Design and management of bird nesting habitat



Design and management of bird nesting habitat
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Design and management of bird nesting habitat:
tactics for conserving colonial waterbird
biodiversity on artificial islands in Hamilton
Harbour, Ontario
J.S. Quinn, R.D. Morris, H. Blokpoel, D.V. Weseloh, and P.J. Ewins
Abstract: Hamilton Harbour, at the west end of Lake Ontario, supports breeding colonies of six piscivorous waterbirds:
double-crested cormorant (Phalacrocorax auritus), black-crowned night-heron (Nycticorax nycticorax), herring gull (Larus
argentatus), ring-billed gull (Larus delawarensis), common tern (Sterna hirundo), and Caspian tern (Sterna caspia). Most
breeding pairs of all species nest on privately owned mainland locations that are subject to multiple industrial and
development activities. The construction of three artificial islands in the eastern basin of Hamilton Harbour in the winter of
1995–1996 presents an opportunity to simultaneously reduce current land-use conflict and maintain the existing avian
biodiversity. Accordingly, we used ecological information to determine suitable substrates and vegetation to satisfy nesting
habitat preferences and to facilitate the occupation of the islands by nesting waterbirds. Our recommendations for habitat
design and management emphasize techniques to prevent displacement of both tern species by ring-billed gulls and of
night-herons by cormorants. We propose long-term management procedures based on aspects of the biology and known
ecological interactions among the various species.
Résumé : Le havre de Hamilton, situé à l’extrémité ouest du lac Ontario, héberge des colonies reproductrices de six oiseaux
aquatiques piscivores : le cormoran à aigrettes (Phalacrocorax auritus), le bihoreau gris (Nycticorax nycticorax), le goéland
argenté (Larus argentatus), le goéland à bec cerclé (Larus delawarensis), la sterne pierregarin (Sterna hirundo) et la sterne
caspienne (Sterna caspia). Chez toutes ces espèces, la plupart des couples de reproducteurs nichent sur la terre ferme, sur des
terrains privés soumis à de multiples activités industrielles ou travaux d’aménagement. La construction de trois îles
artificielles dans le bassin est du havre, à l hiver 1995–1996, est une occasion d’atténuer le conflit qui touche en ce moment
l’utilisation des terres et de préserver en même temps la diversité actuelle de la faune ailée. Nous avons donc utilisé les
données écologiques pour déterminer quels substrats et quels types de végétation conviennent à la nidification, afin de
répondre aux préférences des oiseaux aquatiques et de faciliter l’occupation des êles par ceux-ci. Nos recommandations en
matière de conception et de gestion des habitats privilégient les techniques qui permettent d’éviter le déplacement des deux
espèces de sternes par le goéland à bec cerclé et du bihoreau gris par le cormoran à aigrettes. Nous proposons un mécanisme
de gestion à long terme fondé sur les divers aspects de la biologie des oiseaux et sur les interactions écologiques connues
entre les diverses espèces.
[Traduit par la Rédaction]
Habitat degradation by humans has reduced biodiversity at
global, and often local, levels. Loss of suitable habitat appears
Received November 1994. Accepted May 12, 1995.
J.S. Quinn.1 Department of Biology, McMaster University,
Hamilton, ON L8S 4K1, Canada.
R.D. Morris. Department of Biological Sciences, Brock
University, St. Catharines, ON L2S 3A1, Canada.
H. Blokpoel. Canadian Wildlife Service, Environment
Canada, 49 Camelot Drive, Nepean, ON K1A 0H3,
D.V. Weseloh. Canadian Wildlife Service, Environment
Conservation Branch, Environment Canada, P.O. Box 5050,
867 Lakeshore Drive, Burlington, ON L7R 4A6, Canada.
P.J. Ewins. Canadian Wildlife Service, Environment
Conservation Branch, Environment Canada, 25 St. Clair
Avenue East, Toronto, ON M4T 1M2, Canada.
Author to whom all correspondence should be addressed.
Can. J. Fish. Aquat. Sci. 53(Suppl. 1): 45–57 (1996).
to be the greatest threat to colonial waterbird species in North
America (Parnell et al. 1988). In Hamilton Harbour, Lake Ontario (43°16ʹ′N, 79°46ʹ′W) extensive wetlands along the south
shoreline have been lost to industrial development over the
past 100 yr (Hamilton Harbour Remedial Action Plan (RAP)
Beginning with intermittent breeding colonies in the mid1970s (Dobos et al. 1988), Hamilton Harbour has become one
of the most important colonial waterbird nesting areas in the
Great Lakes basin in the 1990s (Blokpoel and Tessier 1991).
Largely cut off from human and other mammalian interference
by busy fenced highways, the regions locally known as Windermere Basin and piers 25, 26, and 27 (Fig. 1) have provided
nesting habitat for several avian colonial species: doublecrested cormorants (Phalacrocorax auritus), black-crowned
Hamilton Harbour Remedial Action Plan (RAP). 1992.
Environmental conditions and problem definition. Stage I
Report. 2nd ed. Available from Library, Canadian Centre for
Inland Waters, P.O. Box 5050, 867 Lakeshore Drive,
Burlington, ON L7R 4A6, Canada.
© 1996 NRC Canada
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night-herons (Nycticorax nycticorax), herring gulls (Larus argentatus), ring-billed gulls (L. delawarensis), common terns
(Sterna hirundo), and Caspian terns (S. caspia).
Three of these species warrant special concern. Common
terns in the Great Lakes have been declining in numbers and
are listed as endangered, threatened, or of special concern by
most states bordering the Great Lakes (Courtney and Blokpoel
1983; Kress et al. 1983; Blokpoel and Scharf 1991). Caspian
terns have been designated as vulnerable in Canada
(COSEWIC 1992), endangered in Wisconsin, and threatened
in Michigan (Blokpoel and Scharf 1991). Recently, Austen
et al. (1994) recommended adding the black-crowned nightheron to the list of rare species and downlisting the Caspian
tern to “rare” status. The continued existence of these breeding
colonies at Hamilton Harbour is in jeopardy because of development plans for the privately owned property where the birds
nest. Most individuals of all of these species nest on property
owned and under development by the Hamilton Harbour Commissioners.
The Fish and Wildlife Restoration Project of the Hamilton
Harbour RAP has implemented the construction of three wildlife islands in northeastern Hamilton Harbour (Hamilton Harbour RAP 1992; see footnote 2), expected to begin in the
winter of 1994–1995 (Fig. 1). The islands are designed to
accomplish multiple goals, principal among them: (i) to break
wave action to reduce turbidity in the lee of the islands (thus
allowing submerged vegetation growth and increases in associated fish populations) and (ii) to provide protected habitat for
amphibians, reptiles, and especially colonially nesting waterbirds (Hall 19913). Construction of these islands provides a
valuable opportunity to simultaneously reduce land-use conflicts and protect and manage the local waterbird biodiversity.
We propose a management plan for creating habitat on
artificial islands and managing colonially nesting waterbirds
with two distinct though compatible goals. First, we seek to
maintain and monitor the local biodiversity of colonial waterbirds in Hamilton Harbour. Second, we focus special efforts
on the two tern species and the black-crowned night-heron to
conserve these colonial species in the Great Lakes. These goals
can be accomplished by creating nesting habitat and by managing the colonies on the islands, including control of ringbilled gulls, double-crested cormorants, vegetation, and
human disturbance.
Below we describe the ecological characteristics of each of
the major colonial waterbird species that nest at Hamilton
Harbour. We then discuss the resulting conservation issues,
final design features for the newly created habitat, and the
short- and long-term management plans for the area. Some of
the proposed approaches are without precedent and require
further evaluation.
Species accounts
Can. J. Fish. Aquat. Sci. Vol. 53(Suppl. 1), 1996
August as dates when eggs are present in nests, with most
(50%) of the records occurring between 5 June and 5 July.
Data from eastern Lake Ontario suggests somewhat earlier
clutch initiation. At Pigeon and Little Galloo Islands in Lake
Ontario, clutch initiation was well advanced by the third week
of April with 16–35% of the nests present having eggs at
Pigeon (1990–1992; D.V. Weseloh, unpublished data), and
5–10% having eggs at Little Galloo (1985–1986; Weseloh and
Ewins 1994). For management purposes, one may expect cormorant eggs in nests at Hamilton Harbour by the third or
fourth week of April.
Great Lakes cormorants nest on the ground or in trees.
Tree-nesting cormorants build their nests in open and highly
visible sites. Colonies typically initiated nesting in trees, often
moving to the ground when tree-nesting space became limiting
because of overcrowding or loss of trees (Wesoloh and Ewins
1994). Although trees appear to be preferred nesting sites
(Blokpoel and Scharf 1991), tree death and subsequent toppling frequently occurs most likely resulting from the effects
of cormorant guano. Pairs of cormorants that nest on the
ground in the lower Great Lakes seem to prefer elevated sites
such as rock piles, raised beaches, or toppled trees. The first
cormorant nest in Hamilton Harbour was found on the ground
in 1984 (Dobos et al. 1988). Thereafter, cormorants nested
exclusively in mature cottonwood trees (Populus deltoides)
along the western shore of the northernmost confined disposal
facility until 1991 when nesting began in a box elder tree (Acer
negundo) on Farr Island (Fig. 1; Moore et al. 1995). In 1992,
some cormorant pairs nested on the ground in the vicinity of
the cottonwoods.
Numbers of pairs have increased steadily since 1984; in 1993
the colony totalled 685 nests (Moore et al. 1995) making this
colony the fourth largest of 13 colonies in Lake Ontario in the
early 1990s (D.V. Weseloh, personal observations). Numbers
of nesting pairs have increased throughout the Great Lakes at
an average rate of 29% per annum since the early 1970s (Weseloh et al. 1995). This increase is likely due to several factors
including reductions of DDE levels in the food source, reduced
human persecution, and increased prey abundance in breeding
and wintering areas (Price and Weseloh 1986; Blokpoel and
Scharf 1991; Weseloh and Ewins 1994; Weseloh et al. 1995).
Interactions with other species
The acidic faeces produced by double-crested cormorants kills
vegetation beneath their nests. Also, cormorants strip leaves
and small branches for nesting material. As a result of this,
nesting trees die and eventually become unable to support
cormorant nests (Weseloh and Ewins 1994). The destruction
of forest understorey cover by cormorants has necessitated
culling of cormorants on some islands in the St. Lawrence
River that support an eider-down industry (Bédard and Nadeau
1995). We anticipate that the guano-induced destruction of
Double-crested cormorant
Nesting chronology and nest site
Double-crested cormorants nesting in Ontario begin laying
eggs in the latter part of April. Because of their inaccessible
nests in spindly trees at Hamilton Harbour, data on clutch
initiation are lacking. Peck and James (1983) list 27 April – 30
Hall, J. 1991. Fish and wildlife habitat restoration in Hamilton
Harbour and Cootes Paradise, concept summary. Fish and
Wildlife Restoration Committee for Hamilton Harbour and
Cootes Paradise. Available from Library, Canadian Centre for
Inland Waters, Box 5050, 867 Lakeshore Drive, Burlington,
ON L7R 4A6, Canada.
© 1996 NRC Canada
Quinn et al.
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Fig. 1. Map of eastern Hamilton Harbour showing the location of current nesting colonies and proposed
artificial wildlife islands. Produced under licence with the Ontario Ministry of Natural Resources.
© 1996 NRC Canada
Can. J. Fish. Aquat. Sci. Downloaded from by Brock University on 10/05/11
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Carolinian forest cover and ground vegetation on some islands
in the western basin of Lake Erie will soon become a considerable concern (D.V.Weseloh and P.J. Ewins, personal observations).
Ground-nesting cormorants are able to take over active
nests of other colonial species, including herring gulls (P.J.
Ewins, personal observations), and may take up nesting space
that would otherwise be used by other ground-nesting species.
Deterioration of the shrub layer by cormorants nesting overhead would reduce nesting opportunities for black-crowned
Black-crowned night-herons
Nesting chronology and nest site
Data are lacking for Hamilton Harbour. Peck and James
(1983) give 6 May – 9 September as a range of dates when
eggs can be found in night-heron nests and 1–17 June as when
most records occur. Field data from the Lake Ontario – Niagara River shoreline suggest somewhat earlier dates. In the Niagara River during 1991–1993, more than 80% of
black-crowned night-heron nests had eggs on 26 April, although this colony may be unusually early (D.V. Weseloh,
unpublished data). Night-herons that nest on Snake Island in
Lake Ontario had initiated clutches by 24 April each year of
1990–1992. At Hamilton Harbour, we expect night-herons to
start laying eggs by the last week of April.
Colonies are generally located in areas that are relatively
inaccessible to terrestrial predators such as islands, swamps, or
over open water (Davis 1993). Nests typically are built in trees
or shrubs of a variety of species (Davis 1993), and a preference
for concealed sites has been reported (Palmer 1962). Blackcrowned night-herons nested in Hamilton Harbour in the middle branches of cottonwood trees at Pier 27; however, since
1988 they have shifted nesting to sandbar willow (Salix
exigua) thickets along the west dyke of the confined disposal
facilities. Additionally, night-herons have begun nesting in
cottonwood trees along the mainland shore adjacent to Neare
Island and in the box elder tree on Farr Island (Moore et al.
1995). By 1994, night-herons had begun to nest on the ground
among rocks on Neare and Farr islands.
Black-crowned night-herons were first reported nesting in
Hamilton Harbour in a colony of over 100 pairs during the
1950s (Dobos et al. 1988). Fifteen pairs were reported in 1959
and nests were periodically reported thereafter until 1980.
Night-herons were first recorded at their current nesting site
(Pier 27; Fig. 1) in 1975, numbering six pairs (Dobos et al.
1988). The population grew steadily until it reached 212 pairs
in 1987 and then declined somewhat to 134 and 90 pairs in
1993 and 1994, respectively (Moore et al. 1995). This decline
was coincident with an increase in the number of doublecrested cormorants nesting in the cottonwoods and with a shift
in nesting trees from cottonwoods to sandbar willows by the
black-crowned night-herons (Moore et al. 1995). In Southern
Ontario, this species is expanding its range following population declines during the late 1960s and early 1970s (Goodwin 1987); however, there are strong indications of marked
declines since the 1980s in the lower Great Lakes (P.J. Ewins,
Can. J. Fish. Aquat. Sci. Vol. 53(Suppl. 1), 1996
unpublished data). Peck and James (1983) report a total of
8340 nests in 68 colonies across Ontario.
Interactions with other species
Black-crowned night-herons may be excluded from nesting
trees by double-crested cormorants, either by direct competition for suitable branches on which to build the nests, or by
defoliation of shrub layers from cormorant guano (P.J. Ewins,
personal observations). Increases in numbers of cormorants
nesting in Hamilton Harbour during 1988 coincided with a
shift in night-heron nesting locations from the cottonwood
trees, that were increasingly occupied by cormorants, to
nearby sandbar willows (Moore et al. 1995). The direct causes
of this shift are unknown; however, cormorant-induced defoliation of the cottonwood trees may have diminished their
value as night-herons appear to prefer well-concealed nest
sites (H. Blokpoel, personal observations). Additionally, cormorants may have taken over nest sites lower in the cottonwoods, forcing the night-herons out. Ring-billed and herring
gulls are known to predate black-crowned night-heron eggs
(Wolford and Boag 1971; P.J. Ewins, personal observations).
Black-crowned night-herons are known to predate pipping
common tern eggs and recent hatchlings (Hunter and Morris
1976) and immatures were observed depredating common tern
nestlings in Windermere Basin during 1993 (D. Moore, personal communication). Shealer and Kress (1991) reported
adult nocturnal abandonment by common terns in a colony
that was disturbed by black-crowned night-herons. The combination of predation by the night-herons and the nocturnal
desertion of adult terns leaving the nests vulnerable resulted in
reduced tern fledging success.
Herring gulls
Nesting chronology and nest sites
Herring gulls begin nesting as early as 14 April (Peck and
James 1983). In Hamilton Harbour, up to 14% of nests present
had eggs by 12 and 14 April in 1989 and 1986, respectively.
By 20–27 April in 1981 and 1987, up to 30% had eggs present.
Peck and James (1983) also give 14 April as the earliest date
on which eggs have been found in herring gull nests. Hence,
we expect herring gull eggs around mid-April.
Herring gulls in the Great Lakes normally nest on islands,
though some mainland nesting occurs. Nests are built on rock
surfaces, including cliffs, with or without vegetation, sometimes on sand and gravel, and in grassy and weedy areas (Peck
and James 1983). Additionally some nest on trees, buildings,
and other human-made structures. Nests are usually slight to
bulky raised mounds or platforms with variable cups, often
with large bases (Peck and James 1983). Locally, herring gulls
nest along the central and northern dykes of the confined disposal facilities and along the western part of Pier 27. A number
of pairs began nesting on Farr and Neare islands in 1988 and
1989, respectively (Moore et al. 1995). Some isolated pairs
nest among the ring-billed gulls on Pier 26, in the Windermere
Basin, and on the Stelco property occasionally (Fig. 1).
Herring gulls were first reported nesting in Hamilton Harbour
in 1976 when seven nests were recorded (Dobos et al. 1988).
About 300 pairs have nested there each year since 1989
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Quinn et al.
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(Moore et al. 1995), which represents about 5% of the lower
Great Lakes population (Blokpoel and Tessier 1991).
Interactions with other species
Herring gulls sometimes act as predators on a number of other
colonial ground nesters including adult and fledgling ringbilled gulls (K.M. Brown and J.S. Quinn, personal observations) and common tern adults (R.D. Morris, personal
observations) and chicks (Morris et al. 1991). Herring gulls
have been observed predating the eggs and chicks of all Great
Lakes colonial piscivore bird species (Parsons 1971; Quinn
1981; H. Blokpoel, P.J. Ewins, R.D. Morris, J.S. Quinn, and
D.V. Weseloh, personal observations). Two pairs of herring
gulls nesting in the midst of the Hamilton Harbour ring-billed
gull colony in 1992 inflicted heavy losses on adult ring-billed
gulls (J.S. Quinn, personal observations). At one point 45 dead
ring-billed gulls, some with their stomachs removed, were
counted within a 3 m radius of one herring gull nest. Specialist
bird predators have been described in a number of studies of
herring gulls (Morris et al. 1992) and other species.
Ring-billed gulls
Nesting chronology and nest sites
Peck and James (1983) report the earliest egg-laying date in
Ontario to be 14 April. In the areas surrounding the Caspian
and common tern colonies in Hamilton Harbour, the first ringbilled gull eggs observed in 1994 were laid April 17 and April
19, respectively.
Most ring-billed gull colonies in Ontario occur on islands
in the Great Lakes, the St. Lawrence and Ottawa rivers, and
James Bay. Some colonies surround nesting herring gulls,
common terns, or Caspian terns. Nests are typically raised
mounds or platforms variously formed of grasses, plant stalks,
aquatic plants, mosses, sticks, fish bones, or feathers (Peck and
James 1983). Though sometimes using barren sites, nests are
usually positioned in grasses, weeds, shrubs, and other vegetation. Although ring-billed gulls will nest almost anywhere,
providing there are others nesting nearby, those initiating the
earliest nests often choose piles of rubble and sites with much
visual relief such as vegetation, driftwood, rocks, etc. (Ryder
1993). On South Limestone Island in Lake Huron’s Georgian
Bay, a limestone knoll that is vegetated mainly by a low profile
species of cinquefoil (Potentilla norvegica) remains unused by
nesting ring-billed gulls and is available for the later arriving
Caspian terns (J.S. Quinn, personal observations). The gulls
nest in a dense colony among the stands of stinging nettle
(Urtica dioica) surrounding and on the edges of the knoll.
Ring-billed gulls will sometimes nest in open areas with prostrate vegetation (P.J. Ewins, personal observations).
Ring-billed gulls nested in dense subcolonies on Pier 27 in
areas unoccupied by herring gulls and over most of Pier 26 as
well as at the Windermere Basin and Neare and Farr Islands
(Fig. 1). Gull control was attempted at the southern end of Pier
26 and all of Pier 25 by the Hamilton Harbour Commissioners
through extensive scaring operations utilizing crackers, propane cannons, ultrasonic – high pitch broadcasts, and all-terrain vehicles which appeared to reduce the density of nesting
gulls drastically in the area of disturbance, although there were
no control areas with which to compare.
The population of ring-billed gulls has exploded in the Great
Lakes, increasing from about 56 000 pairs in 1976 to about
four times that number in 1990 (Blokpoel and Tessier 1991).
Ring-billed gulls first nested in Hamilton Harbour in 1978
when 78 nests were found. The most recent count, in 1990, put
the Hamilton Harbour colony at 39 621 nests (Moore et al.
1995). This represents approximately 14% of the population
of the lower Great Lakes (Blokpoel and Tessier 1991).
Interactions with other species
Ring-billed gulls nest earlier in the year than either of the tern
species (see Morris et al. 1992), thus reducing tern nest-site
availability. Ring-billed gulls are believed to have caused the
abandonment of common tern colonies in Hamilton Harbour
(Moore et al. 1995) and numerous sites on the lower Great
Lakes (Courtney and Blokpoel 1983). Displacement of
Caspian terns by ring-billed gulls has not yet been demonstrated clearly (Blokpoel and Tessier 1986).
Ring-billed gull predation of eggs and chicks of Caspian
(Quinn 1981; Quinn and Morris 1986) and common terns
(Morris et al. 1992) may be of concern in some cases. Close
observation of Caspian and common tern colonies has revealed that predation by gulls may be due to one or a few
individuals. Recently, in Hamilton Harbour, J. Sirdevan (personal observations) witnessed the complete loss of eggs from
over 15 Caspian tern clutches to predation by the same individual ring-billed gull. The predatory gull followed a characteristic pattern for predation and consumption of the eggs.
Following the shooting of that gull (under CWS permit), egg
losses to that part of the colony stopped.
Common terns
Nesting chronology and nest site
Peck and James (1983) provide 8 May as an early egg-laying
record for the lower Great Lakes. At Hamilton Harbour from
1991 through 1994, first eggs of the year were laid between 5
and 12 May (D. Moore, personal communication). Common
terns normally initiate nesting approximately 3 weeks later
than ring-billed gulls at this and other colonies (Morris et al.
1980, 1992).
Common terns breed on islands and shores and in marshes
of lakes and rivers (Peck and James 1983). Nests are typically
depressions, though some are flat or raised, consisting of simple to elaborate bowls of vegetation (Peck and James 1983).
Nests are built on or in rocks, sand, and gravel, among various
herbaceous plants, among driftwood, etc. Both descriptive
(Blokpoel et al. 1978) and experimental (Richards and Morris
1984) studies of nest locations showed that colonizing birds
preferred to nest near plants or other objects.
Common tern numbers in the Great Lakes have been in decline
since the early 1970s with reductions in numbers of breeding
birds at some sites, and entire colony desertions at others
(Morris and Hunter 1976; Courtney and Blokpoel 1983; Blokpoel and Scharf 1991; Morris et al. 1992). Additionally, the
Great Lakes population appears to be demographically separate from the thriving East Coast population. During the period
from the early 1920s until 1976, only four movements of
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banded birds into the Great Lakes from the East Coast of North
America were recorded (Haymes and Blokpoel 1978), suggesting that the Great Lakes population represents a distinct
genetic stock. The Hamilton Harbour population, numbering
954 nests in 1993 (Moore et al. 1995) represents over 40% of
the lower Great Lakes population, which totalled 2348 nests in
1990 (Blokpoel and Tessier 1991).
Interactions with other species
Common terns are most often the victims in their interactions
with other species dwelling near common tern colonies. They
are subject to loss of eggs, chicks, and nesting habitat by
ring-billed gulls (Morris et al. 1992); predation of chicks and
adults by herring gulls (Morris et al. 1991); and loss of young
by black-crowned night-herons (Hunter and Morris 1976) or
the nocturnal desertion of adult terns due to night-heron disturbance (Shealer and Kress 1991). In addition, available nesting substrate can be lost to earlier nesting ring-billed gulls
(Morris et al. 1992).
Caspian terns
Nesting chronology and nest site
Clutch initiation began in Hamilton Harbour 6 May 1994 (J.
Sirdevan and J.S. Quinn, unpublished data) concurring with
the early egg date listed in Peck and James (1994). Ring-billed
gull egg laying began 14 April in the vicinity of the Hamilton
Harbour Caspian tern colony and ring-billed gull breeding was
advanced well into incubation at the time the first Caspian tern
eggs were laid (J. Sirdevan and J.S. Quinn, unpublished data).
Caspian terns nest in dense colonies in the Great Lakes,
often in association with ring-billed or herring gulls. Their
nests consist of a depression in the substrate scraped out with
their feet and occasionally lined with plant material, pebbles,
and fish bones. J.S. Quinn and J. Sirdevan (unpublished data)
found a clear preference for sand when terns were given a
choice of three substrates: 1 cm crushed stone, pea gravel, and
construction grade sand. Caspian tern colonies typically are in
very open and largely unvegetated areas (Peck and James
1983). When vegetated, the vegetation is typically sparse and
prostrate. H. Blokpoel (personal observations) observed that
Caspian terns on Pigeon, Little Galloo, and Gull islands nested
on the muddy margins of interior ponds after they had dried
out during the spring. These areas were too wet to be used
earlier in the season when ring-billed gulls initiated their nests.
Caspian terns have nested in Hamilton Harbour since 1986,
forming one to three subcolonies each year in the Pier 26–27
areas. In 1993 and 1994, one to six late-nesting pairs nested on
a sand-covered raft designed for Caspian terns and anchored
in the southern confined disposal facility (Fig. 1) near one
subcolony location (Lampman et al. 1996).
Following the establishment of a colony of 48 nests in 1986,
numbers have generally increased and appear to have stabilized at about 300 pairs in one to three subcolonies. The establishment of additional subcolonies and an apparent increase in
nesting density at the main subcolony may have been necessitated by the encroachment of ring-billed gulls and vegetation
in the vicinity of the main colony site located about 100 m
north of the Pier 25 docking facilities (J.S. Quinn, personal
Can. J. Fish. Aquat. Sci. Vol. 53(Suppl. 1), 1996
observations). The Hamilton Harbour subcolonies collectively
represent the fourth largest of five Caspian Colonies in Lake
Ontario in 1994 (Moore et al. 1995). In the Great Lakes,
Caspian tern nesting numbers have increased steadily from
1995 pairs in 1963 (Ludwig 1979), to about 5700 pairs in 1987
(Blokpoel and Scharf 1991). Mean annual increases between
the late 1970s and early 1990s have been highest on Lake
Ontario (23% per annum; Ewins et al. 1994). These increases
have been attributed to a combination of declining contaminant burdens and to increased availability of forage fish
(Ewins et al. 1994). Caspian terns are a species of concern in
the Great Lakes according to Canadian and U.S. agencies
(Blokpoel and Scharf 1991).
Interactions with other species
Caspian terns begin nesting about 2 weeks after ring-billed
gull colonies have become well established (J.S. Quinn, personal observations). Nesting is excluded from areas occupied
by high densities of gulls. Their colonies are typically located
in areas devoid of vegetation that would provide vertical structure or reduced visibility and with few gulls on the immediate
site. While this may reflect some preference on the part of
Caspian terns, recent indirect evidence that ring-billed gulls
may prefer nesting near vertical structure (described below)
suggests the possibility that Caspian terns avoiding tall vegetation may be due to the avoidance of such areas by ring-billed
Predation of Caspian tern eggs and chicks by both ringbilled gulls (Quinn and Morris 1986; P.J. Ewins, personal
observations) and herring gulls (Quinn 1981; P.J. Ewins, personal observations) has been observed. Presumably because of
gape limitations, ring-billed gulls were implicated in taking
only young Caspian tern chicks (Quinn and Morris 1986),
while herring gulls were observed to take chicks of at least 21
days of age (J.S. Quinn, personal observations; Quinn 1981).
Egg predation by ring-billed and herring gulls can be exacerbated by humans disturbing Caspian tern colonies (Quinn
1984; P.J. Ewins, personal observations).
Conservation issues
We have identified five key factors that require short- and
long-term management attention. These include (i) vegetational succession, (ii) habitat modification by birds, (iii) interspecific competition for nesting space, (iv) predation by birds,
and (v) human disturbance. First, successional changes in
vegetation may alter the suitability of nesting habitat for both
ground-nesting and tree-nesting species. Vegetational succession is known to cause changes in occupancy by island-dwelling colonially nesting waterbirds (Parnell et al. 1988).
Successional encroachment of herbaceous plants appears to
inhibit nesting by common terns (Morris et al. 1992). Such
changes in habitat may have a similar effect on Caspian terns,
although it is difficult to discount the confounding influence
of nest-site competition with ring-billed gulls (see below). The
two gull species are better able to cope with advancing successional stages (Blokpoel and Tessier 1986).
Second, activities of the birds themselves can significantly
modify the structure of the habitat. Of the species noted here,
© 1996 NRC Canada
Quinn et al.
Table 1. Summary of substrate and vegetation plans and short-term management plans for the artificial islands.
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Habitat features
Short-term management plans
Overall barren, without topsoil; 200-m knoll on northern end for
Caspian terns; 250-m 2 area topped with mixed sand and gravel with
scattered driftwood and larger stones on southern end for common
Southern third surfaced with topsoil, planted with native trees and
shrubs; northern two-thirds barren; 250-m2 area on northern end for
common terns (see above); 250-m2 area along northern edge of planted
area with “decoy nests” for ground-nesting cormorants; black-crowned
night-herons may be attracted to shrubs and trees
Surfaced with topsoil and planted with trees and shrubs; shrubs planted
along shore to attract black-crowned night-herons
cormorants provide the best example as their acidic guano can
kill vegetation. Such death and resultant toppling of nesting
trees at some breeding sites, including Hamilton Harbour, has
reduced tree-nesting sites (Weseloh and Ewins 1994; Weseloh
et al. 1995). The result of this loss of habitat has been a shift
at some cormorant colonies in the lower Great Lakes to nesting
on the ground, sometimes displacing herring gulls, or in low
bushes, where they may displace black-crowned night-herons.
Third, nest-site competition among ground-nesting species
favours earlier nesters such as ring-billed gulls over later nesting species such as terns. Habitat that is acceptable to earlier
nesting species will be unavailable for later nesters unless they
are sufficiently large and aggressive to take over occupied
nesting sites. Of the species in question, we know of only one,
double-crested cormorants, for which we have data showing
that takeovers of occupied territories occurs, specifically cormorants taking over occupied herring gull nests (P.J. Ewins,
personal observations). Suitable nesting habitat can be preserved for late nesters by blocking access by early nesters (e.g.,
covering with materials to prevent access) or removal or deterrence of the earlier nesting species (see examples for preservation of common tern habitat in Morris et al. 1992).
Fourth, predation by one colonially nesting species on another may reduce the nesting success of the prey species. Eggeating ring-billed gulls have been observed and removed
(under Canadian Wildlife Service permit) after extensive predation of common tern (Morris et al. 1992) or Caspian tern
eggs (J.S. Quinn, personal observations). In both instances,
egg predation stopped after the removal of one to three eggspecialists. Losses of eggs to gulls may be exacerbated by
human disturbance, although such losses during research can
be reduced by the use of “nest domes” to cover vulnerable eggs
(Quinn 1984).
Finally, human disturbance of colonial nesting birds can
reduce breeding success (see Parnell et al. 1988). Such disruptions may result in the temporary abandonment of tern, cormorant, or night-heron colonies leaving the nest contents exposed
to gull predators or inclement ambient conditions. Disturbances during the nestling period may induce the scattering of
mobile offspring and subsequent attacks by conspecific adults
as displaced young attempt to find their way back (Fetterolf
1983). Disturbances on small islands with insufficient cover
during periods of chick mobility may lead to the offspring of
ground-nesting species swimming off and getting lost, drowning, or dying of exposure.
Each of these five factors is taken into consideration in our
Tern nesting habitat to be covered or patrolled
(ring-billed gull nests destroyed) until terns
begin nesting; vegetation on Caspian tern site to
be controlled, if necessary
Common tern area covered or patrolled (to restrict
or remove ring-billed gull nests); cormorant nests
removed from trees
Double-crested cormorants will be discouraged
from nesting
discussion of habitat design and management procedures that
Habitat design
The design and management of the islands will require the
cooperation of avian biologists and parties associated directly
with the Fish and Wildlife Habitat Restoration Project team of
the Hamilton Harbour RAP. The following discussion represents our proposal for management and fine-scale island design given the rudimentary plan determined previously by the
Fish and Wildlife Habitat Restoration Project team, following
recommendations pertinent to waterbirds from three of us
(R.D.M., J.S.Q., and D.V.W.). The three artificial islands will
be suitable to differing extents for the six colonial species
because of substrate and vegetation differences (Table 1). The
location and structural design of the islands has been finalized
(Mar-land Engineering Ltd. 19944). Each of the islands has
been designed so that they will not be washed over during
storm events. The basic landscaping of the islands is in the
advanced planning stages and we assume will not change substantially (Fig. 2). The southern island will be surfaced with
topsoil and planted with native trees and shrubs. The centre
island will remain devoid of topsoil on its northern end while
the southern portion will be covered with topsoil and planted
with native shrubs. The northern island will remain devoid of
topsoil and will not be planted.
The challenge and intent of the design plan is to encourage
the terns and night-herons while restricting the dominating
impacts of ring-billed gulls and double-crested cormorants.
All authors are fully aware of the risks posed by investigators
to eggs and chicks of breeding waterbirds (see for example
Parnell et al. 1988; Brown and Morris 1995), and will conduct
management procedures appropriately. For readability, we
present our proposed management plan as if already approved
and as if we will be responsible for implementation.
We will accommodate double-crested cormorants on the
centre island using a strategy intended to reduce cormorant
impact on night-herons and cormorant damage to trees. This
island will provide 250 m2 of ground-nesting habitat, sufficient for about 250–500 pairs of cormorants, assuming an area
of 0.5–1.0 m2 per pair. Ground-nesting will be encouraged in
the middle region, north of the vegetated part of the island
Mar-land Engineering Ltd. 1994. Project No. 410-201. c/o Mr.
J. Hall or Hamilton Harbour Commissioners, 605 James Street
North, Hamilton, ON L8L 1K1, Canada.
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Fig. 2. Site plan for three wildlife islands near the northeastern shoreline of Hamilton Harbour, with proposed nesting sites for
colonially nesting waterbirds.
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Quinn et al.
(Fig. 2). To allow easy access to the colony site by cormorants
we recommend that short ridges be constructed along the east
and west shore adjacent to the colony site. The provision of
these ridges, providing a 60° slope down to the water will
facilitate cormorant take-offs and will increase the suitability
of the site. In addition, the plan calls for the erection of five
artificial nesting structures that will be placed temporarily in
the area designated for cormorant ground nesting. These accessories are intended to attract cormorants to nest in the designated area before trees on the islands are large enough for
cormorant nesting. Nesting structures described by Meier
(1981) consist of platforms supported by a wooden pole. In our
plan, each pole would extend 6 m above the island’s surface
(Meier (1981) describes 9 m poles) and would provide eight
platforms. To encourage ground-nesting by cormorants at the
new location, we intend to move existing and previously used
cormorant ground nests from the current site in Hamilton Harbour to the area under the poles. As the platforms are occupied
by nesting cormorants, we expect pairs to begin nesting on the
ground near the nesting structures. Once a group of groundnesting pairs has been well established, we will remove one or
two of the poles from the island each autumn until none remain. The removal of poles will improve the aesthetic appearance of the islands and should facilitate our attempts to
discourage tree nesting by cormorants on the wildlife islands.
To reduce damage from cormorants breaking branches for nest
building, and to reduce the likelihood of tree-nesting in later
years, we propose that a 5-m buffer zone between the cormorant colony and the vegetated part of the island be maintained
by removal of nests prior to egg laying. If cormorants begin
damaging the young trees, we will provision them with sticks
for nesting. If necessary, young trees will be covered with
netting to prevent damage by cormorants. Once trees have
matured it may become necessary to discourage tree nesting
by removing cormorant nests from trees as early as possible.
For aesthetic reasons and to encourage other tree nesters including black-crowned night-herons, we hope to encourage
tree growth. However, it is possible that cormorants will prove
too destructive. Aside from Meier’s (1981) report describing
just the artificial nesting structures, there is no known precedent for our approach.
Predation of cormorant eggs by ring-billed and herring
gulls can be a serious problem during human disturbance;
cormorants flush from their nests and gulls fly in and eat the
unprotected eggs. Predation by gulls can be reduced dramatically by restricting researchers’ colony visits to periods of
darkness (K. Stromberg, U.S. Fish and Wildlife Service, personal communication).
Habitat suitable for black-crowned night-herons will be
created on the southern and centre islands. Plantings of redosier dogwood (Cornus sericea); peach leaf willow (Salix
amygdaloides), previously used as nesting habitat by blackcrowned night-herons; and other low shrubs will provide sufficient cover and structure for nesting black-crowned
night-herons. The creation of stands of these shrubs may encourage night-heron nesting and is likely to discourage predation of night-heron eggs and chicks by gulls. Cormorants are
less likely to nest in these small shrubs, specifically chosen for
planting on the islands because of their small growth form
when mature.
Ground-nesting species with particular habitat require-
ments are the focus of our habitat design considerations for the
surfacing of the artificial islands. Since both gull species are
effective at establishment and maintenance of colonies, we
assume that they will nest without the need for any special
habitat modification.
The design and management challenge posed by groundnesting gulls and terns is to restrict gull nesting from areas
targeted for terns. The habitat designed for use by common
terns will need to be protected from occupancy by nesting
gulls. Experimental studies by Richards and Morris (1984) and
observational studies by Blokpoel et al. (1978) both demonstrate that common terns prefer to nest on small-sized gravel
or chip substrate, often near plants, rocks, and logs. Furthermore, terns nesting on such enhanced substrates had larger
clutches and higher hatching and fledging success than those
nesting on adjacent substrates without these (Richards and
Morris 1984). This difference was attributed to the greater
visibility of eggs and chicks to predators on the sites lacking
structure provided by logs and plants. We propose to create
approximately 250 m2 of common tern habitat at the northern
tip of the unplanted portion of the centre island as well as at
the southern tip of the north island. Assuming 0.25 m2 per pair
(R.D. Morris, personal observations), this is sufficient space
for 1000 pairs of common terns at each colony site. The area
will be surfaced with a mixture of sand and gravel, with driftwood and larger stones scattered over the area and low herbs
such as Potentilla simplex or P. canadensis planted over the
area. The habitat will be kept free of nesting gulls by regular
daily or twice daily human patrols from early April (when
gulls start building nests) until mid-May (when common terns
begin nesting). During these patrols, all gull nests on the terndesignated area will be destroyed.
In concert with new common tern habitat created on the
islands, we strongly recommend the protection and encouragement of the common terns already nesting on the “spur dyke
island” in the Windermere Basin. Management procedures include covering of the entire dyke substrate with plastic sheeting early each season and destruction of any gull nests started
on the dyke. In 1993 and 1994, common terns nesting on the
spur dyke island in Windermere Basin were more successful
than those nesting on the mainland surrounding the Windermere Basin (H. Blokpoel, K. Lampman, and D. Moore,
personal observations). If possible, the construction of new
spur dyke islands in Windermere Basin would provide important additional habitat for common terns. In addition to maintaining the spur dyke for occupancy by common terns, we plan
to anchor tern rafts in the Windermere Basin to increase insular nesting space for common terns as a short-term measure.
These small rafts have been used successfully by small numbers of nesting common terns elsewhere in the Great Lakes
(Dunlop et al. 1991; Morris et al. 1992; H. Blokpoel, personal
Previous studies of Caspian terns suggest specific habitat
design components. During experiments for substrate preference with Caspian terns at Hamilton Harbour in 1994 (J.S.
Quinn and J. Sirdevan, unpublished data), ring-billed gull nest
construction was interrupted immediately upon discovery. The
nesting material was disturbed and scattered away from the
immediate vicinity of the nest site by a field assistant who
patrolled the site of the 1993 main Caspian tern colony several
times a day (under Canadian Wildlife Service permit) starting
© 1996 NRC Canada
Can. J. Fish. Aquat. Sci. Vol. 53(Suppl. 1), 1996
Table 2. Proposed methods to manage the six main species of colonial waterbirds on artificial islands in Hamilton Harbour.
Double-crested cormorant
Black-crowned night-heron
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Herring gull
Ring-billed gull
Common tern
Caspian tern
Encouragements to nesting
Install nesting platforms; move ground nests from
mainland site to island site
Plant shrubs; install decoys; remove cormorant nests from
trees and shrubs
Provide gull-free substrate; install visual relief objects;
install decoys; broadcast sounds; install chick shelters
Provide gull-free substrate; create dry sandy knoll with
low-profile vegetation; install decoys; broadcast sounds
from 5 April until 11 May. Nest initiations were common and
persistent throughout the area, which had begun to be invaded
by a number of plant species. However, the site of the substrate
preference experiment, which was devoid of vegetation, was
virtually ignored by the gulls as a nesting area. Over the entire
period of nest removal, only two gull nests were initiated on
the 9 × 9 m plot despite the frequent initiation of numerous
gull nests in the vegetated area within 2 m of the experimental
plot (J.S. Quinn and C. Pekaric, personal observations). In
sharp contrast, 19 of the first 20 Caspian tern nests were initiated on the experimental plot, and only one on the existing
partly vegetated substrate (J. Sirdevan and J.S. Quinn, unpublished data). These data suggest that substrate devoid of tall
vegetation may serve to attract Caspian terns and repel ringbilled gulls.
Caspian terns will be encouraged to nest on a 200-m 2 knoll
raised about 1 m above the highest part of the north island. The
knoll will be surfaced with a mixture of mainly sand and some
pea gravel, planted with a low-lying xerophytic plant such as
P. simplex. The knoll should be well drained and unsuitable
for herbaceous plants with significant moisture requirements.
Under these conditions we expect the knoll to be the least
vegetated part of the island and so reduce the tendency for
ring-billed gulls to nest on the knoll. Controlled experiments
planned to be undertaken in 1995 by J.S. Quinn and J.
Sirdevan will evaluate the effect of a lack of vegetation on
ring-billed gull and Caspian tern nesting. If necessary, gulls
will be prevented from nesting on the knoll by patrolling and
removing gull nests or by placement of plastic sheeting over
the area until early May when Caspian terns are about to nest.
We expect that the knoll will be eventually surrounded by
nesting gulls.
Attraction to sites
With the provision of suitable habitat, we assume that pairs of
the six species of breeding waterbirds will colonize the islands
naturally; there will be no effort to restrict their access to
mainland nesting sites unless the Hamilton Harbour Commissioners forecast construction on existing mainland colony
sites. Tern habitats on the islands will be groomed and prepared and decoys placed along with speakers and tape recordings of tern calls. Decoys have been used successfully to
attract least terns (S. antillarum; Kotliar and Burger 1984), and
decoys and tape recordings have been used to attract nesting
common terns, arctic terns (S. paradisaea; Kress 1983), little
blue herons (Egretta caerulea), and cattle egrets (Bubulcus
ibis; Dusi 1985). Prior attempts to attract Caspian terns with
Discouragements to nesting
Remove nests from trees and shrubs
Removal of nests in tern-nesting areas
Removal of nests in tern-nesting areas
decoys and playback were met with equivocal results (D.V.
Weseloh, P.J. Ewins, and H. Blokpoel, personal observations).
There will be no further effort to dissuade birds from nesting
at mainland sites for 4–6 yr unless plans to develop the nesting
sites are forecast within this time frame.
In the event that colonization does not occur over a period
of 4–6 yr, or if imminent development dictates, we will dissuade birds from nesting on the mainland while attracting
them to their respective prepared sites. Assuming advance notice of development plans, we will attempt to move at least
Caspian and common tern pairs in the year preceding development.
Common terns attempting to nest on all mainland areas
surrounding Windermere Basin will be disturbed on a regular
basis from their first arrival in mid-April. With the cooperation
of the Hamilton Harbour Commission and required Canadian
Wildlife Service permits, regular truck traffic will be rerouted
to run along the dykes separating settling ponds, and human
patrols will supplement disturbance caused by trucks. As the
spur dyke island fills with nesting pairs, we anticipate displacement of remaining pairs to the new island sites.
Within 5 days of the first Caspian tern clutch initiations
researchers will cover the largest of the mainland Caspian tern
subcolony sites with polyethylene and continue broadcasting
the playback at the proposed new colony site. We anticipate
that Caspian terns, ready to oviposit, will be attracted to the
proposed Caspian tern site.
Short-term island management
Short-term management goals will be to maintain gull- and
cormorant-free nesting habitat for terns and night-herons (Table 2). Caspian and common tern breeding areas will be covered with plastic sheeting and be regularly patrolled to prevent
ring-billed gull nest establishment. We will minimize our control procedures on gulls at other locations unless other species
are being compromised by nest-site competition or extensive
predation. As they have been shown to reduce gull predation
on common tern chicks, shelters (Burness and Morris 1992)
will be placed adjacent to all nests at the tern colonies prior to
egg hatching. During researcher visits to the colonies, nest
domes will be placed over tern nests for the protection of eggs
and young chicks (Quinn 1984). Excessive levels of gull predation on the eggs or chicks of terns will be considered on a
case by case basis. When successful nesting of significant
numbers of pairs is known to be compromised, predatory individual(s) will be eliminated under permit. Finally, the control of noninvestigator human disturbance will require the
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posting of signs on the islands, at local marinas, and at public
boat ramps during the breeding season. Patrolling of the islands may be necessary to enforce signing.
Long-term management and philosophical considerations
The establishment of breeding colonies on artificial islands in
Hamilton Harbour will not automatically maintain waterbird
biodiversity in the region; continuing efforts to monitor and
manage the islands are essential. Routine censusing of all colonially nesting species on the islands, in concert with detailed
studies of the breeding success of less abundant species (especially the terns and night-herons), will provide an early warning of developing problems. If problems are detected, careful
studies would be implemented to determine the primary factors and possible solutions. We recommend that research-related disturbance be minimized on the islands during the first
few years of colony establishment.
In the event that the ring-billed gull or double-crested cormorant populations become sufficiently large to threaten the
continued existence of other colonial species, long-term population control is recommended. Although labour intensive, the
annual application of mineral oil to the eggs of all clutches in
the control area is a measure intended to reduce recruitment
into the population and would be expected to cause the numbers to decline over the period of several years (Morris and
Siderius 1990; Christens and Blokpoel 1991). Previous work
with the technique showed that ring-billed gull pairs continued
incubating their oiled clutch, containing eggs with dead embryos, for up to 6 weeks beyond the usual incubation period
(Morris and Siderius 1990). These pairs did not renest during
that breeding season and so recruited no young into the population. Such drastic action would require careful consideration
but would need to be set into action early to avoid the loss of
species at risk. This need for early action underscores the
necessity of monitoring the success of some species.
One important, yet often neglected, component of conservation addresses the issue of overabundance (Garrott et al.
1993). Species that can adapt and thrive in human-altered environments can be devastating to less adaptive and sensitive
species. Accordingly, conservation biologists must be prepared to confront the problem of overabundance of some species to avert numerical losses in others. Extreme views on the
ethical justifiability of management contrast the passive with
the proactive approach to human interaction with nature. Anthropomorphists such as Norton (1986) hold that humans are
the source of all value and that the value of non-human lifeforms ought to be determined as favourable or nonfavourable
to objectives established by the human manager. Conversely,
non-anthropocentrists such as Katz (1985) state that species
reductions or replacements that occur by natural processes are
acceptable; artificial reductions or replacements (i.e., management of resources) are not acceptable as they cannot be justified on philosophical grounds. The recent dramatic population
number increases of ring-billed gulls and double-crested cormorants are attributable largely to human activities (Blokpoel
International Joint Commission. 1991. Review and evaluation
of the Great Lakes RAP: remedial action plan program 1991.
International Joint Commission, Windsor. Available from
Library, Canadian Centre for Inland Waters, Box 5050,
867 Lakeshore Drive, Burlington, ON L7R 4A6, Canada.
and Tessier 1986, 1988; but see Southern 1987, 1988; Ewins
1994; Weseloh et al. 1995). This being the case, management
that reduces the impact of inflated numbers of ring-billed gulls
and double-crested cormorants may be viewed as partially
correcting a human-caused problem. Here we adopt a philosophy of management for the expressed purpose of maintaining
colonial nesting species diversity both locally and in the Great
Lakes. We accept the responsibility of actions that may be
detrimental to ring-billed gulls or double-crested cormorants
to the benefit of other species.
Concluding remarks
Efforts directed towards habitat creation on artificial islands,
establishment of bird colonies on those islands and their maintenance on currently used sites, and appropriate long- and
short-term management of Hamilton Harbour’s colonial nesting birds will help to maintain colonial waterbird species diversity. Continuing studies are expected to provide data for
refining the approaches suggested here. Hamilton Harbour’s
RAP appears to be ahead of most of the other 18 RAPs in the
43 Areas of Concern designated by the International Joint
Commission (19915). The techniques and strategies tested here
will provide information that should be applicable to other
areas with colonial waterbird populations undergoing remediation in human-altered aquatic ecosystems.
We thank the Hamilton Harbour Commissioners for their cooperation, assistance, and permission to conduct our studies on
their property. Thanks to Mark Taylor for providing details of
the wildlife islands, to Jane Sirdevan for providing some of the
data on ring-billed gulls and Caspian terns, and to David
Moore and Karen Lampman for access to unpublished material. John Hall provided additional information on the Fish and
Wildlife Habitat Restoration Project. Thanks to Joe Minor and
Steve Boyle for preparing the figures. We gratefully acknowledge the efforts of agencies and the public for participation in
stakeholder meetings as required by the Environmental Assessment Review Process. Our current and previous research
on colonial nesting waterbirds has been supported by the Tricouncil Green Plan (McMaster EcoResearch Program; J.S.Q.
and R.D.M.), Natural Sciences and Engineering Research
Council (J.S.Q. and R.D.M.), the World Wildlife Fund (Wildlife Toxicology Fund; J.S.Q.), and Environment Canada
(Clean Up Fund and Great Lakes 2000; H.B., D.V.W., and
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1. James S. Quinn, Jane Sirdevan. 1998. Experimental measurement of nesting substrate preference in Caspian terns, Sterna
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