1. No category

What is causing the decline of little
penguins (Eudyptula minor) on Granite
Island, South Australia?
Report to the South Australian Department for Environment and Heritage,
Wildlife Conservation Fund and the Nature Foundation SA
1
2
2
Natalie M Bool , Brad Page and Simon D Goldsworthy
1
University of Adelaide, North Terrace Adelaide SA 5009
South Australian Research and Development Institute (SARDI), 2 Hamra
Avenue, West Beach SA 5024
2
What is causing the decline of little penguins
(Eudyptula minor) on Granite Island, South
Australia?
South Australian Research and Development Institute
SARDI Aquatic Sciences
2 Hamra Avenue
West Beach SA 5024
Telephone: (08) 8207 5400
Facsimile: (08) 8207 5481
http://www.sardi.sa.gov.au/
Disclaimer
Copyright South Australian Research and Development Institute 2007.
This work is copyright. Except as permitted under the Copyright Act 1968
(Commonwealth), no part of this publication may be reproduced by any process,
electronic or otherwise, without the specific written permission of the copyright
owners. Neither may information be stored electronically in any form whatsoever
without such permission.
Printed in Adelaide, July 2007
SARDI Aquatic Sciences Publication Number F2007/000288-1
SARDI Research Report Series No. 217
Authors: Bool NM, Page B, and Goldsworthy SD
Reviewers: Dr Shane Roberts and Dr Adrian Linnane
Approved by: Dr. T. Ward
Signed:
Date: 12 July 2007
Circulation:
Public Domain
2
Table of Contents
1 Executive Summary ................................................................................................. 5
2 Background .............................................................................................................. 6
3 Introduction .............................................................................................................. 8
Aims and approach of the project ..........................................................................11
4 Methods ................................................................................................................. 12
Burrow monitoring ..............................................................................................12
Date of laying and fledging.................................................................................13
Analysis of reproductive success .......................................................................13
Chick growth rate ...............................................................................................13
Census of the breeding population.....................................................................14
Historical data.....................................................................................................14
Foraging ecology....................................................................................................15
Data analyses.....................................................................................................15
Spatial overlap in foraging area..........................................................................16
Little penguin diet ...............................................................................................17
Diet of New Zealand fur seals ................................................................................18
5 Results ................................................................................................................... 21
Reproductive success ............................................................................................21
Predation by rats ................................................................................................21
Inter-annual breeding success at Granite Island................................................21
Inter-annual breeding success at West Island ...................................................21
Chick growth.......................................................................................................22
Population census during the breeding season .................................................23
Foraging ecology................................................................................................24
Summary of foraging behaviour .........................................................................25
Foraging parameters ..........................................................................................25
Oceanography....................................................................................................27
Overlap of foraging areas between Granite and West Island penguins .............29
Penguin Diet...........................................................................................................29
Size and weight of prey ......................................................................................31
Comparison of diet between colonies ................................................................31
New Zealand fur seal diet...................................................................................32
Number of New Zealand fur seals hauled-out....................................................33
6 Discussion.............................................................................................................. 34
Reproductive parameters .......................................................................................34
Impacts of tourism and predation by rats on little penguins ...................................36
Predation by New Zealand fur seals ......................................................................37
Foraging parameters ..............................................................................................38
Similarity in prey diversity and meal sizes..............................................................40
Chick growth rates .................................................................................................41
The future of the little penguin populations at Granite and West Islands ...............42
7 Conclusions............................................................................................................ 45
8 Recommendations for future research................................................................... 47
9 References............................................................................................................. 48
10 Acknowledgements .............................................................................................. 55
3
List of Figures
Figure 1. The location of Granite and West Islands and other islands
mentioned in the text. ...............................................................................7
Figure 2. The number of active little penguin burrows during the breeding
season reported at Granite Island from 2001-2006. ...............................24
Figure 3. Circular histogram of the mean direction of travel of breeding little
penguins from Granite (blue) and West Islands (red) in 2006.................25
Figure 4. Summary of the inferred foraging routes undertaken by little
penguins at Granite Island (red) and West Island (blue) based on satellite
tracking in 2006. .....................................................................................26
Figure 5. The proportional time spent foraging in oceanic areas of 1 x 1 km
grids by 10 little penguins from Granite Island. Proportional time spent in
each grid is indicated by colour: where red represents regions where
penguins spent more time followed by orange, yellow, green and finally
blue areas where penguins spent relatively little time.............................26
Figure 6. The proportional time spent foraging in oceanic areas of 1 x 1 km
grids by 8 little penguins at West Island. Proportional time spent in each
grid is indicated by colour: where red represents regions where penguins
spent more time followed by orange, yellow, green and finally blue areas
where penguins spent relatively little time...............................................27
Figure 7. The average number of New Zealand fur seals hauled-out at Seal
Rocks, West Island and Granite Island between March-October 2006...33
List of Tables
Table 1. Measures of reproductive success for Granite and West Islands from
1990 – 2006. Data from 1990 - 2005 were obtained from the Granite
Island Little Penguin Monitoring Group (N. Gilbert and R. Brandle unpubl.
data). ......................................................................................................22
Table 2. Growth parameters of 12 little penguin chicks at Granite Island in
2006........................................................................................................23
Table 3. Growth parameters of 11 little penguin chicks at West Island in 2006.
................................................................................................................23
Table 4. Summary of the foraging parameters of little penguins at Granite and
West Islands in 2006. .............................................................................28
Table 5. Summary data of the oceanographic characteristics of the areas that
the little penguins foraged in from Granite and West Islands in 2006. ....29
Table 6. The prey consumed and their relative numerical abundance (NA)
and frequency of occurrence (FO) in stomach contents of little penguins
from the Granite and West Island colonies in 2006. The number of
samples examined, the number with prey remains and the minimum
number of individuals consumed were 37, 37 and 191 respectively. ......30
Table 7. The mean, minimum and maximum lengths (mm) and weights (g) of
anchovies, southern sea garfish and pilchards consumed at Granite and
West Islands. ..........................................................................................31
Table 8 The percent numerical abundance (NA) and frequency of occurrence
(FO) of prey types found in New Zealand fur seal scats at Granite and
West Islands. The number of scats examined, the number with prey
remains and the minimum number of individuals consumed are 29, 29
and 98, respectively................................................................................32
4
1 Executive Summary
The number of little penguins at Granite Island has been declining since the
early 1990s. Large numbers of tourists visit Granite Island to view the
penguins and the island is inhabited by non-native black rats (Rattus rattus)
and water rats (Hydromys chrysogastes) whereas nearby West Island does
not have any tourists nor land-based predators. Data on the little penguin
population size, breeding success, diet composition, foraging behaviour and
predation by both rats and New Zealand fur seals (Arctocephalus forsteri)
were collected at Granite and West Islands to determine the likely onshore or
offshore factors that may be causing the decline in the Granite Island
population. In 2006, the percentage of chicks that fledged was significantly
less at Granite Island (37%) compared to West Island (54%). Although there
was little overlap (9%) in the foraging areas used by penguins at each site,
there were no significant differences in prey species consumed or the mass of
stomach contents at Granite (45.0 ± 33.1 g) (mean ± SD) and West Islands
(46.0 ± 35.0 g). Both sites showed a similar delayed onset of breeding in 2006
and population surveys in 2006 indicated that both sites might be in decline,
suggesting that tourism and predation by rats are not solely responsible for
the decline in the Granite Island population. New Zealand fur seals consumed
adult penguins at both sites, indicating that they may be causing part of the
decline at Granite and West Islands and accordingly, ongoing monitoring of
fur seal diet is required.
5
2 Background
The little penguin population at Granite Island, South Australia is accessible
by a causeway from the township of Victor Harbour, and forms an important
local tourism attraction, with 30,000 tourists undertaking nightly guided tours
each year. Since monitoring began in 2,000, the number of little penguins
coming ashore have decreased by between 8-16 % per annum, and the
number of breeding pairs has declined from about 800 in 2001 to 300 in 2005
(N. Gilbert unpubl. data; R. Morcom unpubl. data). The cause(s) of these
declines are currently unclear, but are significant enough to jeopardise the
sustainability of the little penguin tourism business on the island. In addition to
significant numbers of tourists, Granite Island has populations of water rats
(Hydromys chrysogastes) and introduced black rats (Rattus rattus). Cats
(Felis catus) and foxes (Vulpes vulpes) are also occasionally found on the
island.
Little penguins also breed on West Island (6 km west of Granite Island),
where the population was estimated at be about 4,000 penguins in 1990/91,
with 2,029 being banded between 1990 and 1992 (M. Waterman unpubl.
data). The island is relatively unmodified and does not have any introduced
predators, and it is not visited by tourists. The current status of the little
penguin population there is unknown.
This study was undertaken at Granite Island (35º 65’ E, 138º 62’ S) and West
Island (35º 37’ E, 138º 35’ S) from March to November 2006. Granite Island is
a South Australian Recreation Park connected to the mainland by a
permanent causeway, which facilitates access to the island (Fig. 1). The park
receives up to 70,000 visitors per year (Department for the Environment and
Heritage unpubl. data). The waters around Granite Island are characterised
by shallow sea grass beds around the northern end of the island, with water
depths being a maximum of 16 m. (N. Gilbert unpubl. data).
6
West Island is a marine reserve 6 km west of Granite Island (Fig. 1). Access
to the island by the general public has been prohibited for 30 years (Shepherd
and Womersley 1970). The waters on the southern side of the island drop to
29 m, with a maximum depth of 5 m on the northern slopes. There are no
introduced terrestrial mammals on the island. The current number of little
penguin breeding pairs on West Island is unknown. West Island is a haul-out
site for non-breeding New Zealand fur seals. New Zealand fur seals also
reside on both the nearby island of Seal Rocks, which is located 2 km south of
Granite Island, and on a breakwater, which extends to the east of Granite
Island.
Figure 1. The location of Granite and West Islands and other islands
mentioned in the text.
The little penguin is a generalist predator, with clupeoid species being the
preferred prey (Gales and Pemberton 1990; Cullen et al. 1992). During the
breeding season, relatively short foraging trips of up to 20 km per day are
undertaken (Klomp and Wooller 1988; Weavers 1992; Ropert-Coudert et al.
2006). In contrast, post breeding foraging trips in preparation for the annual
moult can extend up to hundreds of kilometres from the breeding grounds
7
(Reilly and Cullen 1983; Weavers 1992). In South and Western Australia, the
first clutch is typically laid around April, with second and replacement clutches
occurring from August (N. Gilbert pers. comm.). Little penguins lay two eggs
per clutch and double breeding is common (Johannesen et al. 2003). It
typically takes 95 days from the lay date for chicks to fledge (Stahel and
Gales 1987). Eggs are incubated for approximately 36 days and chicks fledge
when they are around 59 days post hatching (Knight and Rogers 2004).
Chicks are guarded by at least one adult for approximately three weeks
following hatching, with foraging alternating between the two adults during the
guard-stage. After this period both adults forage at the same time to provision
their chicks.
3 Introduction
Seabirds are highly conspicuous components of the marine ecosystem and
thus typically have a high conservation profile (Wanless et al. 1998). The
decline of seabird populations has prompted numerous studies to examine
the interactions between these apex predators and their environment in an
effort to determine the cause of population decline (Bertram 1995; Catard et
al. 2000; Rindorf et al. 2000). Investigations into seabird decline have
highlighted that both natural and anthropogenic factors can affect seabird
populations at sea and on land (Lewis 1981; Monaghan et al. 1992; Furness
and Camphuysen 1997; Robinson et al. 2005). For example, populations of
seabirds nesting in the Peruvian upwelling system have declined in response
to guano harvesting and are further threatened because of reduced food
availability due to physical oceanographic changes and pressures from
commercial fisheries (Jahncke et al. 2004). Changes in food availability during
the breeding season (Norman and Ward 1992), predation (Burger and
Gochfield 1994), tourism (Giese 1996), habitat alteration, guano and egg
harvesting (Reilly 1994), and oil pollution (Goldsworthy et al. 2000) can have
short and long term impacts on seabird populations. Such pressures can lead
to population decline, particularly if they reduce reproductive success, survival
8
and/or recruitment, as has been indicated by the decrease of some little
penguin (Eudyptula minor) colonies in southern Australia (Dann 1992; Dann
and Norman 2006).
Seabirds may be disproportionately susceptible to several threats because
they have to cope with hazards on land and at sea (Edgar et al. 2005).
Seabirds are marine animals whose foraging grounds are spatially separated
from their breeding sites on land (Boersma 1998; Yorio 2000). This has led to
the evolution of a number of life-history characteristics that are in tune with
change in seabird’s marine environment (Yorio 2000). For example, seabirds
are long lived, have delayed maturity, experience low fecundity and are often
wide ranging (Yorio 2000), which means that populations are particularly
sensitive to changes in ocean productivity and environmental perturbations
such as the El- Nino and La Nina Southern Oscillation events (ENSO) (Lewis
1981; Barbraud and Weimerskirch 2001; Adams et al. 2004).
The marine environment is a dynamic system, in which food resources are
patchily and widely distributed (Weimerskirch et al. 2005). However, the ability
of seabirds to track their prey is restricted during the breeding season
because of the need to return to land to provision young and defend nesting
territories (Hunt et al. 1986; Ainley et al. 1998; Boersma 1998; Weimerskirch
and Cherel 1998; Catard et al. 2000; Forero et al. 2002). The energetic costs
of locating foraging grounds are critical elements to reproductive success
(Numata et al. 2000). Therefore, it is possible to detect changes in food
availability by monitoring changes in foraging habits using satellite telemetry
and examining reproductive parameters (Chastel et al. 1993; Bryant et al.
1999; Rey and Schiavini 2005). Measures of breeding success therefore
integrate information about the status of environmental conditions that affect
the availability of prey near seabird breeding colonies (Safina et al. 1988;
Diamond and Devlin 2003; Velarde et al. 2004; Rey and Schiavini 2005).
However, interactions between predators and their prey are typically complex
and tracking such relationships is difficult in a dynamic ecosystem (Botsford et
al. 1997; Croxall and Wood 2002; McCann et al. 2003; Doherty et al. 2004).
9
Seabird populations are not only sensitive to fluctuations in prey but also to
changes in predation rates (Long and Gilbert 1997; David et al. 2003). Seals
are known predators of seabirds and their numbers are increasing around
seabird colonies in Australia and Africa, following the end of exploitation by
sealers (Wickens et al. 1992; Shaughnessy et al. 1995; Ainley et al. 2005;
Mecenero et al. 2005; Page et al. 2005). It is possible that seabirds in these
areas increased in number and distribution in the absence of large
populations of seals (Crawford et al. 1989). However, now that many seal
populations are recovering, predation on seabirds has become prevalent,
such as in South Australia and in the Benguela ecosystem where New
Zealand fur seals (Arctocephalus forsteri) and Cape fur seals (Arctocephalus
pusillus pusillus) prey on penguins, gannets and cormorants (David et al.
2003; Page et al. 2005; Johnson et al. 2006). Although predators may have
negative impacts on seabird populations, quantifying these impacts is
challenging, because it is difficult to obtain information on the nature of the
interactions between seals and their prey, which occur at sea (Croxall and
Prince 1987; Descamps et al. 2005).
On land, seabirds often form large colonies because suitable nesting areas
without large numbers of predators are sparse (Burger and Gochfield 1994).
Seabirds are vulnerable on land to predators because they are less agile
there due to their physical adaptation to foraging at sea (Crawford et al.
1989). Little penguins and other seabirds are particularly vulnerable to
introduced predators, which may be able to enter burrows to eat eggs and
chicks (Dann 1992). The introduction of exotic predators, in particular rats
(Rattus spp), has led to the extinction of numerous animal species on islands
(Lyver 2000; Martin et al. 2000; Stapp 2002; Votier et al. 2004; Descamps et
al. 2005).
The current distribution of the little penguin, is smaller than indicated by
historical records, possibly because of habitat alteration and predation (Stahel
and Gales 1987; Gales and Pemberton 1990; Reilly 1994; Rogers et al.
1995). For example, little penguins recently became locally extinct at two sites
in Victoria, probably because of predation by red foxes (Vulpes vulpes) and
10
domestic pets (Dann and Norman 2006). In addition, a breeding colony of little
penguins on Kangaroo Island became extinct following the expansion of New
Zealand fur seals into their breeding area between 1992 to 2000
(Shaughnessy and Dennis 2001; Page et al. 2005). Consequently, little
penguin colonies in the vicinity of fur seal colonies and their haul-out sites and
those little penguin colonies, which are accessible to introduced predators,
are threatened by predation (Dann 1992; Weerheim et al. 2003).
At several breeding colonies throughout their range in southern Australia and
New Zealand, little penguins attract large numbers of tourists, which are of
significant importance to local economies (Dann 1992). However, disturbance
caused by tourism can adversely affect penguin breeding success (Yorio et al.
2001; Carpenter et al. 2004) by causing adults to desert their nests (Bolduc
and Guillemette 2003), or by causing chicks to receive fewer meals (Wilson et
al. 1991). For example, little penguins breeding in areas disturbed by tourists
had lower breeding success compared to those breeding in undisturbed areas
on Penguin Island in Western Australia (Klomp et al. 1991).
Aims and approach of the project
The aim of this study was to investigate factors that may be causing the
decline in the little penguin population at Granite Island. The approach was to
compare the breeding success and foraging ecology of little penguins
breeding at Granite and West Islands, and determine if differences between
sites could be explained by land-based factors including rat predation and
tourism, or at sea factors including prey availability and predation by New
Zealand fur seals. The numbers of breeding pairs at each site was also
surveyed in order to determine if recent declines in numbers were restricted to
the Granite Island population, or were part of a general decline in little
penguin numbers in the region.
11
4 Methods
Burrow monitoring
Data on little penguin breeding were collected on Granite and West Islands
from May to November 2006. Active breeding burrows were identified by the
presence of adult birds with eggs or chicks. Burrows were visited
approximately every fortnight on Granite Island, but less frequently on West
Island. In all, 73 burrows on Granite Island and 42 burrows on West Island
were monitored. Each burrow was numbered and marked with flagging tape
for individual identification.
The sex of adults was determined by measurement of bill length (vertical
thickness of the bill at the nostrils) and bill width (lateral thickness of the bill at
the nostrils) using callipers and equations developed by Gales (1988). Birds
were weighed in a cloth bag using a Salter spring balance (2kg x 10 g). Small
chicks were weighed in a smaller cloth bag using Salter 200 g x 1 g hanging
scales. Individuals were tagged by inserting glass encapsulated TIRISTM
(Texas, USA) transponders, which are individually encoded, subcutaneously
midway down the back. Tiris transponders were read using an AllflexTM
reader. Chicks were tagged once they had reached seven weeks of age
and/or had lost ≥90% of their mesoptyle down (Johannesen et al. 2003). To
determine whether rats were preying on chicks, all recently-killed chicks,
which were found dead in or near a burrow, were collected and examined for
signs of predator tooth marks.
12
Date of laying and fledging
The date of lay was determined by either recording the presence of the egg if
the burrow was checked within two weeks of the egg being laid or was
calculated by using the hatching or fledging date, following the methods of
Knight and Rogers (2004). Hatching date was recorded if the chick was seen
piping its egg or if the chick still had its eyes closed. If hatching was not
recorded and the chick was observed to fledge, the date of lay was
ascertained by back counting 95 days from the fledging date. A chick was
determined to have fledged if it was greater than 7 weeks of age and had >90
% of its adult plumage.
Analysis of reproductive success
Four measures of breeding productivity were calculated: 1) hatching success the number of chicks hatched per egg laid; 2) fledging success - the number
of chicks fledged per chick hatched; 3) egg success – the number of chicks
fledged per egg laid, and 4) the number of chicks fledged per breeding pair
(Rogers et al. 1995).
Chick growth rate
Growth parameters were calculated for chicks that had either fledged or were
very close to fledging (Rogers et al. 1995). Growth parameters were not
calculated for chicks in previous years because neither laying, hatch nor
fledge dates were available. The individual chick growth data were
summarised by fitting a logistic curve, because this was found to be the best
fit for penguin chick growth rates (Wienecke et al. 2000).
The logistic curve here is based on the one presented by O’Connor (1984)
and Wienecke et al. (2000):
S=
A
1− e−k(a−t )
Equation 1
where S is the parameter measurement, body mass (g) at age “a” days since
hatching; A is the asymptote where growth ceases, e is the base of the
natural logarithms, k is the rate at which the data drives the curve to
asymptote, and t is the age at which maximum growth occurs.
13
Census of the breeding population
To provide an indication of the size of the breeding population on both islands
a population census was carried out.
Granite Island
To coincide with the peak of nesting activity, the census was carried out on
August 11 and August 18, four weeks after several nests with eggs were
found (following the methods of Gilbert unpubl. data). Burrows were marked
with talcum powder to avoid double counting. The status of each burrow was
recorded as 1) active but empty (denoted by the presence fresh guano and/or,
tracks, nesting materials, recent burrow excavations), 2) burrow active (with
adult and or eggs and chicks present), 3) inactive, presence of cobwebs and
the absence of the above criteria. The annual rate of decline was calculated
by fitting an exponential curve to the census data.
West Island
The census on West Island was carried out over a period of two consecutive
days (September 20-21). Burrow status was determined using the same
criteria as those employed on Granite Island.
Historical data
To compare breeding success across locations and years, data were obtained
from the Granite Island Penguin Monitoring Group (N. Gilbert and R. Brandle
unpubl. data). Granite Island annual census data since 2001 are also used in
this study to compare the number of active burrows during the breeding
season across years (N. Gilbert unpubl. data).
14
Foraging ecology
To determine the location of foraging areas used by penguins from each
colony, satellite transmitters were deployed on adult birds that were rearing
chicks. The Cricket KiwiSat 202 (Sirtrack, Havelock North, NZ) satellite
transmitters are embedded in epoxy resin, are hydro-dynamically shaped and
weigh 28 g, which is less than 5% of a penguin’s body mass. Transmitters
have a saltwater switch that deactivates the transmitter when it is submerged.
Transmitters were glued directly onto feathers on the centre of the penguins
back, to coincide with its centre of gravity (Healy et al. 2004), using Loctite
401 glue, and were further secured with a single cable tie. Once the device
was attached, the penguin was released back into its burrow. Penguins were
recaptured on shore following a single foraging trip and the tracking device
was removed. If the penguin was caught before it returned to its burrow, its
catch was removed from its stomach using the water offloading technique
(Wilson 1984; Gales 1987) (refer to diet section, below).
Data analyses
Satellite derived location data from transmitters were provided through
Service Argos Inc, France. Accuracy of the location data is categorised into
six classes 1) Class 3: <150m, 2) Class 2: 150-350m, 3) Class 1: 350- <
1000m 4) Class A, and
5) B have no accuracy assigned, 6) Class Z: no location calculated (Sterling
and Ream 2004). The inaccuracy of Z positions, resulted in these being
discounted from analyses. Position data were filtered to exclude location
points which exceeded the maximum horizontal speed of 8.02 km/h for little
penguins, using the R statistical software (version 2.0.1, R Development Core
Team, R Foundation for Statistical Computing, Vienna) and the Time-Track
package (version 1.0-9, M. D. Sumner, University of Tasmania, Hobart). This
maximum horizontal speed is based on travel speeds of little penguins in
South Australia (A. Wiebkin unpublished data). Time-track also extrapolated
new positions between the actual locations based on a constant swim speed
between locations, so that there was a position at 15 minute intervals.
15
The foraging behaviour of each penguin was summarised to describe several
behavioural parameters: (1) the total distance travelled, (2) the maximum
distal point travelled in a straight-line from the colony (3) the mean bearing
travelled from the colony, and (4) the average speed km/h travelled (the
distance between interpolated locations, divided by the duration).
To describe the differences between penguin foraging in relation to physical
oceanographic parameters the: (1) mean and (2) median bathymetry, (3)
mean bathymetric gradient (change in depth in metres for each horizontal
kilometre), (4) median directional bearing of the bathymetric gradient (slope)
were calculated using the MapInfo program (MapInfo Corporation, Troy, NY,
USA) and the Vertical Mapper (GIS software) extension. The bathymetry
values were calculated based on the distance to the nearest nodes and were
assigned to each 15 minute interval. Bathymetry values were interpolated
based on the distance to the nearest bathymetry reading, taken from 1 km by
1 km depth readings (GeoScience Australia).
Spatial overlap in foraging area
To examine the degree of spatial overlap in the foraging areas used by birds
from Granite versus West Islands, the proportions of time spent in each 1 km
grid cell was explored using the niche overlap index (O), developed by
n
Schoener (1968): O = 1 − 0.5 × ∑ p1 j − p2 j
Equation 2
j =1
Where p1j and p2j are the proportion of time spent in the jth 1 x 1 km grid cell
by birds from each colony. If the 2 groups foraged in the same areas in
identical proportions, the index would equal 1. If entirely different areas were
used, the index would equal 0. One penguin which did not return to the
colony, was both included and excluded in this analysis, to examine whether
its foraging trip influenced the extent of spatial overlap between birds from the
two colonies.
16
Little penguin diet
To obtain diet samples, birds were caught as they returned to the colony.
Penguins were selected at random and as a result their breeding status
(breeding or non-breeding bird) was not known. Birds were held in an
enclosure until they were stomach flushed following the techniques outlined
by Wilson (1984) and Gales (1987). This was done by inserting a duodenal
tube Unomedical TM 6 mm diameter into their stomach to apply water using a
hand-held pump. After being lavaged the penguin was fed with 40 ml of a
blended pilchard mix and was given 40 ml of Vy-Trate TM diluted to 10% to
prevent dehydration. Birds were kept in the enclosure for 30 minutes to
ensure they were not harmed by the procedure. Each bird was weighed, a
TIRIS encapsulated transponder was inserted and flipper band numbers (if
present) were noted.
Samples were drained and weighed to the nearest 0.5 g. Large fleshy
remains were sieved to allow extraction of sagittal otoliths and cephalopod
beaks. All otoliths and cephalopod beaks were removed from the sample and
stored in vials. Fish species were identified by comparison to an otolith
reference collection (S. D. Goldsworthy and B. Page unpubl. data).
Cephalopods were identified from upper and lower beaks following standard
techniques (Lu and Ickeringill 2002; Chiaradia et al. 2003). Otoliths were
paired to estimate the minimum number of fish consumed and any odd
otoliths were assumed to represent additional individual fish. The minimum
number of cephalopods was estimated by counting pairs of lower and upper
beaks. Percentage of larval fish was estimated when present in a sample.
Otoliths in this case were too small to be extracted for identification.
To estimate the length and mass of fish consumed, sagittal otolith length was
measured using a binocular microscope, which was fitted with a graticule, to
the nearest 0.1 mm, using the Image- Pro Plus TM (Media Cybernetics, Inc)
image analysis program. Digestion of stomach contents causes degradation
of otoliths (Gales 1988), and hence can reduce the size of the otoliths and
may result in underestimates of fish length and mass. Therefore, only otoliths
that had not been affected by digestion were measured. The small number of
17
otoliths and cephalopod beaks collected from most prey species and the
unavailability of regressions for some prey species meant that analyses of
prey size were restricted to anchovies (Engraulis australis), southern sea
garfish (Hyporhamphus melanochir) and pilchards (Sardinops sagax).
The diet composition was summarised in 3 ways 1) the frequency of
occurrence (FO) (the proportion of samples collected that contained a specific
taxon), 2) percentage numerical abundance (NA) (percentage of the total prey
items made up by each prey taxa) and 3) wet mass. Estimated fish mass
(EFM) and fish length were calculated using otolith length versus fish length
regressions, which were calculated from the equations in Gales and
Pemberton (1990).
The niche overlap index was used to identify the extent to which the same
prey species were consumed by penguins from West and Granite Islands
(Equation 2) Schoener (1968), where p1j and p2j are either the percent
biomass or percent numerical abundance of the jth prey taxa for each colony.
If the 2 groups consumed the same prey in identical proportions, the index
would equal 1. If entirely different prey were taken, the index would equal 0.
Diet of New Zealand fur seals
Scats of New Zealand fur seals were collected from the non-breeding haul out
sites on the Granite Island break-water and from West Island from March to
November 2006. Each scat was placed separately into a plastic bag and was
taken back to the laboratory and stored at –20°C until examined. Scats were
soaked in hot soapy fresh water for at least one day to allow extraction of prey
remains. Individual scats were sieved using water in 1.0 and 0.5 mm sieves.
Otoliths, feathers and crustacean carapaces were stored dry and cephalopod
beaks were kept in 70% ethanol.
Scats from both islands were pooled to give an indication of seal diet in the
region, because the sample sizes were not large enough to describe patterns
at each island. Prey remains such as bird feathers, fish otoliths, crustacean
18
carapaces and cephalopod beaks were identified to the lowest taxonomic
group possible using a reference collection (S. D. Goldsworthy & B. Page
unpubl. data). To determine the importance of prey taxa in the diet, two
standardised measures were assessed 1) percentage numerical abundance:
NA) and 2) percentage scats with a particular prey item or frequency of
occurrence: FO. Estimation of the minimum number of individuals in each
sample was carried out by counting the: upper and lower cephalopod beaks,
left and right otoliths and eye lenses. Scats that contained penguin feathers
were assumed to contain one individual only. Where otoliths or bones were
too eroded to enable identification they were assigned to the unidentified prey
class. Octopods were identified to genus and squid beaks were identified to
Family level.
To provide an estimate of the number of New Zealand fur seals hauled-out in
the Granite and West Island region, counts were made on each visit to both
Granite (March – October) and West Islands (May – October). Because Seal
Rock is approximately 2 km from Granite Island, the number of seals present
was counted using binoculars (June – October). Where multiple counts were
conducted in a month the average number of seals counted is presented.
To determine whether there were any significant differences in reproductive
success, foraging behaviour and diet of little penguins between colonies
ANOVA were used if the data were normally distributed. When examining the
difference in prey size and mass between colonies a Levene’s test for equality
of variances was used, prior to analysis. If the data were not normally
distributed Mann-Whitney tests were used, for which Z approximations are
presented. Means are presented as ± standard deviation and all t tests are
two-tailed, unless stated, the α level of statistical significance was accepted at
0.05. Initial analyses indicated that data collected from male and females, in
all cases, did not differ significantly, so these data were pooled for analyses.
To explore correlations between foraging and oceanographic variables a
Spearman’s correlation coefficient r was used. Differences in the mean
direction of foraging trips among the two groups were tested using the circular
19
statistics package Oriana (V 2.02c, Kovach Computing Service, Pentraeth,
Wales, UK). A two- tailed Pearson parametric correlation coefficient was used
to examine the relationships between dietary variables.
To examine differences in diet composition and foraging behaviour of little
penguins at Granite and West Islands, Analysis of Similarities (ANOSIM),
using PRIMER® (Plymouth Marine Laboratory, UK) was used. ANOSIM ranks
the dissimilarities of the average of all groups using the Bray-Curtis
dissimilarity testing procedure (Quinn and Keogh 2002). To compare the rank
between-group and within-group variation a global test statistic (r) and pairwise R values were calculated. R is scaled from +1 to -1. Groups are
considered significantly different when R values are significantly greater than
0 indicating that within group variations are less than those between groups.
Negative R values indicate that within group dissimilarities are greater than
those between groups. The null hypothesis cannot be rejected when the R
value does not differ significantly from 0 indicating no significant difference
between groups.
20
5 Results
Reproductive success
In all, 62 and 28 breeding attempts by little penguin pairs at Granite and West
Islands respectively were included in the final analyses. The onset of breeding
was approximately 2 and 3 months later than average at both colonies. The
breeding season was also asynchronous at Granite Island with the first eggs
laid in June, with a peak hatching period in August. At West Island the onset
of the breeding season was later than at Granite Island. At West Island, the
peak of breeding activity (approximately 40 breeding burrows) occurred in
September, whereas in early August there were only 10 burrows with eggs
and two burrows with chicks.
Predation by rats
It was possible to confirm that rats killed 4 chicks at Granite Island. An
additional 15 dead chicks were found near their burrows with wounds inflicted
by rats, but in these cases it was not possible to determine if the chick had
been scavenged by the rat, because they had been dead for >1 night.
Inter-annual breeding success at Granite Island
Breeding success was variable across years with 0.3 to 1 chicks fledged per
pair (Table 1). On average 0.58 ± 0.23 chicks fledged per pair across all years
(n=9). The lowest number of chicks fledged per pair (0.30) occurred in 1991
and 2001. Breeding success from 2002 to the present (0.60-0.70 chicks
fledged per pair) did not fluctuate as much as in previous years (0.30-1.00
chicks fledged per pair).
Inter-annual breeding success at West Island
At West Island breeding success varied from 0.4 to 0.9 chicks fledged per pair
with an average of 0.78 ± 0.20 chicks fledged per pair across all years (n=5)
(Table 1). In years where comparative data were available in 1991, 2002,
2003 and 2006, breeding success was greater at West Island than at Granite
21
Island. This difference was most apparent in 1991 when only 0.3 compared to
0.8 chicks were fledged per pair at Granite and West Islands respectively.
Hatching success (F = 0.39, df = 1, P > 0.05), egg success (F = 1.80, df = 1, P
> 0.05), and the number of chicks fledged per pair (F = 1.2, df = 0.1, P > 0.05)
did not vary significantly between colonies. However, fledging success did
vary significantly (37.0% at Granite Island, 54.3% at West Island; F = 7.43, df
= 1, P < 0.05) (Table 1).
Table 1. Measures of reproductive success for Granite and West Islands from 1990
– 2006. Data from 1990 - 2005 were obtained from the Granite Island Little Penguin
Monitoring Group (N. Gilbert and R. Brandle unpubl. data).
Island
1990
1991
1995
1996
1999
2000
2001
2002
2003
2004
2005
2006
Mean
Granite Island
Hatching success
Fledging success
Egg success
No. of fledglings per pair
76.6
67.3
51.6
1.0
49.0
27.6
13.5
0.3
85.5
52.1
44.6
0.9
-
48.1
-
39.1
-
72.2
17.9
13.0
0.3
54.3
52.6
28.6
0.6
85.0
41.2
35.0
0.7
66.0
43.5
28.7
0.6
78.8
39.0
30.8
0.6
83.0
37.0
30.0
0.6
71.7 ± 13.3
39.8 ± 13.1
28.0 ± 12.6
0.6 ± 0.2
-
62.8
66.7
41.9
0.8
-
57.0
38.8
22.1
0.4
-
73.0
-
-
71.8
64.3
46.2
0.9
72.9
62.8
45.8
0.9
-
-
76.1
54.3
41.3
0.8
68.1 ± 7.9
60.0 ± 12.0
39.4 ± 9.9
0.8 ± 0.2
West Island
Hatching success
Fledging success
Egg success
No. of fledglings per pair
Chick growth
Chicks at Granite and West Islands reached their maximum growth rate at a
similar age and the rate of growth was similar for chicks at both sites in 2006
(Tables 2 & 3). The age at which maximum growth occurred t (Z = -1.01, P >
0.05), the maximum amount of growth per day for a chick (Z = -1.81, P >
0.05), the rate of growth since hatching k (F = 1.15, df = 1, P > 0.05) and the
asymptote weight (Z = 1.63 P > 0.05) did not differ significantly between
colonies.
22
Table 2. Growth parameters of 12 little penguin chicks at Granite Island in
2006.
Chick
1
2
3
4
5
6
7
8
9
10
11
12
Average
Asymptotic
mass
r
k
t
Max growth rate g/d
812
837
1091
862
980
884
870
1230
1165
819
1075
1430
1005
0.97
0.99
0.98
1.00
0.99
0.99
0.99
0.98
0.99
1.00
0.99
0.95
0.98
9.7
11.8
35.7
24.4
33.4
9.5
26.4
21.8
12.0
46.4
43.0
1.2
22.9
11
11
18
29
36
14
22
21
13
5
20
5
17
36
27
180
28
34
34
35
90
32
21
29
135
57
Table 3. Growth parameters of 11 little penguin chicks at West Island in 2006.
Chick
1
2
3
4
5
6
7
8
9
10
11
Average
Asymptotic
mass
r
k
t
Max growth rate g/d
639
870
992
440
614
771
722
895
764
1190
981
807
0.99
0.97
0.99
0.99
0.99
0.99
0.99
0.99
0.99
0.99
0.99
0.99
0.3
31.7
1.4
12.5
15.7
16.1
13.9
17.0
14.7
36.6
27.9
17.1
17
15
17
21
13
19
17
9
12
28
23
17
90
220
47
24
14
23
20
16
29
24
16
48
Population census during the breeding season
On Granite Island, the number of burrows occupied by adult little penguins
has been declining since 2001 (Fig. 2). There were 774 occupied burrows in
2001, which subsequently dropped to 414 and 294 active burrows in 2002
and 2003, respectively. However, in 2004 the number of active burrows
increased to 542. The mean rate of decline in the number of active burrows at
Granite Island was 15.6 %/year. No data on the number of active burrows are
available for West Island, prior to 2006. In 2006, 120 active burrows were
counted.
23
No. of active burrows
800
y = 679.89e-0.1453x
R2 = 0.4743
600
400
200
0
2001
2002
2003
2004
2005
2006
Year
Figure 2. The number of active little penguin burrows during the breeding
season reported at Granite Island from 2001-2006.
Foraging ecology
Satellite transmitters were deployed on 6 male and 4 female penguins on
Granite Island during July, and 5 male and 3 female penguins were tracked
from West Island during September 2006. The average direction of travel,
foraging routes and proportion of time spent in area are presented in Figure 3,
4 and 5, respectively. One of the penguins from West Island failed to return to
the colony after undertaking a foraging trip. The transmitter failed after four
days at sea. Therefore, part of the data on its foraging trip was excluded from
some analyses: trip duration, maximum distance from the colony, and total
distance travelled. In total, 146 and 280 unfiltered satellite positions were
obtained from Granite Island and West Island penguins, respectively. The
filter removed 8 (0.5%) and 64 (23%) locations from Granite and West Islands
respectively, thus the average number of filtered locations per day were: 14
and 27 for Granite Island and West Island penguins respectively. The mass of
adult birds at Granite (mean: 1260 ± 21 g) and West Islands (mean: 1130 ± 13
g) did not differ significantly (F = 1.86, P > 0.05) (Table 4), nor did the mass of
chicks (Granite Island mean: 281 ± 126; West Island mean: 321 ± 118; F =
1.019, P > 0.05). However, when all behavioural and oceanographic
parameters were compared across islands, ANOSIM detected a significant
24
difference in foraging behaviour between Granite and West Island little
penguins (one-way ANOSIM, P1000 = 0.01, Global R = 0.204).
Summary of foraging behaviour
Most penguins from Granite Island travelled away from the colony on a
median bearing of 135 (SE) degrees and travelled total distances of between
20 and 60 km (mean: 40.9 ± 3.9 km) (Table 5). Penguins from West Island
travelled at a median bearing of 207 (SW) degrees and travelled total
distances between 35 and 242 km (mean: 76.3 ± 28.1) (Fig. 3). The direction
of travel did not differ significantly for little penguins at both colonies (WatsonWilliams F-test, F = 0.18, df = 9, P > 0.100) (Fig.4).
0
4
3
2
1
270
4
3
2
1
1
2
3
4
90
1
2
3
4
180
Figure 3. Circular histogram of the mean direction of travel of breeding little
penguins from Granite (blue) and West Islands (red) in 2006.
Foraging parameters
The duration of foraging trips did not differ significantly between Granite Island
(0.54 ± 0.05 days) and West Island (0.97 ± 1.12 days) (Z = -1.39, P > 0.05).
Penguins from West Island reached a greater maximum distance (mean: 18.7
± 4.8 km) from the colony compared to Granite Island birds (mean: 10.0 ± 1.0
km) (Z = -2.57, P < 0.010) (Table 4). The average speed travelled by West
Island penguins (mean: 3.6 ± 0.4 km/h) and Granite Island penguins (mean:
3.4 ± 0.3 km/h) did not differ significantly (Z = -0.08, P > 0.05). Total distance
travelled was negatively correlated to bird mass (r = -0.63 P < 0.01). The
25
maximum distance from the colony was positively correlated to total distance
travelled (r = 0.56, P < 0.05), average speed (r = 0.54, P < 0.05) and chick
mass (r = 0.48, P < 0.05) (Table 4).
Figure 4. Summary of the inferred foraging routes undertaken by little
penguins at Granite Island (red) and West Island (blue) based on satellite
tracking in 2006.
Figure 5. The proportional time spent foraging in oceanic areas of 1 x 1 km
grids by 10 little penguins from Granite Island. Proportional time spent in each
grid is indicated by colour: where red represents regions where penguins
spent more time followed by orange, yellow, green and finally blue areas
where penguins spent relatively little time.
26
Figure 6. The proportional time spent foraging in oceanic areas of 1 x 1 km
grids by 8 little penguins at West Island. Proportional time spent in each grid
is indicated by colour: where red represents regions where penguins spent
more time followed by orange, yellow, green and finally blue areas where
penguins spent relatively little time.
Oceanography
There were no significant differences in water depth used by little penguins
from Granite Island (median depth: 38.4 m and mean depth: 34.1 ± 2.4 m)
and West Island (median 38.1 and mean depth: (39.5 ± 1.2 m) (F = 10.46, P =
0.05), nor in bathymetric gradient between Granite Island (0.34 ± 0.04 per km)
and West Island (0.34 ± 0.04 per km) (F = 0.18, P > 0.05) (Table 5).
27
Table 4. Summary of the foraging parameters of little penguins at Granite and West Islands in 2006.
Penguin
no.
665
707
787
844
671
688
705
726
821
972
7a
624
681
23
124
596
647
674
Colony
Month
Granite
Granite
Granite
Granite
Granite
Granite
Granite
Granite
Granite
Granite
West
West
West
West
West
West
West
West
July
July
July
July
July
July
July
July
July
July
Sep
Sep
Sep
Sep
Sep
Sep
Sep
Sep
Penguin
sex
F
F
F
F
M
M
M
M
M
M
F
F
F
M
M
M
M
M
Mass at
retrieval (g)
1060
1100
1500
1000
1000
1180
1235
1200
1450
1300
1260
1000
1180
950
1240
1150
Females Granite (mean ± sd, median)
Males Granite (mean ± sd, median)
Females West (mean ± sd, median)
Males West (mean ± sd, median)
Juvenile males (mean ± sd, median)
a
Incomplete foraging trip because the transmitter failed at sea.
Foraging trip
commenced
26/07/2006
11/07/2006
24/07/2006
12/07/2006
25/07/2006
14/07/2006
11/07/2006
15/07/2006
09/07/2006
23/07/2006
12/09/2006
17/09/2006
19/09/2006
12/09/2006
13/09/2006
15/09/2006
12/09/2006
18/09/2006
Foraging trip
duration (days)
0.5
0.5
0.5
0.5
0.7
0.5
0.5
0.5
0.5
0.6
3.0
0.6
0.6
0.6
0.5
3.5
0.5
0.5
0.5 ± 0.01, 0.5
0.5 ± 0.1, 0.5
1.4 ± 1.4, 0.6
1.1 ± 1.3, 0.5
Total distance Median bearing
travelled (km)
(degrees)
37.0
164
49.4
171
20.4
48
50.8
142
49.9
181
60.8
157
37.3
142
25.1
76
36.5
133
42.6
138
190.8
115
50.8
223
43.7
217
46.2
202
78.7
174
242.3
206
35.2
223
37.3
207
39.4 ± 14.1, 43.2
42.0 ± 12.3, 39.1
95.1 ± 82.9, 50.8
87.9 ± 88.0, 46.2
153
140
216
205
Maximum distance
from colony (km)
6.3
11.8
5.0
10.2
13.2
11.9
8.0
6.3
13.1
14.4
63.0
18.1
10.4
13.8
15.5
47.5
14.7
11.3
Mean horizontal
speed (m/s)
3.1
4.0
1.7
4.6
3.0
5.0
4.0
2.1
3.5
3.0
3.5
3.6
3.2
3.3
6.2
2.9
3.1
2.9
8.3 ± 3.2, 8.3
11.1 ± 3.2, 12.5
30.5 ± 28.4, 18.1
20.6 ± 15.1, 14.7
3.4 ± 1.3, 3.6
3.4 ± 0.1, 3.3
3.5 ± 0.2, 3.5
3.7 ± 1.4, 3.1
28
Table 5. Summary data of the oceanographic characteristics of the areas that
the little penguins foraged in from Granite and West Islands in 2006.
Colony
Granite
Granite
Granite
Granite
Granite
Granite
Granite
Granite
Granite
Granite
West
West
West
West
West
West
West
West
Penguin
No.
665
844
707
787
726
705
821
688
671
972
624
681
7
647
674
23
596
124
Mean Median Mean gradient
Penguin depth depth (change in depth
Month
sex
(m)
(m)
(m) per horiz. km)
July
F
20
27
0.4
July
F
21
23
0.4
July
F
30
36
0.2
July
F
6
5
0.6
July
M
13
15
0.4
July
M
21
22
0.3
July
M
27
30
0.2
July
M
29
35
0.6
July
M
29
35
0.3
July
M
30
36
0.1
Sept
F
25
27
0.4
Sept
F
28
35
0.7
Sept
F
36
39
0.1
Sept
M
24
27
0.9
Sept
M
31
35
0.3
Sept
M
32
35
0.2
Sept
M
32
33
0.2
Sept
M
33
36
0.2
Granite females (mean)
Granite males (mean)
West females (mean)
West males (mean)
19
25
28
31
27
32
31
35
0.3
0.8
0.3
0.4
Overlap of foraging areas between Granite and West Island penguins
Niche overlap indices calculated that the foraging areas used by the birds
from Granite versus West Islands overlapped by 9.0%. When the incomplete
foraging trip of bird 7 from West Island was included the proportion of spatial
overlap in foraging areas was slightly less, 7.0%. This suggests that there was
little spatial overlap in the foraging space of little penguins from each island in
2006.
Penguin Diet
In all, 57 birds were captured from July to October to obtain diet samples from
Granite Island (n = 43) and West Island (n = 14). Of these birds, only 25 (58
%) from Granite and 11 (78 %) from West Island had prey remains in their
stomachs. The average wet mass of samples was 45.0 ± 33.1 g (range: 8.5 –
113.7 g) at Granite Island and 46.0 ± 35.0 g (range: 10.0 g – 125.2 g) at West
29
Island. There was no significant difference in the mean sample mass between
the two islands (F = 0.12, df = 1, P > 0.05). Mean mass of stomach contents
was not correlated with body mass (r = -0.24, P> 0.05).
A total of 421 otoliths were found and measured, and classified to be in either
pristine or poor condition. This resulted in the exclusion of 332 otoliths from
the length and weight analyses because they were judged to be too eroded.
The prey species in this study comprised nine species of fish and two species
of cephalopods. In all, there were 191 individual prey items identified. The
most frequently detected prey for penguins at both colonies were anchovies,
with a FO value of 31.3% and 32.0% at Granite and West Island colonies,
respectively. Anchovies occurred in the diet in all months. The next most
frequent prey species were southern sea garfish 20.3% and 12.0%; pilchards
14.1% and 8.0%; Gould’s squid (Nototodarus gouldi) 9.4% and 12.0%; and
unidentified fish larvae 7.8% and 8.0% at Granite and West colonies
respectively. Other species of fish occurred but they were infrequently
recovered and typically found in low quantities (Table 6).
Table 6. The prey consumed and their relative numerical abundance (NA)
and frequency of occurrence (FO) in stomach contents of little penguins from
the Granite and West Island colonies in 2006. The number of samples
examined, the number with prey remains and the minimum number of
individuals consumed were 37, 37 and 191 respectively.
Prey type
NA
FO
Granite Is. West Is. Granite Is. West Is.
Anchovy (Engraulis australis )
South sea garfish (Hyporhamphus melanochir )
Pilchard (Sardinops sagax )
Unidentified larvae
Gould's squid (Nototodarus gouldi ) lower beaks
Tommy rough (Arripis georgianus )
Unidentified fish
Sandy sprat ( Hyperlophus vittatus )
Spotted pipe fish (Stigmatopora argus )
Redbait (Emmelichthys nitidis )
Snapper (Pagrus auratus)
Blue mackerel (Scomber australasicus )
Red cod (Pseudopycis bachus )
Calamari squid (Sepioteuthis australis ) lower beaks
Total
50.0
11.6
4.6
17.9
6.5
2.0
4.2
0.0
2.2
0.0
0.0
0.0
0.7
0.2
100
42.5
4.6
6.5
17.2
8.8
7.4
2.8
5.6
0.0
1.9
1.9
0.9
0.0
0.0
100
31.3
20.3
14.1
7.8
9.4
1.6
6.3
0.0
6.3
0.0
0.0
0.0
1.6
1.6
100
32.0
12.0
8.0
8.0
12.0
8.0
4.0
4.0
0.0
4.0
4.0
4.0
0.0
0.0
100
30
Size and weight of prey
Table 9 shows the mean, minimum and maximum lengths and weights of
anchovies, southern sea garfish and pilchards consumed by little penguins in
this study. The estimated length of each prey species did not differ
significantly between colonies: anchovy (t = 0.66, df = 18, P > 0.05); garfish (t
= 0.38, df = 7, P > 0.05), and pilchard (t = -0.958, df = 9, P > 0.05). The
heaviest fish was a garfish weighing 24.3 g with most fish weighing under 10
g. Only garfish exceeded 12 cm in length (89 % of fish) with the longest fish
being 21 cm. There were no significant differences in the mass of the different
prey between colonies, anchovy (t = 1.15, df = 18, P > 0.05), garfish (t = 0.93,
df = 7, P > 0.05) and pilchard (t = -1.033, df = 9, P > 0.05).
Table 7. The mean, minimum and maximum lengths (mm) and weights (g) of
anchovies, southern sea garfish and pilchards consumed at Granite and West
Islands.
Species
Anchovy
Anchovy
Southern sea garfish
Southern sea garfish
Pilchard
Pilchard
Colony
Granite
West
Granite
West
Granite
West
Length
(mm)
84.1
79.8
131.7
116.5
64.4
75.9
SD
12
6.9
34.4
3.2
15.7
22.7
Mass
(g)
6
4.7
5.2
2.2
3.2
5.5
SD
2.4
1.4
4.4
0.2
2.6
4.8
Max
length
124
89.9
219.8
118.8
94.4
100.5
Min
length
55.7
97.8
66.9
112.5
41.9
52.8
Max Min
mass mass
20.4 1.3
6.9
2.8
24.6 0.3
2.4
1.9
8.9
0.7
10.9 1.5
Comparison of diet between colonies
The species and number of prey consumed by little penguins were not
significantly different within and between colonies (one-way ANOSIM P1000 =
0.447, Global R = -0.03). Based on the niche overlap index, there was high
overlap in both the numerical abundance and frequency of occurrence of
species consumed between Granite and West Islands (80% and 70%
respectively). The differences in diets between colonies were accounted for
by differential consumption of anchovies (numerical abundance: 14 %,
frequency of occurrence: 1.4%) southern sea garfish (13%, 16%), tommy
rough (10%, 12%), sprat (10%, 7.8%), pilchards (3.6%, 11%) and pipe-fish
(12%, 4.3%) (Table 8).
31
New Zealand fur seal diet
In all, 34 samples were collected, of which 29 (83%) contained prey remains.
Fish were found in 20 (68%) of the samples containing remains, penguin
feathers were found in 12 (41%) samples and octopod beaks were found in 3
samples (10 %) and 1 sample had a squid beak (>1 %). In total, 5 species of
fish, 1 squid family, 1 Octopus genus, and penguins were identified. The
minimum number of individual prey identified was 98. In all, 80 individual fish,
12 penguins and 6 squid were consumed. Yellow-eyed mullet was the most
prevalent prey item (FO 35.8%), followed by; little penguin (FO 30%), garfish
(FO 7.5%), Octopus sp. (10%), cardinal fish (FO 5.0%), barracouta (FO
2.5%), unidentified fish (FO 2.5) and Ommastrephidae (FO 2.5%).
Table 8 The percent numerical abundance (NA) and frequency of occurrence
(FO) of prey types found in New Zealand fur seal scats at Granite and West
Islands. The number of scats examined, the number with prey remains and
the minimum number of individuals consumed are 29, 29 and 98, respectively.
Prey type
NA
FO
Yellow Eyed Mullet (Aldrichetta forsteri )
Little penguin (Eudyptula minor )
35.8
33.7
35.0
30.0
Garfish (Hyporhamphus melanochir )
Octopus sp.
Red Bait (Emmelichthys nitidus )
Cardinal (Vincentia conspersa )
Baracouta (Thyrsites atun )
Unidentified Fish
Ommastrephidae
Total
6.9
6.2
5.8
4.9
3.4
2
1.1
100
7.5
10.0
5.0
5.0
2.5
3
2.5
100
32
Number of New Zealand fur seals hauled-out
New Zealand fur seals were present throughout the region for the duration of
this study. Seal Rocks consistently had the greatest number of fur seals, with
a mean maximum of 32 ± 2.2 seals in June and a minimum of 13 ± 1.1 seals
in October (mean 22.4 ± 7.9 per month). At West Island the mean maximum
number of seals 12 ± 3.5 peaked in September and was lowest in May 5 ± 1.4
(mean 8 ± 2.5 per month). Overall, there were fewer fur seals on the Granite
Is than in the other locations, a mean maximum of 7 ± 1.4 in July and a
minimum of 1 ± 1.2 in Aug and Sept (mean 2.6 ± 1.8 per month) (Fig. 8).
35
Seal Rock
30
West Island
25
Granite Island breakw ater
20
15
10
5
S
t
ep
te
m
be
r
O
ct
ob
er
A
ug
us
Ju
ly
Ju
ne
M
ay
M
ar
ch
0
Figure 7. The average number of New Zealand fur seals hauled-out at Seal
Rocks, West Island and Granite Island between March-October 2006.
33
6 Discussion
This study investigated factors that may be responsible for the population
decline in little penguins at Granite Island. The breeding success of little
penguins at Granite and West Islands were compared, as were aspects of the
diet and foraging behaviour. Results comparing the reproductive success and
foraging ecology of each population, the current size of each population and
the importance of little penguins in the diets of New Zealand fur seals are
discussed below.
Reproductive parameters
Although the number of active burrows fluctuated at Granite Island, overall the
number of active burrows has decreased since 2001. The number of active
burrows found in 2006 at West Island was much less than expected given the
large number of birds banded there in the early 1990s. However, the number
of active burrows at West Island was underestimated in 2006 because some
areas were inaccessible (approximately 50 m by 50 m), although there did not
appear to be many birds entering this area. Another census is required to
confirm that the West Island population has declined, although findings from
this study indicate that the population has declined significantly since the early
1990s.
Seabirds typically breed when their prey peak in abundance in near-shore
waters, which minimises the time adults spend commuting between foraging
grounds and the colony (Grémillet et al. 2004). Therefore, when prey
abundance is poor near the colony the onset of breeding can be delayed
(Dann et al. 2000). The breeding season in 2006 was delayed by 2 and 3
months at Granite and West Islands respectively. Interestingly,
commencement of breeding at nearby Troubridge Island (Fig. 1) was also
delayed in 2006 (A. Wiebkin unpubl. data). The Bonney Upwelling, which
nourishes this region, was reported to be poor in 2006 because of a lack of
winds from the SE (P. Gill and A. Levings pers. comm.). Although the
34
distribution of little penguin prey was not measured in this study, it is possible
that the peak abundance of their prey was delayed or relatively small,
because the poor upwelling resulted in depressed primary production (Ward
and Staunton-Smith 2002).
Such factors may have influenced the delayed onset of breeding of little
penguins breeding at these colonies this year. The late finish of the moult this
year may have also influenced the late onset of breeding. At Granite Island,
the moulting period was longer than usual in 2006 (N. Gilbert pers. comm.),
which has been associated with a late onset of breeding at other little penguin
colonies (Reilly and Cullen 1983). When egg laying is delayed, the ability of a
little penguin to raise a second clutch is reduced because they abandon their
dependent chicks in preparation for the annual moult, which potentially halves
the annual breeding success (Perriman et al. 2000; Knight and Rogers 2004;
Robinson et al. 2005).
Both colonies showed marked inter-annual variation in breeding success,
however little penguins at West Island produced relatively more fledglings per
pair. Inter-annual variation in breeding success is common among penguins
and in other seabirds and has been linked to food availability and predation
(Bertram 1995; Rindorf et al. 2000; Wienecke et al. 2000). Overall, the
breeding success at Granite and West Islands was relatively poor, compared
to Philip Island (Victoria) and Lion Island (New South Wales), where the
respective average numbers of chicks fledged was 1.0 ± 0.4 and 0.88 ± 0.18
per breeding pair per year (Rogers et al. 1995; Dann et al. 2000). Based on
rankings system to assign to breeding success at Phillip Island (Robinson et
al. 2005), little penguins at Granite and West Islands had poor breeding
success in 66% and 25% of the years presented in this study, respectively.
In 2006, the success of earlier breeders was lower than those that bred later
in the season, in contrast to the findings of other studies (Chiaradia and Kerry
1999). This finding is presumably because earlier breeders typically have
shorter incubation and foraging trips, that reduces the time between meals for
35
chicks and leads to greater fledging success (Reilly and Cullen 1981).
Although, this observation may not be as pronounced when food availability is
poor early in the breeding season. The findings presented here based on the
timing of breeding, indicates that prey may have been in relatively low
abundance early in the breeding season.
Impacts of tourism and predation by rats on little penguins
Interestingly, the breeding success of penguins at Granite Island is
comparable to that at Penguin Island, where penguins are also a tourist
attraction (Klomp et al. 1991). The colonial nesting of seabirds in large
aggregations in often scenic coastal locations attracts many tourists (Yorio et
al. 2001). To accommodate the needs of the expanding tourism industry on
Granite Island, a restaurant, gift shop and penguin information centre were
built in 1995 and vehicles regularly use the area where little penguins come
ashore.
At Granite Island little penguins nest in burrows and they may therefore be
less sensitive to disturbance than those that nest on the surface. This implies
that disturbance when little penguins are returning to their burrows and when
they return to the sea is most critical. Penguins are highly faithful to their
landing sites, where disturbance delays breeding birds from coming ashore
and may deter young birds from breeding in these areas (Weerheim et al.
2003). Klomp et al. (1991) demonstrated that little penguins breeding in areas
with many tourists had lower breeding success (26%) compared to those
breeding in areas of low tourism impact (40%). At Granite Island, considerable
effort has been made to reduce disturbance to the little penguins nesting in
the areas frequented by tourists. Nest boxes and board-walks have been
constructed and access to the island is restricted at dusk to people taking part
in an organised penguin tour. Despite these efforts, it is possible that tourists
have negative impacts on the Granite Island population.
Numerous species of seabirds are negatively affected by predation by
introduced rat species (Martin et al. 2000; BirdLife International 2004).
36
Predation by rats was shown to reduce breeding success by 24% at a little
penguin colony in New Zealand (Perriman et al. 2000). At Granite Island, rats
preyed on relatively small chicks, post incubation, rather than on eggs, which
has been the case in other seabird studies (Igual et al. 2006). Chicks being
incubated by their parents are less vulnerable to predation by rats, because
adult penguins could keep rats at bay. In many cases, it was not possible in
the current study to distinguish whether chicks had died as a result of wounds
inflicted by rats or if rats had scavenged dead chicks, as has been noted in
other studies (Cunningham and Moors 1994; Stapp 2002). In addition, when
chicks disappeared from their burrows it was not possible to determine the
cause of death. Daily checks of burrows would be required to determine how
many chicks were killed by rats, which was not feasible in the current study. In
addition, extensive rat baiting was carried out during the penguin breeding
season in 2006 on Granite Island, which may have reduced the number of
chicks killed by rats. In previous years, the number of chicks killed has been
observed to be greater (N. Gilbert pers. comm.). Despite the absence of
tourists and terrestrial predators on West Island, there appears to have been
a substantial decline in the number of birds nesting on this island as well. This
indicates that, in addition to onshore predation and disturbance caused by
tourists at Granite Island that other perturbations may be placing pressure on
both populations.
Predation by New Zealand fur seals
This study confirmed that little penguins formed a significant part of the diet of
New Zealand fur seals around Granite and West Islands. Little penguins were
the second most prevalent prey item suggesting predation pressure could be
important. Based on these findings, if we conservatively set the number of
resident fur seals in the region at 20, and if they consume a single penguin
per week, more than 1000 penguins would be consumed in a year. Given that
the number of little penguins at Granite and West Island probably only number
in the hundreds, this estimate of fur seal predation would not be sustainable.
A more conservative estimate of fur seal predation of 20 seals eating one
penguin per month would equate to 240 adult penguins being taken a year.
37
The average number of resident seals is likely to be greater than 20 (if
surveys in this study are an indication), and based on the dietary data
predation rates on penguins could be even higher. However, the importance
of little penguins in the diet of New Zealand fur seals from nearby Seal Rocks,
the largest haul-out in the region, is unknown. Predation of adult birds during
the breeding season has dual impacts because not only is an adult lost, but its
clutch is also likely to be abandoned.
Fur seals prove a threat to birds by direct consumption and by competition for
breeding spaces. For example, the consumption of little penguins by New
Zealand fur seals around Kangaroo Island is thought to have caused the local
extinction of a little penguin breeding colony at Cape Gantheaume (Page et
al. 2005). Whereas, competition for space with Cape fur seals in recent years
has resulted in several seabird colonies being displaced on islands in Namibia
(Crawford et al. 1989). Populations of New Zealand fur seal on Kangaroo
Island are currently increasing by 12.5% per annum, and have increased
almost 10-fold in size over an 18-year period (based on data presented in
Shaughnessy 2006). As populations of New Zealand fur seals continue to
recover, predation rates are likely to increase, escalating the risk of more
localised extinctions, including populations at Granite and West Islands.
Foraging parameters
The results obtained in this study provide insights into the foraging behaviour
of little penguins during the chick raising period. Unexpectedly, there was very
little spatial overlap in the foraging areas used by penguins from the two
colonies. Fish stocks are known for their heterogenous distribution and
therefore, the utilisation of different foraging areas by penguins from each
island may reflect movement of fish during the breeding season
(Weimerskirch et al. 2005) and the differences in timing of satellite tracker
deployments on birds from the two islands.
Although, penguins may reduce competition for prey with con-specifics from
nearby islands by using foraging areas that do not overlap. Aggregations of
breeding seabirds may deplete local food resources (Ashmole 1963; Birt et al.
38
1987), which increases the separation of breeding and feeding habitats and
amplifies the cost of commuting to provision dependent young, which remain
at the central place (Orians and Pearson 1979). Ideally, seabird colonies
should be sufficiently spaced to reduce intra-specific competition for prey
(Cairns 1989), however, the availability of islands suitable for breeding often
occur in groups (Forbes et al. 2000). There are many examples of how intracolony competition can result in different foraging strategies. For example,
magellanic penguins (Spheniscus magellanicus) foraged close to their
colonies and consumed fish that were of relatively poor quality rather than
feeding in distant waters, which were used by penguins from other colonies
(Forero et al. 2002). In contrast, cape gannets (Morus capensis) at nearby
colonies in South Africa exhibited complete separation in foraging areas,
despite the islands being within a cape gannets foraging range (Grémillet et
al. 2004). West and Granite Islands are located well within the foraging
ranges of little penguins breeding on either island. Considering the small size
of the populations it is unlikely that there is density dependent localised prey
depletion and resultant foraging separation around the islands.
With the exception of two birds from West Island (one of which did not return
and is not considered further), all little penguins undertook <16 hr foraging
trips, which were within 20 km of the colony. Little penguins forage by sight
and are therefore only able to forage during daylight, and their arrival and
departure times from their colonies reflect this restriction (Ropert-Coudert et
al. 2006). The average maximum distance travelled in a day was greater at
Granite (41 km) and West Islands (48 km) than little penguins at Phillip Island
(33 km) (Weavers 1992), indicating that little penguins in this study travelled
relatively further in search of prey. Little penguins have been shown to make
longer foraging trips and travel further when prey availability is relatively low
(Hobday 1992; Weavers 1992). The greater travel speeds and travel
distances of little penguins in this study suggest that the prey availability
around Granite and West Islands was reduced during in 2006.
39
The average travel speed in this study at Granite Island (3.4 ± 0.3 km per
hour) and West Island (3.6 ± 0.4 km per hour) was also greater than at Phillip
Island (1. 5 ± 1.1 km per hour (Weavers 1992). Marine animals exploiting
patchy environments would be expected to move slowly in areas where
resources are plentiful and quickly move through areas where resources are
scarce, because searching for plentiful patches is typically more beneficial
than remaining in sparse ones (Pinaud and Weimerskirch 2005). These
findings support the conclusions of relatively low prey availability around
Granite and West Islands during 2006. Although, considering that little
penguins from West Island on average travelled further in search of prey, yet
they had greater reproductive success suggests that in the current season
increased effort in searching for food did not lower reproductive success at
West Island compared to Granite Island.
Similarity in prey diversity and meal sizes
The relatively late breeding season, low breeding success, long duration and
distance of foraging trips, which were documented in this study, indicate
relatively poor prey availability. Although prey availability was not measured,
the diet of penguins on both islands was quantified, and provides information
on the size of individual prey available and the size of meals fed to chicks.
However, seabird diet studies must be interpreted with caution, because they
are subject to biases as a result of differential rates of digestion, which
depend on: 1) stomach fullness, 2) the time the food has spent in the stomach
and 3) the size of the prey (Gales 1988; Gales and Pemberton 1990).
Consequently, otoliths can be eroded to varying degrees, which can affect
estimates of the size of individual prey consumed, and some otoliths may be
completely digested. Although otoliths that were completely digested cannot
be accounted for, the impact of these biases was likely to be minimal as only
otoliths that were in pristine condition were used to estimate the size of prey
consumed.
As expected, there was significant overlap in the diet of penguins at both
islands, despite the relatively disjunct foraging habitats used by these
40
populations. The diet of little penguins at Granite and West Islands reflects the
generalist foraging behaviour of little penguins at other locations (Montague
and Cullen 1988). Fish were the most common prey in the diet, with only two
squid species being recovered. Fish are also usually the most frequently
consumed prey of little penguins in Tasmania and Victoria (Gales and
Pemberton 1990). Although many species of fish and squid were eaten,
anchovies (80-100 mm in length) occurred most frequently, which is similar to
the findings of other dietary studies on little penguins in South Australia (A.
Wiebkin unpubl. data).
Relatively little is known about small pelagic fishes and their distributions and
abundance in South Australia (Ward et al. 2001; Ward and Staunton-Smith
2002; Ward et al. 2006). Anchovies and pilchards dominate the clupeoid
assemblages in temperate Australia and both species form large schools,
occupy similar habitats and undergo comparable fluctuations in abundance
(Ward et al. 2001). Little penguins typically prey on juvenile anchovies and
pilchards, which are less than 1 year of age (Hobday 1992). The availability of
suitably sized prey exploited by little penguins is likely to be linked to the
strength of coastal upwellings, which are thought to strongly influence the
breeding success of small pelagic fish (Ward et al. 2006). The peak in
anchovy and pilchard spawning generally occurs from January to April in
South Australia, which is associated with the timing of coastal upwelling that
brings nutrient rich waters, giving rise to relatively high primary, secondary
and tertiary production (Ward et al. 2006). In years of strong coastal
upwelling, penguins at Phillip Island attain better body conditions and breed
earlier, possibly because of increased prey availability (Mickelson et al. 1992).
Therefore, it is possible that the timing of breeding of little penguins at Granite
and West Islands is also influenced by the strength of regional coastal
upwellings.
Chick growth rates
Breeding success and chick growth are typically correlated to the meal sizes
fed to chicks (Barrett et al. 1987; Olsson 1997; Dahdul and Horn 2003;
Ramos et al. 2006). Older chicks are preferentially fed when food availability
41
is poor to improve the chances of fledging at least one healthy chick, this
validates that chick survival can be reduced by limited food resources (Radl
and Culik 1999). Meal sizes fed to chicks at Granite and West Islands (45 g
and 46 g) were smaller than those fed to chicks at nearby Troubridge Island
(Fig 1), (315 g, A. Wiebkin unpubl. data), but comparable to those in
Tasmania (31-47 g) (Gales and Pemberton 1990) and Victoria (80 g)
(Montague and Cullen 1988). The amount of food that a chick is fed
determines its fledging mass and ultimately influences its survival. At
Troubridge Island the average fledging mass was 1300 g (A. Wiebkin unpubl.
data), whereas at Granite and West Islands the asymptotic mass was 955 g
and 800 g respectively.
Breeding success only indicates the number of chicks that fledge, whereas
mass at fledging can be a better predictor of survival to adulthood because
chicks, which fledge with higher body mass, have greater survival rates
(Giese et al. 2000). Therefore, the lower body mass at fledging of chicks in
the current study may result in poor survival post fledging. The lower fledging
success at Granite Island was not related to either meal sizes nor chick
growth rates, indicating that differences in breeding success between the two
islands were not due to a difference in the capacity of adult little penguins to
deliver food to their chicks. In fact this study highlights the similarities in the
prey species eaten by penguins at both sites.
The future of the little penguin populations at Granite and West Islands
Based on estimates of the size of the breeding population of little penguins at
Granite and West Islands conducted during this study, both sites appear to be
in a state of decline. This result, plus the fact that both sites displayed a
similar late commencement of breeding during the 2006 season, and because
little penguins targeted similar prey species, suggests that both populations
are responding to factors in their marine environment in similar ways. The
decline of both little penguin colonies suggests that either predation pressures
or changes in food availability may be causing reductions in adult little
penguin survival at both colonies. However, this study has identified strong
42
evidence in support for a ‘predation hypothesis’, where recent declines in little
penguin populations are attributed to increasing predation pressure from
recovering New Zealand fur seal populations.
In addition, the potential impacts of land-based pressures from either rat
predation or disturbance by tourists at Granite Island cannot be discounted.
Although no differences were detected in meal masses, chick growth rates,
foraging trip duration and distance to foraging sites between Granite and West
Island, the breeding success was significantly lower at Granite Island,
possibly because of lower chick survival. Evidence of young chicks being
removed from burrows and killed by rats, suggested that land-based predation
of chicks may be a factor responsible for lower reproductive success at
Granite Island. However, the potential role of disturbance from tourism in
reducing breeding success could not be discounted in this study.
To understand how the combined factors of predation, prey availability and
tourism may affect population declines at Granite and West Islands, it would
also be necessary to investigate how variations in survival and fecundity affect
their population demography (Gardali et al. 2000; Jenouvrier et al. 2005).
Seabird populations are considered to be most vulnerable to a reduction in
adult survival because they have low fecundity and because an increase in
adult mortality has an immediate impact on population size (Lewison et al.
2004). Because rats are only capable of preying on small unattended little
penguin chicks, they are not likely to be impacting on the Granite Island
population as negatively as predation by fur seals. Therefore, predation by
New Zealand fur seals is likely to have a major impact on the population sizes
of both colonies.
However, low fecundity and reduced reproductive success and/or poor
juvenile survival over the long-term can lead to population decline. For
example, a population of emperor penguins (Aptenodytes forsteri) declined by
50% during the 1960s because of climate change (Jenouvrier et al. 2005).
The population did not recover, because reproductive success for the next 30
years was poor. Given the continued poor breeding success at Granite Island,
43
it is possible that recruitment will limit population recovery (Thompson and
Ollason 2001). Therefore, it is important to bolster the reproductive success
and subsequent recruitment of juveniles into the breeding population at
Granite Island to ensure the perpetuity of this colony.
44
7 Conclusions
This study identified that the decline in little penguin numbers were not
restricted to Granite Island, and they appear to also be decreasing at
neighbouring West Island. This suggests that declines could be due primarily
to changes in the marine environment that affect penguin survival. The very
significant levels of predation by local New Zealand fur seals, as supported by
the importance of little penguins in their diet, could account for much of the
observed decline, especially as their populations (and presumable predation
pressure on little penguins) have increased rapidly over the last 20 years.
Reductions in prey availability may also be affecting the status of little penguin
populations, but there was not evidence to support this in the current study.
Breeding success of little penguins at Granite Island was significantly lower
than at West Island, potentially because of lower chick survival. This may be
the result of predation of young chicks by rats. Further research is needed to
quantify the impact rat predation has on overall reproductive success at
Granite Island. Although the role of disturbance by tourism could not be
discounted, results suggest that land-based factors are also contributing to
population declines in the Granite Island population. As such, the rates of
decline in penguin populations are likely to be greater at Granite Island
compared to West Island. Effort should be directed to continuing population
surveys at both sites, and at reducing the impact of land based predators and
tourism at Granite Island.
The results from this study indicate that no single factors are responsible for
the little penguin population decline at Granite Island. The relatively short
duration of this project meant that it was not possible to quantify how each
factor (adult predation, nestling predation, low nesting effort or nestling
mortality due to a lack of food) or the combination of these factors have
contributed to the rate of population decline and this remains to be
investigated. These findings indicate that because the population is still
45
declining, the number of tourists allowed to go on nightly penguin tours should
not increase to further limit the impacts of tourism. Tourist impacts present a
difficult challenge for wildlife managers, but the eradication of rats may
increase recruitment. New Zealand fur seals and water rats are protected
species and therefore the management of their interactions with little penguins
also presents a challenging management issue. The inter-play between
tourism, predation, fluctuating oceanographic productivity and the relationship
between prey availability and foraging success and reproductive success are
complex challenges that face little penguins at these sites. Small increases in
the impacts of any one of these factors on little penguins may ultimately be
sufficient to progress these little penguin subpopulations to local extinction.
46
8 Recommendations for future research
Carry out additional population surveys at West Island to determine whether the little
penguin population there is in decline.
Continue monitoring the survival of chicks on both islands to determine post fledging
survival.
Continue baiting of black rats on Granite Island and monitor penguin burrows
frequently to provide a better estimate of the number of little penguin chicks killed by
both black and water rats.
Continue diet studies on both little penguins and New Zealand fur seals.
Carry out studies on the interactions between little penguins and tourists as the little
penguins return to the shore in the evening. This will help to differentiate between the
impact of rats and tourists on breeding success at Granite Island.
47
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10 Acknowledgements
The Fisheries Research and Development Corporation, the Nature
Foundation of South Australia and the SA Department for the Environment
Wildlife Conservation Fund provided funding for this project.
David Paton also provided academic advice and guidance on project design.
Thanks to the fellow students in the GAB lab at SARDI for all of their help and
encouragement. Especially Annelise Wiebkin, Al Baylis, Luke Einoder and
Lachie McLeay. They showed me the ropes, gave advice on field techniques
and data analyses, and they answered my calls for satellite downloads well
into the night. I would especially like to thank Annelise who provided advice
on field techniques and she was a wealth of knowledge when it came to little
penguins.
Field work was carried out with the assistance of a number of eager
volunteers, including Sam Adams Denise Bool, Leanne Trott, Augi Facelli,
Beiha Malen Yanez, Toni Milne, Dave Turner, Heidi Valstar, Alex Pembshaw
and Clare Adams.`
The dedicated staff and volunteers at Granite Island, Natalie Gilbert, Nick,
Dorothy, Keith and Ruben provided assistance in the field, warm cups of
coffee and plenty of encouragement. In particular, Natalie Gilbert provided me
with her historical data, also passed on her knowledge and expertise of the
little penguins on Granite and West Islands. I am very grateful for her support
and assistance this year.
55