich and Ward 2006).
Anchovy populations during this period
comprised primarily 0+ and 1+ age-cohort fish.
This population structure is consistent with the
view that bays like PPB are important nursery
areas for juvenile anchovy. The lack of older
(>1+ age) fish during the annual surveys is also
consistent with the observation that adult fish
are largely absent during the winter months,
only entering the Bay from October onwards to
spawn during the warmer months (Blackburn
1950, Arnott and McKinnon 1985, Hoedt et al.
1995).
The importance of bays and inlets as important
spawning areas for anchovy has generally been
inferred from presence of eggs and larvae from
spring to late summer (Arnott and McKinnon
1985, Jenkins 1986, Hoedt and Dimmlich 1995,
Acevedo et al. 2010) and the absence of anchovy
eggs and larvae in offshore waters (Hoedt and
Dimmlich 1995, Dimmlich et al. 2004). Anchovy
larvae have been detected in PPB from late
November to early January during the period
2008-2011 (Acevedo et al. 2008, 2009, 2010;
Jenkins and Kent 2011). This study estimated
that 0+ anchovy were spawned from September
to April, with peak spawning occurring from
late November to early January. This peak
spawning period coincides with the sampling
period for the CDBMP egg and larval surveys
(Jenkins and Kent 2011). The timing of peak
spawning periods is likely to be critical to the
survival of anchovy larvae, and may have a
significant impact on annual recruitment
variability (Methot 1983). According to Jenkins
and Kent (2011), the relatively high inter-annual
variation in abundance of anchovy eggs and
larvae in PPB during the period 2008-11 is likely
to be linked to environmental fluctuations that
affect phytoplankton productivity, and therefore
influence spawning and larval survival.
The significance of this life-history strategy is
not well understood, but is likely to have
evolved to maximize egg, larval and juvenile
survival (Bellier et al. 2007). Food limitation is
often cited as an important source of mortality
in the early life of fish, and hence a regulator of
recruitment and year-class strength (Islam and
Tanaka 2009). Post-hatch mortality rates for
juvenile anchovies may be as high as 95%
(Scharf et al. 2002), and growth rates for juvenile
anchovy are known to be correlated with both
plankton availability and temperature (Takasura
and Aoki 2006). The diets of larval and juvenile
anchovies comprise primarily zooplankton
(Islam and Tanaka 2009), and spawning in PPB
coincides with a period of higher primary
productivity in the Bay associated with
seasonally higher ambient water temperatures
and pulses of nutrients (Spooner et al. 2011).
7
The life-history strategy for anchovy to some
extent contrasts with other pelagic fish such as
pilchards (Sardinopsis sagax) that are also
temporary residents of PPB. Like anchovy,
juvenile pilchards utilize PPB as a nursery,
entering the bay as mostly 0+ and 1+ juveniles in
late spring/early summer, but do not attain
sexual maturity within the Bay (Neira et al. 1999,
Jenkins and McKinnon 2006). In contrast to
anchovy, pilchards spawn primarily in offshore
coastal shelf waters (Dimmlich et al. 2004) and
rarely in PPB (Neira et al. 1999). Ward et al.
(2001) have suggested that the spatial
segregation of spawning areas for anchovy and
pilchards in South Australian coastal waters
may have evolved as a result of competition
avoidance between these two species.
Diet
Calanoid copepods such as Paracalanus sp. were
the major component of the identifiable diet of
anchovy collected from PPB at night during
May-June 2011, and comprise a major part of the
diets of anchovy examined elsewhere (James
1987, Islam and Tanaka 2009). Anchovy are size-
selective zooplanktivores (James 1987), but it is
unknown if anchovy demonstrate such selective
feeding in PPB. Paracalanus sp. is the dominant
zooplankton species recorded in PPB over
winter (Kimmerer and McKinnon 1985) and
would be expected to be a major component of
the diets of anchovy in the Bay. By comparison,
other common species of zooplankton recorded
from PPB such as the cyclopoid copepod Oithona
sp. and cladoceran Podon spp. were less
frequently recorded in the stomachs of anchovy
examined in this study. More detailed analysis is
required to examine the selectivity of anchovy
diets in PPB.
Anchovy are visual predators and feeding
activity appears to be greatest during the day
(Tudela and Palomera 1997, Taylor et al. 2007)
when anchovy are most active. Most studies of
anchovy diets have been undertaken on fish
collected during the day when fish are
schooling. Schooling behaviour enhances
foraging success, increases hydrodynamic
efficiency and predator avoidance. In one of the
few studies of diurnal feeding patterns
completed for anchovy, Tudela and Palomera
(1997) found that only small quantities of larger
prey such as decapod larvae and mysids were
ingested at night. Tudela and Palomera (1997)
found that anchovy stomachs were most full in
the evening between 1500 and 2100 hours and
most empty in the early morning between 0300
8
and 0600 hours indicating a lag between the
time of ingestion and assimilation of food in the
stomach (Tudela and Palomera 1997).
The present study (May-June 2011) found a
similar relationship between stomach fullness
and time elapsed during the night. The
presence of small numbers of prey items
associated only with the benthos, such as
gammaridean amphipods (particularly
ampeliscids), cumaceans and harpacticoid
copepods, indicates that some feeding occurs at
night whilst anchovy are aggregating near the
benthos. The presence of readily identifiable,
freshly ingested, specimens of Paracalanus sp. in
some of the anchovy stomachs may also suggest
that some calanoid copepods are also ingested at
night.
Distribution
Anchovy in PPB displayed a consistent spatial
pattern of distribution during the present study,
from 2008 to 2011. Anchovy were primarily
distributed in the central and eastern part of
PPB, and largely absent from the north-west of
PPB, the Geelong Arm and Corio Bay. In
addition to variation at the baywide scale,
acoustic surveys undertaken using sonar from
2009 to 2011 showed that anchovy aggregations
were also patchily distributed at smaller spatial
scales within PPB (i.e. at the scale of km).
The spatial distribution of pelagic schooling
species such as anchovy is characteristically
patchy, reflecting the variable distribution of
their primary zooplankton prey, at both larger
and smaller spatial scales (Giannoulaki et al.
2005). Little is known about zooplankton
dynamics at smaller spatial scales in PPB. At the
baywide scale, zooplankton are likely to
respond to large spatial patterns in primary
productivity within PPB that reflect the sources
of nutrients that stimulate phytoplankton
growth (Harris et al. 1996).
The presence of anchovy aggregations in the
central and eastern region of PPB is adjacent to
two major sources of nutrients into PPB, the
Yarra and Patterson Rivers. These contribute
90% of the total nitrogen load from catchment
sources into PPB (EPA unpub. data; E2Port
catchment model), and the river plumes and
subsequent mixing zones are associated with
areas of high phytoplankton and zooplankton
productivity in PPB (EPA 2011). Regions of high
zooplankton productivity are likely to be
spatially displaced from the original source of
nutrients because there is a temporal lag
between the delivery of nutrients, there population in PPB maybe highest over the
subsequent transport away from the source, and summer months following the influx of adult
eventual uptake and assimilation by anchovy.
phytoplankton, and in-turn, zooplankton. Water
PPB supports the largest commercial anchovy
quality modelling shows that the region
fishery in Victoria (Jenkins and McKinnon 2006).
between Beaumaris and Mornington and
The commercial fishery in PPB caught only 17
offshore of Western Treatment Plant (WTP)
tonnes of fish in 2010/11, the lowest annual catch
exhibit high zooplankton productivity following
since 1990/91 (Figure 3). Despite falling
storm events that deliver pulses of nutrients to
commercial catches, catch rates (per unit effort)
PPB (EPA 2011). It is unclear why anchovy
have remained relatively stable since 1995/96.
populations were absent from the western part
Commercial catches have declined as a
of PPB, an area which receives high nutrient
consequence of a steady decline in fishing effort
loadings from the WTP, although freshwater
(i.e. total days fished), rather than changes to
outflow from the WTP is far less seasonal than
anchovy abundance (Figure 3).
the freshwater catchments.
According to the Detailed Design for this sub-
The Detailed Design for this sub-program
program (PoMC 2010) it was intended to
(PoMC 2010) provides for statistical analysis of
monitor interannual trends in anchovy
anchovy distribution, where patterns by site and
abundance estimated over the four annual
depth are apparent. No such patterns were
surveys. It is now clear that variance in annual
observed in the data during the survey period
abundance estimates for PPB is so great that it
(2008-11).
has not been possible to detect an overall trend
during the survey period (2008–11).
Abundance
Anchovy populations in PPB exhibited Interactions with other species
consistent recruitment in each year from 2008 to
PPB supports three other species of clupeoid
2011. The relative size of the 0+ age-cohort in
fish: pilchards, sandy sprat and blue sprat; all
each year was similar to or larger than the
potential competitors of anchovy for food (Ward
existing juvenile cohort (comprising most 1+
et al. 2001). In addition to clupeoids, PPB
fish) recruited in the previous year (see Figure
supports a range of other small planktivorous
8). This suggests that the biomass of the over-
fish, including yellowtail scad that were
wintering population of juvenile anchovy has
abundant in May-June 2011. In May-June 2011,
remained relatively stable over the period 2008
anchovy accounted for 45% of the total catch of
to 2011, despite high variation in annual
pelagic fish, sandy sprat 21% and blue sprat
estimates of total biomass (299 – 1222 tonnes)
0.02%. Yellowtail scad and pilchard were both
from the present study.
locally abundant and accounted, for 12% and 5%
Jenkins and Kent (2011) reported a trend of respectively of the total pelagic fish caught. In
declining anchovy egg production and larval comparison to previous years (2008–2010)
abundance from 2007/2008 to 2010/2011, but pelagic fish species such as pilchards, sandy
concluded that this was likely due to variation sprat and yellowtail scad were far more
in wind driven currents affecting the spatial abundant in 2011 (Parry et al. 2008, 2009, Hirst et
distribution of eggs and larvae in the Bay al. 2010).
relative to sampling areas, or the timing of
Blue jellyfish (Catostylus mosaicus) were very
sampling relative to spawning, rather than a
abundant in PPB and comprised a majority of
decline in anchovy spawning per se. Indeed, it
the catch in 2008 and 2009 (Parry et al. 2008,
was also concluded that inter-annual variation
2009). Dietary analyses suggest that large
in abundance of anchovy larvae and eggs during
jellyfish species occupy a similar trophic level to
the period 2008-11 was relatively high, likely
that of small pelagic species such as anchovy
due to environmental fluctuations that influence
and pilchards (Brodeur et al. 2008), and may also
spawning and larval survival (Jenkins and Kent
consume large numbers of the eggs and larvae
2011).
of pelagic fish (Lynam et al. 2005). In regions
Total anchovy biomass in PPB may be at its where small pelagic fisheries have collapsed,
lowest during early winter when the population jellyfish have been implicated in the slow
is dominated by juvenile anchovy, following the recovery of populations of small pelagic fish
departure of adult anchovy to Bass Strait. such as anchovy (Lynam et al. 2006).
Conversely, the biomass of the anchovy

9
There is little evidence that anchovy recruitment • Calanoid copepods are the major component
was impacted during 2008 and 2009 when of the diet of juvenile anchovy during May-
jellyfish abundance was high in PPB. Blue July, and feeding appears to be most active
jellyfish were practically absent from the catch earlier in the evening following sunset
in the 2011 survey, and only 70 kg jellyfish were • Anchovy are an important food source for
caught compared with 17 tonnes in the 2009 many predatory fish and seabirds, and
survey. provide an important trophic link in PPB
Ecologically, anchovies are an important link between zooplankton productivity and
between zooplankton productivity and higher trophic levels.
predators such as squid, fish and seabirds in
PPB. Anchovy are important prey items for
Crested Terns (Chiaradia et al. 2002) and Little
Penguins (Montague and Cullen 1988, Hobday
et al. 1992, Chiaradia et al. 2003). Little Penguins
have been found to feed mostly on juvenile
pilchards from spring to summer, and anchovy
from autumn to winter, reflecting changes in the
availability of preferred prey (Montague and
Cullen 1988). The decline in the PPB pilchard
population in the mid to late 1990s has probably
increased the ecological importance of anchovy
in the Bay (Jenkins and McKinnon 2006).
Species formerly dependent on pilchards as a
major food source have switched to a diet of
other pelagic species, including anchovy (Bunce
and Norman 2000).

Conclusion
The distribution and age-class structure of
anchovy in PPB during May–June 2011 were
consistent with the results of previous annual
surveys (2008–2010) and the known attributes of
anchovy populations in other parts of Australia.
The results of the 2011 survey are also consistent
with the conceptual population and life history
model developed for anchovy in PPB (Hirst et al.
2010). This model is further complemented by
additional information on diet, feeding activity
and spawning season generated during 2011.
The model features the following characteristics:
• PPB is an important spawning and nursery
area for anchovy within its Victorian
distribution
• Anchovy population in PPB is dominated by
juvenile (0+ and 1+ ) anchovy during May-
July, with few adult (>1+) fish present
• It is believed that adult fish move out of PPB
following spawning in summer to occupy
offshore waters during winter months
• Anchovy in PPB exhibit an elongated
spawning season, extending from September
to April, with peak spawning estimated to
occur from late November to early January

10
 500 2000
A

400 1600
Total catch (tonnes)

catch rate (kg/day)

300 1200

200 800

100 400

0 0
1975 1985 1995 2005 2015
year
600 600
B
500 500
Total catch (tonnes)

Effort (days)
400 400

300 300

200 200

100 100

0 0
1975 1985 1995 2005 2015
year
Figure 3. Total catch (bars), A) catch rate (kg/day), and B) effort (days) for the commercial anchovy
fishery in PPB 1978/79 to 2010/11

11
Table 2. Conceptual understanding of anchovy population in PPB gained from CDBMP surveys 2008–2011 and the literature
Parameter Finding Evidence Implications

Population structure Anchovy populations dominated by 0+ and 1+ Anuual ageing from 2008-2011 Consistent with model that PPB is an important
juvenile fish in May-July nursery area for juvenile anchovy
No resident adult population (>1+ fish) in PPB over-
winter
Adult anchovy appear to be transient population,
migrating in and out of PPB
Timing of migration of adult anchovy uncertain, but
may be inferred from presence of egg/larvae and
estimated spawning dates (see below)

Spawning Spawning occurs from September to April
Peak spawning estimated to occur from late
November to early January
Presence of anchovy eggs/larvae over this period
(Jenkins and Kent (2011), Jenkins (1986), Neira and
Sporcic (2002))
Daily ageing for 0+ fish from 2008-2011
Estimated from age (days) of 0+ fish
Similar to bays and estuaries in other parts of
southern Australia
Critical period for early life of anchovy larvae
Coincides with period of CDBMP egg/larval surveys

Distribution Anchovy occur primarily in central and eastern part Trawl and acoustic surveys 2008-2011 Pattern consistent 2008-2011 (NB. Geelong Arm not
of PPB surveyed in 2008). Cause/s of this large-scale spatial
pattern not well understood, but may be linked to
areas of high zooplankton productivity in PPB

Distribution highly patchy at smaller-spatial scales
Diet Diet of anchovy in PPB dominated by calanoid
copepods
Feeding activity appears to be reduced at night
Acoustic surveys 2009-2011
Anchovy stomach contents 2011
Pattern of anchovy stomach fullness 2011
Consistent with ecology of schooling, pelagic
species. Spatial patchiness likely to be related to
distribution of primary diet, zoolankton.
Consistent with literature
Majority of feeding occurs during the day when
anchovy are schooling in surface to mid waters

Abundance Recruitment consistent from 2008-2011
Ecological significance Anchovy are an important link between
zooplankton productivity and predatory fish,
seabirds
PPB anchovy are the major prey species for seabirds
such as Little Penguins and Crested Terns
Size of 0+ cohort consistent relative to the size of the No evidence of high recruitment variability for the
cohort (mostly 1+ fish) recruited in previous year period 2008 to 2011, despite high interannual
biomass estimates and declining egg and larval
densities
Anchovy stomach contents 2011 Ecological significance of anchovy in PPB may have
increased since the decline of the pilchard
population in the mid 1990s

12
Acknowledgements
Thanks to all staff for their cheerful approach to
the long and difficult hours worked throughout
the night during this project.
Thanks particularly to David Guillot whose
expertise in net design and trawling was crucial
to the success of this project, and the skipper of
the Western Alliance Andrew Kelly. Paul McCoy
assisted in the wheelhouse, and Fabian Trinnie
Kade Mills, Peter Young, Matt Koopman and
Andrew Brown helped sort and measure fish
throughout the night. Thanks also to Pam
Oliveiro for dissecting anchovy otoliths for
ageing and Peter Young for preparing GIS maps.

13
References
Acevedo, S., Jenkins, G. and Harris, A. (2008).
Baywide Egg and Larval Survey Sub-Program.
Milestone Report No. 1 (2008). Fisheries Victoria
Technical Report No. 19. Department of Primary
Industries, Queenscliff, Victoria, Australia.
Acevedo, S., and Jenkins, G. (2009). Baywide Egg
and Larval Survey Sub-Program. Milestone
Report No. 2 (2009). Fisheries Victoria Technical
Report No. 40. Department of Primary Industries,
Queenscliff, Victoria, Australia.
Acevedo, S., Jenkins, G. and Kent, J. (2009).
Baywide Egg and Larval Survey Sub-Program.
Milestone Report No. 3 (2009). Fisheries Victoria
Technical Report No. 65. Department of Primary
Industries, Queenscliff, Victoria, Australia.
Acevedo, S., Jenkins, G. and Kent, J. (2010)
Baywide Egg and Larval Surveys Sub-Program,
Milestone Report No. 4 (Nov. 2009–Jan. 2010).
Fisheries Victoria Technical Report Series No.
111, September 2010. Department of Primary
Industries, Queenscliff, Victoria, Australia.
Arnott, G.H., and McKinnon, A.D. (1985). The
spawning of the anchovy, Engraulis australis
antipodum Gunther, in relation to temperature
and salinity in the Gippsland Lakes. Australian
Journal of Marine and Freshwater Research 36, 433-
439.
Bellier, E., Planque, B., and Petitgas, P. (2007).
Historical fluctuations in spawning location of
anchovy (Engraulis encrasicolus) and sardine
(Sardina pilchardus) in the Bay of Biscay during
1967-73 and 2000-2004. Fisheries Oceanography 16,
1-15.
Blackburn, M. (1950). A biological study of the
anchovy, Engraulis australis (White), in Australian
waters. Australian Journal of Marine and Freshwater
Research 1, 3–84.
Brodeur, R. D., Suchman, C. L., Reese, D. C.,
Miller, T. W., and Daly, E. A. (2008). Spatial
overlap and trophic interactions between pelagic
fish and large jellyfish in the northern California
Current. Marine Biology 154, 649-659.
Bunce, A., and Norman, F. I. (2000). Changes in
the diet of the Australasian gannet (Morus
serrator) in response to the 1998 mortality of
pilchards (Sardinops sagax). Marine and Freshwater
Research 51, 349-353.
14
Chiaradia, A., Dann, P., Jessop, R., and Collins, P.
(2002). The diet of Crested Tern (Sterna bergii)
chicks on Phillip Island, Victoria, Australia. Emu
102, 367-371.
Chiaradia, A., Costalunga, A., and Knowles, K.
(2003). The diet of Little Penguins (Eudyptula
minor) at Phillip Island, Victoria, in the absence of
a major prey - Pilchard (Sardinops sagax). Emu
103, 43-48.
Dimmlich, W. F., Breed, W. G., Geddes, M., and
Ward, T. M. (2004). Relative importance of gulf
and shelf waters for spawning and recruitment of
Australian anchovy, Engraulis australis, in South
Australia. Fisheries Oceanography 13, 310-323.
Dimmlich, W.F., and Ward, T.M. (2006).
Ontogenetic shifts in the distribution and
reproductive patterns of Australian anchovy
(Engraulis australis) determined by otolith
microstructure analysis. Marine and Freshwater
Research 57: 373-381.
EPA (2011). Port Phillip and Western Port
receiving water quality modelling: nutrient
dynamics and primary production. Publication
1379.
Giannoulaki, M., Machias, A., Somarakis, S., and
Tsimenides, N. (2005). The spatial distribution of
anchovy and sardine in the northern Aegean Sea
in relation to hydrographic regimes. Belgian
Journal of Zoology 135, 151-156.
Harris, G., Batley, G., Fox, D., Hall, D., Jernakoff,
P., Molloy, A., Murray, B., Newell, J., Parslow, G.,
Skyring, G., and Walker, S. (1996). Port Phillip
Bay Environmental Study Final Report. CSIRO,
Canberra, Australia.
Hirst, A.J, White, C.A., Parry, G.D., Heislers, S.,
Werner, G.F. and Spooner, D. R. (2010). Baywide
Anchovy Sub-program Milestone Report No. 3
(2010). Fisheries Victoria Technical Report Series
No. 114 November 2010. Department of Primary
Industries, Queenscliff, Victoria, Australia.
Hirst, A.J, White, C.A., Heislers, S., Werner, G.F.
Young, P. and Spooner, D. R. (2011). Baywide
Anchovy Sub-program Progress Report No. 4
(2011). Fisheries Victoria Technical Report Series
No. 146 August 2011. Department of Primary
Industries, Queenscliff, Victoria
Hobday, D.H. (1992) Abundance and distribution heavily fished ecosystem. Current Biology 16, 492-
of Pilchard and Australian Anchovy as prey 493.
species for the Little Penguin Eudyptula minor at
Methot, R. D. (1983). Seasonal variation in the
Phillip Island, Victoria. Emu 91: 342-354.
survival of larval northern anchovy, Engraulis
Hoedt, F.E., and Dimmlich, W.F. (1995). Egg and mordax, estimated from the age distribution of
larval abundance and spawning localities of the juveniles. Fisheries Bulletin 81, 741-750.
anchovy (Engraulis australis) and pilchard
Montague, T. L., and Cullen, J. M. (1988). The
(Sardinops neopilchardus) near Phillip Island,
diet of the Little Penguin Eudyptula minor at
Victoria. Marine and Freshwater Research 46, 735-
Phillip Island, Victoria. Emu 88, 138-149.
743.
Neira, F. J., Sporcic, M. I., and Longmore, A. R.
Hoedt, F. E., Dimmlich, W. F., and Dann, P.
(1999). Biology and fishery of pilchard, Sardinops
(1995). Seasonal variation in the species and size
sagax (Clupeidae), within a large south-eastern
composition of clupeoid assemblages in Western
Australian bay. Marine and Freshwater Research 50,
Port, Victoria. Marine and Freshwater Research 46,
43-55.
1085-1091.
Neira, F. J., and Sporcic, M. I. (2002). Use of
Islam, S., and Tanaka, M. (2009). Diet and prey
ichthyoplankton ecology to evaluate ecosystem
selection in larval and juvenile Japanese anchovy
changes: a case study in a large, semi-enclosed
Engraulis japonicus in Ariake Bay, Japan. Aquatic
Australian bay. Marine and Freshwater Research 53,
Ecology 43, 549-558.
339-354.
James, A. G. (1987). Feeding ecology, diet and
PoMC (2010). Anchovy study– Detailed Design -
field-based studies on feeding selectivity of the
CDP_ENV_MD_016 Rev 3. Port of Melbourne
Cape anchovy Engraulis capensis Gilchrist South
Corporation.
African Journal of Marine Science 5, 673-692.
Parry, G. and Stokie, T. (2008). Baywide Anchovy
Jenkins, G.P. (1986). Composition, seasonality
Study Sub-Program. Milestone Report No. 1
and distribution of ichthyoplankton in Port
(2009). Technical Report No. 23. Fisheries
Phillip Bay, Victoria. Australian Journal of Marine
Victoria, January 2009. Department of Primary
and Freshwater Research 37, 507–520.
Industries, Queenscliff, Victoria.
Jenkins, G. and Kent, J. (2011) (2010) Baywide
Parry, G., Hirst, A., Stokie, T., Green, C., White,
Egg and Larval Surveys Sub-Program, Milestone
C., Heislers, S. and Werner, G. (2009). Baywide
Report No. 5 (Nov. 2010–Jan. 2011). Fisheries
Anchovy Study Sub-Program. Milestone Report
Victoria Technical Report Series No. 145,
No. 2. Technical Report No. 73. Department of
September 2011. Department of Primary
Primary Industries, Queenscliff, Victoria.
Industries, Queenscliff, Victoria.
Parry, G., Hirst A., and White, C. (2010) Baywide
Kimmerer, W. J., and McKinnon, A. D. (1987). A
Anchovy Study Sub-Program – Sonar Report.
comparative study of the zooplankton in two
Fisheries Victoria Internal Report No. 25.
adjacent embayments, Port Phillip and
Department of Primary Industries, Queenscliff,
Westernport Bays, Australia. Estuarine, Coastal
Victoria.
and Shelf Science 21, 145-159.
Scharf, F. S., Buckel, J. A., and Juanes, F. (2002).
Jenkins, G.P, and McKinnon, L. (2006). Channel
Size-dependent vulnerability of juvenile bay
Deepening Supplementary Environmental
anchovy Anchoa mitchilli to bluefish predation:
Effects Statement – Aquaculture and Fisheries.
Does large body size always provide a refuge?
Internal Report No. 77, Primary Industries
Marine Ecology Progress Series 233, 241-252.
Research Victoria, Queenscliff.
Spooner, D., Walker, T. I., Acevedo, S., and
Lynam, C. P., Heath, M. R., Hay, S. J., and
Morris, E. (2011). Port Phillip Bay Environmental
Brierley, A. S. (2005). Evidence for impacts by
Data Review - Biophysical Assessment of
jellyfish on North Sea herring recruitment.
Climate Change. Fisheries Victoria Internal
Marine Ecology Progress Series 298, 157-167.
Report no. 27, Queenscliff, Victoria
Lynam, C. P., Gibbons, M. J., Axelsen, B. E.,
Takasuka, A., and Aoki, I. (2006). Environmental
Sparks, C. A. J., Coetzee, J., Heywood, B. G., and
determinants of growth rates for larval Japanese
Brierley, A. S. (2006). Jellyfish overtake fish in a
anchovy Engraulis japonicus in different waters.
Fisheries Oceanography 15, 139-149.

15
Tudela, S., and Palomera, I. (1997). Trophic
ecology of the European anchovy Engraulis
encrasicolus in the Catalan Sea (northwest
Mediterranean). Marine Ecology Progress Series
160, 121-134.
Ward T.M., Hoedt F., McLeay L.J., Dimmlich
W.F, Jackson G., Rogers P.J. and Jones K. (2001)
Have the recent mass mortalities of Sardine in
Australia facilitated an expansion of the
distribution of Engraulis australis?. Marine Ecology
Progress Series 220, 241–251.
Ward T.M., Staunton-Smith J., Hoyle S. and
Halliday, I.A. (2003) Spawning patterns for four
species of predominantly temperate pelagic
fishes in sub-tropical waters of southern
Queensland. Estuarine, Coastal and Shelf Science
56, 1125–1140.
Ward, T. M., Mcleay, L. J., Dimmlich, W. F.,
McClatchie, S., Matthews, R., Kampf, J., and Van
Ruth, P. D. (2006). Pelagic ecology of a northern
boundary current system: effects of upwelling on
the production and distribution of sardine
(Sardinops sagax), anchovy (Engraulis australis)
and southern bluefin tuna (Thunnus maccoyii) in
the Great Australian Bight. Fisheries Oceanography
15, 191-207.

16
Appendix 1. Materials and Methods
The methods used in the present study are
described in PoMC (2010), Parry et al. (2009, 2010)
and Hirst et al. (2010). Additional methods not
otherwise reported are described below.
Abundance and biomass
Mean anchovy biomass in PPB was estimated for
the region in which anchovies occurred (Figure
4), based on mean catch per tow (kg/10 min tow)
across the region.
The total biomass in PPB was estimated for the
region in which anchovies occurred (km2) using
the mean biomass estimate (kg/10 min tow) on
the assumption that anchovy biomass is
relatively uniform across this region (Parry et al.
2009).
It was also assumed that anchovies were
captured with 100% efficiency within a swept
area of approximately 36,000 m2 (door spread of
approx. 36 m x mean tow length of approx. 1.01
km).
Distribution
Information on the distribution of anchovies in
PPB was augmented by analysis of sonar images
recorded every 2 minutes whilst the survey
vessel was underway between fixed-sampling
sites. This information was collected and
analysed using the methods described in Parry et
al. (2010) and Hirst et al (2010b).
Age
Annual
The age structure of anchovies in PPB was
determined using otoliths from 14-16 randomly–
selected fish at each of 31 fixed sites where
anchovies were collected by trawl in 2011 (Table
3; n.b. no anchovy collected at Sandringham 22
m). A total of 493 fish were aged in 2011 using
the ageing protocol described by Parry et al.
(2009).
Daily increment
Daily ageing of 0+ age-cohort fish (< 90 mm)
collected in 2010 and 2011 was determined using
daily increment analysis (Parry et al. 2009).
Overall, a total of 105 0+ age-cohort fish were
aged in the present study from a range of sites in
PPB from annual surveys between 2008–2011
(2008: 20 fish; 2009: 27 fish; 2010: 29 fish; 2011: 29
fish).
Differences in growth rates between years were
examined using a test for homogeneity of
regression slopes. This test was done by fitting a
model that related total length (mm) to year, age
(covariate) and the interaction between year and
age using analysis of covariance (ANCOVA). The
latter term tests for the H0 of equal regression
slopes between years.
Spawning dates for 0+ fish recruited in 2008–2011
was estimated by back-calculating the hatching
dates of fish in days and including an incubation
period for the eggs of 2–3 days (Ospina-Alvarez
et al. 2011).
Peak spawning period in each year was
estimated from the mode of the 0+ age-cohort in
each year using a method similar to Methot
(1983). The approximate spawning date for this
modal length was calculated using linear
regressions of daily age against length in each
year (Table 4). All regressions were significant at
P<0.001 and explained between 56 and 84% of the
variance between the two variables.
Anchovy diets
The diets of anchovy in PPB in 2011 were
analysed by examining the stomach contents of
fish collected from eight sites. Fish were fixed in
10% formalin solution in the field and stomachs
dissected, and the contents analysed in the
laboratory. Stomach contents were sorted, and
prey items identified to the lowest possible
taxonomic level. Prey items that were fully
digested were recorded as ‘unidentifiable
digested material’, whereas many prey items
were partially digested, thereby enabling
identification to family or order level only. The
volume (mm3) of prey items was calculated by
estimating the area and depth occupied by each
item. A qualitative (%) estimation of stomach
fullness was also recorded for each fish.
Differences in the composition of anchovy
stomach contents between sites in 2011 were
analysed using similarity-based multivariate
statistical techniques (Clarke and Warwick 2001).
Variation in diet composition between sites was
analysed using analysis of similarity (ANOSIM)
tests based on Bray-Curtis indices of similarity.
17
The unidentifiable digested material category
was excluded from analysis because this category
provided no information on the diet of anchovy
collected from different sites. Differences
between sites were represented visually using
non-metric multi-dimensional scaling (nMDS)
ordination. All analyses were undertaken in the
PRIMER multivariate statistical package.

18
Table 3. Number of anchovies aged at each site for annual PPB surveys in 2008–2011.
Transect depth 2008 2009 2010 2011
Aspendale 12 30 26 16
17 16
22 27 16
Beacon Point 17 29 16
22 30 16
Beaumaris 7 16
12 30 30 27 16
17 16
22 30 30 26 16
Central North 22 30 26 16
Central South 22 6 26 16
Davey Point 7 30
12 30 30 26 16
17 30 30 26 16
22 16
Dromana 7 30 16
12 29
17 29 30 26
22 29 31 26
Geelong 12 26
Geelong Arm N3A 7 16
Hobsons Bay 7 30
12 30
17 30 16
22 30 26 16
Mornington 7 30
12 29 16
17 29 30 16
22 29 30 25 16
Point Cook 7 16
12 25
17 25
22 28 26 16
Geelong Arm S2 26
Sandringham 17 26 16
22 26
Seaford 12 16
17 15
22 30 31 26 16
St. Leonards 7 16
12 14
17 16
22 30 30 26 16
Total 501 518 545 493

Table 4. Parameters for linear regressions between length (mm) and daily age (days) for anchovy aged
using daily growth increments in 2008, 2009, 2010 and 2011 (age (days) = slope*length + constant).

Year n slope constant r2

2008* 20 2.486 27.734 0.56
2009 27 2.915 -17.693 0.82
2010 29 2.949 -44.609 0.84
2011 29 3.288 -69.316 0.70

*anchovy samples collected in two different weeks (Parry and Stokie 2008)

19
Appendix 2 Results
Results for the 2008, 2009 and 2010 annual
surveys were presented in Parry and Stokie
(2009), Parry et al. (2009) and Hirst et al. (2010).
Preliminary results for the 2011 annual survey
were presented in Hirst et al. (2011).
Quality Control
The acoustic survey indicated that 77% of
clupeoid fish echoes occurred within the region
swept by the trawl net (i.e. within 3 m of the
seabed) (Table 5). In 2011, anchovy accounted
for 61% of the total clupeoid biomass caught.
Anchovies
Abundance and biomass
A total of 231,711 anchovies weighing 795 kg
were caught during the 2011 survey.
The mean biomass of anchovies caught within
this region was 24.7 + 1.3 kg/10min tow,
excluding, an isolated trawl (Geelong N3A site)
taken in the Geelong Arm. The total biomass of
anchovies in PPB during 2011 was estimated as
523.8 ± 27.6 tonnes (Table 6).
Distribution
The distribution of anchovies in 2011 was
concentrated in the centre and east of PPB in an
area covering approximately 762 km2 (Figure 4).
Anchovies were largely absent from Hobsons
Bay, Corio Bay and the Geelong Arm (with the
exception of the N3A site).
Anchovies occurred in an area of PPB covering
653 km2 in 2010, 742 km2 in 2009 and 660 km2 in
2008, noting the Geelong Arm was not sampled
in 2008 (Figure 5). Anchovies were found further
north in 2009 than 2008, 2010 or 2011; and
further west in 2010 and 2011 compared with
2008 and 2009.
Table 5. Vertical distribution of clupeoid fish within 5 m of the seabed: cumulative % of fish echoes at
1 m intervals. *n=1014 images
Depth range (m) cumulative % of fish
echoes*
0-1 33.7
1-2 58.4
2-3 77.2
3-4 90.4
4-5 100.0
20
Anchovies were found in a small region
(approx. 27 km2) in the Geelong Arm in 2010
and at the Geelong N3A site in 2011. A region
could not be delineated in the Geelong Arm in
2011 on the basis of information for only a single
site. Anchovies were largely absent from the
southern most transect, Dromana, in 2011.
Underway acoustic surveys of clupeoid
distribution in PPB showed a similar pattern to
that inferred from the fixed-site sampling design
for anchovy distribution (Figure 6). These
clupeoid aggregations were dominated by
anchovy in 2011, and are indicative of anchovy
distribution within PPB.
Clupeoid aggregations were patchily distributed
throughout the primary region of distribution
inferred from the fixed-site sampling design
(Figure 6). Dense patches were common
throughout the deeper, central region of PPB
and offshore at Sandringham, Aspendale,
Frankston and Portarlington. By comparison,
aggregations were largely absent from Hobson
Bay, the coast between Point Cook and Point
Wilson (including the eastern end of the
Geelong Arm), Corio Bay, and offshore of
Dromana. Acoustic sampling confirmed the
presence of anchovies in the vicinity of the
Geelong N3A site (anchovies comprised 100% of
the catch at this site) (Figure 6).
Population structure
Precision
Repeat readings of annual increments for 124
fish from the 2011 survey produced an IAPE
(index of average percent error) of 2.95%.
Annual age estimates
Anchovy populations in PPB in 2008, 2009 and
2010 comprised 0+, 1+, 2+ and 3+ age cohorts. In
2011 only 0+, 1+ and 2+ age cohorts were
recorded (Table 7). The dominant age cohorts in
all years were 0+ and 1+ fish (Table 7, Figure 7).
Anchovies belonging to the same age cohort
displayed considerable variation in length in
2011, and from year-to-year (Figure 7). The
length of anchovy in 2011 for:
• 0+ age-cohort varied from 50 to 99 mm
• 1+ age-cohort from 77 to 124 mm
• 2+ age-cohort from 86 to 117 mm.
The mean lengths of anchovies in the 1 + and 2+
age-cohorts were generally smaller (Table 7) and
comprised fewer individuals > 100 mm than in
previous surveys (2008–2010)(Figure 8).
Distribution of annual age-cohorts
There were differences in the distribution of
anchovy of different age-cohorts within PPB. In
2011, anchovy from different age-cohorts were
distributed throughout PPB (Figure 9):
• 0+ aged anchovies were more prevalent in
the central part of PPB
• 1+ aged anchovies were more prevalent
around the periphery of PPB, particularly
from Beaumaris to Mornington
• 2+ aged anchovies comprised only a small
proportion of fish aged at all sites (< 25%).
Daily age estimates
ANCOVA indicated there was no significant
difference between growth rates for 0+ age-
cohort fish in different years based on daily
growth increments (ANCOVA; F3,97=1.254,
P=0.295; Figure 10).
Spawning dates for the 0+ aged fish collected
during surveys from 2008 to 2011 ranged from
(Figure 11):
• September 2007 to February 2008 for fish
collected in June-July 2008
• September 2008 to April 2009 for fish
collected in June 2009
• September 2009 to March 2010 for fish
collected in June 2010
• October 2010 to March 2011 for fish collected
in May-June 2011.
Peak spawning period was estimated from the
modal length of each 0+ age-cohort sampled
from 2008 to 2011 (see Figure 8). Peak spawning
ranged from late November 2007, for 75 mm fish
collected in June-July 2008, to early January 2009
for 65 mm fish collected in June 2009 (Table 8).
Anchovy collected in 2010 and 2011 were
calculated to have peak spawning period
occurring in December 2009 and 2010,
respectively.
Diet
Stomach fullness varied between anchovy
sampled at eight sites across PPB (Table 7).
Stomachs from fish collected from Hobsons Bay
22 m, Seaford 22 m, Geelong N3A 7 m and St
Leonards 22 m were >60% full, whereas
stomachs from fish collected from Central south
22 m, Davey Point 17 m, Dromana 7 m and
Mornington 22 m were by comparison relatively
empty (i.e. <30% full).
The relationship between % stomach fullness
and nominal time (24 hour) indicated that
stomachs tended to be fuller earlier in the night
following sunset than during the early morning
prior to sunrise (Figure 12).
The identifiable component of the diet of
anchovies was dominated by calanoid
copepods, most probably Paracalanus sp. (at least
40 of 79 fish stomachs analysed) as no other
calanoid copepod species were identified among
the stomach contents. The other main
component of the diets of anchovies recorded
was a range of benthic invertebrates, including
gammarid and caprellid amphipods,
harpacticoid copepods and cumaceans. The
majority of the material found in the stomachs of
anchovy collected in 2011 was unidentifiable
digested material. It is typical for the stomachs
of small pelagic fish with high metabolisms to
contain a high % of digested material. It is
assumed that the composition of this material
occurs in direct proportion to recently ingested
and hence identifiable material (Tudela and
Palomera 1997).
There was a high degree of overlap between the
diets of fish collected from different sites within
PPB, with the exception of Dromana 7 m (Figure
13). Fish stomachs at Dromana 7 m contained
the smallest volume of material examined (Table
7) and contained no calanoid copepods, but
were dominated by benthic prey, including
gammarid and caprellid amphipods.
Other pelagic species
Abundance and Biomass
The biomass and abundance of all fifteen pelagic
species caught in the anchovy survey in 2011 are
shown in Table 10. The most abundant pelagic
species in the water-column in 2011 were
21
anchovies, with 795 kg caught. Sandy sprat
(total catch = 364.7 kg), yellow-tailed scad (217.6
kg) and pilchards (93.7 kg) were the next most
abundant pelagic species caught. Only 70.9 kg of
blue jellyfish (Catostylus mosaicus) were caught
in 2011. C. mosaicus was the most abundant
pelagic species encountered in previous surveys
(2008–2010), with over 17 tonnes caught in 2009.
Distribution
The baywide distributions of the next three most
abundant pelagic species in 2011, sandy sprat,
yellow-tailed scad and pilchards, are shown in
Figure 14.
Sandy sprats, yellow-tailed scad and pilchards
were found throughout the Bay. Sandy sprats
were more abundant in the north-east part of
PPB offshore of Beaumaris. Yellow-tailed scad
and pilchards were patchily distributed, with
the majority of the total catch for these two
species caught at only two sites during the 2011
survey (Figure 14).

22
Figure 4. Distribution of anchovy biomass (kg/10 minute tow) in the 2011 annual survey in PPB.
Polygon indicates the approximate area in which anchovies were recorded by trawl or sonar at fixed
sites during 2011.

Table 6. Mean biomass of anchovies /trawl tow (+ se) for the region identified in the 2011 survey
(Table 2, Figure 2), approximate area occupied by anchovies in PPB, and estimated biomass of
anchovies in PPB (+ se tonnes) in the 2011 survey.

Region Mean biomass (kg/10 2 Estimated biomass

Area (km )
min tow) (tonnes)*
Region (excluding 24.7 ± 1.3 762 523.8 ± 27.6
Geelong Arm N3A)

*based on a swept area of approximately 36,000 m2

23
Figure 5. Polygons showing the approximate distribution of anchovies in PPB during annual surveys
conducted in 2008–2011.

24
Figure 6. Acoustic survey 2011: percentage area of fish echoes within 2 m of seabed for underway
sampling between fixed sites. The polygon shows the primary region of anchovy distribution inferred from fixed
sampling conducted in May–June 2011.

25
 25
2008
20
15
10
5
0
30 40 50 60 70 80 90 100 110 120 130 140

25
2009
20
15
10
5
Number

0
30 40 50 60 70 80 90 100 110 120 130 140

25
2010
20
15
10
5
0
30 40 50 60 70 80 90 100 110 120 130 140

30
2011
25
20
15
10
5
0
30 40 50 60 70 80 90 100 110 120 130 140
Length (mm)

Age 0+ 1+ 2+ 3+

Figure 7. Age-length frequency distributions of anchovies from annual surveys in PPB during June-
July 2008, June 2009, June 2010 and May-June 2011.

26
 3000 2008
2500
2000
1500
1000
500
0
30 40 50 60 70 80 90 100 110 120 130 140

20000
2009
15000

10000

5000
Number

0
30 40 50 60 70 80 90 100 110 120 130 140

3000 2010
2500
2000
1500
1000
500
0
30 40 50 60 70 80 90 100 110 120 130 140

16000 2011
12000

8000

4000

0
30 40 50 60 70 80 90 100 110 120 130 140
Length (mm)

Figure 8. Length-frequency distributions of anchovies from annual surveys in Port Phillip Bay 2008–
2011. Number is the projected number of anchovies in each size category (1 mm increments) based on the number of
fish measured as a proportion of the total number of anchovy caught at each site (see Parry et al. 2009). Note, y axes
differ between years. The arrow indicates the modal length of the 0+ age-cohort in each year used in calculations of peak
spawning period (Table 8).

27
Table 7. Mean length (mm) and standard error of anchovies of different ages in PPB in the period June–July 2008, 2009, 2010 and 2011.
N: number of anchovies aged for each age cohort.*
Age 2008 2009 2010 2011
N length N length N length N length
mean se mean se mean se mean se
0+ 201 79.7 0.7 234 76.0 0.9 167 76.1 0.9 166 73.8 0.8
1+ 283 104.2 0.6 217 97.4 0.5 284 100.4 0.6 274 92.2 0.4
2+ 15 106.4 1.7 52 105.4 1.1 70 105.1 1.0 45 97.6 1.1
3+ 2 119.5 5.5 1 115.0 0.0 9 107.2 2.5 0
Total 501 504 530 485
*Not all otoliths collected were aged. In 2009 n = 14, in 2010 n= 15 and in 2011 n=8 otoliths could not be read.

28
 0+ age cohort, 2011 1+ age cohort, 2011
#
#
# #

# # # #
# #
#
# #

# # #
# # # # # #
#

#
# #
#
# #
# #
# # #
#
#
# # # #
# #
# #
S
#
##
#
# #
#
##
#
# #
# #
# #

#
S 0 #
#
S 0 #

# 0.1 - 25
# 0.1 - 25
# 25.1 - 50
# 25.1 - 50
# 50.1 - 75
# 50.1 - 75
# 75.1 - 100
# 75.1 - 100

2+ age cohort, 2011

#
S

#
S

#
S #

# #
S
#
S
# # #
S #
S
S#
# # #
#
# #
#
# #
S
# #
#
# #
#
#

#
#

#
S 0 #

# 0.1 - 25
# 25.1 - 50

Figure 9. Distribution of age cohorts surveyed in May-June 2011; % 0+, 1+, and 2+ fish aged at each site.

29
 120

2008
100 2009
2010
2011
length (mm)

80

60

40

20
0 50 100 150 200 250 300
estimated age (days)

Figure 10. Estimated growth based on daily growth increments of anchovy in PPB for 0+ age-cohort
fish collected in 2008, 2009, 2010 and 2011 surveys. Lines = linear regressions between length and age in each
year. Analysis shown in Parry et al. (2009) only includes fish <90 mm length for the purpose of validating the 0+ age-
cohort.

30
 Jul
Aug
Sep
Oct
Nov
Month

Dec
Jan
Feb
Mar
Apr
May
Jun
2008 2009 2010 2011
year
Figure 11. Range of spawning dates (months) encountered for 0+ age-cohort fish aged using daily
growth increments in 2008, 2009, 2010 and 2011.

Table 8. Peak spawning period estimated from age (days) of 0+ anchovy at the modal length (mm) in
2008, 2009, 2010 and 2011.

Year mode* (length mm) age (days)** spawning date

2008 75 216 late November 2007

2009 65 174 early January 2009
2010 75 179 December 2009
2011 70 163 December 2010

*mode derived from length-frequency distribution (Figure 8)

**age estimated from linear regressions (Table 4)

31
Table 9. Mean length (mm), stomach fullness (%) and number of prey items recorded for fish
examined at 8 sites in May-June 2011.

site depth Time of N mean fish length mean % full mean no. prey
sampling (mm) items
Central South PPB 22 1:44:00 10 79.6 28.5 4.2
Davey Point 17 0:20:00 10 89.0 18.5 2.1
Dromana 7 20:41:00 9 88.9 10.6 1.1
Hobsons Bay 22 22:01:00 10 73.5 75.5 3.5
Mornington 22 4:10:00 10 94.4 22.4 3.4
Geelong Arm N3A 7 2:16:00 10 90.2 63.5 4.1
Seaford 22 21:01:00 10 88.9 60.0 2.0
St. Leonards 22 22:17:00 10 89.4 63.0 3.3
Total 79

Table 10. Summary of anchovy stomach content analysis undertaken in 2011.

Species/taxon Taxnonmic group mean volume Frequency of

(mm3) occurrence*

Luciferidae uid. Decapod crustacea 0.04 1

Decapoda unid. 0.40 1
Iphimediidae unid. Gammaridean amphipod 0.50 1
Gammaridean unid. Gammaridean amphipod 0.90 1
Ampeliscidae unid. Gammaridean amphipod 90.00 1
Podon spp. Cladoceran crustacea 0.20 2
Ectinosomatidae unid. Harpacticoida copepod 0.01 3
Oithona sp. Cyclopoida copepod 0.01 3
Cestoda unid. Platyhelminthes worms 0.04 3
Caprellidae unid. Caprellid amphipod 0.27 3
Cumacea unid. Cumacean crustacean 0.78 3
Corycaeidae unid. Poecilostomatoida copepod (ectoparasites) 0.01 4
Mysidae unid. Mysid crustacean 3.15 4
Ostracoda unid. Ostracod crustacean 0.69 5
Harpacticoid copepods unid. 0.01 8
Paracalanus sp. Calanoid copepod 0.03 21
Calanoida unid. Calanoid copepod 0.03 40
Unidentifiable digested material 2.57 79

*n = 79 fish

32
 100

80
Fullness (%)

60

40

20

0
0 3 6 9 12 15 18 21 24
Time (24 hours)
Figure 12. Relationship between mean (± se) anchovy stomach fullness (%) and nominal sampling
time (24 hour scale) for anchovies collected from 31 May–4 June 2011. A quadratic function has been
fitted to the data following Tudela and Palomera (1997). Vertical lines indicate position of sunrise
(approx. 7.25 am) and sunset (5.10 pm).

2D Stress: 0.09 site

Central South PPB 22
Davey Point 17
Dromana 7
Hobsons Bay 22
Mornington 22
N3A Geelong Arm 7
Seaford 22
St. Leonards 22

Figure 13. Non-metric multidimensional scaling (nMDS) plot displaying the compositional similarity
of anchovy stomach contents of fish collected at eight sites in 2011. n = 10 fish stomachs site-1

33
Table 11. Biomass and abundance of all pelagic species caught in the 2011 survey.
Species Frequency (no. Biomass (kg) Number
shots)
Australian Anchovy, Engraulis australis 32 795.0 231711
Sandy Sprat, Hypherlophus vittatus 20 364.7 96073
Yellowtail Scad, Trachurus novaezelandiae 21 217.6 6149
Pilchard, Sardinops sagax 26 93.7 3265
Blue jellyfish, Catostylus mosaicus 5 70.9 37
King George Whiting, Sillaginoides punctata 9 68.3 420
Southern Calamari, Sepioteuthis australis 30 57.7 749
Snapper, Chrysophyrus auratus 19 43.5 256
Barracouta, Thrysites atun 26 30.8 282
Silverbelly, Parequuela melbournensis 6 6.6 142
Luminous bay squid, Loliolus noctiluca 19 2.2 242
Warehou, Seriolella robustus 4 1.2 4
Blue Sprat, Spatelloides robustus 2 0.4 118
Silver Dory, Cyttus australis 6 0.4 6
Arrow squid, Notodarus gouldi 1 0.3 1

34
 Sandy sprats 2011 Yellow-tailed scad 2011
#
S #
S

# #
S

# # #
#
S

# #
S
#
S #
S

# # #
S

# #
#
S #
# #
# # #
S
# #
#
S #
S

#
S
#
S #
# # # #
# #
#

#
S
# # # #
#
S #
#
S #
S # #
# S
S # #
S #
#
S #

# #

#
S #
# #

#
S 0 # #
S 0 #

# 0.01 - 1 # 0.01 - 1
# 1.1 - 10 # 1.1 - 10
# 10.1 - 50 # 10.1 - 100
# 50.1 - 100 # 100.1 - 200

Pilchards 2011
#

#
S
#
# #
S

#
# #
# # #
#
#
#
#
S
#
S
# #
#
S
# #
#
# #
# #
#
S

#
#
#

#
S 0 #

# 0.1 - 1
# 1.1 - 10
# 10.1 - 50
# 50.1 - 100

Figure 14. Distribution and biomass (kg/10 min tow) of the three most abundant pelagic species (other
than anchovy) caught during the anchovy survey in 2011.

35
Appendix 3 Data Files
Electronic data files are as follows: Ageing data contained in the following
electronic data file:
• Catch.xls, Shot.xls, lengthfrequencies.xls.
• Anchovy_age_2011.xls
• Underway_sonar_2011.xls
• Fixed_sonar_catches_2011.xls
• Metadata descriptions are contained in
README_catch.doc, README_shot.doc
and README_lengthfreq.doc,
README_underway_sonar.doc,
README_fixed_sonar_catches.doc

36