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Full text of "Merlin, Falco columbarius"

BLM LIBRARY 



88034470 



T-N-271 

Filing Code $#§*. £,6 Ot> 

Date Issued June 1975 




TECHNICAL NOTE 



U.S. DEPARTMENT OF THE INTERIOR - BUREAU OF LAND MANAGEMENT 



HABITAT MANAGEMENT SERIES FOR UNIQUE OR ENDANGERED SPECIES 

by Stephen A. Trimble, Biologist 
Conservation Library 
Denver Public Library 

Report No. 15 

Merlin 
Falco columbarius 



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84.2 
.L35 
no .271 



10/ copies of Technical Notes are available from DSC, Federal Center Building 50, Denver, Colo., 80225 



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BlM LIBRARY 

SC-653, BLOG. 50 '::>%;*'. •■•■ 

DENVER FEDERAL CENTER ;'V 

P. 0. BOX 25047 

DENVER, CO 80226-0047 







POHEWORD 

This Technical Note series on wildlife is designed to provide a 
literature review and summary of current knowledge pertaining to 
endangered and other wildlife species occurring on public lands. 
We in the Bureau of Land Management have recognized the need for 
basic wildlife information in order to do an effective job in 
land-use planning. Sound planning must identify the negative 
aspects as well as the positive benefits of any proposed land 
management decision or program. It is our hope, too, that this 
series will also prove useful to others --be they land managers, 
students, researchers or interested citizens. 



M 



Director 

Bureau of Land Management 

Department of the Interior 






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TABLE OF CONTENTS 

Page 

Species Description 1 

Distribution 6 

Status and Population Trend 10 

Life History ..... 12 

Daily activity patterns 12 

Food habits 13 

Hunting behavior 15 

Courtship 16 

Nest site fidelity and nesting density 17 

Egg-laying and incubation 17 

Juvenile development 18 

Territoriality and interspecific relations .... 20 

Vocalizations 22 

Migration and wintering behavior 23 

Productivity 24 

Pesticide relationships 25 

Habitat Requirements 27 

Breeding requirements 27 

Wintering requirements 29 

Limiting Factors 29 

Habitat destruction 29 

Chemical contamination 30 

Other mortality factors 30 

Species and Habitat Management Recommendations .... 31 

1. Research needs: Alaska 31 

2. Pesticide contamination 31 

3. Habitat protection and improvement 32 

4. Falconry 32 

Protective Measures Instituted 32 

A. Legal or regulatory 32 

B. Captive rearing 33 

Current Research 33 

Authorities 34 

Summary 34 

Literature Cited 37 

Additional References 41 



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SPECIES DESCRIPTION 

The merlin, or pigeon hawk, occurs throughout much of the 
northern hemisphere. Brown and Amadon (1968) name eight races 
worldwide. In North America, three distinct geographic races 
of this small, swift-flying falcon breed in three ecologically 
distinct regions: the taiga, Pacific coastal forest, and 
prairie-parkland. Although the American Ornithologist's 
Union (A.O.U.) Checklist (1957) recognizes four subspecies, 
Temple (1972a) and Brown and Amadon (1968) recognize only three 
of these forms, corresponding with the above-mentioned breeding 
populations. In this report, I follow Temple's taxonomic 
revision. 

Falco columbarius columbarius includes all North American taiga- 
breeding birds (merging the A.O.U. subspecies columbarius and 
bendirei , forms similar in phenotype and ecology). Falco 
columbarius suckleyi breeds in the Pacific coastal forest. 
And F. c_. richardsonii refers to the prairie-parkland merlins. 
Although the A.O.U. does not recognize common names for sub- 
species, to avoid constant repetition of Latin subspecifics 
I will refer to the three subspecies above as the taiga merlin, 
black merlin, and Richardson's merlin, respectively. 

Merlins exhibit the typical falcon silhouette--long, pointed 
wings and a longish tail. They are much smaller than the large 
falcons--about 25 to 34 centimeters long, with a wingspread of 
about 60 centimeters (Peterson, 1961). Like most other fal- 
coniforms, female merlins surpass males in size. Despite 
subspecific size variations, the degree of sexual size dimorphism 
remains constant in all populations. Females average larger 
than males in most measurements—about 20 millimeters longer in 
the wing (averaging 213.1 mm versus 192.2 mm), and about 11 mm 
longer in the tail (averaging 128.57 mm versus 117.8 mm) (pooled 
data for all North American merlins; Temple, 1972a). Males 
have a proportionately wider wing than females (Temple, 1970). 
Adult and immature merlins also differ in size: juveniles have 
longer flight feathers than adults, thus affording young birds 
lighter wing loading while they develop their powers of flight. 
(Wing loading is the ratio of weight to wing surface area.) 

Five male taiga merlins averaged 162 grams in weight (Brown and 
Amadon, 1968) . Two taiga females weighed 200 grams and about 
227 grams (Swartz, 1972). In a group of Newfoundland and 
Atlantic Coast migrants, 47 males averaged 159 grams, and 69 
females averaged 218 grams (Temple, personal communication). 
Female Richardson's merlins weigh about 227 grams (Oliphant, 
1974). A wintering female black merlin weighed 218 grams 
(Slipp, 1942). 



Merlins display marked sexual dichromatism, particularly in 
adult plumage. Subspecies vary, but in general, adult males 
(for a "typical" adult male F. c_. columbarius , see Figure One) 
have slaty blue-gray dorsal and crown plumage, varying from 
light gray to near-black, and marked by a mottled buff nuchal 
collar and the distinct black shaft stripes of individual 
feathers. Breast, flanks, and abdomen are buffy, boldly 
streaked vertically with cinnamon to black, with the widest 
streaks on the flanks. Few streaks mark the clear buff-colored 
throat and sides of the head. Both immature and adult merlins, 
of both sexes, possess dark tails distinctly banded with gray 
or buff. Merlins lack the bold moustachial stripe characteristic 
of many other falcons. The bill is bluish, the eye very dark 
brown; cere, eyelid, tarsi, and toes are yellow, and the claws 
black. 

Adult females differ from adult males primarily in the dark 
brownish color of their dorsal plumage, contrasting with the 
slate-blue of the male. This single distinction suffices for 
separation of adults in the field. Streaking below tends 
toward brown, on a lighter background than the male. 

Immature merlins of both sexes closely resemble adult females in 
dorsal coloration. The rump and upper tail coverts of the 
adult female, however, are slate-brown in contrast to the dark 
brown of the back, whereas the rump and upper tail coverts of 
immatures are the same shade of brown as the back. Many immatures 
have more greenish soft parts than the rich yellows of most 
adults. Immature males and females differ in the color of the 
light tail bands, especially in the central part of their tail 
feathers. Immature males have light gray bands; immature fe- 
males have buffy bands. Merlins acquire definitive adult 
plumage with their first prebasic molt, which lasts from April 
until September of the calendar year following their hatching 
year. These falcons continue to molt once each year, during 
these same months, throughout their lives, although the female 
may molt before the male when breeding (Temple, 1972a, 1972b, 
and pers. coram.; Bent, 1938; Brown and Amadon, 1968; Friedmann, 
1950; Peterson, 1961; Laing, 1938; Eyre and Paul, 1973; 
Oliphant, 1974). 

At hatching, creamy -white and pure white down covers the young, 
except for the bare abdomen. A second, coarser smoke-gray down 
replaces this first plumage. By the age of 18 days, primaries 
and tail rectrices have burst their sheaths. By 25 days the 
young birds are fully feathered (Bent, 1938; Brown and Amadon, 
1968; Friedmann, 1950; Fox, 1964). 




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Merlins vary considerably across their range in both size and 
plumage characters. Richardson's merlins have longer wings and 
tails, on the average, and lighter wing loading, than other 
North American merlins. Presumably, this proves a selective 
advantage for frequent long flights in their open orairie- 
parkland habitat. Black merlins, inhabiting the densest habi- 
tat of the three subspecies, have the heaviest wing loading. 
Taiga populations are intermediate (Temple, 1972a). 

Plumage coloration provides one of the most obvious means for 
separating subspecies. Richardson's merlins (prairie-parkland 
breeders) are lightest in color, black merlins (humid coastal 
forest breeders) are darkest, and taiga merlins intermediate. 
Black merlins derive their common name from the extreme darkness 
of their dorsal plumage. Richardson's merlins differ from 
other populations in having five light tail bands (these figures 
do not include the terminal light band). Black merlins usually 
have four light tail bands, but extreme saturation of tail 
rectrices by dark pigment often eliminates one of these, 
creating an appearance of only three bands. Taiga merlins 
nearly always have four light bands. The uppermost light band 
in all subspecies may be obscured by tail coverts. Crown 
streaking and primary barring also show patterns of geographic 
variation (Temple, 1972a). 

Merlins earn their second widely-used name--the pigeon hawk-- 
through similarity of size and flight, rather than from any 
preference on the merlin's part for consuming domestic pigeons, 
or rock doves ( Columba livia ). Falconers have flown merlins 
for centuries—particularly at small birds such as larks 
(Alaudidae) . Merlins suggest miniature peregrine falcons 
( Falco peregrinus ) in their slate-blue coloration and swift, 
direct flight. Even in normal flight they exceed speeds of 
20 meters/second (45 miles per hour) . In their ability to 
twist and turn while pursuing quarry, they surpass other 
falcons, and challenge the skill of accipiters. Like other 
falcons, merlins attain peak speeds when stooping at prey--at 
perhaps 27 to 40 meters per second (60 to 90 mph) . Merlins 
soar infrequently (Williams and Matteson, 1947; Fox, 1964; 
Brown and Amadon, 1968). Fox (1964) describes a curious 
hovering flight, most characteristic of fledglings begging 
food, with wings extended and tail spread, flapping the distal 
portions of the wing like a kingbird ( Tyrannus sp.). 

Observers in the field may confuse merlins with sharp-shinned 
hawks ( Accipiter striatus) , but the falcon's pointed wings and 
direct flight with deep, regular wingbeats serve to distinguish 
the two. Smaller size and lack of moustachial stripes should 
easily separate merlins from larger falcons. Lack of rufous 



on tail or back, along with indistinct facial markings, dis- 
tinguish merlins from North America's other small falcon, the 
American kestrel ( Falco sparverius ) (see Figure Two). Peterson 
(1961) emphasizes dark dorsal coloration and the distinctly 
banded tail as helpful field marks. 

DISTRIBUTION 

Merlins occur at some time of year in every province of Canada 
and every state of the continental United States. Although 
most wintering birds migrate south, some merlins remain near 
their breeding grounds all year. Breeding ranges for the three 
North American merlin subspecies are considerably more restricted, 
nearly coinciding with the limits of the boreal forest, plus 
the northern Pacific coastal forest and northern Great Plains 
(see Figure Three). 

Falco columbarius columbarius , the taiga merlin, breeds in the 
boreal forest from tree-limit in northern Canada, from 
Newfoundland west to northwestern Alaska, south to south- 
central Alaska through British Columbia east of the Cascade 
and Coast Ranges to north-central and eastern Washington and 
Oregon, and (perhaps) western Montana (Missoula County). The 
boundary then swings north into Canada west of the prairie- 
parkland, east across the taiga, then south into the northern 
parts of North Dakota, Minnesota, Michigan, and New England 
(A.O.U. Checklist, 1957; Ellis, 1974; Brown and Amadon, 1968; 
Jewett, et_ al_. , 1953; Friedmann, 1950). Taiga merlins occasion- 
ally breed as far south as northern California, Idaho (Bingham 
County), southwestern Wyoming (Fort Bridger), Colorado (Summit 
County, Grand County, and Fort Lewis—although it seems more 
likely that these are actually misidentif ied Richardson's 
merlins), and the Wasatch Mountains of Utah (A.O.U. Checklist, 
1957; Williams and Matteson, 1947; Bent, 1938; Bailey and 
Niedrach, 1965; Wolfe, 1946). Most taiga merlins migrate 
south in autumn, wintering across the western states, south to 
the southern states, through Mexico to Central America and the 
West Indies, and as far as Venezuela, Ecuador, and northern 
Peru (A.O.U. Checklist, 1957). 

Richardson's merlin breeds in the prairie-parkland of the 
northern Great Plains from the fringes of the boreal forest 
(northeast of Prince Albert, Saskatchewan; Oliphant, personal 
communication) through the Canadian prairie provinces 
(Alberta, Saskatchewan, and Manitoba) south to northern Montana 
(Lewis and Clark, Valley, Cascade, Toole, and probably 
Choteau Counties), northern North Dakota, and South Dakota (A.O.U. 
Checklist, 1957; Ellis, 1974; Brown and Amadon, 1968; Hunter, 
1967). Richardson's merlins have also nested in eastern Wyo- 
ming, in Albany and Weston Counties (Fox, 1964). Falco 









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columbarius richardsonii winters south through Wyoming and 
Colorado to California (where it is uncommon), northern Mexico, 
and west Texas (Friedmann, 1950). Most migrant Richardson's 
merlins reportedly winter in Wyoming and Colorado (Fox, 1964) . 

The black merlin, F. c_. suckleyi , breeds in the moist coastal 
forests of western British Columbia, Vancouver Island, the 
Queen Charlotte Islands, and the coastal islands of southeastern 
Alaska, inland to the Okanagan Valley and the town of Blue 
River, and north to Atlin and (probably) the upper Yukon River 
drainage (Bent, 1938; Munro, 1946; Laing, 1935; Sutton, 1935; 
Friedmann, 1950; Swarth, 1935). No certain breeding records 
exist for the black merlin within the United States. Bent 
(1938) mentions a possible nesting at Fort Klamath, Oregon, 
another in the Puyallup River valley of Washington, and a set 
of eggs ( suckleyi or columbarius ?) collected at Sitka, Alaska 
in 1887. Black merlins may have nested at Bumping Lake, 
Yakima County, Washington (Jewett, e_t al_. , 1953). Jewett (1939) 
observed what appeared to be a family of recently fledged black 
merlins in early August in Deschutes County, Oregon—well east 
of the Cascades. Gabrielson and Lincoln (1959) report an 
adult female black merlin taken on Sergief Island, north of 
Wrangell, Alaska, on August 19. An extremely dark male merlin 
was taken as a nestling by Pat Redig (of the University of 
Minnesota) in northern Minnesota in 1974 (Oliphant, personal 
communication). Wintering black merlins remain chiefly in the 
coastal area southward through Washington, Oregon, western 
Montana (Ellis, personal communication), and central California 
(Unglish, 1934), to southern California (van Rossem, 1934; 
Miller, 1941; Spaulding, 1947), but also wander southwestward 
as far as New Mexico (Jewett, 1944) and Colorado (Bailey, 1942) 
and north to coastal Alaska (Friedmann, 1950; A.O.U. Checklist, 
1957). Although forty years have passed since Swarth (1934, 
1935) lamented the near-complete lack of nesting data for this 
subspecies, even today we know little more of its habits and 
range. 

STATUS AND POPULATION TREND 

As a species, merlins are not endangered. But the three North 
American subspecies vary dramatically in status. 

Taiga and black merlins are probably not yet in any serious 
population trouble. Both subspecies breed in environments not 
highly altered by man (Temple, personal communication. ; Fyfe, 
personal communication) . Swartz (personal communication) found 
taiga merlins breeding in interior Alaska with good productivity 
and in normal density, which is to say, not very abundant. 
Migration counts in Alaska in 1968 indicated merlin populations 
seemed to be holding their own (Adolphson, 1969). Richardson's 
merlins, however, have suffered population declines. 



10 




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All three subspecies carry high levels of chlorinated hydrocarbon 
residues. For F. c. columbarius and suckleyi , lowering of 
reproductivity due to pesticide poisoning is significant, but 
slight. Richardson's merlin, in contrast, has suffered massive 
reproductive failure over most of its range—which constitutes 
the most contaminated of all North American merlin habitats 
(Temple, personal communication). Fox (1971) documented a 43% 
decrease in hatching success since 1950 for Richardson's merlins 
nesting in the Great Plains grasslands, corresponding with a 
marked decrease in eggshell weight. 

Not only has productivity declined, but the Richardson's breed- 
ing range has contracted rapidly during the past 20 years. Two 
factors have reduced the area of suitable breeding habitat. 
First, Richardson's merlins depend on grassland birds for food. 
As areas of native grassland shrink under the pressures of 
intensive cultivation, Richardson's merlin populations likewise 
shrink. Second, these merlins nest exclusively in trees. With- 
out suitable nest sites, they cannot reproduce. Where native 
grassland and nest sites persist, merlins seem to be maintaining 
their populations, or even slowly increasing (Fyfe, personal 
communication). Oliphant (personal communication) considers 
the Richardson's merlin common in the Saskatoon, Saskatchewan 
area, with good production of young (4-5 fledged in many nests). 
However, on Saskatchewan Christmas counts, wintering merlins 
were down 43% during 1964-1967, compared to 1957-1963, and down 
55% during the same time period in the number of locations 
where the bird was sighted (Fox, 1971). Recent Saskatchewan 
Christmas counts have been high, six birds seen in Saskatoon 
alone in 1973-1974 (Oliphant, personal communication). Temple 
and Fox both regard Richardson's merlin as endangered (Temple, 
personal communication; Fyfe, personal communication; Fox, 
personal communication) . 

Migration counts at Duluth, Minnesota, show that while total 
raptors observed increased by 87% during 1960-1967 (over 1935- 
1942), merlins decreased by 837 . Increased late-season 
observations might bias these data somewhat. At Hawk Mt., 
Pennsylvania, merlin counts for these same years were down 137o 
while total raptors increased 9% (Fox, 1971). Duluth birds 
probably include migrants from the eastern edge of the Rockies 
to central Ontario—central taiga merlins, with perhaps some 
Richardson's. Hawk Mt. birds breed along the Atlantic Coast 
and in Newfoundland, though Hawk Mt. is not on a major merlin 
migration route (Temple, personal communication). Variation 
in merlin counts may reflect the health of merlin populations 
nesting in the differing source areas. 



I 



11 



Ellis (1974) reported eight positive breeding records for 
merlins in Montana. Productivity was high for successful nests, 
but three of seven nests with eggs failed—all Richardson's 
merlins. Merlins have probably long been established in 
Montana, but are so rare as to nearly have gone undetected. 
Ellis believes the Montana population may be declining. 

Although populations of taiga and black merlins seem to be only 
slightly declining on a continent-wide basis, biologists should 
watch them closely. As the taiga of northern Canada and Alaska 
grows ever more civilized while man develops its resources, 
merlins there may fail to reproduce successfully. 

One note of optimism may be found in work by Oliphant (1974) in 
Saskatchewan. He described eight nesting attempts by merlins 
from 1971-1974 within the urban environment of Saskatoon. These 
merlins successfully fledged young in the constant presence of 
thousands of people, dogs, cats, cars, trucks, etc. Their diet 
consisted almost entirely of house sparrows ( Passer domesticus ) , 
rather than native grassland birds. Fox (personal communication) 
reports other consistently successful merlin nestings in 
Canadian prairie cities. 

LIFE HISTORY 

Daily activity patterns 

Merlins are strictly diurnal birds. When nesting, the male 
leaves his roost to hunt early in the morning, usually returning 
quickly with prey for the female and young. Taylor (1914) noted 
two periods of hunting and feeding activity during the day for 
merlins nesting in Britain: daybreak to 11 A.M. and 4 P.M. to 
dusk. Selous (1915) found merlins in Iceland following a 
similar activity pattern, with the early and late periods 
shortened, and a third hunting and feeding period added from 
about 1 P.M. to 4 P.M. 

On the Minnesota-Ontario border, Craighead and Craighead (1940) 
counted three feedings of young merlins between 9 A.M. and 
5 P.M. These feedings usually included an early morning and 
late evening hunt. Lawrence (1949) counted 1 to 2 feedings in 
3 to 5 hours in Ontario. Breckinridge and Errington (1938), 
however, observed a nest of half-grown young in northern 
Minnesota to which the adults brought food 10 times during the 
day. 

Merlins hunt at twilight, at least occasionally. Lawrence 
(1949) noted this behavior in a nesting male, while Johnson 
and Coble (1967) observed merlins actively hunting as late as 




12 



1 



9:30 P.M. in Isle Royale National Park, Michigan. Bats made 
up part of these merlins' diet, which presumably accounts for 
such late-evening hunting. 

In migration, taiga merlins feed early in the day (Bent, 1938), 
and fly more in the afternoon than do other hawks migrating 
along the east coast (Allen and Peterson, 1936). 

Food habits 

Merlins survive on a diet of birds almost exclusively. Brown 
and Amadon (1968) summarize the diet of the species, worldwide, 
as 80% birds, 5% mammals, and 15% insects. Merlins also have 
consumed toads, lizards, and snakes (Bent, 1938). 

Birds taken by merlins range in size from small sparrows 
(Fringillidae) and wood warblers (Parulidae), to teal ( Anas sp.), 
rock doves, and ptarmigan ( Lagopus sp.) (Bent, 1938). Fyfe 
(personal communication) has not recorded a single mammal in 
extensive food data for Canadian-breeding taiga and Richardson's 
merlins. Lawrence (1949) estimated that one breeding pair of 
merlins require 450 birds to support themselves and their 
young during the 2%-month nesting season. Brown and Amadon 
(1968) think this an excessively high estimate, suggesting 
that 3 to 5 kills a day may suffice for the female and three 
young. 

Males evidently prey on smaller species than females, reflecting 
the smaller size of the male falcon. At one Newfoundland nest, 
the male taiga merlin captured all prey during the first week 
after hatching, while the female constantly brooded the nestlings, 
Of 19 prey items brought to the nest during this week, 79% were 
species of birds weighing less than 50 g. From the end of this 
first week until fledging, 27% of 109 food items weighed under 
50 g. During this period the female resumed hunting regularly, 
and the larger species are presumably her prey (Temple, 1972b). 
Alternately, merlins may take larger prey species as food 
demands of their fast-growing young increase (Lawrence, 1949). 

Of 136 prey remains at nests of Newfoundland-breeding taiga 
merlins, three species comprised over half the merlins' diet: 
gray jay ( Perisoreus canadensis ) , American robin ( Turdus 
migratorius ) , and Savannah sparrow ( Passerculus sandwichensis ) 
(Temple, 1972c). Merlins, like other raptors, will concentrate 
on a few prey species if these species prove easy to capture 
owing to local abundance or behavior. Oliphant (1974) found 
that merlins breeding in urban Saskatoon, Saskatchewan, con- 
sumed primarily house sparrows, plus a few cedar waxwings 
( Bombycilla cedrorum ) , tree swallows ( Iridoprocne bicolor ) , and 



13 



horned larks ( Eremophila alpestris ) ; house sparrows accounted 
for over 90% of prey at one nest. Wintering merlins in 
Alberta cities fed largely on Bohemian waxwings ( Bombycilla 
garrulus ) and sparrows (Fyfe, personal communication). A pair 
of Richardson's merlins breeding on the Saskatchewan prairie 
survived on a diet of 53.5% horned larks, 13.3% brown-headed 
cowbirds ( Molothrus ater ) , 6.7% vesper sparrows ( Pooecetes 
gramineus ) , 6.7% song sparrows ( Melospiza melodia ) , 6.7% Baird's 
sparrows ( Ammodramus bairdii ) , and 13.6% chestnut-collared 
longspurs ( Calcarius ornatus ). With the exception of Baird's 
sparrows, these species all were abundant in the merlins' 
hunting territory (Fox, 1964) . 

In Isle Royale National Park, a pair of breeding taiga merlins 
consumed primarily birds, including warblers, sparrows and 
crossbills, red-breasted nuthatch ( Sitta canadensis ), common 
nighthawk ( Chordeiles minor ) , common flicker ( Colaptes auratus ) , 
and an unidentified member of the Rallidae. These merlins 
also preyed on bats (red bats, Lasiurus borealis , and Myotis sp.), 
dragonflies (Odonata), and beetles (Coleoptera) (Johnson and 
Coble, 1967). Taiga merlins migrating past Cape May, New 
Jersey, also had consumed red bats (Allen and Peterson, 1936). 
The few other mammals recorded in merlin diets include pocket 
gophers (Geomyidae) , squirrels (Sciuridae), and field mice 
(Microtus sp.) (Bent, 1938; Allen and Peterson, 1936; Burleigh, 
1972). 

During summer and fall, insects increase in importance in the 
merlin diet. Young falcons, especially, seem to rely on 
insects for food as they develop their predatory skills 
(Williams and Matteson, 1947; Eyre and Paul, 1973; Lawrence, 
1949). 

Jewett (1939) described two immature female black merlins in 
Oregon which had eaten nothing but black ground beetles 
(Carabidae). Other arthropods in merlin diets include cater- 
pillars, butterflies, and moths (Lepidoptera) , grasshoppers 
and crickets (Orthoptera) , spiders (Araneida), scorpions 
(Scorpionida) , and crayfish ( Astacus and Cambarus sp.) (Bent, 
1938; Allen and Peterson, 1936; Fox, 1964). 

Dragonflies, however, make up the most important insect element 
in merlin diets. Again, the young birds in particular seem to 
favor these winged insects. Several authors have marveled at 
the flying skills demonstrated by young merlins hunting dragon- 
flies and other insects (Lawrence, 1949; Street, 1960; Oliphant, 
1974). Their twists and swoops almost resemble the flight of 
swallows (Hirundinidae) , with the notable difference that merlins 
snatch insect prey with their feet. 



14 



At Cape May, New Jersey, large numbers of migrant merlins and 
dragonflies occurred on the same days. Investigation of 41 
stomachs showed that these falcons had eaten more dragonflies 
than anything else (Allen and Peterson, 1936). 

Merlin castings, or pellets, have been noted to contain 
disintegrated feathers, beetles, red bat fur, and avian bone 
chips. Thirty-three taiga merlin castings averaged 25 X 12 mm, 
tapering at one end (Johnson and Coble, 1967; Breckinridge and 
Errington, 1938). 

Hunting behavior 

Forest-nesting merlins occasionally do not hunt in their breed- 
ing territory, since they prefer more open hunting areas. 
Lawrence (1949) recorded eight passerines known to be merlin 
prey that nested safely in the immediate vicinity of a taiga 
merlin nest. Other small birds passed through the territory, 
sometimes very close to the nest, without incident. The breed- 
ing merlins also ignored small mammals, such as squirrels and 
chipmunks ( Tamias sp.). Brown and Amadon (1968) suggested that 
merlins' aggressiveness toward common crows ( Corvus brachyrhynchos ) 
and other raptors near their nest might further benefit 
passerines nesting in the same area. In contrast, a male 
Richardson's merlin in Saskatoon pursued robins and house 
sparrows from his perch near the nest site, though he generally 
hunted at some distance from the nest (Oliphant, 1974). 

Merlins commonly fly low over the ground when hunting, darting 
after prey with rapidly beating wings. These falcons will 
glide short distances, frequently rising and falling, but 
glide and hover less than many other falcons. On the attack, 
merlins fly at avian prey with a direct, very fast dash. If 
this first strike fails, they generally try a series of short, 
twisting stoops from above, shooting up after each attack for 
a few wingbeats and twisting into another stoop. When necessary, 
merlins follow prey into dense cover (Brown and Amadon, 1968; 
Fox, 1964; Lawrence, 1949). High-speed stoops from very high 
above prey, in the manner of the larger falcons, seem less 
common. Laing (1938) described a black merlin along the 
British Columbia coast which struck a black swift ( Cypseloides 
niger ) in a spectacular stoop. The Craigheads (1940) watched 
a taiga merlin take a tree swallow in similar fashion. Merlins 
hunting over water often pursue escaping prey high into the air 
in a ringing flight and then capture the prey in a high speed 
stoop as the victim tries to dive to cover (Temple, personal 
communication). McClure (1957) described a merlin in Japan 
that took a brown-eared bulbul ( Ixos amaurotis ) to the ground 
in a stoop and killed it with a bite at the base of the 
skull. 



15 



Blackbirds (Icteridae) , starlings ( Sturnus vulgaris ), meadow- 
larks ( Sturnella sp.), shrikes ( Lanius sp.), and kingbirds all 
perch conspicuously in open places where merlins can strike 
from behind without warning. In the open, the slow flight of 
meadowlarks, as well as woodpeckers and flickers (Picidae), 
make them vulnerable to direct attack. In forests, merlins 
often concentrate hunting efforts on bright-colored and tree- 
nesting passerines. Merlins fly full-speed into flocks of 
ground -dwelling birds such as horned larks, but seldom make a 
second attempt if the first surprise attack fails. With 
enough room to gain top speed, mourning doves ( Zenaida macroura ) , 
rock doves, and robins can outdistance merlins. Smaller, 
slower birds often escape by abrupt and agile dodging (Bond, 
1936; Lawrence, 1949; Fox, 1964). 

Courtship 

Male merlins arrive on the breeding territory about a month 
before females. Most males arrive in April, but male Richardson's 
merlins may arrive back in Alberta and Saskatchewan as early 
as late February or early March. Further north, taiga merlins 
may not finish migration until May (Bent, 1938; Fox, 1964; 
Brown and Amadon, 1968; Lawrence, 1949; Fyfe, personal communi- 
cation; Beer, 1966). 

On arrival, males make a series of vocal flights from perch to 
perch. After the female's arrival, the male repeats this per- 
formance near her with a very distinctive fluttering flight. 
The wings beat very rapidly and the male calls while flying 
(see page 20 for description of call). Richardson's males have 
demonstrated this same flight pattern and call when flying to 
the female to copulate (Fyfe, personal communication; Brown 
and Amadon, 1968) . 

Female Richardson's merlins evidently do not solicit the males' 
attention. Males, in contrast, call and display both in and at 
the entrance to nests to attract the female to the nest. The 
male's display posture seems aimed at best presenting the blue 
back, upper wings, and head, and at the same time, the full 
tail pattern. Males accomplish this by bowing low and raising 
the tail, while at the same time spreading the tail and slightly 
raising the wings. In this way they present to the female an 
almost solid form of blue combined with the striking contrast 
of the tail bars. In a variation, the male presents to the 
female at one side or the other — usually the left — instead of 
head-on. From the side, the male varies his posture slightly, 
tilting to one side and raising the more distant wing above 
his body, thereby again presenting the back pattern to the 
female (Fyfe, personal communication). 



16 



Nest site fidelity and nesting density 

Merlins use the same general area year after year for breeding, 
but not necessarily the same actual site, particularly if 
young were fledged the previous year (Brown and Amadon, 1968; 
Fox, 1964). Beer (1966) observed defending merlins around a 
single Minnesota nest in six out of fifteen years. Merlins 
nesting in Montana did not reuse the same nest, or nest grove, 
in successive years, although a successful 1974 fledging occurred 
150 meters from a 1973 nest site (Ellis, 1974). In Newfoundland, 
one ground-nesting site on the side of a cliff was used con- 
tinuously for 23 years (Temple, personal communication). 

Nesting density is generally low--probably as a result of low 
merlin populations. The closest concurrent Richardson's 
merlin nestings found in Montana were 17 km. apart (Ellis, 1974), 
while three pairs of this subspecies nested in one 4.8-km. 
stretch of the North Saskatchewan River (Oliphant, personal 
communication). Craighead and Craighead (1940) found a concen- 
tration of merlins nesting on lakes on the Minnesota-Ontario 
border where nests were at least 3.2 km. apart. Raspberry 
Island, an 11. 73-hectare islet off Isle Royale, Michigan, 
supported one pair of merlins and their two young; the adults 
hunted primarily in a spruce bog and along sheltered coves on 
the periphery of the island (Johnson and Coble, 1967). Lawrence 
(1949) estimated that a single pair of taiga merlins hunted in 
a circular area of 1.6 km. radius, with the nest at the center 
of the circle. In Saskatoon, a male hunted up to 3.2 km. from 
its probable nest (Oliphant, 1974). 

Egg-laying and incubation 

Merlins lay clutches of four to five eggs. Taiga merlins 
breeding in Newfoundland in 1969 averaged 4.3 eggs/clutch (N=15) 
(Temple, 1972c). Richardson's merlins on the Canadian prairie 
averaged 4.48 eggs/clutch, the mode being four and the range 
3 to 7 (Fox, 1964). Montana prairie merlins laid five-egg 
clutches in two out of five nestings, and observers noted five 
young at another (Ellis, 1974). Second clutches of Richardson's 
merlins generally number only three eggs (Bent, 1938). Females 
lay eggs at two-day intervals (Fox, 1964, 1971; Brown and 
Amadon, 1968; Williams and Matteson, 1947; Eyre and Paul, 1973). 

At the southern fringes of their range, merlins may lay eggs in 
early April (Brown and Amadon, 1968; Williams and Matteson, 
1947). Egg-laying commences on the Canadian prairies during 
the first half of May, and peaks about May 20 (Fox, 1964; 
Oliphant, 1974). Taiga merlin eggs may be laid in late May or 
June (Brown and Amadon, 1968; Bent, 1938; Craighead and Craig- 
head, 1940). Temple (1972c) found that 20 pairs of Newfoundland 
taiga merlins were incubating eggs between May 24 and July 9. 



17 



Merlin eggs vary in shape from short-ovate to oval to nearly 
elliptical-ovate. They look much like miniature peregrine 
falcon eggs. Egg ground color is cinnamon to light creamy 
buff, and is almost obscured by spots and blotches of burnt 
umber, chocolate, and red-brown. Occasional clutches are 
nearly white. Fifty Richardson's merlin eggs averaged 40.2 mm 
X 31.3 mm, with ranges of 37.0 - 44.5 X 30 - 33.5 mm. Sixty 
taiga merlin eggs averaged 40.5 mm X 31.4 mm, and ranged from 
37 - 44.5 X 30 - 33 mm (Bent, 1938; Wolfe, 1946; Brown and 
Amadon, 1968). Prairie-nesting Richardson's merlins lay 
significantly larger eggs than other merlins (Temple, 1970). 

Partial incubation begins before the female completes the 
clutch, and full-time incubation begins with the laying of the 
last egg. The female does most of the incubating, and the male 
most of the hunting, calling the female off to receive food. 
One male Richardson's merlin in Saskatoon assumed incubation 
duties whenever he brought food to the female, spending as 
long as two hours on the eggs while his mate fed (Oliphant, 
1974). During incubation, male taiga merlins in Newfoundland 
flew from the nest about one-third of the times birds were 
flushed (Temple, 1972b). The Craigheads (1940) noted females 
sharing in hunting duties soon after the young had hatched. In 
observations of captive Richardson's merlins, only the female 
incubated (Fyfe, personal communication). Incubation lasts 
from 28-32 days (Fox, 1964; Brown and Amadon, 1968). 

During the incubation period, the male supplies the female with 
food. She flies off the nest to receive prey, either on a 
perch or in the air. Oliphant (1974) noted that the female 
took prey in the air only after chicks hatched. Prior to hatch- 
ing, she alighted on a perch facing the male and both grasped 
the prey in their beaks. After spreading their wings, pulling 
and calling softly, the male let go and the female flew to her 
favorite plucking perch. 

Immatures can and perhaps regularly do establish territories 
and acquire mates. Three of 20 pairs of taiga merlins breeding 
in Newfoundland in 1969 contained males undergoing their first 
prebasic molt and not yet in definitive adult plumage; all 
three of these pairs fledged young (Temple, 1972c). Bent (1938) 
noted a breeding taiga female in immature plumage which had 
laid a full clutch of eggs. 

Juvenile development 

Young hatch at intervals. The eldest is markedly larger than 
the youngest, whose small size and lack of aggressiveness at 
feedings usually result in the smallest receiving the least 
food. Sixty-two percent of 21 taiga merlin nestlings were 



18 



female (Craighead and Craighead, 1940). Forty-eight percent 
of 30 Richardson's nestlings were female (Fox, 1964). Hatching 
occurs in Alberta and Saskatchewan in early June. In northern 
forests, taiga merlin eggs generally hatch in late June (Brown 
and Amadon, 1968; Lawrence, 1949; Fyfe, personal communication; 
Craighead and Craighead, 1940). 

Adults may feed young on the first day out of the shell. Food 
provision consumes more and more time and energy of adults as 
the young quickly grow. For the first week or two, the female 
broods the young, receiving prey from the male on brief flights 
from the nest. Only the female has been noted brooding. Males 
or females may cache surplus prey on the ground or in tree 
cavities for future use. The male generally transfers avian 
prey to the female intact, who plucks it and often consumes 
the head. Females feed young birds small bits of food, and 
feedings increase in length from five minutes during the first 
week to as long as 25-35 minutes, later in the development of 
the young. Occasionally, the male plucks prey before transfer, 
and very rarely feeds the young himself--this last occurrence 
has been recorded only in taiga merlins in Minnesota by the 
Craigheads (1940). Taiga merlin females in Newfoundland often 
forced the male aggressively from the nest if he lingered after 
delivering food (Temple, 1972b). Even up to the time chicks 
are well-feathered, the female broods them at night. Daytime 
brooding continues until the chicks reach about two weeks of 
age (Lawrence, 1949; Selous, 1915; Breckinridge and Errington, 
1938; Oliphant, 1974; Temple, personal communication; Brown 
and Amadon, 1968; Fox, 1964). 

At hatching, the young are weak and immobile, their eyes closed. 
At three days of age they open their eyes. Activity increases 
steadily, until at 10 days active yawning and wing-flapping 
occur, while the primary and secondary sheaths become visible. 
Three 18-day-old young weighed about 165, 162, and 106 grams. 
At 18-20 days feathers cover almost all the down, wings are 
approaching full development, and the young falcons spend much 
time in preening, sometimes mutual preening. At this age, 
females may be separated from males by their larger feet and 
tarsi and bulkier form. At 25 days young beg food from their 
parents, "play" with nest mates, and sleep in the adult 
position (Fox, 1964; Brown and Amadon, 1968; Lawrence, 1949; 
Taylor, 1914). 

Merlins fly at 25-35 days, averaging just about a month. 
Although they leave the nest at this time, their powers of 
flight are poor, amounting to little more than extended gliding. 
They remain dependent on their parents for food for several 
weeks as they perfect predatory and flying skills. Within a 
week after fledging, young take long training flights, and at 
the end of a second week can successfully catch insects. They 
remain in the general vicinity of the nest during these two 

19 



weeks. Dispersal begins soon after, until young birds reach 
complete independence at about five weeks after fledging (Fox, 
1964; Eyre and Paul, 1973; Oliphant, 1974; Lawrence, 1949; 
Fyfe, personal communication; Brown and Amadon, 1968). 

Territoriality and interspecific relations 

Merlin eggs and nestlings occasionally fall prey to predators, 
but the adults' defense of the nest generally discourages 
territorial trespassers. Avian predation, however, could reach 
considerable proportions if parental behavior is seriously 
disrupted by pesticide residues (Fyfe, personal communication). 
Merlins usually drive away potential predators as soon as they 
penetrate the nesting territory, reacting when the invader is 
as much as .8 km. from the nest. Except for domestic dogs, 
and foxes in Newfoundland (Temple, personal communication), 
observers note merlins attacking only avian predators. Breed- 
ing and wintering falcons have chased red-billed tropicbirds 
( Phaethon aetherus ) , sharpshinned hawks, Cooper's hawks ( Accipiter 
cooperii ) , broad-winged hawks ( Buteo platypterus ) and other 
buteos, golden eagles ( Aquila chrysaetos ) , prairie falcons ( Falco 
mexicanus ) , peregrine falcons, other merlins, American kestrels, 
great blue herons ( Ardea herodias ) , gulls ( Larus sp.), great 
horned owls ( Bubo virginianus ) , crows, Clark's nutcrackers 
( Nucifraga columbiana ) , and common grackles ( Quiscalus quiscula ) . 
Antagonism between merlins and any larger falcon seems to 
intensify if the larger bird took quarry; merlins seem particu- 
larly intolerant of accipiters (Fox, 1964; Lawrence, 1949; 
Bent, 1938; Oliphant, 1974; Fyfe, personal communication; Ellis, 
1974; Allen and Peterson, 1936; Taylor, 1914). Migrating 
merlins frequently steal prey from flying kestrels and sharp- 
shinned hawks by stooping at them and snatching the prey as 
the bird tries to defend itself (Temple, personal communication). 

Merlins, in turn, may be harassed by other species. Bent (1938) 
notes an attack on an Alaskan merlin by seven black-billed 
magpies ( Pica pica ) . Fox (1964) watched his captive male mobbed 
by a party of one robin, a pair of northern orioles ( Icterus 
galbula ) , one common grackle, and several house sparrows. Other 
observers have seen robins attack merlins very aggressively 
(Fox, 1964; Lawrence, 1949). Hummingbirds (Trochilidae) con- 
sistently harassed Bond's (1936) trained female taiga bird, 
sometimes as many as 6 or 7 at once. On the breeding grounds, 
taiga merlins are often attacked by lesser yellowlegs ( Totanus 
flavipes ) if they approach the shorebird's nesting territory 
(Temple, personal communication). 

Throughout their breeding range, merlins are notoriously bold, 
fearless, noisy, and aggressive in defending their nest from 
human intruders. All three subspecies consistently advertise 



20 



nest locations through aerial and highly vocal defense (Bent, 
1938; Lawrence, 1949; Laing, 1938; Wolfe, 1946; Williams and' 
Matteson, 1947; Craighead and Craighead, 1940; Ellis, 1974). 

Generally the male merlin reacts to intruders first, flying out 
to scold them as far away as 1.5 km., and frequently at .8 km., 
from the nest (Ellis, 1974; Fox, 1964; Bent, 1938). Incubating 
or brooding females may remain silent on the nest under very 
close approach--less than 100 meters, or even until the observer 
begins to ascend the nest tree (Ellis, 1974; Bent, 1938). Once 
flushed, female merlins defend the nest more aggressively than 
males, in most cases (Fox, 1964; Lawrence, 1949; Craighead and 
Craighead, 1940). 

Richardson's merlins defend their territory from the date of 
the male's arrival, peaking in intensity when the young are 
about two weeks old. Defensive behavior then diminishes rapidly 
until it disappears about one month after fledging (Fox, 1964). 

One notable exception exists in this pattern of nest defense. 
Merlins nesting in densely populated areas of Saskatoon did not 
defend their nesting territory, but tolerated humans, dogs, 
cats, passing traffic, and large birds (including crows and 
gulls) without apparent concern or vocalization. Only when the 
nest tree was climbed to band the young, and when crows and 
grackles came within 6 meters of the nest, did the adults act 
defensively. These merlins did vocalize frequently when in- 
specting possible nest sites. At least one, and probably two, 
pairs of crows successfully reared young within 61 meters of 
one nest with very little interaction with the merlins. 

An estimated 5,000 to 10,000 people passed within 15 meters of 
two of these nests while the merlins occupied them. The merlins 
may have adapted to their urban environment by limiting nest 
defense to the nest site itself to avoid spending literally all 
day in territorial defense—which would attract still more 
predators to the nest site (Oliphant, 1974). The contrast with 
parental behavior in all other merlin populations also suggests 
pesticide residues as a possible cause of the Saskatoon breeders' 
abnormal behavior (White, personal communication), though Fox 
(personal communication) reports that these birds lay eggs with 
thicker shells and lower residues than nearby rural populations. 
Oliphant (personal communication) has observed some successfully 
breeding pairs in Saskatchewan outside the city that have shown 
little aggression toward territorial intruders. 



21 



Vocalizations 

The merlin's most frequently heard call is given during nest 
defense. Observers have variously described this as a harsh, 
loud, piercing, high-pitched "ki-ki-ki-ki-keeee" (Craighead and 
Craighead, 1940; Brown and Amadon, 1968; Oliphant, 1974); 
a "kla-kla-kla-kla-kla-kla," similar to the call of the American 
kestrel (Bent, 1938); or a "keeeyick-kyick-kyick-kyick-kyick" 
(Lawrence, 1949). Brown and Amadon (1968) attribute this 
"ki-ki-kiee" call to the male, and a lower-pitched "kek-ek-ek- 
ek-ek" to the female. Several other authors have described a 
"kek-ek" call, while not attributing it solely to the female 
(Craighead and Craighead, 1940; Bent, 1938). Oliphant (1974) 
noted that Richardson's merlins used this primary call during 
most male/female interactions, especially when the male brought 
food to the nest. A softer version of the call occurred when 
adults alternated incubatory duties or transferred prey while 
perched on a limb (see page 16 of this report). Female merlins 
have used the primary call while calling in fledged young to 
feed (Street, 1950). 

In courtship flight, the male gives a "chrrrrr" call while 
flying. During copulation the call may be either this same 
"chrrrrr" or a slightly different and higher note. In display- 
ing to the female at the nest, the male gives a high "chip, 
chip, chip" note which the female may answer (Fyfe, personal 
communication) (see also page 14). This "chip" note may be the 
same call described in males by Brown and Amadon (1968) as a 
thin, chirruping "chic-chic-chic" uttered more slowly than the 
rapidly repeated primary call, and by Bent (1938) in black 
merlins in their nest tree with fledged young: a plaintive 
whimpering not unlike the "kik-kik-kik" call of the flicker, 
but much more subdued. 

In calling females off the nest to receive food, males may use 
a series of single, soft "chups" (Oliphant, 1974). In the same 
situation, the Craigheads (1940) noted a male to emit a call 
very similar to the long drawn out food cry of the peregrine 
falcon, but higher pitched. Females solicit food with an extended 
plaintive "eeep-eeep-eeep" (Brown and Amadon, 1968). 

When the female feeds the young during the first week or two 
after hatching, and while she herself feeds, she emits a sharp 
"tick-tick-tick" or a series of "clicks," touching a chick's 
bill with a morsel of food at the same moment (Lawrence, 1949). 
The young, in turn, give a "keeyep-keeyep-keeyep" begging call 
(Lawrence, 1949; Brown and Amadon, 1968). Just prior to and 
after fledging, young merlins may emit a series of single rising 
notes, or "chees," especially at feeding times. Within a week 



22 



-■^■.in^.JM^r wra^ 



of fledging, the young birds space these single calls more 
closely, and begin to group them in calls more like those of 
adults (Oliphant, 1974). 

Migration and wintering behavior 

Richardson's merlins migrate a relatively short distance into 
the southern Great Plains; a smaller segment of the population 
winters on their northern plains breeding grounds. The black 
merlin resides semipermanently in the climatically stable 
coastal forests of the Northwest, and only a small portion of 
the population migrates south of their breeding range. Most 
taiga merlins, in contrast, are highly migratory. Western and 
central taiga-breeding birds migrate past Richardson's and 
black merlin populations in leapfrog fashion to winter in 
Central*and South America. Eastern taiga birds also migrate 
far to the south, many individuals crossing the Gulf of Mexico 
to winter on the Caribbean Islands and in South America. A 
few taiga birds remain in the boreal forest all year (Temple, 
1972a; Beer, 1966). 

During fall migration, merlins tend to follow sea coasts, 
borders of streams and large lakes, and open country, preying 
on small birds also making their way south—although in 
Minnesota merlins seem to scatter, rather than follow well- 
defined migration routes (Brown and Amadon, 1968; Bent, 1938; 
Beer, 1966). Fall migration is much more conspicuous than the 
return trip northward in spring (Brown and Amadon, 1968; Bent, 
1938). Along the New Jersey coast, taiga merlins (along with 
kestrels and peregrines) wander southward over salt meadows in 
preference to wooded areas favored by migrating accipiters 
(Allen and Peterson, 1936). At Fishers Island, New York, they 
favor flying on a southwest wind (Bent, 1938). Merlins fly 
rather high, sometimes in large straggling flocks (Brown and 
Amadon, 1968). In Washington, migrant black merlins prefer 
open country and tidal flats along Puget Sound in both fall and 
early spring (Bent, 1938). Some merlins in Alaska migrate along 
the Alcan Highway, perhaps to take advantage of the abundant 
food in the open pathway through the forest, for thousands of 
Lapland longspurs ( Calcarius lapponicus ) migrate along this 
same route (White, 1969). 

Immature merlins generally migrate before adults, leaving breed- 
ing grounds about one month to six weeks after leaving the 
nest (Bent, 1938; Brown and Amadon, 1968). Young Richardson's 
merlins drift southward from Saskatchewan in mid-August (Fox, 
1964), about as early as any merlins in North America. Migra- 
tion peaks between mid-September and mid-October in most areas. 
Nearly all merlins have left the northern fringes of their 
range by September 15; migration peaks in Minnesota during the 
second and third weeks of September, in South Carolina in 
October (Brown and Amadon, 1968; Beer, 1966; Bent, 1938). Most 

23 



Richardson's merlins arrive on their eastern Colorado wintering 
grounds by the last two weeks of October; immatures migrate 
farther south than adults--as far as Arizona and New Mexico, 
rather often. More adults remain on the Canadian prairies 
through the winter (Fox, 1964; Bailey and Niedrach, 1965). 

Some wintering birds seem to wander at random; others stay for 
some time at a location (such as an urban area) that provides 
abundant prey (Fox, 1964; Oliphant, personal communication). 
The three races may mingle in winter quarters (Eyre and Paul, 
1973; Brown and Amadon, 1968). 

In spring, most merlins pass northward through central North 
America in March (Brown and Amadon, 1968; Fox, 1964; Williams 
and Matteson, 1947). Most taiga-bound birds pass through 
northern Minnesota in April (Beer, 1966). The very last to 
leave the more southern winter ranges head north in late March 
and early April (Bent, 1938). 

Productivity 

Productivity in North American merlin populations has declined 
significantly in the past 20 years. Prior to about 1950 (and 
prior to the widespread use of persistent organochlorine pesti- 
cides), merlins across the continent hatched more young per 
nest than they have since that date. 

The mean clutch of merlins has varied little since the first 
data were recorded. The number of young hatched per nest in 
the northern Great Plains prior to 1950 averaged 4.3 (N=3) , 
from a mean of 4.7 eggs/nest (N=10) (Fox, 1971). In eastern 
Canada, taiga merlins laid 4.2 egg clutches (N=18), and 
hatched 3.8 young per nest with young (N=15) prior to 1947 
(Temple, 1972c). 

Eastern taiga birds have suffered lower productivity since 
those years. From 1950-1969, these merlins laid average 
clutches of 4.5 (N=2), from which they hatched only 2.2 young 
per nest with young (N=6) ; hatching success totaled 48%, down 
from almost 91% prior to 1947 (Fox, 1971; Temple, 1972c). 
Newfoundland merlins in 1969 laid clutches averaging 4.3 eggs 
(N=15), and hatched 3.0 young per nest with young (N=19) 
(Temple, 1972c). Although these Newfoundland birds reached 
only 707o hatching success, they fledged young in 95% of 20 
nests. Central and northern forest-breeding birds have main- 
tained the highest productivity of any North American merlins 
in recent years. From 1960-1969, northern taiga merlins laid 
4.5 eggs/nest (N=2) , hatching 4.0 birds per nest with young 
(N=5)--a hatching success of 897o (Fox, 1971). Merlins breeding 
in forested areas of the Great Plains maintained good production 



24 






of young from 1950-1969, laying 4.1 eggs/nest (N=9), and hatch- 
ing 98% of these--4.0 young per nest with young (N=16) (Fox, 
1971). Only one of two taiga merlin nests recently found in 
Alaska fledged young, but this single nest fledged five birds 
(Adolphson, 1969). 

Richardson's merlins nesting in the Great Plains have decreased 
more alarmingly in productivity. Fox (1964) found that 10 pairs 
of Saskatchewan-nesting merlins laid an average of 4.5 eggs/nest, 
Hatching success, 91% prior to 1950, decreased to 60% for these 
birds, which hatched 2.7 young per nest with young. Fledging 
remained good, however, with 2.6 young fledged per nest. Most 
disturbing, though, is the fact that this population of merlins 
ceased to breed soon thereafter (Fox, 1971). Hatching success 
throughout the Great Plains from 1950-1969 averaged 49% (Fox, 
1971). Richardson's merlins breeding in Montana laid clutches 
of five eggs, but hatched only 3.2 young per nest with young 
(Ellis, 1974). They fledged 3.2 birds per nest with fledglings; 
57.1% of seven nests fledged young. Hunter (1967) reports that 
three merlin nests in South Dakota all failed to fledge young. 
Five of eight (62.5%) Saskatoon nests fledged young; all five 
fledged 4-5 young (Oliphant, 1974). 

These data indicate decreasing productivity in North American 
merlin populations. Fyfe (personal communication) attributes 
lowered Richardson's merlin productivity primarily to desertion 
during incubation and failure of incubated eggs to hatch. 
Temple (1972c) believed that problems in Newfoundland were 
probably due to increased egg loss and decreased hatchability . 
Broken eggs and adult mortality resulted in 2 out of 3 nest 
failures in Montana (Ellis, 1974). 

Pesticide relationships 

Research has linked decreased reproductive success and popula- 
tion declines in bird-eating raptor populations to accumulation 
and concentration of chlorinated hydrocarbon pesticide residues 
through the food chain. Zarn (1974) has reviewed briefly the 
effects of DDT and other organochlorine pesticides on raptors. 
Aberrant behavior and eggshell-thinning seem the most conspic- 
uous symptoms of pesticide poisoning. Specifically, chlorinated 
hydrocarbons seem to inhibit carbonic anhydrase activity in the 
shell gland, resulting in decreased calcium carbonate deposition 
in the shells of developing eggs, and consequently, thinner- 
shelled eggs. 

Merlins, like other bird-eating falcons, are suffering reduced 
productivity probably resulting from accumulation of pesticide 
residues. The lipid of four adult merlins migrating along the 



25 



western shore of Lake Michigan contained an average of 302 ppm 
DDE, and 196 ppm PCB compounds (polychlorinated biphenyls) . DDE 
accounted for 96% of all DDT residues in the birds. The lipid 
of three immature merlins contained 50.3 ppm total DDT, of 
which 49.3 ppm was DDE. PCB ' s averaged 28.6 ppm. All but one 
of the merlins biopsied (both adult and immature) contained 
more DDE than PCB ' s . Peregrines trapped here showed the 
reverse ratio. Pesticide fallout must evidently vary in the 
areas of boreal forest and tundra where these two migrants 
respectively breed (Risebrough, e_t a_l. , 1970). These figures 
may be compared to DDE figures for lipids from adult peregrines 
from the MacKenzie River region (Enderson and Berger, 1968) and 
from Alaska (Cade, e_t al_. , 1968) of 392 ppm and 725 ppm, 
respectively. The merlins have almost the same level of con- 
tamination as the peregrines, and might therefore be expected 
to show eggshell thinning. 

Fox (1971) calculated eggshell weights for 51 merlin eggs 
collected from 1890 to 1969. From 1890 to 1949, eggshells 
from Great Plains and taiga-breeding birds did not differ 
significantly and averaged about the same through those years-- 
1.71 grams. From 1950-1969, merlin eggshells averaged 1.31 
grams, a 23% decrease. This decrease resembles those reported 
by Hickey and Anderson (1968) for declining bald eagle 
( Haliaeetus leucocephalus ) and osprey ( Pandion haliaetus ) popu- 
lations and extirpated peregrine falcon populations in the 
United States. Fox (personal communication) studied 40-50 pairs 
of Richardson's merlins nesting in Alberta in 1974, and judged 
reproductive success 40-50% of that prior to pesticide use, 
while eggshells averaged 15-20%, thinner and pesticide residues 
proved high. Swartz (personal communication) has noted high 
pesticide residues and eggshell thinning in eggs from interior 
Alaskan merlins, although productivity appears fairly normal. 

Merlins breeding in Newfoundland contained 267 ppm (dry weight 
basis) DDE, while five unhatched eggs averaged 40.4 ppm (dry 
weight basis) DDE (Temple, 1972c). These residue counts are 
lower than the breeding peregrines and their eggs and lower 
than the Lake Michigan migrant merlins. No marked delay in the 
breeding cycle of the Newfoundland merlins has occurred, nor 
has clutch size decreased significantly since pre-pesticide 
times. Eggshells, however, have decreased 97<> in thickness, a 
decrease accompanied by reduced productivity. 

In Montana, residues (wet weight basis) in 3 non-viable eggs 
from one Richardson's merlin nest averaged 9.4 ppm DDE, 8.0 ppm 
dieldrin, and 5.6 ppm heptachlor epoxide. PCB ' s , toxaphene, 
and other chlorinated hydrocarbons were undetected (Ellis, 1974). 
Only one egg contained a dead embryo. Pesticide residues 
calculated on a dry weight basis are roughly comparable to 5x the 



26 



level calculated on wet weight basis; lipid fractions are about 
17-20x wet weight (Fox, personal communication). DDE residue 
levels in these Montana eggs thus are similar to DDE levels in 
Temple's (1972c) Newfoundland eggs. But the Montana merlins 
suffered considerably more nest failures--3 out of 7 as compared 
to 1 out of 20 in Newfoundland. Embryonic mortality, in con- 
trast, was high in the Newfoundland population: all five 
unhatched eggs collected for residue samples contained dead 
embryos . 

HABITAT REQUIREMENTS 

Breeding requirements 

Taiga merlins breed in the boreal forest, and within this 
biome choose somewhat open territories, such as forest edges, 
lake shores, bogs, etc. These merlins generally nest adjacent 
to water. All nests described from northern Minnesota were in 
conifers near the edge of large lakes, many on islands in the 
lakes (Beer, 1966). In denser areas of the boreal forest, 
taiga merlins usually nest in abandoned crow or common raven 
( Corvus corax ) nests in coniferous trees. Trees utilized 
include many dominant boreal species: black spruce ( Picea 
mariana ) , red spruce ( Picea rubens ) , white spruce ( Picea glauca ) , 
eastern white pine ( Pinus strobus ) , jack pine (Pinus banks iana ), 
etc. Along the northern edge of their range, on the fringes 
of the tundra, they often nest under tree branches in a scrape 
on the ground (Temple, personal communication; Fyfe, personal 
communication; Johnson and Coble, 1967; Bent, 1938; Craighead 
and Craighead, 1940; Lawrence, 1949). 

Taiga merlins may reline the nest with some combination of 
small twigs, pieces of bark, feathers, grass, soft rootlets, 
and conifer needles. They nest almost anywhere in trees, from 
1.5 meters above ground to the top of 18-meter spruce, usually 
very close to the trunk. Taiga merlins also have nested in 
tree cavities, under the roofs of deserted buildings, and on 
cliff ledges in nests made of bits of rock (Bent, 1938; Craig- 
head and Craighead, 1940; Fox, 1964; Williams and Matteson, 
1947; Lawrence, 1949). 

Few nests of the black merlin have been described in detail. 
These merlins breed in moist Pacific coastal forests, probably 
in areas similar to the open habitat favored by taiga merlins 
in boreal forest. Most observed nests of F. c_. suckleyi have 
been very high in conifers in dense coastal stands of Douglas 
fir ( Pseudotsuga menziesii ) , Sitka spruce ( Picea sitchensis ) , 
western hemlock ( Tsuga heterophylla ) , Pacific silver fir 
( Abies amabalis ), and western redcedar ( Thuja plicata ) . Both 
stick-nests and cavity-nests are known. Black merlins frequently 
nest near large lakes (Temple, personal communication; Laing, 
1938; Bent, 1938). 

2/ 



Richardson's merlin breeds only in the prairie-parkland of the 
northern Great Plains. Within this biome, the subspecies seems 
to prefer isolated groves of trees with open prairie surround- 
ings, mixed woods, and wooded areas along prairie river banks 
and islands (Temple, personal communication; Fyfe, personal 
communication; Fox, 1964). 

F. c_. richardsonii tend to breed where native grassland provides 
adequate prey species, and quaking aspen ( Populus tremuloides ) 
or other trees provide suitable nesting habitat. On the Sas- 
katchewan-Alberta prairie they prefer to nest in deserted 
farmstead shelterbelts and groves of deciduous trees where 
farmland has gone back to grassland. The merlins choose trees 
spaced well apart whose lower 2.4 or 3 meters have been rubbed 
bare of branches, and where undergrowth has been trampled and 
destroyed by cattle. This appears to be a temporary stage in 
the total destruction of trees by cattle, and the number of 
sites that fit this description is steadily decreasing. Fox 
(1964) found about half of 25 nests in Saskatchewan within .8 
km. of water. He also noted that water may be utilized by 
merlins for bathing. Captive merlins have been known to bathe 
in water, although Wing (1950) observed a wild merlin dust 
bathing (Fyfe, personal communication; Fox, 1964, 1971). 

In southern areas of the open grassland, Richardson's merlins 
nest most frequently in deserted crow and magpie nests, and 
seem to have a preference for the latter (Fyfe, personal communi- 
cation). Ninety-two percent of Fox's (1964) nests were old 
nests of these two species. On the same Saskatchewan study 
area, 21 of 25 merlin pairs nested in deciduous trees: aspen, 
poplar ( Populus sp.), box elder ( Acer negundo ) , willow ( Salix sp.), 
and birch ( Betula sp.). The key determinant is the nest rather 
than the species of tree: in areas where crows and magpies 
nest in conifers, merlins will then nest in these nests. All 
of seven Richardson's merlin nests on the Montana prairie were 
in groves of conifers—Douglas fir, ponderosa pine ( Pinus 
ponderosa ) , and limber pine ( Pinus flexilis ) --near flat or 
rolling grassland. All these merlins nested in old corvid 
nests--six of them magpie nests (Ellis, 1974). 

The prairie subspecies occasionally nests in tree cavities and 
in old magpie nests placed in holes in cliffs (Fyfe, personal 
communication). Nests are usually lined with dry inner bark 
of poplar (Bent, 1938). Seven of eight Richardson's merlin 
pairs in Saskatoon nested high in large spruce, with tall 
deciduous trees nearby, always in what appeared to be an old 
crow's nest. Two pairs nested in highly populated areas within 
about 30 meters of major roads with heavy traffic (Oliphant, 
1974). 






28 



No observers have recorded Richardson's merlins nesting on the 
ground, a most important fact considering the serious shortage 
of suitable nest trees in the subspecies' range. A shift in 
nesting patterns to include ground-nesting would greatly in- 
crease habitat available for this bird (Fyfe, personal communi- 
cation), though predator pressure would also increase (Ellis, 
personal communication) . 

Wintering requirements 

Wintering taiga merlins tolerate an amazing range of habitats-- 
from beaches, dunes, and swamps to tropical forests and scrub, 
to farmland and urban areas. Black merlins also range through 
a variety of habitats on their wintering grounds--desert scrub, 
coastal forest, coniferous forest, farmland, and urban areas 
(Temple, personal communication). 

Richardson's merlins, on the other hand, seem more selective in 
choosing wintering grounds, preferring prairie habitat with 
scattered trees, utility poles, or other hunting perches- 
habitat very similar to their breeding habitat (Temple, personal 
communication). Fence posts and lower perches attract the 
falcons even where telephone poles are available (Fox, 1964). 
Wintering Richardson's merlins favor winter wheat stubble in 
Wyoming, and in Colorado prefer very open country on the plains 
or land along or near creek beds overgrown with cottonwoods 
and willows (Fox, 1964) . In Utah, where all three subspecies 
winter, Richardson's merlins occur with greatest frequency in 
open farmland; the other two subspecies show no strong habitat 
preference (Eyre and Paul, 1973). In northern parts of its 
wintering range, Richardson's merlin also frequents urban 
areas; many immature birds, for example, winter in Saskatoon 
(Temple, personal communication; Fyfe, personal communication; 
Oliphant, 1974 and personal communication). 

LIMITING FACTORS 

Three factors presently limit merlin populations. Two are com- 
ponents of the environment required for successful reproduction: 
first, an adequate food base and second, available, and suitable, 
nest sites. The third factor exists in varying strength, and 
results in varying decreases in productivity: pesticide resi- 
dues (Fyfe, personal communication). 

Habitat destruction 

Habitat destruction to date is of minor importance for taiga and 
black merlins. Hence, their supplies of food and nest sites 
remain adequate (Fyfe, personal communication; Temple, personal 
communication). This is not true for Richardson's merlins, 
however. 



29 



An adequate food base for prairie merlins means an adequate 
supply of grassland birds—which can only mean uncultivated 
grassland habitat. Without native grassland, there will be 
few Richardson's merlins (Fyfe, personal communication). Like- 
wise, destruction of habitat reduces the number of adequate 
nest sites—which, for this subspecies, are narrowly defined. 
Fox (1971) documents a steadily decreasing number of suitable 
nest trees on the Saskatchewan prairies. Merlins have ceased 
breeding entirely in some areas of intensive cultivation (Fox, 
personal communication). Habitat destruction continues to 
accelerate as more and more land goes under intensive agriculture, 
putting ever greater pressure on merlins of the prairies. 

Chemical contamination 

Merlins are carrying high pesticide residue levels— organo- 
chlorines, mercury, and PCB ' s (Fyfe, personal communication). 
Thin-shelled eggs and poor overall productivity have been 
noted in varying degrees throughout the range of the species. 
Richardson's merlins, however, breed in the most contaminated 
habitats and predictably their reproductive success has de- 
creased drastically over most of their range. Pesticide 
contamination may be the single biggest problem facing North 
American merlins (Temple, personal communication; Fox, personal 
communication) . 

Little seasonal variation exists in the pesticide content of 
merlin diets, since most of these falcons prey on the same 
species on both breeding and wintering grounds (Temple, personal 
communication). Migration patterns in prey species can, how- 
ever, dramatically affect pesticide levels in prey and predator. 
In Newfoundland, brain tissue of resident gray jays contained 
only .24 ppm (dry wt. basis) DDE, while brains of migrant robins 
and Savannah sparrows contained 3.17 ppm and 2.10 ppm, respect- 
ively. These three species accounted for the bulk of the breed- 
ing merlins' diet (Temple, 1972c). 

Other mortality factors 

Several specific merlin mortality factors have been noted. 
Nestlings or eggs may not survive bad storms (Oliphant, 1974), 
and may also succumb to disease or parasites, such as Simulium 
flies (Ellis, 1974). Adults and young may catch fatal 
Trichomonas gallinae (frounce) from consuming infected doves 
(Stabler, 1969). Before protective legislation, merlins were 
often shot; they allow a close approach when perched and will 
fly quite near a man in the open (Bent, 1938; Lawrence, 1949; 
Jewett, 1948). Temple (personal communication), however, feels 
that shooting no longer poses a serious threat to the species, 



30 



at least as indicated by banding recoveries. Merlins have 
also been known to die on impact with windows in cities 
(Porter and Knight, 1952). 

Museum specimens of Richardson's merlins examined by Fox (1964) 
showed a decrease in the ratio of subadults to adults from 607 o 
in August to 30-407» in January, when molt begins. Over 707» 
of total returns for merlins banded as nestlings occurred during 
their first year of life. 

Human interaction with nesting merlins may have serious and 
harmful effects on reproductive success. Oliphant (1974) 
believed that two Saskatoon Richardson's merlin nests may have 
failed because the nest tree was climbed during the early 
stages of incubation. As far as he knew, no successful nest 
was climbed prior to hatching. Temple, Fox, and Ellis all 
climbed nests which were ultimately successful, however (Ellis, 
personal communication). Bent (1938) notes that if the nest of 
a Richardson's merlin is climbed before an egg is laid, the 
adults will always desert the nest, though merlins do not desert 
easily after the clutch is complete (Fox, personal communica- 
tion). In contrast, a pair of taiga merlins moved their nest 
and two eggs some 36 meters after two near misses by observers 
attempting to collect the adults as specimens (Bent, 1938). 

Falconers trap numbers of merlins annually, and have definite 
impact on the population of the species. Such impact may prove 
positive, since some trapped birds are used in captive breed- 
ing projects. But as the species decreases in abundance, the 
negative impacts of trapping birds from the wild increase. 

SPECIES AND HABITAT MANAGEMENT RECOMMENDATIONS 

1. We know little of merlin distribution, breeding behavior, 
habitat requirements, or reproductive success in Alaska. 
We know virtually nothing about black merlin ecology. 
Breeding population surveys, as well as any black merlin 
research, are essential before we have the basic data 
necessary to formulate management plans. Merlins in 
Alaska, although reproducing adequately now, as far as we 
know, should be watched very closely. Development of the 
state's resources has increased sharply in pace, and wilder- 
ness country has begun to shrink. Merlins no longer remain 
isolated from the drastic changes in habitat wrought by man. 
Managers of public lands have an enormous responsibility to 
carefully consider the effects of each of their decisions on 
merlins, as well as on all wildlife species. 

2. Ban, or at least severely limit, use of persistent pesticides 
in North America and in Central and South American wintering 
areas . 



31 



3. In the northern Great Plains, habitat management for 
Richardson's merlins seems possible. First, land managers 
should give careful consideration to preservation of native 
prairie whenever development alternatives exist. BLM- 
administered public lands as well as National Grasslands, 
National Wildlife Refuges, and National Parks and Monuments, 
are prime candidates for preservation of grassland habitat. 
Without the native grassland, the merlins' food base will 
disappear, along with the merlins themselves soon after 
(White, 1974). 

Merlin nesting sites should also be provided both on grass- 
land preserves and on other managed grasslands. Scattered 
small groves of trees managed for merlin nest sites would 
benefit the falcon, even in agricultural areas and on 
wintering grounds (Temple, personal communication). Con- 
struction of artificial nest sites, as described for 
ferruginous hawks ( Buteo regalis ) in Olendorff, 1973, should 
also be investigated. Anywhere in the northern plains where 
such management occurs, even though perhaps not presently 
within the Richardson's merlin breeding range, could prove 
valuable for the falcon if it shifts its range slightly to 
adapt to the change in habitat. 

White (personal communication) has pointed out that such 
shifts in range can indeed take place. Observers in Utah 
collected one probable Richardson's merlin about 1870, but 
from 1920-1960 only non- richardsonii were seen and collected 
in the state. In about the mid-1960' s, Richardson's merlins 
suddenly increased in frequency in the state, until today 
(1975) about half of the eight to ten merlins trapped 
annually along the Wasatch front are F. c_. richardsonii . 

Fox (personal communication) believes that Richardson's 
merlins have already begun to shift their range in response 
to increased cultivation and habitat loss in rural Saskat- 
chewan and Alberta, and cites recent nestings in Saskatoon, 
Edmonton, Montana, and Wyoming are evidence. He predicts 
an increase in Montana, Wyoming, and North and South Dakota 
nestings . 

4. Carefully regulate capture of merlins for falconry purposes. 
PROTECTIVE MEASURES INSTITUTED 

A. Legal or regulatory 

1. Regulations administered by the U. S. Environmental 
Protection Agency limiting the use of DDT and other 
persistent pesticides within the United States should 



32 



'• 



benefit merlins by reducing pesticide burdens in merlin 
prey. Similar regulations are also in effect in 
Canada. 

2. On March 10, 1972, the United States and Mexico added 
merlins to the list of bird species protected by the 
convention for the Protection of Migratory Birds and 
Game Mammals, originally ratified on February 7, 1918. 

B . Captive rearing 

One method of protecting species endangered by pesticide- 
contaminated environments is to maintain the species in 
captivity until the environment becomes safe, and reintro- 
duction is possible. Merlins bred successfully in captivity 
for the first time in 1974. The following list details 
recent captive merlin breeding projects: 

1. John Campbell, of Black Diamond, Alberta, raised four 
young merlins in 1974--the first successful captive 
rearing of the species (Fyfe, personal communication; 
Swartz, personal communication). 

2. In 1974, Richard Fyfe's (Edmonton, Alberta) captive 
merlins laid fertile eggs, but they didn't hatch; those 
eggs left with the females (rather than placed in an 
incubator) disappeared, perhaps eaten by the adult 
falcon (Fyfe, personal communication). 

3. L. G. Swartz, at the University of Alaska, had a near- 
miss in 1969, when his female merlin died with a full 
clutch of five eggs "on the way" (Swartz, 1972). 

4. Two other captive breeding projects in the United States 
have not been successful to date. The first is run by 

a private individual in California. The second, the 
U. S. Fish and Wildlife Service project at Patuxent, 
Maryland, has succeeded in getting fertile eggs, though 
none have hatched (Fyfe, personal communication). 

CURRENT RESEARCH 

Richard Fyfe and his coworkers of the Canadian Wildlife Service 
conduct merlin research in western Canada. Their work has 
emphasized Richardson's merlins, but concerns taiga merlins as 
well. They have considerable data on merlin courtship and 
breeding biology, population changes and habitat degradation, 
and residue levels (Fyfe, personal communication). L. W. 
Oliphant (Dept. Vet. Anat., Coll. West. Vet. Med., Univ. of 
Sask. , Saskatoon) has recently received funds to conduct merlin 
research in the Saskatoon area (Oliphant, personal communication) 



33 



AUTHORITIES 

1. Dr. Stanley A. Temple 
Laboratory of Ornithology 
Cornell University 
Ithaca, New York 14850 

2. Richard Fyfe (F. c_. richardsonii and columbarius ) 
Canadian Wildlife Service 

Room 1110 

10025 Jasper Avenue 

Edmonton, Alberta T50 156 

Canada 

3. Glen A. Fox (F . c_. richardsonii ) 
Toxic Chemical Section 
Canadian Wildlife Service 
Environment Canada 

Ottawa, Ontario K1A 0H3 
Canada 

4. Dr. L. G. Swartz (Alaska) 
Biology Department 
University of Alaska 
College, Alaska 99701 

5. Dr. David H. Ellis (Montana) 

U. S. Fish and Wildlife Service 

Room B-3 

301 W. Congress St. 

Tucson, Arizona 85701 

SUMMARY 

Merlins, small swift-flying falcons, range through much of the 
northern hemisphere. Three subspecies breed in North America, 
each in an ecologically distinct region. Male merlins have 
blue-gray dorsal plumage, females brownish plumage. Immatures 
resemble adult females. Merlins average 25 to 34 centimeters 
long, with a wingspread of about 60 cm. Small size, a distinctly 
banded tail, dark dorsal plumage, and indistinct facial markings 
distinguish merlins from other falcons. 

Falco columbarius columbarius , the taiga merlin, breeds in the 
boreal forest from Alaska to Newfoundland, and winters south to 
South America. F. c_. richardsonii , the Richardson's merlin, 
breeds in the prairie-parkland of the northern Great Plains, 
wintering mostly in Colorado and Wyoming. The black merlin, 
F_. c_. suckleyi , resides semipermanently in the moist coastal 
forests of western British Columbia, and perhaps, Washington 
and southeast Alaska. Some black merlins winter south as far 
as California and New Mexico. 

34 






All North American merlins have recently decreased in reproduc- 
tive success. Organochlorine pesticide residues have contributed 
to these decreases. Intensive agriculture has decreased both 
the native grassland necessary to support the food base of 
Richardson's merlins and the number of suitable nest-trees. 
Taiga and black merlins, as well, face habitat destruction as 
man develops the natural resources of the northern forests. 

Merlins hunt diurnally, occasionally into twilight. They consume 
birds, almost exclusively, with insects a minor, but significant, 
part of their diet. Mammals are taken occasionally. Merlins 
hunt avian prey ranging in size from sparrows to ptarmigan. 
They attack with a direct, very fast dash, but also may stoop 
from high above prey. 

Most males arrive on the breeding territory in April, about a 
month before females. Males display to females during courtship. 
Merlins generally do not nest in the same site in successive 
years, but may nest in the same area. Nesting density is low. 

Merlins lay clutches of four or five eggs from April to July, 
mostly in May. Females do most, and usually all, of the incu- 
bating. Incubation lasts 28-32 days. Males provide food for 
the incubating female, and later the young as well. Males, 
however, rarely feed the young. Young merlins fledge at about 
one month of age. About five weeks later they reach full 
independence. 

Nest defense by adults discourages most predators. Merlins 
react very aggressively to large birds and raptors near their 
nests, as they do to human intruders. Some merlins nesting 
in urban environments refrain from such aggressive nest defense. 
Merlins vocalize often when breeding, using their primary nest 
defense call most frequently. 

Fall migration is much more conspicuous than spring migration. 
Migration southward peaks between mid-September and mid-October. 
In spring, most merlins pass through central North America in 
March. 

Productivity has declined significantly in North American merlin 
populations during the past 20 years. Desertion during incuba- 
tion, and increased egg loss and decreased hatchability seem 
the major problems. These all derive primarily from organo- 
chlorine pesticide residues in breeding merlins, resulting in 
eggshell thinning, dead embryos, and aberrant parental behavior. 



35 



* 



Taiga merlins breed in open areas of the boreal forest, usually 

in abandoned corvid nests in conifers, but occasionally on the 

ground. Taiga merlins usually nest near water. Black merlins 

favor open habitat in the Pacific coastal forests, nesting very 

high in conifers. Richardson's merlin breeds in old corvid 

nests on the prairie-parkland, in deciduous trees or conifers-- 

wherever the crows and magpies choose to build nests. Suitable 

nest sites for Richardson's merlins on the Canadian prairies t 

are diminishing fast in number. Richardson's merlins do not . 

nest on the ground, an adaptation which would greatly expand 

suitable habitat. 

Wintering taiga and black merlins tolerate a wide variety of 
habitats. Richardson's merlins have narrower tolerance for 
wintering grounds, seeking open farmland and prairie similar 
to their breeding habitat. 

Three main factors limit merlin populations: availability of 
nest sites, adequacy of the food base, and pesticide residue 
levels. All three subspecies depend on these same factors, but 
Richardson's merlin populations have been most pressured, and 
therefore are in the greatest trouble. Other mortality factors 
(disease, weather, etc.) are not as immediately important. 
Nests may fail, however, if the nest tree is climbed during 
incubation. 



36 



• 



LITERATURE CITED 



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Allen, R. P. and R. T. Peterson. 1936. The hawk migrations 
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American Ornithologists' Union. 1957. Check-list of North 
American birds. Fifth edition. Lord Baltimore Press, 
Baltimore. 691 pp. 



Bailey, A. M. 1942. The black pigeon hawk in Colorado. 
Condor 44:37. 

and R. J. Niedrach. 1965. Birds of Colorado. 



Denver Museum of Natural History. Vol 1:246-248. 

Beer, J. R. 1966. The pigeon hawk in Minnesota. The Loon 38(4) 
129-132. 

Bent, A. C. 1938. Life histories of North American birds of 
prey, Vol. II. U. S. Natl. Museum Bull. 170, Wash. D. C. 
pp. 70-90. 

Bond, R. M. 1936. Speed and eyesight of a pigeon hawk. 
Condor 38:85. 

Breckinridge, W. J. and P. L. Errington. 1938. Food habits of 
small falcons in north-central states. Auk 55:668-670. 

Brown, L. and D. Amadon. 1968. Eagles, hawks, and falcons of 
the world. McGraw-Hill, New York. Vol. 11:802-807. 

Burleigh, T. D. 1972. Birds of Idaho. Caxton Printers, 
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Cade, T. J., C. M. White, and J. R. Haugh. 1968. Peregrines 
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Craighead, J. and F. Craighead. 1940. Nesting pigeon hawks. 
Wilson Bulletin 52:241-248. 



Ellis, D. H. 1974. The first breeding records of merlins in 
Montana. (in prep.) 

Enderson, J. H. and D. D. Berger. 1968. Chlorinated hydro- 
carbon residues in peregrines and their prey species from 
northern Canada. Condor 70:149-153. 



37 



Eyre, L. and D. Paul. 1973. Raptors of Utah. Utah Division 
of Wildlife Resources, p. 63. 

Fox, G. A. 1964. Notes on the western race of the pigeon 
hawk. The Blue Jay 22(4) : 140-147 . 

. 1971. Recent changes in the reproductive success 



of the pigeon hawk. J. Wildlife Management 35:122-128. 

Friedmann, H. 1950. The birds of North and Middle America, 
Part XI. Smithsonian Institution/U. S. National Museum 
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Gabrielson, I. and F. C. Lincoln. 1959. The birds of Alaska. 
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Wash. D. C. 922 pp. 

Hickey, J. J. and D. W. Anderson. 1968. Chlorinated hydro- 
carbons and eggshell changes in raptorial and fish-eating 
birds. Science 162(3850) :271-273. 

Hunter, D. 1967. Raptor populations: South Dakota population 
survey. Raptor Research News 1(3) :41. 

Jewett, S. G. 1939. Additional notes on the black pigeon hawk. 
Condor 41:84-85. 



• 



I 
I 
I 
I 



I 
I 
I 



. 1944. 

Condor 46:206. 



1948. 



The black pigeon hawk in New Mexico. 
Another black pigeon hawk from California. 



Condor 50:228. 



, W. P. Taylor, W. T. Shaw, and J. W. Aldrich. 1953. 

Birds of Washington State. Univ. Washington Press. 767 pp. 

Johnson, W. J. and J. A. Coble. 1967. Notes on the food habits 
of pigeon hawks. Jack-Pine Warbler 45(3):97-98. 

Laing, H. M. 1935. Notes on the black pigeon hawk. Auk 52: 
305-307. 

. 1938. Additional notes on breeding of black 



pigeon hawk. Auk 55:525-527. 

Lawrence, L. de K. 1949. Notes on nesting pigeon hawks at 
Pimisi Bay, Ontario. Wilson Bull. 61:15-25. 

McClure, H. E. 1957. Merlin attacks brown-eared bulbuls. 
Auk 74:101. 



c 
I 



* » 



38 



I 



Miller, L. 1941. The black merlin in southern California. 
Condor 43:199-200. 

Munro, J. A. 1946. A record of the black pigeon hawk nesting 
in British Columbia. Condor 48:182. 

Olendorff, R. R. 1973. The ecology of the nesting birds of 
prey of northeastern Colorado. U. S* IBP Grassland Biome 
Tech. Report 211. 233 pp. 

Oliphant, L. W. 1974. Merlins--the Saskatoon falcons. Blue 
Jay 32(3). 8 pp. 

Peterson, R. T. 1961. A field guide to western birds. 
Houghton-Mifflin Co., Boston. 309 pp. 

Porter, R. D. and H. Knight. 1952. Records of the black pigeon 
hawk, Falco columbarius suckleyi , in Utah. Auk 69:84-85. 

Risebrough, R. W., G. L. Florant, and D. D. Berger. 1970. 
Organochlorine pollutants in peregrines and merlins 
migrating through Wisconsin. Can. Field Nat. 84(3) :247-253. 

Selous, E. 1915. A diary of ornithological observation made 
in Iceland during June and July, 1912. The Zoologist 19: 
58-66, 169-174. 

Slipp, J. W. 1942. Black pigeon hawk taken in eastern Wash- 
ington. Condor 44:128. 

Spaulding, E. R. 1947. Black pigeon hawk at Santa Barbara, 
California. Condor 49:244. 



Stabler, R. M. 1969. Trichomonas gallinae as a factor in the 
decline of the peregrine falcon, pp. 435-437 In Peregrine 
falcon populations: their biology and decline. J. J. 
Hickey, ed. Univ. Wisconsin Press. 

Street, M. G. 1960. Pigeon hawk catches dragon-flies. The 
Blue Jay 18(3) :124. 

Sutton, G. M. 1935. Note on the breeding range of the black 
pigeon hawk. Auk 52:79-80. 

Swarth, H. S. 1934. Problems in the classification of north- 
western horned owls. Condor 36:38-40. 

1935. Systematic status of some northwestern 



birds. Condor 37:199-204. 



39 



Swartz, L. G. 1972. Experiments on captive breeding and 

photoperiodism in peregrines and merlins. Raptor Research 
6(2):73-87. 

Taylor, E. W. 1914. The habits of the merlin. Naturalist, 
April 1, 1914:115-120. 

Temple, S. A. 1970. Systematics and evolution of the North 
American merlins. M. S. thesis. Cornell Univ., Ithaca, 
N. Y. 62 pp. 

. 1972a. Systematics and evolution of the North 



American merlins. Auk 89:325-338. 
. 1972b. Sex and age characteristics of North 



American merlins. Bird-Banding 43:191-196. 



. 1972c. Chlorinated hydrocarbon residues and 

reproductive success in eastern North American merlins. 
Condor 74:105-106. 



Unglish, W. E. 1934. The black pigeon hawk in Santa Clara 
County, California. Condor 36:166. 

van Rossem, A. J. 1934. A second record of the black merlin in 
southern California. Condor 36:176. 

White, C. M. 1969. Breeding Alaskan and Arctic migrant popu- 
lations of the peregrine. pp. 45-51 In. J. J. Hickey, ed. 
Peregrine falcon populations: their biology and decline. 
Univ. Wisconsin Press. 

. 1974. Current problems and techniques in raptor 



management and conservation. Trans. 39th N. A. Wildlife 
and Nat. Resources Conf . :301-312 . 

Williams, R. B. and C. P. Matteson, Jr. 1947. Wyoming hawks: 
American merlins or pigeon hawks. Wyoming Wild Life 11(12): 
21-24. 



Wing, L. 1950. Dusting by the merlin ( Falco columbarius ) 
Condor 52:275. 



*.- 



Wolfe, Col. L. R. 1946. Pigeon hawk breeding in Utah. 
Condor 48:97. 



40 




Zarn, M. 1974. Osprey ( Pandion haliaetus carolinensis ) . 

Habitat Management Series for Unique or Endangered Species, 
Report No. 12. Bureau of Land Management, Denver, Colorado. 
41 pp. 



Additional references (not consulted for this report) 

Beebe, F. L. 1955. The black merlin. Falconry News and Notes 
l(6):25-29. 

Newton, I. 1973. Egg breakage and breeding failure in British 
merlins. Bird Study 20(4) :241 -244. 



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