PhD Abstract and Outline: Falk Hüttmann (Ph.D., Diplom-Forstwirt univ.) PostDoc Fellow, Simon Fraser University |
Falk Hüttmann
Centre for Wildlife Ecology, Biology Dept.
8888 University Drive, Simon Fraser University (SFU), Burnaby B.C., Canada V5A 1S6
Tel: 604 291 5618 Fax: 604 291 3496
Email: huettman@sfu.ca
Environmental determination of seabird distribution
by
Falk Hüttmann
Faculty of Forestry and Environmental Management, ACWERN UNB
ABSTRACT
This thesis investigates major aspects of seabird distribution. The general topic of seabird distribution is first reviewed and presented. I then introduce the study area by stratifying the Seascape of the Northwest Atlantic using 20 environmental data sets, a Geographic Information System (SPANS-GIS), and Clustering evaluated by Classification and Regression Trees (Cart). For the first time the physical environment of the study area is described, laying the groundwork for the ‘Seascape Ecology’ approach developed in the thesis.
I evaluated existing knowledge of seabird migration with data from the PIROP (Programme Intégré de recherches sur les oiseaux pélagiques) database using monthly observations of immature and moulting seabirds to track migration. The results allow, for the first time, the identification of flyways using at-sea observations throughout the year.
I then used the 20 environmental data sets, and GIS-overlaid them with standard seabird at-sea observations for winter, including results from Principal Component Analysis to characterize seabird assemblages. Logistic regression identified significant predictors, and Cart developed descriptive models. Unknown to previous knowledge, aspect of sea floor and distance to coast were the most frequent significant predictors of seabird distributions.
For the breeding season I hypothesized that the seabird colony location is the driving force for seabird distribution. The same methods as for the investigation of winter seabird distribution were used. Specific seabird model predictions using Cart were evaluated by the nearest distance to seabird colonies, and by geo-referenced observation residuals. Results showed that breeding seabirds are attached to colony locations. Non-breeders can stay far away from colonies, and colonies can also be located in non-optimal foraging habitat. Distribution modelling allows for filling gaps in previous information on seabird distribution and colony locations.
I investigated scale issues, a major methodological question in seabird distribution research, using again the PIROP data set. The results indicated that scale had no major effect on previous PIROP analyses, but was useful for describing seabird distribution and for seabird management questions.
I have demonstrated that seabird distributions need to be examined throughout the year, and that new understanding can be achieved using abiotic parameters alone. Modern remotely-sensed data sets and statistical approaches offer unparalleled opportunities to track changing responses of seabirds to environmental change.
TABLE OF CONTENTS with page numbers
ABSTRACT ................................................................................................................ ii
PREFACE................................................................................................................... iv
ACKNOWLEDGEMENTS....................................................................................…. ix
TABLE OF CONTENTS........................................................................................… xiii
LIST OF TABLES....................................................................................................... xv
LIST OF FIGURES .............................................................................................….. xvii
CHAPTER 1..................................................................................................................1
General Introduction and Literature review
References....................................................................................................................31
CHAPTER 2................................................................................................................62
Characterizing the marine environment of seabirds of the Northwest Atlantic
References..............................................................................................................…..99
CHAPTER 3..............................................................................................................137
Seabird migration in the Canadian North Atlantic: moulting locations and
movement patterns of immatures
References..................................................................................................................180
CHAPTER 4 ..............................................................................................................213
A Descriptive Model of Environmental Determination of Winter Seabird
Distribution in the Canadian North Atlantic
References...................................................................................................................250
CHAPTER 5...............................................................................................................293
Seabird colony locations and environmental determination of seabird
distribution: A spatially explicit seabird breeding model in the Canadian North
Atlantic
References..................................................................................................................328
CHAPTER 6..............................................................................................................366
Scale questions in seabird research: Selected issues for the Northwest Atlantic
References..................................................................................................................391
CHAPTER 7..............................................................................................................413
Final Discussion
References.................................................................................................................427
VITA
IN CASE SOME MORE BACKGROUND INFORMATION ON THE THESIS TOPIC IS NEEDED,
HERE AN EXCERPT FROM THE ORIGINAL PHD THESIS PROPOSAL:
INTRODUCTION
The question: "why is an animal at this location and not another one ?" is fundamental to
ecology and has
intrigued human beings for a long time. The question can be asked for one individual, a
metapopulation, a single
species, a (trophic) guild (e.g. sharing a foraging method) and for a community or an entire
biomass.
Only 300 from app. 9000 bird species on the world are classified as seabirds, among them is one of the most abundant birds of the world , Wilsons' storm-petrel (Oceanites oceanicus). Detailed information about seabirds at sea is lacking for most of the species. Seabirds are extremely mobile animals and to answer questions on their distribution, techniques such as tagging, banding, telemetry, remote sensing, geographic information systems (GIS) are used in conjunction with modeling, statistics, linear and non-linear mathematics. Understanding distribution is fundamental to conservation biology and species management. If one wants to conserve seabirds at sea in a seabird management plan, one has to understand why they are located where they are, in order to predict where they will be located in the future and which approach has to be chosen. Explaining bird distribution is as well a key question for their ecology and can lead to a better understanding of seabird foraging strategies at sea and their role and position in the food web.
The aim of this PhD research is to characterize and to explain the distribution of seabirds at sea in terms of the characteristics of their marine environment. The study area is the Atlantic region of Eastern Canada, where seabird distribution data have been maintained by various agencies and institutes.
Due to the high number of parameters that might potentially influence distribution - the literature has identified more than 20 parameters thought to be important directly or indirectly for the distribution of seabirds - this is likely a demanding question to answer.
Which reasons are important for the distribution of species on a more or less 2-dimensional surface such as the sea ? This research should bring more light to the questions: What are the reasons and factors ? How are they linked together ? And depending on the resolution scale, which are of the most importance for objectives of study ? The use of a GIS in this research has to be considered as a tool to improve knowledge about seabirds, but not only creating colorful maps. The PhD thesis will investigate whether there is a hierarchy of parameters determining distribution or if the distribution is even randomly determined (which does not necessarily mean that there are no reasons to be randomly distributed).
The results will be discussed with regard to seabird management, assessment of impacts on marine environments and selection of marine protected areas for an international frame; and furthermore will help to evaluate data sets and fill existing gaps in research on seabirds at sea.
RESEARCH APPROACH
The following research areas are thought to be important for this study:
Based on an intense literature review, the following hypothesis and 4 predictions can be concluded: The hypothesis "Distribution of seabirds at sea is non-random, but is associated with different environmental factors at several spatial and temporal scales" will be tested by examining the following 4 predictions:
1. At the largest spatial scale (study area off the Canadian east coast: Gulf of Maine to the
Canadian
Arctic), distribution can be explained by mechanisms caused by physical features such as
bathymetry,wind, water temperature, salinity, weather and pack-ice.
All transects in the PIROP data set for the whole study area of the North Atlantic will be
analysed by
overlaying the available (time and spatial) information, such as profiles from bathymetry, sea
surface temperature,
salinity, windspeed and ice cover.
Bathymetry profiles from the ETOPO5 seafloor map can be received (depths per
latitude/longitude coordinates as
ASCII file) and allow further processing. The depth will be classified into three groups:shallow
(<50m), medium (50m-100m) and deep (>100m).
The sea surface temperature and salinity information per transect will be achieved by analysing the
pixel values from
georeferenced satellite images for the transects from the National Oceanographic Data Centre
(NODC). Temperature
information will be classified into steps of one degree, salinity will be classified in parts 1/1000.
The weather
information, based on a query of the weather/sea condition column of the PIROP watch
information, will be classified
in wind, rain,fog, sun, snow. Additional data have to be evaluated, such as windspeed data
achieved by analysing pixel
values of ERS-1 satellite images.
The ice cover, derived from databases and satellite images, can be grouped into ice, no ice and ice
edge with a 2 km
bufferzone.
The approach taken will consider Multiple Regression, but mainly using the Classification and
Regression Tree
(CART) functions in Splus and Factor Analysis, if enough overlap of data can be found.
For the CART technique tree pruning has to be done before each test. The level of deviance
determines the number
of nodes per tree, allowing for appropriate conclusions. A random selected subdata set will be
used to test the results
for significance, using standard techniques, such as Chi-square tests. Results from each node in
the tree can be plotted
with SPANS GIS, and due to the relational database system, they make also other related factors
obvious. This can
lead to modelling, new hypotheses and findings.
2. At the medium temporal scale (20 years), distribution and habitat use is associated with
the predictable
annual cycle in a seabird's life (e.g. migration, breeding, wintering), reflecting a hierarchy of
distribution
factors. Foraging for food is mainly following traditional knowledge of food sources.
The general biological patterns of seabirds in the North Atlantic are known and will be taken
from the scientific
literature. For instance, shearwaters arrive in the North Atlantic in July and disappear in late
October. Phalaropes
(Phalaropus spec.) pass through the North Atlantic region in spring and fall, Skuas (Catharacta
skua) disperse in
winter along the coast of the eastern Atlantic. Black-legged kittiwakes (Rissa tridactyla) perform
a transatlantic
migration to the wintering grounds off Newfoundland, Puffins (Fratercula arctica) disperse widely
in the southern
North Atlantic. These patterns will become very obvious by analysing the abundance of seabirds
per location (latitude
and longitude coordinates) in the PIROP data set for transects through the fully covered time by
PIROP (1966-1992).
The behaviour column in PIROP, gives further information about the activity of the observed
seabirds, in particular
travelling in certain directions. Analysing transects from north to south (and west to east for
transatlantic migration)
will show the migration fronts, by the presence of the birds and their behaviour (migrating) per
transects.
Database queries of the PIROP database, using standard SQL techniques in dBASE, FoxPro and
other appropriate
programs will be carried out, looking for large flocks/number of seabirds resting at sea, sometimes
provided with
information about feather moult (following the PIROP definitions for moulting) and wintering
(winter plumage, flock
size, resting). Database queries in the behavioural (=> carrying food) and age (=> breeder and
non-breeder) column
of the PIROP data set will be done to answer these questions.
Before a breeding season starts, resting seabirds will be observed near colonies, sitting at sea and
waiting to start
breeding. This will be checked by looking for resting flocks in relation to colony distance. Resting
areas at sea will be
found, a higher number of observations per area represents migration corridors at sea.
Processed maps, plotted observations per months/season and location will make these
observations quite obvious.
Therefore, weighed information per location about the first and the last migratory bird (grouped)
and time of biggest
observation per day will be used. Overlaying the general breeding time cycle per guild will
probably show, that young
birds will be observed with a higher abundance after the season. Floaters, nonbreeders widely
dispersed among the
colonies, will be observed through the whole breeding season. Since they are not involved in the
breeding activities
they show a different behaviour and foraging strategy and can be found farther away from the
colonies.
An approach, using a "GIS bufferzone" of foraging distance of app.100 km around a colony for
breeding birds from
June - August will be used (dates depend on the breeding season and location) to determine the
distance to the colony.
The closest distance to the nearest shore can be achieved by using standard distance functions in
the GIS.
To obtain the results, this approach will use the CART technique, following the technique
described previously.
Since it is believed that these seabird observations are independent from oceanographical
processes, no homogenous
picture will be found with environmental features. Plotting the results over oceanographical
features is expected to show
a high variance.
3. At intermediate spatial scales such as currents, tide waves, fronts (including eddies/gyres),
ice-edges,
shelf areas, shelf edges, deep sea, shallow areas, closest distance to the nearest shore,
distribution is
explained through processes leading to easier prey (plankton, fish) catch ability, summarized
by the
term "trophic relations for food.
The abundance of fish and plankton per unit (currents, tide waves, fronts `including
eddies/gyres`, ice-edges,
shelf areas, shelf edges, deep sea, shallow areas, closest distance to the nearest shore) will be
compared with the seabird
abundance per unit in the spatial and time window of PIROP. The spatial and temporal constraint
for superimposing
the data sets will be limited by the fish- and plankton data. Techniques to allow comparisons such
as averaging,
blocking or Principal Component Analysis (PCA) for several satellite images through a time series
will be used.
Catching results from North American Fisheries Organisation (NAFO) are available for the
ICES/NAFO management
blocks and the PIROP time window . Towing results from St.Andrews and the Bedford Institute
for Oceanography
database will also be implemented.
The stratified seabird data will be overlayed with the bathymetry information from ETOPO5
seafloor maps. Location
of currents will be achieved by analysing SST gradients from satellite images/NODC. The
plankton distribution will
be achieved by analysing CZCS (Coastal Zone Colour Scanner) satellite images. These
information will be analysed
with the help of a Classification and Regression Tree (CART, explained previously), so that
relations of this kind
become quite obvious.
4. At the smallest spatial scale (transect length and 10 min observation period), seabird
distribution is
reflecting the biology (plumage colour, moulting stage, predator avoidance strategy), the
foraging
methods (surface feeders, divers, plunge divers, parasitic feeding, offshore feeders) and habitat
attraction (ice bergs, fishing vessels and floating debris).
Only the transects fulfilling the standard 10min watch length requirement for PIROP will be
analyzed by the
number of observed guilds (surface feeders, divers, plunge divers, parasitic feeders, offshore
feeders), individuals and
behavior. The information about weather, waves, fishing vessels, floating debris during the survey
will be achieved with
a database query from PIROP. The transect information will be split up in full transects and also
in the 10min transect
watches, allowing for a stratification their comparisons. Each of these results will also be
compared by their expected
abundance in this area from the literature, when sufficient data are available. The results, achieved
by using
Classification and Regression Trees (CART) will allow conclusions about the reliability of
observations for different
guilds and for different places and observation conditions so that a correction factor can be
calculated and the "true"
number of birds can be achieved for future surveys in the study area. These results allow as well
for an evaluation of
the PIROP data set and the counting scheme.
Since PIROP covers a coarse resolution a comparison with a smaller resolution will be carried out in the Bay of Fundy, where fieldwork in the form of (ferry) transects is conducted, following the PIROP scheme. Areas of interest are the biological rich upwellings of Brier Island, Grand Manan, the banks (Georges Banks and Brown Banks) and a transect between St.John and Digby. These data will be analyzed in addition with the tide information, available from tidal prediction software. The results will be compared with the results derived from the analysis of the PIROP database for the eastern Canadian coast and the Manomet Bird Observatory data set. The approaches for the selected tests in the other data sets will be repeated. An evaluation of these results will then be carried out. This approach will determine whether matching problems are occurring, how they could be explained biologically and how they could be avoided in an experimental design for seabird counting schemes in the future, so that the best results can be achieved.
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© 1997, 1998, 1999, 2000, 2001 Falk Hüttmann
Last updated: February 28th 2001 by Falk Hüttmann
URL: http://www.geocities.com/CollegePark/Quad/5377/webphdsm.html