Biological Services Program
FWS/OBS-80/01
SELECTED VERTEBRATE
ENDANGERED SPECIES OF THE
SEACOAST OF THE
UNITED STATES
• ' •
^^O wilP^"^^
Fish and Wildlife Service
U.S. Department of the Interior
FWS/OBS-80/01
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
Prepared by ^r W H 0 I
National Fish and Wildlife Laboratory /
U.S. Fish and Wildlife Service I QQCUMENT
10th and Constitution Avenue, N.W. V _-^| . cpTjQN
Washington, D.C. 20560 ^
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
1010 Cause Boulevard
Slidell, Louisiana 70458
This study was conducted in cooperation with the
Office of Endangered Species, U.S. Fish and Wildlife Service
PerfoiTned for
National Coastal Ecosystems Team
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
The Biological Services Program was established within the U.S. Fish
and Wildlife Service to supply scientific information and methodologies on
key environmental issues that impact fish and wildlife resources and their
supporting ecosystems. The mission of the program is as follovis:
• To strengthen the Fish and Wildlife Service in its role as
a primary source of information on national fish and wild-
life resources, particularly in respect to environmental
impact assessment.
• To gather, analyze, and present information that will aid
decisionmakers in the identification and resolution of
problems associated with major changes in land and water
use.
• To provide better ecological information and evaluation
for Department of the Interior development programs, such
as those relating to energy development.
Information developed by the Biological Services Program is intended
for use in the planning and decisionmaking process to prevent or minimize
the impact of development on fish and wildlife. Research activities and
technical assistance services are based on an analysis of the issues a
determination of the decisionmakers involved and their information needs,
and an evaluation of the state of the art to identify information gaps
and to determine priorities. This is a strategy that will ensure that
the products produced and disseminated are timely and useful.
Projects have been initiated in the following areas; coal extraction
and conversion; power plants; geothermal , mineral and oil shale develop-
ment; water resource analysis, including stream alterations and western
water allocation; coastal ecosystems and Outer Continental Shelf develop-
ment; and systems inventory, including National Wetland Inventory,
habitat classification and analysis, and information transfer.
The Biological Services Program consists of the Office of Biological
Services in Washington, D.C., which is responsible for overall planning and
management; National Teams, which provide the Program's central scientific
and technical expertise and arrange for contracting biological services
studies with states, universities, consulting firms, and others; Regional
Staff, who provide a link to problems at the operating level; and staff at
certain Fish and Wildlife Service research facilities, who conduct inhouse
research studies.
PREFACE
The purpose of this series of species accounts is to pro\ide resource managers and the
public with information about Federally listed endangered and/or threatened \ertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data .ue not necessarily equivalent
to critical habitat as defined in the I'.ndangered Species Act of 1973, as anicnded).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS shouki be directed to:
Office of F.ndangered Species
L'.S. Fish and Wildlife Sei"\ice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Sen,ice
iS'ASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
This report should be cited as follows:
National Fish and Wildlife Laboratory. Selected vertebrate endangered species of the sea-
coast of the United States. U.S. Fish and Wildlife Service, Biological Services Program;
FWS/OBS-80/01; March 1980.
Citation of an individual chapter should be made according to this example:
National Fish and Wildlife Laboratory. Selected vertebrate endangered species of the sea-
coast of the United States: Columbian white-tailed deer. U.S. Fish and Wildlife Service,
Biological Services Program; FWS/OBS-80/01.27; March 1980. 6 p.
ENDANGERED SPECIES ACCOUNTS
FWS/OBS-80/Ol.xx
FWS/OBS-80/Ol.xx
01 Red Wolf
02 Blunt-Nosed Leopard Lizard
03 Whooping Crane
04 Light- Footed Clapper Rail
05 San Clemente Loggerhead Shrike
06 Pine Barrens Treefrog
07 Red-Cockaded Woodpecker
08 Ivory-Billed Woodpecker
09 Ocelot
10 Attwater's Greater Prairie Chicken
11 Cape Sable Sparrow
12 Leatherback Sea Turtle
13 Green Sea Turtle
14 Texas Blind Salamander
15 Yuma Clapper Rail
16 Santa Barbara Song Sparrow
17 Eskimo Curlew
18 Southern Sea Otter
19 Morro Bay Kangaroo Rat
20 California Least Tern
21 Kirtland's Warbler
22 HawksbUl Turtle
23 Indiana Bat
24 Atlantic Salt Marsh Snake
25 Dusky Seaside Sparrow
26 Hawaiian Goose (Nene)
27 Columbian White-Tailed Deer
28 Salt Marsh Harvest Mouse
29 Puerto Rican Parrot
30 Kemp's (Atlantic) Ridley Sea Turtle
31 Bachman's Warbler
32 Florida Everglade Kite
33 Puerto Rican Whip-Poor-Will
34 Aleutian Canada Goose
35 West Indian Manatee
36 Island Night Lizard
3 7 Delmarva Peninsula Fox Squirrel
38 Houston Toad
39 American Alligator
40 Brown Pelican
41 Jaguar
42 Gray Bat
43 Okaloosa Darter
44 Eastern Cougar
45 Jaguarundi
46 Florida Panther
47 American Crocodile
48 Key Deer
49 Laysan Duck
50 Red Hills Salamander
5 1 Arctic Peregrine Falcon
52 Mississippi Sandhill Crane
53 Gray Wolf
54 Thick-Billed Parrot
55 San Clemente Sage Sparrow
56 California Clapper Rail
57 American Peregrine Falcon
58 Santa Cruz Long -Toed Salamander
Biological Services Program
FWS/OBS-80/01.1
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE RED WOLF
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubHc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data ;uc not necessarily equivalent
to critical habitat as defined in the I'-ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensiti\c Wildlife
Information System (SVVIS) developed by the U.S. Army Corps oi Engineers in coordina-
tion with the Offices of Endangered Species and Biological Sei-viccs of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to lacilitatc additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of F.ndangered Species
U.S. Fish and Wildlife Sei"vicc
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directcil to:
Information Transfer Specialist
National Coastal PLcosystems Team
U.S. Fish and Wildlife Service
iXASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Eouisi;ma 70458
FWS/OBS-80/01.1
March 1980
SELECTKD VERTEBRAl K ENDANGERED SPECIES
OF THE SEACOAS I OF I HE UNFFED STATES-
THE RED WOLF
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the (Office of Endangered Species
and the
National (Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT: U .S. F ISH 8c W 1 L.DUI FE SE R V IC E
RED WOLF
Canis rufus Audubon and Bachman
KINGDOM Animalia
CLASS Mammalia
ORDER Camivora
FAMILY Canidae
OTHER COMMON NAMES black wolf
DATE
Entered into SWIS to be determined
Updates. 20 December 1977, 14 September 1978,
25 May 1979.
LEGAL STATUS
Federal: Endangered (35 FR 16047, 3 October
1970).
States: Endangered: Delaware, Missouri, Mis-
sissippi, T-exas.
REASONS FOR CURRENT STATUS
Local predator control programs as well as
Federal, State, and local bounty hunters have de-
cimated the red wolf population. By the 1920's,
wolves were virtually extirpated east of the Missis-
sippi River, and in Kansas, Oklahoma, and most
of Texas (Nowak 1972). Extermination was ac-
complished by den hunting, steel trapping, poison
baiting, and shooting.
Predator control has had a much greater
impact on wolf populations than on the coyote
(Canis latrans) populations because bounties were
higher for wolves and people feared and hated
wolves much more than coyotes. The result was
that a few widely separated wolves remained
among many coyotes. The few remaining wolves
began to mate with coyotes and a hybrid swarm
resulted which is today being replaced by pure
coyotes (C. Carley personal communication). At
present, there are probably no remaining popula-
tions of red wolves in the wild, although some
genetically pure individuals may still exist (R.
Nowak personal communication).
Any remaining red wolves are now restricted
to coastal marshes and prairies which are being
lost to industrial and urban expansion and to agri-
cultural development (Riley and McBride 1972).
Other pressures include exploration and deve-
lopment of oil fields in Texas and Louisiana
which have made remote areas more accessible to
hunters and trappers (Pimlott and Joshn 1968).
Some red wolf habitat occurs in hunting preserves
and each year a few are killed by hunters (Riley
and McBride 1972).
The few remaining red wolves are known to
be physically weakened by diseases and parasites
(Red Wolf Recovery Team 1973). Riley and
McBride (1972) found heartworms {Dirofilaria
immitis) to be present in all specimens examined.
Infestation increases with age due to constant ex-
posure to mosquito vectors. Animals over 3 years
of age are usually heavily parasitized, reducing
their tolerance to stress (Riley and McBride
1972). Other internal parasites include hook-
worms (Ancylostoma) which often cause death in
pups (Paradiso and Nowak 1972), and in adults
leads to anemia and conditions which foster low-
level infections (Lowery 1974). Tapeworms (Tae-
nia) and spiny-headed worms of the phylum
Acanthocephala are also found in red wolves, as is
the sarcoptic mange mite [Sarcoptes scabiei)
(Riley and McBride 1972).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The red wolf is dog-like, averaging about 165
cm in total length for males and 145 cm for fe-
males. Weights of 14 specimens from Chambers
County, Texas, ranged from 19 to 28 kg (Riley
and McBride 1972).
Pelage color is variable from tawny to grayish;
muzzle is light with an area of white around the
lips extending up the sides, leaving the bridge
with a tawny to cinnamon coloration. Light areas
also occur around the eyes on many red wolves.
The Red Wolf Recovery Team has established
minimum sizes for the discrimination of red
wolves from coyotes and coyote X wolf hybrids:
male red wolves weigh between 22 and 36 kg, are
more than 134 cm long, have a hind foot length
of more than 22.8 cm, an ear length of at least 12
cm, and stand at least 68 cm high at the shoulder.
Female red wolves weigh between 19 and 31 kg,
are more than 129 cm long, have a hind foot
length of more than 22 cm, an ear length of at
least 11.4 cm, and stand at least 66 cm high at the
shoulder (McCarley and Carley 1979).
Pure coyotes are considerably smaller and
more "fox-like" than red wolves.
Photographs appear in Carley (1975), Horan
(1977), Stevens (1977), Soileau (1977), and Ne-
ville (1978).
RANGE
Red wolves are presently restricted to Cham-
bers, Jefferson, and Galveston Counties of south-
eastern Texas and adjoining Cameron and Calca-
sieu Parishes of Louisiana (Carley 1975).
They formerly occurred from central Texas
eastward to the coasts of Florida and Georgia,
and along the Mississippi River Valley north to
central Illinois and Indiana (Hall and Kelson
1959).
RANGE MAP
Shaded areas on the following page indicate
former and present distribution (C. Carley per-
sonal communication).
STATE/COUNTIES
Louisiana: Calcasieu, Cameron.
Texas: Chambers, Galveston, Jefferson.
HABITAT
Red wolves formerly inhabited dense moun-
tain and bottomland forests, as well as coastal
prairies and marshes (lUCN 1966, Pimlott and
Joslin 1968, Lowman 1975). They are now
restricted to coastal prairies and marshes with
scattered pine islands (Riley and McBride 1972),
although Nowak (1972) indicates that red wolves
move to inland forests during the spring and sum-
mer months.
The primary habitat requirement appears to
be heavy vegetative cover. Cover provided along
bayous and in overgrown fallow fields supplies
the primary resting and denning areas. Wolves
forage out into open fields and marshes using
access roads, dikes, canal levees, and cattle walk-
ways (Carley 1975).
Photographs of the habitat may be found in
Riley and McBride (1972).
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FOOD AND FORAGING BEHAVIOR
Prey includes a variety of invertebrates and
vertebrates. Young and Goldman (1944) and
Riley and McBride (1972) indicate that the marsh
rabbit {Sylvilagus aquaticus), nutria {Myocaster
coypus), and carrion are the major food items.
Other foods consist of white-tailed deer {Odo-
coileus virginianus), rodents, domestic stock,
waterfowl, fish, grasshoppers, beetles, and vege-
tation (Nowak 1972, Riley and McBride 1972).
Wolves feed primarily at night, foraging op-
portunistically for small prey alone or in small
groups (Riley and McBride 1972). They typi-
cally travel in family groups.
SHELTER REQUIREMENTS
See nesting or bedding.
NESTING OR BEDDING
Historically, red wolves denned in hollow
tree trunks, along stream banks, and old holes of
other animals (Nowak 1972, Riley and McBride
1972, Lowman 1975). The dens were usually ob-
scured by brush and vegetation, but afforded the
occupants a view of the surrounding terrain.
Water tables are probably too high for ground-
nesting in the coastal marshes where wolves still
occur, and nesting in tall vegetation has been re-
ported in these areas (C. Carley personal com-
munication).
RITUAL REQUIREMENTS
The role of howling in the social behavior is
not fully understood. The voice of the red wolf is
described by Riley and McBride (1972) and
McCarley (1978).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Nowak (1972) reported that the territory is
generally circular with a diameter attaining 64 km.
Carley (1975), using radio tracking, found that
males range over an area of approximately 116
km^ and females over a somewhat smaller area.
Lowman (1975) reports that the home range of
adult red wolves varies from 65 to 130 km^ .
POPULATION NUMBERS AND TRENDS
Numbers are reduced or extirpated over most
of the range (Pimlott and Joslin 1968). McCarley
(1962) recorded species extant in only a few
places in western and southern Louisiana.
R. M. Nowak (personal communication) sums
up the trend: Steady decline since the coming of
the white man; accelerated since large-scale hy-
bridization began about 1920; pure populations
apparently survived until about 1920. The Recov-
ery Team (RWRT personal communication) esti-
mates probably fewer than 50 pure red wolves in
the wild.
McCarley and Carley (1979) assert that the
red wolf will soon be extinct in the wild, and C.
Carley (personal communication) states that the
species will probably be extinct in the wdld by
1981.
REPRODUCTION
Mating occurs in January and February with
pups born in March, April, and May (Nowak 1972,
Riley and McBride 1972, Lowman 1975). Litters
range from 3 to 12 with an average of 6 or 7
(Nowak 1972,Lowery 1974).
Both parents participate in rearing the young
(Riley and McBride 1972, Lowman 1975), and
yearlings are often found in the vicinity of the
dens, and may help in rearing young.
After the young reach 6 weeks of age, they
spend considerable time away from the den in
well-covered beds (Riley and McBride 1972, Low-
man 1975). Most die before the age of 6 months,
with hookworms reported to be the major cause.
Full size is attained in 1 year; sexual maturity by
3 years (Nowak 1972, Lowman 1975). Life ex-
pectancy is about 5 years in the wild, and greater
in captivity.
Red wolves are more sociable than coyotes
but less so than gray wolves {Cams lupus). It is
not unusual to find three or more wolves tra-
veling throughout the range as a group (Riley and
McBride 1972).
MANAGEMENT AND CONSERVATION
Depletion of the red wolf was first reported in
1962. The U.S. Fish and Wildlife Service confirmed
this in 1968 and further determined that two sub-
species, Canis rufus rufus and C. r. gregoryi, pro-
bably existed, but that C. r. floridanus (inhabiting
the eastern part of the range) was extinct. More
recently, C. r. rufus (western part of range) was
deemed extinct also (Carley 1975).
Efforts by the U.S. Fish and Wildlife Service
in 1969 involved removing depredating animals
from private lands. This served two purposes: (1)
established rapport with owners of remaining red
wolf habitat; and (2) facilitated removal of hy-
brids and coyotes while relocating red wolves
(Carley 1975).
In 1973, biologists were assigned to implement
the Red Wolf Recovery Plan. Removal of depre-
dating animals was continued, but red wolves
captured were treated for various infirmities and
released (often radio -tracked) or transferred
to captive breeding centers. An effort was ini-
tiated to maintain a buffer zone between red
wolves and coyotes. This was determined to be
impossible because of the difficulty of main-
taining such an extensive buffer zone and hybrids
were already present (Carley 1975).
Carley (1975) asserts that red wolves can be
preserved only by relocation. Exclusion of coy-
otes and hybrids from the remaining range is an
insurmountable problem (Carley 1975).
Relocation experiments were initiated in late
1976 on Bulls Island, South Carolina. Although
there have been numerous problems with the pro-
gram, a pair was successfully relocated in January
1978. This island was chosen for a number of
technical reasons, but with no intent to start a
viable population (Department of the Interior
1972, 1977a, 1977b; R. M. Nowak personal
communication). The experiment was successfully
completed in November 1978, when the pair was
recaptured and returned to the captive breeding
program.
In 1977, there were 29 recognized pure adults
and 13 young in a breeding pool in Point Defi-
ance Zoo in Tacoma, Washington (R. M. Nowak
personal communication).
The Recovery Team (RWRT 1973) provided a
step-down plan for restoring the red wolf to non-
endangered status. The four major objectives are
"(1) to restore surviving red wolf subspecies in
their present ranges to desirable population levels;
(2) to maintain an adequate captive red wolf gene
pool; (3) to reestablish surviving red wolf subspe-
cies in additional locations within their historic
range; and (4) to determine the location and
abundance of each surviving red wolf subspecies
population." Specific goals include stopping of
unauthorized killing by man, developing a posi-
tive public attitude, preventing genetic contami-
nation, developing landowner tolerance, improv-
ing and protecting red wolf habitat, controlling
debilitating pathogens and parasites, and control-
ling detrimental effects of environmental contam-
inants (RWRT 1973).
A new Recovery Team was formulated in
1978.
AUTHORITIES
George R. Abraham (Recovery Team)
State Supervisor
U.S. Fish and Wildlife Service
271-273 Agriculture Center
Louisiana State University
Baton Rouge, LA 70803
Curtis Carley
Wildlife Biologist
U.S. Fish and Wildlife Service
Albuquerque, NM 87103
Russell Clapper
Refuge Manager
Anahuac National Wildlife Refuge
AnahuacTX 77514
George G arris
Refuge Manager
Cape Romain National Wildlife Refuge
Awendaw, SC 29429
Joe L. Herring (Recovery Team)
Chief, Division of Game
Louisiana Wildlife and Fisheries Commission
Baton Rouge, LA 70804
Howard McCarley
Austin Universtiy
Department of Biology
Sherman, TX 75090
Ronald M. Nowak
Office of Endangered Species
U.S. Fish and Wildlife Service
Washington, D.C. 20240
Mary Anne Neville (Recovery Team)
Protected Species Coordinator
Georgia Department of Natural Resources
270 Washington Street
Atlanta, GA 30339
Dave Peterson (Recovery Team Leader)
U.S. Fish and Wildlife Service
900 San Marco Boulevard
Jacksonville, FL 32207
PREPARER'S COMMENTS
Investigators (McCarley 1962, Lawrence and
Bossert 1967, Paradiso 1968, Pimlott and Joslin
1968, Nowak 1970, Paradiso and Nowak 1971)
have raised questions concerning the taxonomic
status of the red wolf. Paradiso and Nowak
(1971) and Gipson et al. (1974), using a multi-
variate analysis of skull morphology, concluded
that the red wolf was indeed a valid species.
There are numerous gaps in the knowledge of
the biology of this species which must be attribu-
ted to its depleted numbers, secretive habits, and
limited accessibility.
LITERATURE CITED/SELECTED
REFERENCES
Carley, C. J. 1975. Activities and findings of the
red wolf recovery program from late 1973 to
1 July 1975. U.S. Fish Wildl. Serv. 211 pp.
Freeman, R. C. 1976. Coyote X dog hybridiza-
tion and red wolf influence in the wild Cants
of Oklahoma. M.S. Thesis. Oklahoma State
Univ., Stillwater. 62 pp.
Gipson, P. S., J. A. Sealander, and J. E. Dunn.
1974. The taxonomic status of wild Cants in
Arkansas. Syst. Zool. 23(1):1-11.
Goldman, E. A. 1937. The wolves of North Ame-
rica. J. Mammal. 18(l):37-45.
Hall, E. R., and K. R. Kelson. 1959. The mam-
mals of North America. Vol. 2. Ronald Press,
New York. 547-1,083 pp.
Harper, F. 1927. Mammals of the Okefenokee
Swamp region of Georgia. Boston Soc. Natur.
Hist.Proc. 38(7):191-396.
Horan, J. 1977. Return of the red wolf. Defen-
ders 52(1):16-19;
lUCN (International Union for the Conservation
of Nature and Natural Resources). 1966. Red
data book. Morges, Switzerland.
Lawrence, B., and W. H. Bossert. 1967. Multiple
character analysis of Canis lupus, latrans, and
familiaris, with a discussion of the relation-
ships of Canis niger. Am. Zool. 7(2):223-232.
Lowery, G. H., Jr. 1974. The mammals of Louisi-
ana and its adjacent waters. Louisiana State
Univ. Press, Baton Rouge. 564 pp.
Lowman, G. E. 1975. A survey of endangered,
threatened, rare, status undetermined, peri-
pheral, and unique mammals of the south-
eastern National Forests and grasslands.
USDA For. Serv. Contract 38-2601.
McCarley, H. 1962. The taxonomic status of wild
Canis (Canidae) in the south-central United
States. Southwest. Natur. 7(3-4); 227-235.
. 1978. Vocalizations of red wolves {Canis
rufus).]. Mammal. 59(l):27-35.
McCarley, H., and C. J. Carley. 1979. Recent
changes in distribution and status of wild red
wolves (Canis rufus) . U.S. Fish and Wildl. Serv.
Endangered Species Report 4. 8 pp.
Mech, L. D. 1974. Canis lupus. Mammalian spe-
cies 37. Am. Soc. Mammalogists. 6 pp.
Neville, M. A. T. 1977. In South Carolina, another
transplant runs into trouble. Natl. Wildl.
15(5):10-11.
. 1978. Counterfeit wolves and lonely
islands. Sierra Club Bull. 63(2):22-23.
Nowak, R. M. 1967. The red wolf in Louisiana.
Defenders Wildl. News 42(l):60-70.
. 1972. The mysterious wolf of the south.
Natur. Hist. 81:50-53, 74-77.
. 1974. Red wolf, our most endangered
mammal. Natl. Parks Conserv. Mag. Aug:9-12.
-. 1975. The cosmopolitaJi wolf. Natl.
Rifle Assoc. Coserv. Yearb. 76-83.
Ogilvie, P. W. 1970. Interim report on the red
wolf in the United States. Int. Zoo. Yearb.
(10):122-124.
Paradiso, J. L. 1965. Recent records of red wolves
from the Gulf Coast of Texas. Southwest Nat.
10(4):318-319.
. 1968. Canids recently collected in east
Texas, with comments on the taxonomy of
the red wolf. Am. Midi. Natur. 80(2):529-534.
Paradiso, J. L., and R. M. Nowak. 1971. A report
on the taxonomic status and distribution of
the red wolf. U.S. Fish Wildl. Serv. Spec. Sci.
Rep. Wildl. 145. 36 pp.
. 1972. Canis rufus. Mammalian Species
22. Am. Soc. Mammalogists. 4 pp.
Pimlott, D. H., and P. W. Joslin. 1968. The status
and distribution of the red wolf. Trans. N.
Am. Wildl. Natur. Resour. Conf. 33:373-384.
RWRT (Red Wolf Recovery Team). 1973. Red
wolf recovery plan. Draft. U.S. Dep. Inter.
Bur. Sport Fish. Wildl., Office Endangered
Species and International Activities. 32 pp.
Riley, G. A., and R. T. McBride. 1972. A survey
of the red wolf {Canis rufus). U.S. Fish Wildl.
Serv. Spec. Sci. Rep. Wildl. 162. 15 pp.
Shaw, J. A., and P. A. Jorden. 1977. The wolf
that lost its genes. Natur. Hist. 86(10):80-88.
Soileau, C. 1977. Epitaph for a canine. Louisiana
Conservationist 29(7-8): 14-17.
Stevens, J. T. 1977. Almost gone. Texas Parks
and Wildlife 35(5):2-7.
U.S. Department of the Interior. 1973. Threa-
tened wildlife of the United States. Compiled
by Office of Endangered Species and Interna-
tional Activities. Bur. Sport Fish. Wildl.
Resour. Publ. 114. U.S. Gov. Printing Office,
Washington, D.C. 239 pp.
U.S. Department of the Interior. 1976. Red wolf
released on Bulls Island, Cape Romain Na-
tional Wildlife Refuge, South Carolina, and
recaptured. U.S. Fish Wildl. Serv. News Re-
lease. 27 December 1976. 3 pp.
U.S. Department of the Interior. 1977a. New red
wolf pair to arrive at Cape Romain National
Wildlife Refuge, South Carolina. U.S. Fish
Wildl. Serv. News Release. 30 June 1977. 3 pp.
U.S. Department of the Interior. 1977b. Substi-
tute pair of red wolves shipped to Cape Ro-
maine National Wildlife Refuge, South Caro-
lina. U.S. Fish Wildl. Serv. News Release,
IJuly 1977. 1 pp.
Young, S. P., and E. A. Goldman. 1944. The
wolves of North America. 2 parts. Am. Wildl.
Inst., Washington, D.C. 630 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, Fl 32601
Biological Services Program
FWS/OBS-80/01.2
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
BLUNT NOSED LEOPARD LIZARD
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this scries of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or tlireatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data are not necessarily equivalent
to critical habitat as defined in tiie I'.ndangered Species Act of 197.S, as amended).
This scries of accoimts is intended to complement the compuieri/ed Sensiti\e Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of I-^ndangercd Species and Biological Sei-viccs of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should he directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directcil to:
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Senice .
.\ASA-Slidell Computer Coinplc;\
1010 Cause Blvd.
Slidell, Louisiana 70458
FWS/OBS-80/01.2
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
BLUNT NOSED LEOPARD LIZARD
A Cooperative Effort
by the
National Fish and WildHfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
BLUNT-NOSED LEOPARD LIZARD
Crotophytus (Gambelia) silus
KINGDOM Animalia
CLASS Reptilia
ORDER Sauria
FAMILY Iguanidae
OTHER COMMON
NAMES Blunt-nose leopard lizard
DATE
Entered into SWIS To be determined
Updates 9 November 1979
LEGAL STATUS
Federal Endangered (32 FR 4001, 11 March
1967).
States Endangered: California (21 May
1977)
REASONS FOR CURRENT STATUS
This lizard has become increasingly difficult
to find throughout most of its range because of
agricultural development and urbanization. The
few remaining areas of prime habitat, along the
western side of the San Joaquin Valley, are under-
going rapid development following the recent
completion of a major new aqueduct (the Califor-
nia Aqueduct) (Erode 1978, Bury 1972, Monta-
nucci 1965, Snow 1972, U.S. Dep. Int. 1973).
Off-road vehicle recreation is damaging the re-
maining habitat in some nonagricultural areas
(Erode 1978). Overgrazing and agricultural pest
control may be detrimental to C. (G.) silus popu-
lations (Anon 1978).
PRIORITY INDEX
Not assigned.
DESCRIPTION
A robust lizard with a long, slender, cylindri-
cal tail, C. (G.) silus has a large head with a short,
blunt snout; adults measure 89 to 127 mm in
snout-to-vent length. The dorsal ground color is
gray or brown, and the dorsum has broad, distinct
whitish bands interspaced with dusky spots. The
throat has dark gray blotches. Undersides of the
tail and thighs are white to yellowish; during the
breeding season, the males are salmon or rust
color ventrally or all over the body except the
head (Erode 1978; Bury 1972; Montanucci 1965,
1967, 1970; Snow 1972; Tollestrup 1979). Fe-
males have a breeding color consisting of a single,
or, occasionally, a double row of red-orange spots
on the flanks and sides of the face and a continu-
ous wash of the same color on the undersurface
of the tail and thighs. (Tollestrup 1979). Monta-
nucci (1965, 1967, 1970) describes distinction in
color, pattern, and size between valley floor, foot-
hill, and ecotonal hybrid populations, but Tolles-
trup(1979) did not find such differences.
RANGE
The species occurs only in California. It was
once found throughout the San Joaquin valley
and adjacent foothills from about latitude 37°3l'
north southward into San Luis Obispo County.
It now occurs at scattered locations in the valley,
east to the Sierra foothills, south to the Tehachapi
Mountains, and along the east slopes of the Coast
Range foothills, including the Carrizo Plain and
lower Cuyama Valley (Erode 1978; Montanucci
1965, 1970; Snow 1972), at elevations below
600 m.
RANGE MAP
Known distribution is shown on the accom-
panying map (Anon 1978).
STATES/COUNTIES
California Fresno, Kern, Kings, Madera, Mer-
ced, San Eenito, San Luis Obispo,
Santa Barbara, Tulare, Ventura.
HABITAT
It prefers open habitat with scattered low
bushes, occurring on sparsely vegetated plains,
alkali flats, low foothills, canyon floors, large
washes, and arroyos; it is usually found on sandy
substrates and sometimes on coarse, gravelly soil
and hardpan (Montanucci 1965, 1970).
FOOD AND FORAGING BEHAVIOR
This lizard is an active predator and an oppor-
tunistic feeder, subsisting primarily on large
insects and small lizards. Montanucci (1965) re-
ports seasonal and regional variation in diet,
depending on the availability of insect and lizard
prey. Insects taken include locusts (Trimeratropis
calif ornica), grasshoppers (Melanoplus sp.), ci-
cadas (Okanagana triangulata, O. pallidula),
crickets {Acheta assimilis), and a wide variety of
orthoptera, lepidoptera, and coleoptera species.
Lizards eaten include Uta stansburiana, Phryno-
soma coronatum, small Sceloporus magister, and
Cnemidophorus tigris, as well as juveniles of its
own species. Small amounts of vegetable matter
are also taken (Montanucci 1965, 1967). Tolles-
trup (1979) found that C. (G.) silus feeds princi-
pally on grasshoppers, crickets, beetles, cicadas,
flies, and spiders; no vertebrate prey was found in
this large sample of stomachs (N = 142).
Foraging habits vary relative to habitat and
available prey (Montanucci 1967). Lizards will
leap into the air or into shrubbery in pursuit of in-
sects. Stalking and pursuit are employed in
hunting both insect and lizard prey. Montanucci
(1965) reports that C. (G.) silus will dig up lizards
{Uta stansburiana) that have taken refuge in holes
or loose sand.
SHELTER REQUIREMENTS
Montanucci (1965) notes that population
densities correlate to some extent with abundance
of mammal burrows. Abandoned or occupied
burrows of kangaroo rats {Dipodomys sp) and
abandoned squirrel burrows [Spermophilus sp.),
gopher burrows {Thomomys bottae), and badger
dens {Taxidea taxus) are used for escape cover
and permanent shelter. In areas where mammal
burrows are scarce, adult C. (G.) silus construct
shallow, simple chambered tunnels under exposed
rocks or earthen banks. Immature lizards use
rock piles, trash piles, brush, etc. for temporary
cover (Montanucci 1965).
. / Stanislaus Co^>j- n^ Moriposa Co.
'Merced Co. ^\ .' — •
'MERCEUX^/ I
LOCATOR MAP
Estimated Original
Range of BNLL
0
0
Santa Barbara Co. • ~~ 'T
I \
' Ventura Co. \
10 20 30 40 Miles | \
16 32 48 64 Kilometers I \
Distribution of the blunt-nosed leopard lizard.
NESTING OR BEDDING
POPULATION NUMBERS AND TRENDS
Nesting habits of C. (G.) silus are almost
identical to those of Gambelia wislizenii (see
Parker and Pianka 1976). Females lay eggs in
June and July, in burrows at about 50 cm depth.
A chamber is excavated or enlarged, entrance tun-
nels sealed, and eggs are laid one at a time and
lightly covered with sand or soil. The exit tunnel
is then plugged from without.
RITUAL REQUIREMENTS
Males establish and defend home burrows, but
appear to have overlapping home ranges (Monta-
nucci 1965). The degree to which territoriality is
exhibited is probably correlated with abundance
and availability of food and cover; where mammal
burrows are abundant, individual home ranges
and home burrows are difficult to detect (Monta-
nucci 1965). Recently, Tollestrup (1979) found
that males defend home ranges, not just burrows.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Montanucci (1965) reported that several fac-
tors limit the distribution of C. (G.) silus: ;
1. Agricultural practices
2. Flooding
3. Chaparral
4. Steep or extensive rocky areas.
Tall grass may hamper foraging and rapid loco-
motion, and may account for the relative scarcity
of C. (G.) silus on the east side of the San Joa-
quin Valley (Montanucci 1965). K. Tollestrup
(pers. comm.) reported that tall grass improves
foraging because insect abundance increases and it
gives the lizards cover for escape from predators.
Scarcity of C. (G.) silus in tall grass areas on the
east side of the San Joaquin Valley is probably
due to the fact that human settlements in this
region and farming and grazing of the land date
back many years. Also, in this area, it is common
practice to plow and burn rangelands in the fall,
which decreases or eliminates populations of C.
(G.) silus. Entomologists working in Kern County
reported finding leopard lizards killed by insecti-
cides (DDT and Malathion) (Montanucci 1965).
J. M. Sheppard {in Snow 1972) estimates a
density of 300 to 400 Uzards per square mile of
optimum habitat near Maricopa, Kern County.
Since it is unlikely that all habitats occupied are
optimum, he considered the mean density of
lizards to be 100/mi' (38.6/km' or 0.4/ha).
Tollestrup (1979) estimated that there were 3.5
Uzards per ha at Pixley National Wildlife Refuge,
which is the highest known density. Populations
of C. (G.) silus are low (0.5 to 1 per acre or about
1 to 2.5 per ha) under optimum conditions
(Anon. 1978).
REPRODUCTION
The reproductive cycle varies slightly with
environmental conditions. Males will often mate
with several females. Mating occurs from late
April to early June; clutches of 2 to 5 eggs are
laid in June or July; some females may have two
clutches per year (Montanucci 1965). Incubation
time is estimated at about 57 days. Young (42 to
47 mm in snout-to-vent length) appear from July
30 through September. Sexual maturity in both
sexes occurs between 9 and 18 months (Mont-
anucci 1965, 1967; Tollestrup 1979).
MANAGEMENT AND CONSERVATION
Habitat for the blunt-nosed leopard hzard is
rapidly being lost throughout its range. In the
last 100 years, the natural wildlands of the San
Joaquin Valley have decreased from 3 million ha
to about 200,000 ha (Dickl977). Suitable habitat
(salt-brush desert scrub) on the west side of the
valley is now being developed for agriculture,
since completion of the California Aqueduct.
Also, off-road vehicles have denuded parts of this
region.
R. R. Montanucci ( in Snow 1972) recom-
mended the establishment of a national grassland
preserve in the southwestern part of the San Joa-
quin Valley as the most rapid means of assuring
partial protection of this species. Erode (1978)
recommended protective management programs
on the remnants of suitable lizard habitat on
public lands (the Naval Petroleum Reserve near
Taft, the Kern and Pixley National Wildlife Re-
fuges, Los Padres National Forest, and National
Resources Land). These lands should be managed
to maintain and enhance habitat suitable to the
blunt-nosed leopard lizard. Both refuges recognize
the importance of their lands to the survival of
the lizard; the Pixley National Wildlife Refuge
was used as a study area on the ecology of this
species (Tollestrup 1979). Also near Pixley, the
U.S. Forest Service will manage a tract of grass-
land comprising about 325 ha for this lizard
(Erode 1978). Essential habitat on private land
could be protected by agreement with land-
owners.
The California Department of Fish and Game
has conducted field surveys and initiated coopera-
tive studies with the U.S. Forest Service, Universi-
ty of California at Berkeley, and several colleges
to further determine the distribution and status
of this lizard. A Blunt-Nosed Leopard Lizard Re-
covery Team has been formed to provide needed
coordination of effort to protect this species
(Anon. 1978, Erode 1978).
AUTHORITIES
John M. Erode (Recovery Team Leader)
California Department of Fish and Game
1701 Nimbus Road
Rancho Cordova, California 95670
Richard R. Montanucci
Department of Zoology
Clemson University
Clemson, South Carolina 29631
Kristine Tollestrup
Department of Biology
University of Chicago
1103 E. 57th Street
Chicago, Illinois 60637
PREPARER'S COMMENTS
Recent taxonomic changes for this lizard
warrant explanation to prevent confusion and
promote consistent use of currently accepted
nomenclature. The relationshiops and systematic
status of the lineages of crotaphytaform lizards
(leopard and collared lizards) of North America
have been undergoing revision by herpetologists
as new data and techniques become available.
Montanucci. (1970) presented evidence of genetic
and ecologic differentiation between the leopard
lizards of the San Joaquin Valley and those of
more southern and eastern populations. This evi-
dence supports the recognition of the blunt-nosed
leopard lizard as a distinct species (Gambelia si-
lus), rather than a subspecies of the longnose leo-
pard lizard (G. wislizenii).
Montanucci et al. (1975) present biochemical
evidence indicating that leopard lizards are suffici-
ently different from the closely related collared
lizards (genus Crotaphytus) to be considered a
separate genus [Gambelia).
LITERATURE CITED/SELECTED
REFERENCES
Anonymous. 1978. Blunt-nosed lizard recovery
plan. Part I. (draft). 12 pp.
Erode, J. 1978. Blunt-nosed leopard lizard {Crota-
phytus silus). Pages 24-25 in At the Cross-
roads: a report on California's endangered and
rare fish and wildlife. Calif. Dep. Fish Game
Biannual Rep. 103 pp.
Bury, R. B. 1972. Status report on California's
threatened amphibians and reptiles. Calif.
Dep. Fish Game, Inland Fisher. Rep. 72-2. 31
pp.
Dick, D. 1977. Habitat disappearing for the leo-
pard lizard. Outdoor Calif. 38(6):37-38.
Montanucci, R. R. 1965. Observations on the
San Joaquin leopard lizard Crotaphytus
wislizenii silus Stejneger. Herpetologica
21(4):270-283.
1967, Further studies on leopard lizards,
Crotaphytus wislizenii. Herpetologica 23(2):
119-126.
1970. Analysis of hybridization between
Crotaphytus wislizenii and Crotaphytus silus
(Sauria: Iguanidae) in California. Copeia 1970
(1):104-123.
Montanucci, R. R., R. W. Axtell, and H. C.
Dessaur. 1975. Evolutionary divergence among
collard lizards (Cryophytus), with comments
on the status of Gambelia). Herpetologica
31(3):336-347.
Parker, W. W., and E. R. Pianka. 1976. Ecological
observations on the leopard lizard (Crotophy-
tus wislizenii) in different parts of its range. Berkeley.
Herpetologica 32(1):95-114.
U.S. Department of the Interior. 1973. Threatened
Snow, C. 1972. Blunt nosed leopard lizard: Cro- wildlife of the United States. Bur. Sport Fish.
taphytus situs. Bur. Land Manage. Tech. Note Wildl. Res. Publ. 114. 289 pp.
6601. 13 pp.
PREPARED/UPDATED BY
Tollestrup, K. 1979. The ecology, social structure,
and foraging behavior of two closely related National Fish and Wildllife Laboratory,
species of leopard lizards, Gambelia silus and 1300 Blue Spruce Drive
Gambelia wislizenii. Ph. D. Thesis, Univ. Calif. Fort Collins, Colorado 80524
Biological Services Program
FWS/OBS-80/01.3
March 1980
Selected Vertebrate
Endangered Species
Of the Seacoast of the
United States-
THE WHOOPING CRANE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACi:
The purpose of this scries of species accounts is to provide resr>urcc managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life histt)ry, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Fndangered Spci ies Act of ]97'>, as amended).
This series of accoinits is intended to complement the compuieri/.cd Senbiti\c Wildlife
Information System (SWIS) developed by the U.S. Army C:orps of Kngineers in coordina-
tion with the Offices of f.ndangered Species and Biological Sei-vices of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should he direc ted to:
Office of F.ndangered Species
U.S. Fish and Wildlife Seivice
Interior Building
Washington, D.C. 202 10
Suggestions or questions regarding this report should be dirt
Information Transfer Specialist
National Coastal P^cosystems I cam
U.S. Fish and Wildlife Service
NASA-Slidell ConipLitcr Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
d u
FWS/OBS-80/01.3
March 1980
SELECl i:i) VI.Rl KBRATK ENDANGERED SPECIES
OF IHE SEACOAS r OF 1 HE UNITED S lATES-
THE WHOOPING CRANE
A Cooperative Effort
by the
National Fish and WildHfe Laboratory,
the Office of Endangered Species
and the
National (Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald \V. VVoodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fisii and Wildlife Service
U.S. Department of the Interior
1^^
"a '^ ^-^ '
CREDIT: W. F. KUBICHEK
WHOOPING CRANE
Grus americana Linnaeus
KINGDOM Animalia
CLASS Aves
ORDER Gruiformes
FAMILY Gruidae
OTHER COMMON
NAMES Great white crane
DATE
Entered into SWIS to be determined
Updates 18 July 1978, 25 September 1978.
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 March
1967; 35 FR 8491, 2 June 1970).
States: Endangered: Colorado, Idaho, Mon-
tana, Nebraska, New Mexico, Texas.
REASONS FOR CURRENT STATUS
There are only 75 whooping cranes in the
wild and 27 in captivity (WCRT 1977). They
once nested widely over northern North America;
nesting is now restricted to a small area in the
southern MacKenzie District and northeastern
Alberta, Canada. Breeding habitat and wintering
habitat have been reduced as a result of drainage,
agriculture, the Gulf Intracoastal Waterway, and
human settlements. Whooping cranes avoid areas
of human disturbance even if the habitat is other-
wise suitable (R. Erickson personal communica-
tion).
Cranes were hunted before legal protection
was instituted. Losses were particularly acute on
the Gulf Coast wintering grounds (Alleh 1952).
Accidental shooting probably still occurs when
whooping cranes are mistaken for snow geese
(Chen hyperborea) and sandhill cranes [Grus
canadensis), both legal game in some areas
(WCRT 1977).
Natural factors such as large nesting territories
and a very low reproductive rate of one chick per
year make recovery slow. The biannual migration
covering over 3,800 km usually accounts for some
mortahty (Allen 1962). Stormy weather during
migration poses a serious threat to the few
remaining whooping cranes. The nonmigratory
Louisiana population was reduced 50% during a
hurricane in 1940 and never recovered (Allen
1952).
There remains the possibility of hazardous ma-
terials being spilled on the wintering grounds
along the heavily industrialized Texas Gulf Coast.
PRIORITY INDEX
Not assigned.
DESCRIPTION
Whooping cranes are the tallest birds in North
America. Males are almost 1.5 m high, with a
wingspan of 2.4 m and an average weight of 7 kg.
Females are slightly smaller, with an average
weight of 6 kg. Weight can vary 25% seasonally,
with the maximum reached in winter.
The adult plumage is velvety white except for
the head and portions of the wings and feet.
Forehead, crown, anterior part of occiput, lores,
and cheeks are bare except for sparse black hair-
like feathers. The skin in this area is warty or
granulated, and a reddish color. The bill is a dull
pink, proximally changing to dull brownish in the
middle and dirty yellow on the tip. The alulae,
10 primaries, and 10 greater upper coverts are a
slatey black. The feet are black with flesh-colored
pads.
Juvenal plumage is a variable mixture of white
and buff or brown. The adult plumage is attained
in about 1 year. The sexes are similar.
Vocalizations range from a purring call near
young to a low groaning call when approached by
humans, to a shrill, bugle-hke 'kerloo! ker-lee-oo.'
The latter is known as a unison call and is a defi-
nite indicator of pair bonding (R. Erickson per-
sonal communication).
Photographs appear in Allen (1952), McNulty
(1966), Novakowski (1966), Walkinshaw (1973),
Lowery (1974), Kuyt (1976a, 1976b), Muir
(1976), numerous drawings appear in Allen
(1952).
RANGE
The natural population now nests only in
Wood Buffalo National Park, southern MacKenzie
District, and northeastern Alberta, Canada. It
winters on the Gulf Coast of Texas, in Aransas
National Wildlife Refuge and adjacent peninsulas
and islands. Its migration route is a nearly
straight-line path through east central Texas, cen-
tral Oklahoma, central Kansas and Nebraska,
western and central North and South Dakota,
northeastern Montana, southeastern Saskatche-
wan, and northeastern Alberta.
An experimentally transplanted disjunct
population will hopefully nest at Grays Lake Na-
tional Wildlife Refuge, Idaho, and winter in the
Rio Grande Valley of New Mexico. The migration
route passes through southeastern Idaho, north-
eastern Utah, southwestern Wyoming, western
Colorado, and north-central New Mexico (WCRT
1977).
The crane formerly bred from its present
range south through the prairie provinces and
northern prairie states, including Illinois, Iowa,
Minnesota, and North Dakota, and wintered from
the Carolina coast to central Mexico. A geo-
graphically isolated breeding population, believed
to be nonmigratory (now extirpated), formerly
lived in southestern Louisiana.
RANGE MAP
Wood Buffalo— Aransas population, Grays
Lake Experimental Population and Critical Habi-
tat from WCRT (1977) and Federal Register (43
FR 20938, 15 May 1978).
Wood Buffalo
National Park
N
I.
340
scale in km
WOOD BUFFALO- ARANSAS POP.
▲ NESTING LOCATION
• MAJOR MIGRATION STOPS
D WINTER LOCATION
O CRITICAL HABITAT
NWR NATIONAL WILDLIFE REFUGE
SWA STATE WILDLIFE AREA
WMD WATERFOWL MANAGEMENT DISTRICT
i : APPROXIMATE MIGRATION ROUTE
Pocasse NWR
IOWA
MO.
Kirwin NWR
Cheyenne Bottoms SWA
Quivirg NWR
-^alt\Plains NWR
Wichita NWR
Aransas NWR
Map depicts the nesting and wintering locations of the Wood Buffalo-Aransas populations of the
whooping crane and its migration route. Critical habitat is designated.
MONTANA
N
k
A Melville
IDAHO
Grays^
Lake<^
NWR
0
250
scale in km
A Cora
WYOMING
▲ Lonetree
Jensen %
Ouray NWR •
GRAYS LAKE EXPERIMENTAL POP.
▲ SUMMER LOCATIONS
• MAJOR MIGRATION STOPS
D WINTER LOCATION
O CRITICAL HABITAT
NWR NATIONAL WILDLIFE REFUGE
SWA STATE WILDLIFE AREA
COLORADO
% Ridgeway
Monte Vista(«
Aionnosa NWR
NEW /MEXICO
D Bernardo SWA
S) Bosque Del
Apache NWR
Map depicts summer and winter locations and migration stops for the Grays Lake experimental
population of the whooping crane. Critical habitat is designated.
STATES/COUNTIES
Wood Buffalo-Aransas Population
Winter
Texas Aransas, Calhoun.
Known Migration Stops
Kansas Barton, Reno, Stafford.
Montana Roosevelt, Sheridan
Nebraska Adams, Blaine, Brown, Buffa-
lo, Custer, Dawson, Franklin,
Gosper, Hall, Harlan, Kearney,
Keya Paha, Loup, Phelps,
Rock, Sherman, Thomas,
Valley, Webster.
North
Dakota Burke, Burleigh, Divide,
Dunn, Emmons, McKenzie,
McLean, Morton, Mountrail,
Penville, Sioux, Ward, Wil-
liams.
Oklahoma Alfalfa, Comanche.
South
Dakota Campbell, Corson, Dewey,
Haakon, Hughes, Potter, Stan-
ley, Sulley, Walworth, Ziebach
Grays Lake Experimental Population
Winter
New
Mexico Luna, Socorro, Valencia.
Mexico Chihuahua.
Known Migration Stops
Colorado Alamosa, Canejos, Ouray, Rio
Grand.
Utah Uintah.
Summer
Idaho Bonneville, Caribou.
Montana Sweet Grass.
Utah Uintah.
Wyoming Sublette, Uintah.
HABITAT
The nesting grounds are marshy areas with
many potholes, which are generally shallow and
have soft, muddy bottoms. The pH is usually
between 7.6 and 8.3. Potholes are separated by
land areas with black spruce [Picea mariana),
tamarack (Larix laricitia), and willows (notably
Salix Candida), forming the canopy. The under-
story includes dwarf birch {Betula glandulosa).
bearberry (Arctostaphylos uva-ursi), and several
species of lichen. The ground is deeply carpeted
with sphagnum moss. Emergent vegetation in the
potholes used by cranes is mainly bulrush (Scir-
pus validus) (Novakowski 1966).
The Wood Buffalo-Aransas population winters
on salt-marsh flats in coastal lagoons on the Gulf
of Mexico. Allen (1952) differentiated three
types used by cranes: (1) permanent pond, lake,
ditch, or bayou open to bay tides at all seasons;
(2) semipermanent ponds connected by narrow
bayous to permanent water areas; (3) ephemeral
ponds that offer superior feeding places. Typical
vegetation includes salt grass {Distichlis spicata),
saltwort {Batis maritima), glasswort {Salicornia
sp.), sea ox-eye [Borrichia frutescens), needle
cordgrass (Spartina spartinae), and saltmarsh
cordgrass [Spartina alterniflora) (Allen 1952).
Cranes will occasionally forage into the oak
{Quercus) brush region (Allen 1952), especially
after a fire.
The major requirement for whooping crane
habitat selection, particularly during migration,
is an open expanse for nightly roosting. The
birds typically use sand or gravel bars in rivers and
lakes (43 FR 36588, 17 August 1989).
Historic habitat includes interior tall-grass
prairies, sea-rim and brackish marshes, and
higher interior grasslands (Allen 1952). Some of
these habitat types are still used during migration.
FOOD AND FORAGING BEHAVIOR
Cranes are omnivorous, taking vegetation, in-
sects, amphibians, reptiles, and small mammals
(Bent 1926). The whooper is more aquatic than
the sandhill crane and has a greater preference for
animal foods (Allen 1952). Prey species are listed
in Allen (1952) and Novakowski ( 1966).
Summer foods were studied by Novakowski
(1966) and were determined to be primarily large
nymphal or larval forms of insects, and crustace-
ans. Terrestrial foods such as berries were taken
when abundant (Novakowski 1966).
A greater variety of foods are taken in the
wdnter, and include grains, acorns, insects, marine
worms, crustaceans, mollusks, fishes, amphibians,
reptiles, and, very occasionally, birds (Allen
(1952).
During migration, cranes feed while on mud
flats and sandbars. Allen (1 952) suggests that crust-
aceans, fishes, amphibians, and reptiles in the shal-
lows compose the bulk of the diet. Recent data in-
dicate that during fall migration, cranes often feed
in recently harvested grain fields, eating insects
and wasted grain, especially corn.
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
Nests, 15 to 60 cm above the water, are made
of rushes [Scirpus validus), sedge [Carex sp.), or
cattail {Typha latifolia). Nests are restricted to
areas with limited terrestrial accessibility, and are
not usually reused in subsequent years; but a new
nest is generally built in the same area (Kuyt
1976a). Allen (1952) describes in detail nests and
their construction in historic breeding localities.
RITUAL REQUIREMENTS
Premigration behavior in December or January
includes a restless shifting of habitual winter terri-
tories, increasing in frequency of dancing displays
and circling flights which eventually result in
actual movement northward (Allen 1952, WCRT
1977). Generally the cranes leave Aransas National
Wildlife Refuge between the end of March and
the middle of April. They arrive at Wood Buffalo
National Park 18 to 28 days later (Allen 1956,
WCRT 1977).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Climate plays a vital role in whooping crane
survival. Extreme dryness has little effect on nest-
ing success, but can force young and parents to
travel farther in search of food (Novakowski 1966)
and render nesting areas more accessible to terres-
trial predators (E. Kuyt, personal communica-
tion). Departure of cranes on spring migration is
assisted by strong prevailing southeast winds
(WCRT 1977). However, a hurricane or large
storm could interfere with migration (Allen 1952).
The average territory size is about 162 ha on
the wintering ground. In tlie Sass River area,
Wood Buffalo National Park, the radii of the nest-
ing territories average 0.9 km. Rarely, however,
are nests of two different pairs closer than 1.3
km, and when they are closer, nesting success is
below average (Kuyt 1976a).
POPULATION NUMBERS AND TRENDS
The numbers have increased since a low of 21
birds in 1941 -1942. The 1977 count of whooping
cranes was 75 birds in the wild (Aransas Refuge -
Wood Buffalo National Park had 69 birds, and
experimental Rocky Mountain flock had 6) and
27 in captivity (Patuxent Wildlife Research Center
had 22; International Crane Foundation, 3; and
San Antonio Zoo, Texas, 2) (WCRT 1977).
The potential for population growth in the
wild is quite low for many reasons (Stevenson
and Griffith 1946). Major reasons are that only
one young survives and there is high juvenile
mortality during migration. Erickson (1961)
suggests that although numbers have increased,
there has not been an increase in rate of annual
production per breeding pair. Thus, while the
average annual production per pair has remained
essentially the same, an increasing proportion of
the older birds is returning to Aransas without
progeny. Novakowski (1966) also suggests that
early separation of young from adults during
spring migration may account for high losses of
young of the year.
Allen (1952) estimated that the historical
(1870) population was about 1,500 birds.
REPRODUCTION
Usually two, smooth, buff eggs with dark
blotches are layed in late April or early May. The
incubation period is 29 days. Incubation is shared
by both parents, with one parent generally inclin-
ed to spend more time on the nest (Allen 1952).
Parents and young leave the nest site 2 to 3 days
after hatching, and spend the summer traveling
from pond to pond feeding (E. Kuyt, personal
communication). The family unit is maintained
through the winter, with the young gradually
gaining more independence. Just before the migra-
tion, the parents begin to drive the young away,
but do not separate entirely until enroute to the
nesting grounds (Allen 1952, Novakowski 1966).
MANAGEMENT AND CONSERVATION
Early accounts of a superabundance of
whooping cranes probably caused a delay in rec-
ognizing and responding to the decline in popula-
tion. Many people probably misidentified the
more numerous sandhill crane (Allen 1952).
Efforts to prevent whooping crane extinction
have been considerable and have succeeded mar-
ginally in increasing the total numbers in the wild.
The sequence of events is detailed in Allen (1952)
and WCRT (1977).
The Migratory Bird Act of 1918 provided
legal protection of migratory birds and paved the
way for cooperation in whooping crane manage-
ment between the United States and Canada.
Aransas National Wildlife Refuge was established
in 1937 to protect whooping cranes, waterfowl.
and other coastal Texas wildlife. The refuge lies
between San Antonio Bay and Aransas Bay on
the Texas Gulf coast, and in 1937 was the winter-
ing ground for 60% of the whooping cranes
(WCRT 1977).
The American Ornithologists' Union wrote a
report in 1944 calling for the National Audubon
Society and U.S. Fish and Wildlife Service to
determine the status of whooping cranes, and to
stop the trend toward extinction. The Wilson Soci-
ety made the same request in 1944 (Allen 1952).
In 1945, the Cooperative Whooping Crane
Project, involving the U.S. Fish and Wildlife Ser-
vice and the National Audubon Society, was set
up to investigate means to protect and restore the
species.
Robert Allen's 1952 monograph, The Whoop-
ing Crane, a product of 39 months of study, out-
lined early management needs.
The nesting grounds were discovered in 1954
in Wood Buffalo National Park, which had been
created in 1922 as a preserve and management
area for bison.
The Cooperative Whooping Crane Project was
replaced by the Whooping Crane Advisory Group,
appointed by the Director of the Bureau of Sport
Fisheries and Wildhfe in 1956 to provide advice to
the Bureau on the crane's problems.
The Standing Committee of the International
Wild Waterfowl Association created the Whooping
Crane Conservation Association in 1961. This
association has both Canadian and United States
membership and is privately funded. The associa-
tion encourages artificial propagation to increase
numbers, and promotes public education to
protect the cranes on migration. The International
Crane Foundation is another private organization
whose goals are to conserve cranes worldwide
(WCRT 1977), and also contributes to the whoop-
ing crane project.
Many efforts have been made to reverse the
trend towards extinction. In 1946, food habit
studies were initiated. Shordy afterwards, tech-
niques for keeping abreast of population numbers
were developed, e.g. monitoring flyways. Warning
signs were posted on the Intracoastal Waterway,
which bounds the Aransas Refuge. Oil drilling in
the area of the refuge was handled in a coopera-
tive spirit and cattle grazing was found not to
compete with cranes (Allen 1952). Public educa-
tion played a large role in reversing the downward
trends (Walkinshaw 1973).
The role played by the Canadian Wildlife
Service of the Canadian Department of Environ-
ment is equal in importance to that of the U.S.
Fish and Wildlife Service. This group has con-
ducted surveys since 1954 to record events on the
breeding grounds. Other important functions
include (WCRT 1977):
1. Fire patrol and fire suppression
2. Prohibition of public access into or low-fly-
ing aircraft over the area.
3. Resistance to attempts by commercial inter-
ests to develop the breeding areas.
4. Conduct periodic censuses.
A migration monitoring program was estab-
lished by the U.S. Fish and Wildlife Service in the
fall of 1975 to :
1. Attempt to prevent exposure to disease out-
breaks and other unusual hazards.
2. Alert key areas along flyways.
3. Provide information to be used to determine
major stop-over areas.
4. Provide information for determining critical
habitat.
The National Audubon Society has a network
that is coordinated with the U.S. Fish and Wildlife
Service program; and several States along the fly-
way assist in crane protection.
A recovery team was approved in November
1975, and has produced a draft Recovery Plan
(WCRT 1977). The primary objective of their
plan is to remove the whooping crane from its
endangered status. The requirements necessary to
reach this objective are:
1. Increase the Wood Buffalo— Aransas popula-
tion to at least 40 nesting pairs.
2. Establish at least two additional, separate and
self-sustaining populations consisting of at
least 20 nesting pairs.
Additional research is advised on food habits
and avEiilability in relation to climatic conditions,
spatial requirements, and territorial behavior in
an expanding population. Additional data are also
needed on the impact of increasing human activi-
ties in the habitat area.
The Canadian Wildlife Service and U.S. Fish
and Wildlife Service removed 50 eggs from Wood
Buffalo in 1967, 1968, 1969, 1971, and 1974 and
attempted to hatch them at Patuxent Research
Center, Maryland. As of 1975, 19 whoopers have
been raised in this effort. The goal of producing
second-generation captive birds for release has not
yet been achieved (Kuyt 1976a).
In 1975, 14 eggs were taken from Wood Buf-
falo nests and transplanted to foster sandhill
crane parents at Grays Lake National Wildlife
Refuge, Idaho. The goal of this effort is to estab-
lish an additional isolated population so that a
single calamity could not destroy the entire popu-
lation (Kuyt 1976a). Eggs were also transplanted
in 1976 and 1977. As of 1977, six birds have
been successfully raised and now migrate with the
sandhill cranes (WCRT 1977).
Other management suggestions (Allen 1952)
which have not been achieved include inclusion of
Mustang Lake into the Aransas refuge, closure of
it to fishing, and conversion of the lake to the
predredged condition. Allen (1952) also suggested
extending the boundaries of the refuge, and this
has been accomplished in part by the National
Audubon Society's leasing of lands adjacent to
the refuge (WCRT 1977).
Critical Habitat was designated in 1978 (43
FR 20938; 15 May 1978) and includes these areas:
Colorado Monte Vista National Wildlife Refuge
(all), and Alamosa National Wildlife
Refuge (all).
Idaho Grays Lake National WildUfe Refuge
(all plus a 1.6- km perimeter).
Kansas Quivira National Wildlife Refuge (all),
Cheyenne Bottoms State Waterfowl
Management Area (all).
Nebraska A strip along the Platte River in
Dawson, Gosper, Kearney, Buffalo,
and Phelps Counties.
New
Mexico Bosque del Apache National Wildlife
Refuge (part).
Oklahoma Salt Plains National Wildlife Refuge
(all).
Texas Aransas National Wildlife Refuge (all),
northeast to Pringle Lake, southeast
to Gulf Intracoastal Waterway plat-
form channel marker 25, and north-
west to the refuge.
Eight additional areas have been proposed as
critical habitat (43 FR 36588; 17 August 1978):
Kansas Kirwin National Wildlife Refuge.
Montana Medicine Lake National Wildlife
Refuge.
Nebraska A section of the Nebraska River in
Brown, Burleigh, Divide, Dunn, Em-
mons, McKenzie, McLean, Morton,
Mountrail, Penville, Sioux, Ward, and
areas in Campbell and Williams
Counties.
AUTHORITIES
David Blankinship (Recovery Team)
National Audubon Society
721 Pine Street
Rockport,TX 78382
Roderick Drewien
Bosque del Apache National Wildlife Refuge
Box 278
San Antonio, NM 87832
Ray Erickson (Recovery Team)
Assistant Director
Patuxent Wildlife Research Center
Laurel, MD 20810
Conrad Fjetland
c/o U.S. Fish and Wildlife Service
5796 Bickel Church Road
Pickerington, OH 43147
Harold D. Irby (Recovery Team)
Program Director
Migratory Game
Texas Parks and Wildlife Department
4200 Smith School Road
Austin, TX 78744
Ernie Kuyt
Canadian Wildlife Service
9942 108th Street
Edmonton, Alberta
T5K 2JS
Ross Lock (Recovery Team)
Non-Game Specialist
Nebraska Game and Parks Commission
2200 North 33rd Street
Lincoln, NE 68503
Dave Olsen (Recovery Team Leader)
Division of Wildlife Refuges
U.S. Fish and Wildlife Service
Department of the Interior
Washington, D.C. 20240
L. H. Walkinshaw
4691 Timberlane Road
Lake Wales, FL 33853
PREPARER'S COMMENTS
None.
LITERATURE CITED/SELECTED
REFERENCES
Natl.
South
Dakota
Conson, Dewey, Haakon, Hughes,
Potter, Stanley, Sully, Walworth, and
Ziebach Counties.
Allen, R. P. 1952. The whooping crane.
Audubon Soc. Res. Rep. 2. New York.
. 1954. Additional data on the food of the
whooping crane. Auk. 71(2):198-199.
. 1956. A report on the whooping crane's
northern breeding grounds. Natl. Aud. Soc.
Supple. Res. Rep. 3.60 pp.
Archibald, G. W. 1976. Crane taxonomy as
revealed by the unison call. Pages 225-251 in
J. C. Lewis, ed. Proc. Intl. Crane Workshop.
Oklahoma State Univ. Publ. and Printing.
355 pp.
Bard, F. G. 1956. Whooping cranes in migration,
1956. Blue Jay 14 (2):39-42.
. 1958. Whooping cranes, 1958. Blue Jay
16(1):11-14.
Blankinship, D. R. 1976. Studies of whooping
cranes on the wintering grounds. Pages 197-
206 in J. C. Lewis, ed. Proc. Intl. Crane
Workshop. Oklahoma State Univ., Publ.
Printing. 355 pp.
Bent, A. C. 1926. Life histories of North Ameri-
can marsh birds. Dover Publ., Inc., New York.
392 pp.
Drewien, R. C. 1975 & 1976. Unpublished quar-
terly progress reports 1-8, whooping crane
transplant experiment. Idaho Coop. Wildl.
Res. Unit, Univ. Idaho, Moscow.
Erickson, R. C. 1961. Production and survival of
the whooping crane. U.S. Bur. Sport Fish
Wildl. Serv. Rep. 29 pp.
. 1966. Production and survival of the
whooping crane. U.S. Fish Wildl. Serv. Admin.
Rep.
. 1975. Captive breeding of whooping
cranes at the Patuxent Wildlife Research
Center. Pages 99-114 in R. D. Martin, ed.
Breeding Endangered Species in Captivity.
Academic Press, New York. 420 pp.
. 1976. Whooping crane studies at the
Patuxent Wildlife Research Center. Pages 166-
176 in J. C. Lewis, ed. Proceedings Intl. Crane
Workshop. Oklahoma State Univ. Publ. Print-
ing. 355 pp.
. 1979. Whooping crane (Grus americana).
In R. C. Erickson and Ruhr, eds. Crane
research around the world. Cosponsored by
U.S. Chapter of LC.B.P., Intl. Crane Found.,
and U.S. Fish Wildl. Sen'.
Hahn, P. 1963. Where is that vanished bird. Royal
Ontario Museum, Univ. Toronto, Toronto.
Hovrard, J. A. 1954. Aransas, a national wildlife
refuge. U.S. Dep. Inter., Fish Wildl. Serv. 11:
1-12.
Kepler, C. B. 1976. Dominance and dominance-
related behavior in the whooping crane. Pages
177-196 in J. C. Lewis, ed. Proc. Intl. Crane
Workshop. Cosponsored by U.S. Chapter
I.C.B.P., Intl. Crane Found., and U.S. Fish
Wildl. Serv.
Kuyt, E. 1968. Collection of whooping crane eggs
from Wood Buffalo National Park. Trans Fed-
Prov. Wildl. Conf. 32:30-35.
. 1975. Nest site fidelity, productivity,
and breeding habitat of whooping cranes.
Wood Buffalo National Park, Northwest
Territories, Canada. In Crane Res. Around the
World, Internl. Crane Found.
. 1976a. Whooping cranes: the long road
back. Natur. Can. (Ottowa) 5(2):2-9.
. 1976b. The continuing story of the
whooping crane. Pages 109-111 in Canada's
threatened species and habitats. Symposium
on Canada's threatened species and habitats,
Canadian Nature Federation and World Wild-
life Fund, Ottowa, 20-24 May 1976.
. 1976c. Recent clutch size data for
whooping cranes, including a three-egg clutch.
The Blue Jay 34(2):82-83.
Lowery, G. H., Jr. 1974. Louisiana birds. Pub-
lished for Louisiana Wildlife and Fisheries
Commission by Louisiana State Univ. Pres.
651 pp.
McNulty, F. 1966. The whooping crane; the bird
that defies extinction. Dutton, New York.
190 pp.
Muir, D. 1976. Whooping crane summer: filming
the great white cranes. Natur. Can. (Ottowa)
5(2):25-32.
Novakowski, N. S. 1966. Whooping crane popula-
tion dynamics on the nesting grounds. Wood
Buffalo National Park, Northwest Territories,
Canada. Can. Wildl. Serv. Res. Rep. 1.20 pp.
Pratt, J. J. 1961. Transactions of the whooping
crane conservation conference. Modem Game
Breeding, April 1961.
Stevenson, J. O., and R. E. Griffith. 1946. Winter
life of the whooping crane. Condor 48(4):
160-178.
U.S. Department of the Interior. 1938-present.
Whooping crane census. Informal reports by
the refuge manager, Aransas National Wild-
life Refuge, Austwell, Texas.
U.S. Fish and Wildlife Service. A great white bird.
50-minute film, color. National Audiovisual
Center, GSA, Washington, D.C.
WCRT (Whooping Crane Recovery Team). 1977.
Whooping crane recovery plan. Technical
review draft. U.S. Dep. Inter., Fish Wildl.
Serv. 136 pp.
Walkinshaw, L. H. 1973. Cranes of the world.
Winchester Press, New York. 370 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesvnie, FL 32601
10
Biological Services Program
FWS/OBS-80/01.4
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
LIGHT FOOTED CLAPPER RAIL
K'r^^r^^
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoasl of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivident
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed lo:
Office of Endangered Species
U.S. Fish and Wildlife Ser\'ice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report shoidd be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.4
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
LIGHT FOOTED CLAPPER RAIL
A Cooperative Effort
by the
National Fish and WildHfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-SHdell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT: SANFORD WILBUR
LIGHT-FOOTED CLAPPER RAIL
Rallus longirostris levipes Bangs (1899)
KINGDOM Animalia
CLASS Aves
ORDER Gruifonnes
FAMILY Rallidae
OTHER COMMON NAMES King Rail (part),
California Clapper Rail (part), Southern Cali-
fornia Clapper Rail, Bangs Rail, Light-footed
Rail, Southwestern Clapper Rail, Los Angeles
Clapper Rail (Grinnell and Miller 1944)
DATES
Entered into SWIS to be determined
Update to be determined
LEGAL STATUS
Federal: Endangered (42FR36427, 14 July
1977). Protected Migratory Bird Treaty
Act, (40 Stat. 755; 16 USC 703-711,
3 July 1918) as amended 3 Dec. 1969.
PL91-135.
State: California - Endangered
REASONS FOR CURRENT STATUS
Formerly common in aU coastal marshes with-
in its geographic range, but now, because of drain-
age and/or filling of some marshes and pollution
with chemicals and debris/?. /. levipes is much more
restricted in occurrence and fewer in numbers
(Willett 1912,1933,GrinneIleta]. 1918, Stephens
1919, Edwards 1922). It was not as well protected
from hunters as the California clapper rail, and its
range is being reduced by reclamation of marshes
(Bent 1926).
Overharvesting may have occurred in some
areas, but reduction in populations can be at-
tributed almost entirely to loss of habitat. The
area of the salt marsh between Santa Barbara and
the Mexican Border is estimated to have been
10,500 ha, while at present there are approxi-
mately 3,440 ha (Speth 1971), much of which
has been degraded by water pollution, water di-
version, and restriction of tidal flow. Of the 4 ma-
jor habitat areas, only Anaheim Bay appears to be
relatively safe from future habitat destruction.
Los Penasquitos has dried up and does not sup-
port rail, Upper Newport is "safe" under State
management, and Tijuana Slough is vulnerable,
but FWS plans to preserve it. Particularly hard hit
were areas known to have supported large popula-
tions of rails such as San Diego Bay, reduced from
990 to 146 ha; Mission Bay, from 970 to 8.5 ha;
and the Los Angeles-Long Beach area, from 2750
to 28 ha. Thus, entire populations have been ex-
terminated (Recovery Team 1977).
PRIORITY INDEX
32
DESCRIPTION
Rallus longirostris levipes is a chicken-size
bird with a gray-brown back, tawny breast, ver-
tical dusky and white bars on its flanks, and a
white patch under its short, upcocked tail; it has
long legs, toes and bill and is a strong runner and
a weak flyer. Although similar to R. I. obsoletus,
it is slightly smaller; with a more slender bill and a
back darker, browner or more olive (less grayish)
in tone, with narrower and less black marking; its
breast is a richer cinnamon color and the stripe
over the eye more whitish, less rusty (Bangs 1899,
Bent 1926, van Rossem 1929, Ridgway and
Friedmann 1941).
The size of the adult male is: wing, 154.5 - 167
(av. 161.9) mm; tail, 62.5 - 69 (66.7); exposed
culmen, 56-61 (58.9); tarsus, 53-60.5 (56.9);
middle toe without claw, 50-54 (51.2^. The adult
female is: wing, 138 - 155.5 (147.3); tail, 57 - 67
(62.6); exposed culmen, 51.5-58 (54.2); tarsus,
47-51 (49.5); middle two without claw, 41-48
(44.9) (Oberholser 1937) Weight - male, av. 306
gm.; female, av 248 gm. (Ohmart and Smith 1973).
Its eggs are drab, cream-colored, and sparsely
marked with purple 44.6 x 31.0 mm (Bent
1926).
The light-footed clapper rail has a confusing
taxonomic history. Originally thought to be a
King Rail [Rallus elegans) by Henshaw (1876);
then the same as California Rail [Rallus obsoletus),
which at that time was considered a distinct spe-
cies by Belding (1883); then thought to be a
separate species [Rallus levipes) by Bangs (1899);
then a subspecies of the California Rail [Rallus
obsoletus levipes) that was still considered a dif-
ferent species from either King or Clapper Rail by
van Rossem (1929); then united with the King
Rail as Rallus elegans levipes by Peters (1934); it
was finally classified a clapper rail subspecies Ral-
lus longirostris levipes (Oberholser 1937), which
designation was accepted by Ridgway and Fried-
mann (1941), the American Ornithologists Union
(1957), and subsequent authorities.
RANGE
The light-footed clapper rail ranges the coastal
salt marshes from Santa Barbara County, Cali-
fornia, south to San Quintin Bay, Baja California
1977), which is the originally described range of
Cooke (1914), Grinnell et al. (1918) and Bent
(1926). Later evaluations (Grinnell and Miller
1944, American Ornithologists Union 1957)
found no Santa Barbara records after 1875, so set
the northern limits at Point Mugu and Hueneme,
Ventura County. Confusion over distinction of
levipes from another subspecies farther south in
Baja California resulted in placing the southern
breeding limit of levipes at Ensenada, Baja Cali-
fornia (Friedmann et al. 1950, American Ornith-
ologists' Union). Study of more adequate speci-
men samples extended the breeding range of
levipes southward to San Quintin Bay (S. Wilbur
ms.). Distribution is markedly interrupted be-
cause of discontinuous habitat (van Rossem 1929;
Grinnell et al. 1918, Edwards 1922, Dawson
1924).
Areas that have recent records of light-footed
clapper rails are: Santa Barbara Co. - Carpinteria
Marsh (or El Estero or Sandyland Slough); Ven-
tura Co. - Mugu Marshes (Pacific Missile Range);
Orange Co. - Anaheim Bay marshes. Upper New-
port Bay; San Diego Co. - Tijuana River estuary;
Sweet Water Marsh ; Otay River Slough; Marine
Biology Study Area; San Diego River; Mission
Bay; Los Penasquitos; San Elijo Lagoon (Wilbur
1974). Despite published reference to the contrary
(Friedmann et al. 1950, American Ornithologists'
Union 1957, van Rossem 1947), there is no evi-
dence of migration or of wandering from home
marshes after the breeding season (Wilbur and
Tomlinson 1976).
RANGE MAP
The past and present distribution is shown on
tlie following page.
LIGHT-FOOTED CLAPPER RAIL
Past and Present Distribution
Goleta Slough
Carpinteria Marsh
Mugu Lagoon'
■"^-^
%.
Bolsa Chi CO Lagoon
Upper Newport Bay »
Santa Margarita River
Batiquitos Lagoon
San Elilo Lagoon
San Dieguito River
Los Penasquitos Lagoon
San Diego River - Mission Bay
San Diego Bay
Tijuana Estuary
La Mision - La Salina
Bahia de Todos Santos
Bahia de San Quintin
Past and present distribution of the Hght-footed clapper rail.
STATES/COUNTIES
California: Orange, San Diego, Santa Barbara,
Ventura.
HABITAT
The light-footed clapper rail is found in salt-
water marshes traversed by tidal sloughs, where
cordgrass {Spartina foliosa) and pickleweed {Salt-
cornia) are the conspicuous plants. One nesting is
recorded for an inland marsh grown to reeds in
Los Angeles Co. (Willett 1906). In Tijuana Estuary,
22 of 34 nests were built in cordgrass. Nest den-
sity is greater in tall cordgrass (0.52 nests per ha.
of habitat). Generally speaking, the highest den-
sities of rails appear to be in those marshes with
the most cordgrass Qorgensen 1975). Clapper
rails require a healthy tidal salt marsh environment
with cordgrass or pickleweed for nesting and es-
cape cover; abundant food in the form of crabs,
clams and related invertebrates; and tidal flats
interspersed with saltmarsh vegetation as a feed-
ing area. These conditions prevail in coastal salt
marshes that have a tidal prism adequate to pre-
serve a normal salinity range and prevent stagna-
tion. If suitable physical environment is available,
other factors seem to have little influence. Preda-
tion by other animals is seldom a limiting factor,
and raUs seem to tolerate a high level of human
occupancy of their habitat provided such use does
not result in habitat degradation or loss of birds
(Recovery Team 1977).
FOOD AND FORAGING BEHAVIOR
Striped shore crabs (Pachygrapsus crassipes)
and purple shore crabs [Hemigrapsiis oregonensis)
are regularly eaten by the clapper rail. Remains
found in pellets cast by rails incorporated fiddler
crab [Uca crenulata), beach hopper [Orchestoidea
sp.), California hornshell [Certhidea californica),
and a gastropod [Melampus olivaceus). Probably
(like other clapper rails) the light-footed clapper
rail eats various bivalve molluscs such as clams
of the genus Macoma (Jorgensen 1975).
SHELTER REQUIREMENTS
Dense growth of either pickleweed or cord-
grass for nesting and escape cover (Recovery Team)
is required by the clapper rail.
NESTING OR BEDDING
The clapper rail's nest is a loose arrangement
of plant stems on high ground in a salt marsh,
well concealed in dense or tall vegetation, usually
Spartina. One nest measured 28 by 18 cm on the
outside, with a cavity 13 cm across and 1.3 cm
deep (Bent 1926, Edwards 1922). Nests
are also frequently placed in cordgrass (Spartina).
Those in Spartina most often are placed directly
on the ground, while those in cordgrass are ele-
eated 10 to 45 cm. Nests are constructed of what-
ever vegetation is available at the site and are
placed so as to be well concealed. The nests are
bouyant and will float with the tide Qorgensen
1975).
RITUAL REQUIREMENTS
Probably pairs are spaced by aggressive terri-
torial defense as in other subspecies of clapper
rails.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None other than those mentioned elsewhere.
POPULATION NUMBERS AND TRENDS
Estimates now indicate a total population of
about 250 birds on the basis of work in Santa
Barbara and Ventura counties (V. Basham, un-
published), at Anaheim Bay (B. Massey, C. Collins,
J. Lindell), and at Tijuana Estuary Qorgensen
1975). Tot£il population estimates are distributed
as follows: Santa Barbara County, Carpinteria,
10; Orange County, Anaheim Bay, 40-50; Upper
Newrport Bay 40-50; San Diego County, Los
Penasquitos Lagoon 5, Mission Bay 8, San Diego
River 5, San Diego Bay area 40-50, Tijuana Estu-
ary 75-85; Baja California (occur but population
size undetermined)* Bahia de Todos Santos, La
Mision and La Salina may have rails also (Recovery
Team 1977).
REPRODUCTION
The young of the clapper raU are able to swim
on the day of hatching.
Nesting occurs from mid-March to 1 July,
with extreme dates for complete clutches 18
March to 31 July. Egg dates (56 records) are from
18 March to 11 June (Bent 1926).
Most egg laying occurs from early April to
early May, with 3 to 11 eggs per clutch, usually 5
to 9. The incubation period averages 23 days
Qorgensen, 1975).
Both sexes are believed to incubate (Bent
1926). At Tijuana Estuary 86% of 28 active nests
successfully hatched at least 1 egg. There were
losses due to eggs being washed away by high
tides and failure to hatch for unknown reasons.
Chicks were found dead in several nests apparently
from drowning, although young are said to be
able to swim at day of hatching (Bent 1926).
Nests have been destroyed by rats. At least some
renesting occurs and some pairs may raise second
broods.
MANAGEMENT
A recovery team was appointed and a recovery
plan prepared in January 1977. The light-footed
clapper rail is endangered because its range is
limited to a relatively small remnant of suitable
marsh habitat, and this remnant continues to de-
crease. Arresting the decline of the rail population
requires stopping all loss and degradation of exist-
ing habitat. Even then, population will remciin
threatened because each segment is so small it
could easily be eradicated by pollution, disease,
predation, or other local catastrophes.
The recovery plan proposed by the Light-
Footed Clapper Rail Recovery Team includes:
(1) protecting all existing habitat; (2) increasing
its carrying capacity and stability, thereby in-
creasing the size of each population unit; and
(3) creating and stocking new habitat. The objec-
tive of the recovery plan is to develop and maintain
a breeding population of at least 400 pairs of light-
footed clapper rails well distributed geographi-
cally. This can be done by preserving and restor-
ing approximately 1620 ha. of tidal saltmarsh
habitat in 15 or more coastal marshes. Addition-
ally, efforts should be made to have occupied
salt marsh habitat preserved in Baja California.
An action program to achieve these objectives
includes: (1) preserving and increasing existing
populations by preserving existing habitat, pre-
venting development or degradation, increasing
amount of habitat, and reducing effect of other
limiting factors; (2) reestablishing former rail
populations by restoring tidal flow to previously
inhabited marshes, restoring cordgrass-pickleweed
vegetation by planting if it fails to volunteer, and
transplanting rails to new sites; (3) monitoring
rail populations to determine progress of manage-
ment and status of the populations; and (4) carry-
ing on a program of education and planning ad-
vise directed toward preservation of rail habitat
(Recovery Team 1977). Saving Tijuana Marsh is
the chief priority for recovery of the light-footed
clapper rail (Sanford Wilbur pers. comm.).
AUTHORITIES
Sanford R. Wilbur
U.S. Fish and Wildlife Service
1190 E. Ojai Avenue
Ojai, California 93023
Ronald M. Jurek
California Dept. of Fish and Game
1416 9th Street
Sacramento, California 95814
Ronald Hein
California Department of Fish and Game
Long Beach, California 90840
Charles T. Collins
Department of Biology
California State University at Long Beach
Long Beach, California 90840
R. Guy McCaskie
1310 14th Street
Imperial Beach, California 92032
PREPARER'S COMMENTS
Because of the very small populations and
limited suitable habitat that appears to be rapidly
diminishing, the light-footed clapper rail appears
to be in extreme jeopardy. Only vigorous efforts
to preserve a number of unpolluted salt marshes
with free-flowing tidal currents will be effective
in saving this subspecies from extinction.
LITERATURE CITED/SELECTED
REFERENCES
American Ornithologists' Union. 1957. Check-list
of North American birds. 5th ed. Baltimore,
Md. AOU. 641pp.
Bangs, O. 1899. A new rail from southern Cali-
fornia, Proc. New Eng. Zool. Club 1:45-46.
Belding, L. 1883. Catalogue of a collection of
birds made at various points along the western
coast of Lower California, north of Cape St.
Eugenis. Proc. U.S. Nat. Mus. 5:527-532.
Bent, A. C. 1926. Life histories of North American
marsh birds. U.S. Nat. Mus. BuU. 135:272.
Cooke, W. W. 1914. Distribution and Migration of
North American rails and their allies. U.S.
Dept. of Agric. Bull. 128:50 pp.
Dawson, W. 1924. Birds of California. Vol. 3. p.
1533.
Edwards, H. A. 1922. Some notes on the Light-
footed Rail. Oologist 39:60-61.
Frey, H. W., R. F. Hein, and J. L. Spruill. 1970.
Report on the natural resources of Upper
Newport Bay and recommendations concern-
ing the Bay's development Calif. Dept. Fish
and Game. 68 pp.
Friedmann, H., L. Griscom and R. T. Moore.
1950. Distributional Checklist of the birds of
Mexico. Pac. Coast Avifauna 29:202 pp.
Grinnell, J. 1898. Birds of the Pacific slope of Los
Angeles County. Pasadena Acad. Sci. Pub. 2.
. 1915. A distributional list of the birds of
of California. Pac. Coast Avifauna 11. 217 pp.
Grinnell, J. and A. H. Miller. 1944. The distribu-
tion of the birds of California. Pac. Coast Avi-
fauna 27:128.
Grinnell, J., H. C. Bryant, and T. I. Storer. 1918.
The Game birds of California, p. 289.
Henshaw, H. W. 1876. Report on the ornithology
of the portions of California visited during the
field season of 1875. Ann. Rep. Geographic
Survey west of the 100th Meridian. Appendix
H8 of Appendix JJ. pp. 224-278.
Jorgensen, P. D. 1975. Habitat preference of the
Light-footed Clapper Rail in Tijuana Marsh,
California. M.S. thesis, San Diego State Uni-
versity. 115 pp.
Light-footed Rail Recovery Team. 1977. Light-
footed Clapper Rail recovery plan. Office of
Endangered species, U.S. Fish and Wildlife
Service, Wash., D.C.
McCaskie, G. 1972. Southern Pacific Coast Re-
gion. Am. Birds 26:903.
Mudie, P., B. Browning and J. Speth. 1974. The
natural resources of Los Penasquitos Lagoon
and recommendations for use and develop-
ment. Calif. Dept. Fish and Game, Coastal
wetlands Series no. 7. 75 pp.
Oberholser, H. C. 1937. A revision of the Clapper
Rails {Rallus longirostris Boddaert) Proc. U.S.
Nat. Mus. 84:313-354.
Ohmart, R. D. and R. W. Smith. 1973. North Am-
erican Clapper Rails {Rallus longirostris) liter-
ature survey, with special consideration being
given to the past and current status ofyuman-
sis. Report in fulfillment of Bur. Reclamation
contract 14-06-300-2409.
Peters, J. 1934. Check-list of birds of the world.
Vol. 2. Cambridge, Harvard University Press,
p. 159.
Ridgway, R. and H. Friedmann. 1941. Birds of
North and Middle America. Part IX. Bull. U.S.
Nat. Mus. 50:75.
Romero, P. D. 1972. Anaheim Bay study, July
1970 to June 1971. Calif. Dept. Fish and
Game. 22 pp.
Sams, J. R. and K. Scott, Jr. 1959. Birds of San
Diego County, California. San Diego Soc. Nat.
Hist. Occas. Pap. No. 10.
Sexton, C. W. 1972. Clapper Rails at Upper New-
port Bay, California. Unpub. Ms. U. of Calif,
at Irvine.
Speth, J. W. 1971. The status of coastal wetlands
in southern California. Paper presented at an-
nual meeting. Western Section of The Wildlife
Society. 19 pp.
Speth, J., R. Fordice, R. Hein, and P. Giguere.
1970. Th*" natural resources of Golata Slough
and recommendations for use and develop-
ment. Calif. Dept. Fish and Game. 42 pp.
Stephens, F. 1919. An annotated list of the birds
of San Diego County, California. Trans. San
Diego Soc. Nat. Hist. 3:1-40.
U.S. Fish and Wildlife Service. 1973. Threatened
WildUfe of the United States. U.S. Dept. of
Interior, Resource Pub. 114. 289 pp.
van Rossem,A.J. 1929. The status of some Pacific
Coast Clapper Rails. Condor 31:213-215.
. 1947. Comments on certain birds of Baja
California, including descriptions of three new
races. Proc. Biol. Soc. Wash. 50:51-58.
Warner, D. S. and R. W. Dickerman. 1959. The
status oi Rallus elegans tenuirostris in Mexico.
Condor 61:49-51.
Wilbur, S. R. 1974. The status of the Light-footed
clapper Rail. Amer. Birds 28:868-870.
Wilbur, S. R. and R. E. Tomlinson. 1976. The
literature of the Western Clapper Rails. USFWS
Spec. Sci. Rep. - Wildlife. 194. 31 pp.
Willett, G. 1906. The Southern Clapper Rail
breeding on fresh water. Auk 23:432.
. 1912. Birds of the Pacific slope of south-
em California. Pac. Coast Avifauna 7. 122 pp.
. 1933. A revised list of the birds of south-
wfestem California. Pac. Coast Avifauna 2 1 :
52.
Biological Services Program
FWS/OBS-80/01.5
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
SAN CLEMENTE LOGGERHEAD SHRIKE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or ihrcatcned vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data are not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
iNASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.58
FWS/OBS-80/01.5
March 1980
SKLKCri:i) VIR 1 KBRAir. KNDANGERED SPKCIES
OF i hi: skacx)as 1 of ihk united S IA IES-
SAN CLEMENTE LOGGERHEAD SHRIKE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National CoastaJ Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Scrxicc
U.S. Department of the Interior
PHOTO OF CLOSELY RELATED SUBSPECIES
SAN CLEMENTE LOGGERHEAD SHRIKE
Lanius ludovicianus meamsi Ridgway
KINGDOM Animalia
CLASS Aves
ORDER Passerifonnes
FAMILY Laniidae
OTHER COMMON
NAMES California shrike, island shrike,
San Clemente shrike, Meams' shrike,
Santa Barbara Island Shrike.
DATE
Entered into SWIS To be determined
Update To be determined
LEGAL STATUS
Federal: Endangered: FR 42 40685, 11 Aug.
1977. p. 40685.
States: Protected by California law.
REASONS FOR CURRENT STATUS
Destruction over large areas of the dense,
brushy vegetation required for shrike territory
headquarters has undoubtly reduced the number
of possible wintering and breeding areas consider-
ing the aggressive territorial behavior of the birds.
This has reduced the reproductive potential of L.
I. mearnsi to the point where it has almost the
lowest ratio (30%) of immatures to adults, of any
subspecies except L. I. anthonyi, another island
race (Miller 1931).
PRIORITY INDEX
None assigned.
DESCRIPTION
The San Clemente loggerhead shrike is a
medium-sized bird, slightly smaller than a robin,
gray above, white below and on the rump, with a
black mask over the eyes. Black wings and tail,
each have patches of white. Of all the subspecies
of L. ludovicianus, mearnsi is the most isolated
and is among the most sharply characterized (Mil-
ler 1931).
Overall length is 224 mm (av. of 10 adults)
(Meams 1898). It is similar to L. I. anthonyi of
Santa Cruz Island, but the upper tail coverts are
abruptly white; there is more white on the scap-
ulars, the white spot at the base of the primaries
is larger, and the underparts of the body much
less strongly tinged with gray. In the white upper
tail coverts and the greater extent of white on
posterior scapulars and at the base of the primaries,
it is similar to L. I. gambeli, but the gray of the
upper parts is very much darker (quite as dark as
L. I. anthonyi), with much less white at the base
of primaries and on the lateral rectrices (Ridgway
1904).
L. I. mearnsi may be readily distinguished
from nelsoni by its much darker upper parts,
more conspicuously white rump, shorter wing,
smaller amount of white on primaries, and much
smaller bill.
Measurements: Wing length, av. 97.64 mm;
white on primaries 53.8% of wing length; tail
length av. 100.28 mm; white tip on outer tail
feather 28.1% of tail length; bill length, av. 27.60
mm; middle toe av. 13.20 mm; hind toe av. 9.92
mm. The only race with which meanisi integrades
is anthonyi to the north (Miller 1931).
RANGE
This shrike is a permanent resident on San
Clemente Island, off southwestern California
(American Ornithologists' Union 1957).
RANGE MAP
The range map on the following page shows
the location of San Clemente Island, this shrike's
total range.
STATES/COUNTIES
California: Los Angeles.
HABITAT
San Clemente Island, the southernmost of the
California Channel Islands, is approximately 34
km long and 2.4 to 6.4 km wide. The nearest
island, Santa Catalina, is about 34 km north. The
nearest point on the mainland is about 80 km to
the northeast.
Goats were introduced not later than 1827,
and by 1840 had formed a dense population. By
1877, large numbers of sheep grazed there also
(Raven 1963).
The island has been under jurisdiction of the
U.S. Navy since 1934, when all ranching presum-
ably terminated.
The topography is dominated by a plateau
with a steeply sloping east side and gently sloping
west side, covered with introduced grasses except
in places completely denuded by goats. Trees and
shrubs grow only on the bottoms and sides of the
canyons. On the east side, ironwood {Lyonotha-
mus), island oak (Quercus toinentella), and
lemonade bush {Rhus integrifolia) are the most
abundant species. The west side canyons are
mostly denuded, but have some growth of toyon
(Heteromeles), lemonade bush, and island cherry
[Prunus ilicifolia lyonii) (Stewart and Clow 1974).
Shrikes are found in washes, ravines and mesas,
where there are either scattered tall bushes such
as toyon and wild cherry, or low thorny scrub
and cactus patches (Grinnell and Miller 1944).
The presence or absence of thorny bushes
seems to have little effect on determining the hab-
itat of American shrikes (presumably including
L. I. mearnsi). Impaling devices are so varied as to
be available in some form in almost any type of
habitat except entirely barren areas (Miller 1931).
SHELTER REQUIREMENTS
Dense tall brush or low tree growth is essential
for roosting and nesting in each territory, whether
breeding or wintering (Miller 1931).
NESTING AND BEDDING
In each territory there is a headquarters where
the roosting place or, if a breeding territory, the
nest is located. The headquarters provides good
lookout perches, feeding facilities, and some sort
of brushy growth for shelter at night. The requisite
for roosting places seems to be some support
above the ground within a screen of overhanging
branches. Roosts are marked by conspicuous fecal
deposits.
For nesting, shrikes prefer dense bushes or
small, thickly grown trees at medium heights,
rarely less than 1 m or more than 7 m off the
ground (Miller 1931). Howell (1917) speaks of
nests of mearnsi placed about 0.6 m from ground
in a "certain kind of thorny bush." This state-
ment applies to the more barren parts of San Cle-
mente. Where there are canyons on the island,
Grinnell (1897) and Howell (1917) reported this
shrike nesting in bushes on the steep canyon sides.
Typical nests described by Grinnell (1897) and
Linton (1908) were composed of dry twigs, weed
stems, and grasses, thickly lined with rootlets and
sheep's wool, and well concealed in thick bushes.
SAN FRANCISCO
CALIFORNIA
'0^=^
LOS ANGELES
* ^
/
?^
SAN CLEMENTE ISLAND
Total range of the San Clemente loggerhead shrike
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
In regions where nesting sites are scarce, con-
siderable modification of the winter and fall feed-
ing territories probably takes place when they are
converted into breeding territories. Presumably in
reorganizing breeding territories into fall and win-
ter feeding areas, one or the other of the adults
keeps possession of the nesting locality, while the
other members of the family seek territories un-
occupied by shrikes, and which, in some cases,
may be suitable only for winter and fall occu-
pancy. Size and shape of territories depends on
the vegetative types present, the concentration of
food supply, the provision of nest sites, the local
abundance of the species and local physical bar-
riers (Miller 1931), as well as on the age, sex, and
physical condition of the bird.
POPULATION AND TRENDS
No more than 16 individuals were seen from
1-5 and 8-9 May 1974 (Stewart and Clow 1974).
Specific records were: 2 May (4); 4 May (5-6); 5
May (1 feeding fledgling) (Stewart and Clow 1974).
Formerly, L. I. mearnsi was rated by some
authors as "fairly common," but the total popula-
tion has always been small (Grinnell and Miller
1944). Long ago, it was considered tolerably com-
mon; that is, 2 or 3 could be generally seen during
an hour's walk, but they were very shy and hardest
to secure of any bird on the island (Grinnell 1897 ;
Meams 1898). They were reported fairly well
distributed over the whole island, but extremely
shy by Linton (1908). In especially favored little
canyons, several pairs would congregate. Two
pairs were found breeding not 100 m apart, while
a third was found within 0.4 kilometers (Howell
1917).
Causes of shrike mortality on San Clemente
are unknown, although some, particularly of
nestlings and juveniles, is undoubtedly caused by
predatory birds and mammals. Percentage of first
year birds in samples of winter and spring popula-
tions is only about 36%, by far the smallest per-
centage of all races except anthonyi, another
island form (most races have 50% or more im-
matures). This indicates a relatively poor repro-
ductive rate in the island populations. It may indi-
cate a lower population loss than mainland birds,
but it also shows the vulnerability of island birds
to any change in mortality rate or reproductive
potential (Miller 1931).
FOOD AND FORAGING BEHAVIOR
Shrikes hunt quite late in the evening and
early in the morning, at least in warm weather.
They are opportunists, living on the most abun-
dcmt and readily obtainable supply of animal food,
including all kinds of insects and other arthropods,
small reptiles, birds, and mammals that they can
capture. One was observed carrying a young
house finch with the adult house finch in pursuit
on 5 May 1974 (Stewart and Clow 1974). If there
is an infestation of a particular kind of insect,
shrikes will concentrate on that food (Miller 1931).
The method of hunting is to perch on objects
from 6 in (15.24 cm) to 6 ft (1.83 m) above the
ground where prey may be seen clearly. Oc-
casionally the bird hops about in search of animals.
If prey is not secured from a certain post within a
minute or two, it moves on to another part of the
territory. Passive hunting has been noted com-
monly during a large part of the day at times
other than when feeding young. A less common
method of feeding is capturing insects in the air
(MUler 1931).
Dead prey is impaled on a thorn, twig, splin-
ter, or other sharp structure, or eaten almost im-
mediately, depending on its size. If the shrike is
hungry when large prey is impaled, it eats all it
can— as much as 7 g at one feeding. Then the re-
mainder is left hanging for later feedings, which
usually continue until the last morsel is eaten.
The practical value to shrikes of impaled food
older than a few days is slight, as dry or spoiled
food is not eaten (Miller 1931).
REPRODUCTION
A set consists of five or six eggs. Eggs vary
from dull white to either light neutral gray or
buff, covered with small spots of neutral gray,
yellowish brown and umber, with occasional fine
black scrawlings near the large end.
Several sets have been found in March, and
young are commonly found out of the nest at the
end of March, indicating that eggs are present in
February. Incubation usually starts with the lay-
ing of the next-to-last egg; it is performed solely
by female. The male feeds the female during in-
cubation, either on or off the nest. Incubation
lasts about 16 days. Twenty days is the normal
time for young to remain in nest. Parents con-
4
tinue to feed young until about the 35th day
(Miller 1931).
Some birds raise two broods in one season. A
female whose mate was killed did not find a new
mate while feeding young older than 16 days
(Miller 1931).
It is not certain whether the male seeks the
territory of the female or vice versa for breeding,
but probably the former (Miller 1931).
Specific instances of finding nests with eggs
or young have been described by Bent (1950:182),
Linton (1908), and Howell (1917).
MANAGEMENT
The only practical action to take for the
benefit of the San Clemente loggerhead shrike
would be to remove or confine all of the livestock,
(particularly goats) on the island, to permit the
regrowth of dense patches of tall brush. This must
be accomplished over considerable areas to permit
adequate spacing of territories— at least 100 m
apart for each individual bird at all times of the
year. Replanting of native woody species of plants
in clumps for territorial headquarters would has-
ten rehabilitation. Until livestock can be removed,
exclosures to protect woody plants from over-
browsing might help.
AUTHORITIES
Robert M. Stewart
William C. Clow
Point Reyes Bird Observatory
Box 321
Bolinas, Calif. 94924
Lee Jones
Dept. of Zoology
University of California
Los Angeles, Calif. 90024
Jan Larson
Naval Ocean Systems Center
Code 4105
SanDiego,CaHf. 92152
PREPARER'S COMMENTS
It would appear that the widespread destruc-
tion of dense patches of tall brush spaced widely
enough to accomodate this shrike's aggressive ter-
ritorial behavior and requirements for nesting and
observation perches is the main reason for its
poor reproduction and evident decline. Correc-
tion of overbrowsing by livestock, particularly
goats, is the only hope for survival.
LITERATURE CITED/SELECTED
REFERENCES
American Ornithologists' Union. 1957. Check-list
of North American Birds, 5th Ed. Baltimore,
Md. Amer. Omith. Union.
Bent, A. C. 1950. Life histories of North Ameri-
can wagtails, shrikes, vireos, and their allies.
U.S. Nat. Mus. Bull. 197:180-182.
Grinnell, J. 1897. Report on the birds recorded
during a visit to the islands of Santa Barbara,
San Nicolas, and San Clemente, in the spring
of 1897. Pasadena Acad. Sci. Publ. 1:1-21.
Grinnell, J. and A. H. Miller. 1944. The distribu-
tion of the birds of California. Pac. Coast Avi-
fauna 2 7 .
Howell, A. B. 1917. Birds of the islands off the
coast of southern California. Pac. Coast Avi-
fauna 12:88.
Linton, C. B. 1908. Notes from San Clemente
Island. Condor 10:82-86.
Mearns, E. A. 1898. Description of two new birds
from the Santa Barbara Islands, Southern
CaUfornia. Auk 15:258-264.
Miller, A. H. 1931. Systematic revision and natural
history of the American shrikes (Lanius).
Univ. Calif. Publ. Zool. 38:11-242.
Raven, P. H. 1963. A flora of San Clemente
Island, California. Aliso 5:289-347.
Ridgway, R. 1904. The Birds of North and Middle
America. U.S. Nat. Mus. Bull. 50, pt. 3:252.
Stewart, R. M. and W. C. Clow. 1974. Part I. San
Clemente Island, in The status of the Song
Sparrow and Bewick's Wren on San Clemente
Island and Santa Barbara Island, California,
by R. M. Stewart, J. Small, W. C. Clow and
R. P. Henderson. Report to Endangered
Species Off. U.S. Fish and Wildl. Serv. by
Point Reyes Bird Observatory.
Biological Services Program
FWS/OBS-80/01.6
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE PINE BARRENS TREEFROG
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the F,ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of PLndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of F-ndangercd Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Ser\ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.6
March 1980
SELFXTED VKR I EBRA IE ENDANGERED SPECIES
OF THE SEACOAST OE IHE UNITED STATES-
THE PINE BARRENS TREEFROG
A Cooperati\c Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National (Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. VVoodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1 010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Ser\ice
U.S. Department of the Interior
PINE BARRENS TREEFROG
Hyla andersonii Baird
States: Endangered: Florida, South Carolina.
Threatened: New Jersey.
KINGDOM Animalia
CLASS Amphibia
ORDER Salientia (Anura)
FAMILY Hylidae
OTHER COMMON
NAMES Anderson treefrog
DATE:
Entered into SWIS To be determined
Updates 14 October 1976
5 April 1977
1 June 1979
LEGAL STATUS
Federal: Endangered (Florida population only)
(43 FR 18109; 5 Apr. 77).
REASONS FOR CURRENT STATUS
The species occurs in small, isolated popula-
tions scattered along Atlantic and Gulf Coastal
Plains (relict distribution). Many of these sites
are undergoing rapid alteration, including
drainage, development and agricultural mod-
ifications. The resultant loss of habitat con-
stitutes the most serious threat to the Pine Bar-
rens treefrog.
PRIORITY INDEX
Not assigned.
DESCRIPTION
The animal is green dorsally with a narrow
yellow or white stripe along the dorsolateral
edge. Belly is white. A lavender or plum-colored
band extends along the sides from the nostrils
to the hind limbs. Axilla and normally con-
cealed undersides of hind limbs are light orange
or orange spotted. Adults are 30 to 50 mm
(snout-vent length).
Illustrated in color in Dickerson (1969),
Leviton (1972), Conant (1975), Means and
Longden (1976), and Means (1976b). Black
and white photographs in Wright and Wright
(1949). Eggs illustrated by Livezey and Wright
(1947). Noble and Noble (1923) illustrated the
tadpole stages. Gosner and Black (1967) and
Means and Longden (1976) provide audiospec-
trograms of the breeding call.
RANGE
The species distribution has three principal
foci: 1) the Pine Barrens of central New Jersey;
2) the upper Coastal Plain and parts of the lower
Coastal Plain of North and South Carolina; and
3) Okaloosa, Santa Rosa and Walton counties in
the western Florida Panhandle. A single speci-
men from Richmond County, Georgia, and the
holotype from Anderson, South Carolina, may
represent a fourth distributional center. There
is no indication that the distribution is contin-
uous between New Jersey and Florida.
Although individual populations have been
extirpated and total numbers have no doubt
been reduced, the geographic distribution has
probably changed little in historic times.
RANGE MAP
Known distribution is presented on an ac-
companying map.
STATES/COUNTIES
Florida Okaloosa, Santa Rosa, Walton.
Georgia Richmond (?).
New Jersey Atlantic, Burlington, Camden,
. Cape May, Cumberland, Glouces-
ter, Middlesex, Monmouth,
Ocean, Salem.
N. Carolina Bladen, Cumberland, Duplin,
Harnett, Hoke, Johnston, Jones,
Lee, Lenoir, Moore, Onslow,
Pende, Richmond, Sampson,
Scotland, Wayne.
Pennsylvania Delaware (?)
S. Carolina Chesterfield.
HABITAT
Only the breeding habitat is known. Chorus-
ing Pine Barrens treefrogs usually associate with
evergreen shrubs in sphagnaceous seepage bogs
on hillsides below pine-oak ridges. In New Jer-
sey and North Carolina, Atlantic white cedar
[Chamaecyparis thyoides) is a dominant tree
species (Wright and Wright 1949). Means and
Longden (1976) note the absence of cedar but
the constant dominance of titi trees (Cliftonia
and Cyrilla) in Florida localities. The breeding
habitat can best be characterized as shrubby
bogs.
Means and Longden (1976) describe the
habitat of the Pine Barrens treefrog in Florida,
and Means (1976b) provides photographs of
Florida breeding localities. Habitat photographs
for New Jersey and North Carolina are in Noble
and Noble (1923) and Wright (1932).
FOOD AND FORAGING BEHAVIOR
Noble and Noble (1923) reported that the
food habits are not specialized. The treefrogs
eat anything of small size moving in their vi-
cinity, including grasshoppers, beetles, and ants.
Tadpoles feed on algae (Gosner 1959).
SHELTER REQUIREMENTS
Nothing is known about specific require-
ments. Probably like other small treefrogs, they
use small crevices or the undersides of leaves
during the day, and forage at night.
NESTING OR BEDDING
Spawning occurs in sphagnaceous shrub
bogs (see Habitat, above).
RITUAL REQUIREMENTS
Males call during the breeding season from
low bushes and occasionally from the ground.
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WEST LDMCITUftI
The range of the Pine Barrens treefrog is indicated by shading on this map.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Little is known about the species beyond
its specialized and unique breeding habitat des-
cribed above.
POPULATION NUMBERS AND TRENDS
No population estimates are reported.
Means (1976b) reported that since 1972, two
known breeding sites have been rendered un-
suitable because of clearing for improved pas-
ture. A. J. Bullard (personal communication)
reported some North Carolina breeding local-
ities also have recently been destroyed.
Since the discovery of the Pine Barrens tree-
frog in Florida (Christman 1970), some 47
breeding congregations have been located in
three West Florida counties (P. Moler, personal
communication). These are all small seepage
bogs, and none has been found to contain more
than a dozen calling frogs, with most having
fewer than four (P. Moler, personal communica-
tions).
REPRODUCTION
Treefrogs breed from May (April in Florida)
to August. Eggs are laid singly on the bo t-
tom or attached to sphagnum (Wright and
Wright 1949). Eggs hatch in 3 days and may
number up to 200 or more per female. Tadpoles
transform during the summer and frogs probably
reach sexual maturity in 1 year.
Nothing is known concerning natural longe-
vity or survival rates, although a captive survived
7 years (A.J. Bullard, personal communication).
MANAGEMENT AND CONSERVATION
No management or conservation measures
have been instituted other than legal protection
against taking, possessing or molesting the spe-
cies. The relict, disjunct habitat should be pro-
tected from development.
Many of the known breeding localities in
Florida are located within Eglin Air Force Base
and Blackwater River State Forest. Means
(1976b) suggests purchase of some of the re-
maining breeding localities in Florida to prevent
habitat destruction.
Critical Habitat has been designated in
Okaloosa County, Florida (42 FR 58754; 11
Nov. 77): (1) NW'/4 Sec. 35, T4NR22W; (2)
NE'/4 Sec. 27, T4NR22W; (3) SWA Sec. 26,
T5NR23W; (4) NWA Sec. 34, T4NR23W; (5)
NW'/4 Sec. 32, T4NR22W; (6) NWA Sec. 12,
T4NR22W; (7) NE'A Sec. 11, T4NR22W.
AUTHORITIES
A.J. Bullard, Jr.
103 Smith Chapel Road
Mt. Olive, NC 28365
D. Bruce Means
Tall Timbers Research Station
Route l,Box 160
Tallahassee, FL 32303
Paul Moler
Wildlife Research Lab.
Florida Game & Fish Water Fish. Comm.
Gainesville, FL 32601
PREPARER'S COMMENTS
The disjunct distribution of the Pine Barrens
treefrog makes it of considerable interest from
an evolutionary and biogeographic point of
view; it is the least known treefrog in the United
States. There is a pressing need for basic distri-
butional and biological information.
LITERATURE CITED/SELECTED
REFERENCES
Bullard, A. J. 1965. Additional records of the
treefrog, Hyla andersonii, from the coastal
plain of North Carolina. Herpetologica 21
(2):154-155.
Christman, S. P. 191 Q. Hyla andersonii m Flor-
ida. Quart. J. Florida Acad. Sci. 33(1):80.
Conant, R. 1975. A field guide to reptiles and
amphibians of eastern and central North
America. 2nd ed. Houghton Mifflin Co.,
Boston. 429 pp.
Dickerson, M. C. 1969. The frog book. Dover
Publ.Inc, New York. 253 pp.
Gosner, K. L. 1959. Systematic variations in
tadpole teeth with notes on food. Herpeto-
logical5(40):103-120.
Gosner, K. L. and I. H. Black. 1967. Hyla
andersonii Baird. Cat. Am. Amphib. Rep-
tiles. 54.1-54.2.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Ave., Room 250
Gainesville, FL 32601
Leviton. A. 1972. Reptiles and amphibians of
North America. Doubleday and Co., New
York. 250 pp.
Livezey, R. L., and A. H. Wright. 1947. A sy-
noptic key to the salientian eggs of the
United States. Am. Midi. Natur. 37(1): 179-
222.
. 1976a. Pine barrens treefrog. In
Hillestad, H. O., D. B. Means, and W. W.
Baker, eds. Endangered and threatened
vertebrates of the southeastern United
States. Tall Timbers Res. Stn., Misc. Publ. 4.
. 1976b. Endangered species: Pine bar-
rens treefrog. Florida Natur. 49(5): 15-20.
. 1979. Pine Barrens treefrog. Pages 3-4
in R. W. McDiarmid ed.. Rare and endan-
gered biota of Florida, Vol. 3, Amphibians
and reptiles. Univ. Presses of Florida, Gaines-
ville.
Means, D. B.,andC. T. Longden. 1976. Aspects
of the biology and zoogeography of the
pine barrens treefrog [Hyla andersonii) in
northern Florida. Herpetologica 32(2): 117-
130.
Noble, G. K., and R. C. Noble. 1923. The An-
derson treefrog {Hyla andersonii Baird):
Observations on its habits and life history.
Zoologica ll(18):416-455.
Wright, A. H. 1932. Life-histories of the frogs
of Okefenokee Swamp, Georgia. Macmillian
Co., New York. 497 pp.
Wright, A. H., and A. A. Wright. 1949. Hand-
book of frogs and toads of the United States
and Canada. Comstock Publ. Assoc. Cornell
Univ. Press, Ithaca, N.Y. 640 pp.
Biological Services Program
FWS/OBS-80/01.7
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE RED COCKADED WOODPECKER
Fish and Wildlife Service
U.S. Department of the Interior
prefacp:
The purpose of this series of species accounts is to provide resource managers and the
pubhc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aie not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as ainciuicd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Kndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SUIS should he directed to:
Office of Endangered Species
U.S. Fish and Wildlife Sewice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal F,cosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
FWS/OBS-80/01.7
March 1980
SliLKCn:!) VI:R 1 EBRA 1 K KNDANGERP:D SPKCIES
OF IHi: SKACOAS I OF 1 HE UNFFED S FA TES-
THE RED COCKADED WOODPECKER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems i'eam,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidcll, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and U'iUllifc Ser\ice
U.S. Department of the Interior
RED-COCKADED WOODPECKER
Picoides borealis Vieillot
KINGDOM Animalia
CLASS Aves
ORDER Piciformes
FAMILY Picidae
OTHER COMMON NAMES none
DATE
Entered into SWIS to be determined
Updates 17 October 1976, 1 March 1977
LEGAL STATUS
Federal: Endangered (35 FR 16047, 13 Octo-
ber 1970).
States: Endangered: Florida, Georgia, Missis-
sippi, South Carolina, Tennessee,
Texas. Extirpated: Missouri.
REASONS FOR CURRENT STATUS
The primary reason for the current status of
the red-cockaded woodpecker is a decrease in
quantity and quality of suitable habitat, primarily
due to the short-term-rotation timber management
currently being practiced in the Southeast. Short-
term-rotation prevents the development of ma-
ture, diseased pine trees that are necessary for
roosting and nesting.
PRIORITY INDEX
Not assigned.
DESCRIPTION
The red-cockaded woodpecker is small, 18 to
20 cm long, with awingspanof 35 to 38 cm. The
cap and nape are black, surrounding a large, white
cheek patch. Wings and back are black, horizon-
tally barred with white. Underparts are whitish
with dark spots on the flanks. The adult male has
a small red streak, the 'cockade,' on each side of
the black cap, but this is rarely visible except
when the bird is displaying. The most distin-
guishing field mark is the black cap and nape sur-
rounding the large white cheek patch.
The species is illustrated in Robbins et al.
(1966). Black and white photos appear in Mur-
phey (1939) and Jackson et al. (1976b).
RANGE
Present range (nonmigratory species) includes
the following areas: Alabama, southern Arkan-
sas, Florida, Georgia, Cumberland Plateau region
of Kentucky, Louisiana, southeastern Maryland,
Mississippi, eastern North Carolina, southeastern
Oklahoma, South Carolina, eastern Tennessee,
eastern Texas, and southeastern Virginia.
Former range included the entire States of
Alabama, Arkansas, Florida, Georgia, Kentucky,
Louisiana, Maryland, Mississippi, Missouri, New
Jersey, North Carolina, Oklahoma, South Caro-
lina, Tennessee, Texas, and Virginia (Jackson
1971).
In the Southeast, there are 76.7 million ha of
commercial forest in major pine types. Approxi-
mately 10.4 milHon ha, of which 0.9 million ha
are in public ownership, are suitable for red-cock-
aded woodpeckers (Czuhai 1971).
RANGE MAP
Known distribution is indicated by shading.
Dots represent populations on publicly owned or
managed lands; large dots represent at least 10
clans, while small dots represent 1 to 9 clans.
STATES/COUNTIES
Alabama Baldwin, Bibb, Calhoun, Chilton, Clay,
Cleburne, Covington, Escambia, Hale,
Jefferson, Lawrence, Macon, Perry,
Shelby, St. Clair, Talledega, Tusca-
loosa, Winston.
Arkansas Ashley, Calhoun, Clark, Columbia,
Hempstead, Lafayette, Monroe, Oua-
chita, Polk, Scott, Union.
Florida Alachua, Baker, Bay, Brevard, Char-
lotte, Citrus, Clay, Columbia, Duval,
Franklin, Glades, Gulf, Hernando,
Highlands, Lee, Leon, Levy, Liberty,
Marion, Martin, Okaloosa, Okeecho-
bee, Osceola, Palm Beach, Pasco, St.
Johns, Santa Rosa, Wakulla, Walton.
Georgia Appling, Baldwin, Brantley, Charlton,
Chattahoochee, Clarke, Clinch, Deca-
tur, Floyd, Glynn, Grady, Harris, Jas-
per, Jenkins, Jones, Pierce, Putnam,
Screven, Tattnall, Telfair, Thomas,
Toombs, Turner, Ware, Washington,
Wilkes.
Kentucky Laurel, McCreary.
Louisiana Allen, Beauregard, Bienville, Bossier,
(Parishes) Caddo, Calcasieu, Catahoula, Clai-
borne, Grant, La Salle, Morehouse,
Natchitoches, Ouachita, Rapides, St.
Tammany, Tangipahoa, Union, Ver-
non, Washington.
Maryland Dorchester, Worcester (?).
Mississippi Choctaw, Copiah, Franklin, Harrison,
Hinds, Jackson, Jones, Lafayette, Lau-
derdale, Leake, Madison, Marion,
Noxubee, Oktibbeha, Pearl River,
Scott, Smith, Stone, Wayne, Wilkin-
son, Winston.
North
Carolina
Beaufort, Bertie, Bladen, Brunswick,
Cumberland, Gates, Harnett, Hertford,
Hoke, Hyde, Jones, Montgomery,
Moore, Northampton, Onslow, Pam-
lico, Perquimans, Pitt, Richmond,
Wake, Wayne.
Oklahoma Bryan, Latimer, Le Flore, McCurtain,
Pittsburg, Pushmataha.
South
Carolina
Aiken, Barnwell, Beaufort, Berkeley,
Calhoun, Charleston, Chesterfield,
Clarendon, Colleton, Darlington, Dil-
lon, Dorchester, Edgefield, Florence,
Georgetown, Hampton, Horry, Lau-
rens, Lee, Lexington, Orangeburg,
Richland, Sumter, Williamsburg.
Tennessee Blount, Campbell, Cumberland,
Morgan.
Texas Angelina, Cass, Cherokee, Hardin,
L» if.
CONIC PROJECTION
Shading on this map depicts the present range of the red-cockaded woodpecker in southeastern
United States. Dots represent populations on public lands, small dots, 1-9 clans; large dots, 10 or
more clans.
3
Houston, Jasper, Montgomery, Nacog-
doches, Newton, Polk, Sabine, San
Augustine, San Jacinto, Shelby, Trini-
ty, Tyler, Walker.
Virginia Prince Georges (?), Southampton (?),
Surry, Sussex (?).
HABITAT
Mature to over-mature southern pines are the
best habitat for roosting and nesting. Longleaf
{Pinus palustris), loblolly {Pinus taeda), shortleaf
(Pinus echinata), slash {Pinus elliottii), and pond
(Pinus rigida) pines are used, depending on local-
ity (Thompson and Baker 1971). Younger pines
(10 to 21cm d.b.h.) sometimes are used for forag-
ing, especially after an unusually hot burning of
the area which results in killed and weakened
trees which offer high-grade feeding areas (Beck-
ett 1974). Fire, an essential element of this habitat,
prevents thick understory. Red-cockaded wood-
peckers are discouraged by dense stands of pine
saplings and thick hardwood understories.
Although over-mature pines are required for
nesting and roosting, the red-cockaded woodpeck-
ers will forage in a variety of habitat types
depending on food availability and proximity to
cavity-tree sites.
The Florida Game and Fresh Water Fish Com-
mission (1976) and Nesbitt et al. (1978), have
studied fall habitat usage in a Florida flatwoods
community. Percent of total foraging time spent
in each habitat type was:
Pond /slash pine flatwoods 43.8
Longleaf pine flatwoods 38.2
Slash pine plantation 9.4
Bayhead/pond borders,
cypress domes and others 6.5
Roosting areas 2.2
FOOD AND FORAGING BEHAVIOR
The red-cockaded woodpecker feeds chiefly
on wood-boring insects, ants, beetles, grubs, grass-
hoppers, crickets, and caterpillers (Murphy 1939).
Analysis of 99 stomach content samples from the
Southeast revealed a diet of 84% insects and 16%
plant material (Beal 1911). Plant fruits eaten
include (Murphy 1939, Ligon 1970, Baker 1971a):
Wax myrtle (Myrica cerifera)
Magnolia {Magnolia grandi flora)
Poison ivy {Rhus radicans)
Wild grape ( Vitis sp)
Pokeberry {Phytolacca americana)
Blueberry {Vaccinium spp.)
Wild cherry {Prunus serotina)
Black gum {Nyssa sylvatica)
Woodpeckers prey on the com earworm {Helicov-
era armigeva) when corn fields are nearby. (Baker
1971a).
Foraging on a tree trunk it uses the bill and/or
feet to pry off pieces of bark, exposing insects.
One technique involves backing down the tree,
flaking off bark with the feet while catching prey
with the bill.
SHELTER REQUIREMENTS
Roosting requires a cavity in a living pine tree.
A sample of 259 cavity trees in east central Missis-
sippi ranged from 40 to 116 years in age, with a
mean of 75.9 years (Jackson 1977). Usually the
tree is infected with red heart {Fames pini), a
fungus disease which weakens the heartwood
(Affeltranger 1971). Jackson (1977) suggests a
cavity can provide an infection site for red heart
disease. After the heartwood is weakened, an
excavation can be completed. A tree may have
several roosting cavities.
The distinguishing characteristic of a cavity
tree is the flow of sap that glazes the tree around
the opening. Birds initiate and maintain sap flow
by pecking out chunks of bark and cambium,
forming 'resin wells.' Active sap flow may pre-
vent predators and/or competitors from entering
the roost cavity (Ligon 1970, Dennis 1971, Jack-
son 1974).
NESTING OR BEDDING
The adult male's roost cavity is used for nest-
ing. The bottom of the gourd-shaped cavity is
covered with chips and debris; sap often smears
onto the eggs (Murphey 1939).
RITUAL REQUIREMENTS
Mating behavior, as observed by Crosby
(1971a): the female landed near the male and
raised her tail. The male mounted from her left
side, fell, separated, and resumed feeding after 3
seconds.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Clan (see Reproduction) home-range require-
ments depend on three variables: clan size, habitat
type, and season. Crosby (1971b) followed two
color-marked adult pairs in north central Florida
during the spring. This study produced an average
estimate of 17.2 ha per adult pair. Baker (1971b),
observing color-marked individuals in north Flo-
rida, estimated home range of an eight-bird clan
to be 65.6 ha during the summer.
Lay and Russell (1970) divided known areas
by the number of clans present to estimate home
ranges of 26.7 ha and 67.7 ha per clan in two
eastern Texas forests. Beckett (1974) used the
same technique to estimate 86.2 ha per clan in a
South Carolina forest.
Skorupa and McFarlane (1979) compared
summer and winter home range requirements of
two adult pairs. Results indicated a 112% and
71% increase in winter forage range requirements.
Fall home range requirements in central Florida
were determined by the Florida Game and Fresh
Water Fish Commission (1976) and Nesbitt et al.
(unpublished). A bird in each clan was equipped
with a miniature radio transmitter. Results indi-
cated an average home range of 69.8 ha.
POPULATION NUMBERS AND TRENDS
Estimates of total population range from
3,000 to 10,000 (U.S. Department of the Interior
1973).
The number of colonies (groups of cavity
trees) on public lands is between 2,800 and 3,600;
of these, an estimated 2,500 are active (see Man-
agement and Conservation).
Subpopulations exist on ecological islands of
suitable habitat. The fractioning of distribution
results in reduced gene flow between subpopu-
lations, as well as reduced ability to disperse
into and occupy suitable habitats that might be-
come available in the future (Jackson 1976).
Colonies were surveyed in 10 southeastern
States in 1969-70. Thompson (1976) reported the
status of 312 colonies resurveyed in 1973-74.
Only 271 remained active, a 13.3% loss, at an an-
nual loss rate of 3.5%. For the 4-year period,
losses on Federal, State, and private lands were,
respectively, 8.7%, 27.3%, and 22. 9%. The largest
factor in this reduction was timber harvest.
Clearcutting and short-term-rotation timber
management have virtually eliminated the species
from Kentucky, where Jackson et al. (1976b)
found red-cockaded woodpeckers at only two
locations.
As trends continue toward short-term timber
rotation throughout the Southeast, the species
will become increasingly endangered.
REPRODUCTION
The species nests during April, May, and June.
Clutch size is usually two to four eggs. The incu-
bation period is approximately 10 days; duration
of the nestling stage is about 27 days (Jackson et
al. 1976a, W. W. Baker personal communication).
Usually one or two young are fledged. Lon-
gevity and survival rates are not known.
A family unit is called a 'clan,' and consists of
two pair-bonded adults, the young of the year,
and sometimes several 'helpers' that are young of
previous years.
MANAGEMENT AND CONSERVATION
A Recovery Team has been appointed and a
draft recovery plan submitted to the U.S. Fish
and Wildlife Service for review.
Jackson (1976) suggests that properly man-
aging interstate highway right-of-ways might re-
unite fragmented populations.
The following tabulation presents population
estimates of species numbers on publicly owned
or managed lands. Unless otherwise noted, all
data were obtained by personal communication
with the listed observers. (Key: NF = National
Forest; NWR = National Wildlife Refuge; SF =
State Forest; SF = State Park; WMA = WildHfe
Management Area.)
State
Location, estimate, and reference
Alabama Conecuh NF; 22 colonies; O. Stewart
Talledega NF; 30 colonies; O.Stewart
Tuskegee NF; 3 colonies; O. Stewart
W. B. Bankhead NF; 6 colonies; O.
Stewart
Felsenthal NWR; 20-30 colonies; J.
Howe
Arkansas Ouachita NF; no estimate available.
Florida Apalachicola NF; 227 known; 647
estimated;D. Bethancourt
Aucilla WMA; present; no estimates.
S. Stafford
Austin Gary Memorial Forest (Univ.
of Florida) ; present; exact number not
known; D. Hirth
Big Cypress Swamp; 1 cl£m;J. Kern
Blackwater River SF;approx. 25 colo-
nies; J. Bethea
Camp Blanding; 20-25 active colonies;
J. Schatz
Gary SF; approx. 6 colonies; J.
Bethea
Eglin Air Force Base; approx 55 colo-
nies; W. Alford,
Fisheating Creek WMA; at least 2 ac-
tive colonies ;T. Breault
Johnathan Dickinson SP; 1 active co-
lony; J. Stevenson
J. W. Corbett WMA; 20-30 birds; N.
Eichholz
Nassau WMA; present, number not
known; S. Stafford
Ocala NF; 30 colonies known, 48 es-
timated; D Bethancourt
Osceola NF; 32 known active colonies
50 estimated; pers. observation.
Saint Marks NWR. 3 active colonies;
J. White
Secil Webb WMA; more than 20 active
colonics; L. Campbell
Three Lakes WMA; at least 5 colonies;
R. McCracken
Withlacoochee SF; approx. 27
colonies; J. Bethea
Georgia Baldwin SF; present, exact no. not
known; J. Hammond
Dixon Memorial SF; present, exact
no. not known; J. Hammond
Ft. Benning; at least 80 colonies; J.
Medcaff
Fort Gordon; 4 birds (est.); H. Ford-
ham
Oconee NF; 24 active colonies; T.
Richards
Okefenokee NWR; 33 colonies; J.
Eadia
Piedmont NWR; 70 colonies; S.
Pagans
Daniel Boone NF; 2 or 3 active colo-
nies; W. Williams.
Louisiana D'Arbonne NWR; present, exact no.
not known; J. Howe
Kisatchie NF; 354 colonies; R. Wilson
Fort Polk; at least 200 colonies; R.
Aycock
Maryland Blackwater NWR; 20 birds (est.) B.
Julian
Mississippi Bienville NF; 101 colonies; G. Sirmon
Copiah County WMA; present, exact
no. not known. W. Turcotte
DeSoto NF; 88 colonies; G. Sirmon
Homochitto NF; 84 colonies; G. Sir-
mon
Marion County WMA; present, exact
no. not known. W. Turcotte.
Natchez Trace Pkwy.; 1 active colony;
W. Turcotte
Noxubee NWR. 71 colonies; T.
McDaniel
North
Carolina Blanden Lakes SF; at least 2
colonies; S. Taylor
Crotan NF; 3-35 active colonies; B.
Sanders
Fort Bragg; over 100 active colonies;
B. Sanders
Mattamuskcet NWR; 6 colonies; J.
Roberts
Pee Dee NWR; Present, exact no. not
known; J. Hollowman
Uwharrie NF; 1-3 active colonies; B.
Sanders
Oklahoma McCurtain County; 48-53 active colo-
nies (Wood 1975).
South
Carolina Carolina Sandhills NWR; over 90 colo-
nies; M. Hurdle
Cheraw SP; 16 colonies; R. Hendrick
Francis Marion NF; 500 active colo-
nies (est.) D. Urbston
Givhans Ferry SP; at least 1 colony;
R. Hendrick
Harbison SF; 3 colonies; J. Tiller
Huntington Beach SP; 1 colony; R.
Hendrick
Lee SP; at least 1 colony; R. Hendrick
Little Pee Dee SP; 5 colonies; R.
Hendrick
Manchester SF; 50 colonies; J. Tiller
Sand Hills SF; 55 colonies; M. Hurdle
Santee NWR; 4 colonies; C. Strickland
Santee State Resort; 3 colonies; R.
Hendrick
Savannah River Plant, U.S. Dept. of
Energy; 14 colonies; D. Roth.
Sesquicentennial SP; at least 2 colo-
nies; R. Hendrick
Sumter NF; 15 active colonies; D.
Urbston
Tennessee Catoosa WMA; 6 colonies; B. Yambert
Great Smoky Mountains NP; sightings
in Cade's Cove area; J. Collier
Texas Angelina NF; min. 46 colonies, max.
60; D. Gates
Davy Crocket NF; min. 245 colonies;
max. 29 1;D. Gates
Sabine NF; min. 104 colonies, max.
120; D. Gates
Sam Houston NF; min. 87 colonies,
max. 225; D. Gates
AUTHORITIES
W. Wilson Baker (Recovery Team)
Tall Timbers Research Station
Route l,Box 160
Tallahasse, FL 32303
Ted Beckett
Magnolia Gardens
Route 4
Charleston, SC 29407
Vernon Carter (Recovery Team)
Regional Forester
Fish and Wildlife Service
17 Executive Park Drive, N.E.
Atlanta, GA 30329
Thad Cherry (Recovery Team)
Wildlife Research Supervisor
Weyerhauser Company
P.G. Box 1391
New Bern, NC 28560
Melvin Hopkins (Recovery Team)
National Forests in North Carolina
P.O. Box 2750
Asheville,NC 28802
Jerome A.Jackson (Recovery Team Leader)
Department of Zoology
Mississippi State University
P.O. Box Z
Mississippi State, MS 39762
Daniel W. Lay
Texas Parks and Wildlife Department
Box 4608, SFA
Nacogdoches, TX 75961
Michael R. Lennartz
Department of Forestry
Southeastern Forest Experiment Station
Clemson University
Clemson,SC 29631
Teddy E. Lynn, Jr.
International Paper Company
Georgetown Woodlands Region
Georgetown, SC 29440
Robert W. McFarlane
Savannah River Ecology Lab
P.O. Drawer E
Aiken, SC 29801
Richard L. Thompson
Florida Management Biologist
Fish and Wildlife Service
P.O. Box 190
Tallahassee, FL 32302
PREPARER'S COMMENTS
Cavities produced by red-cockaded wood-
peckers are used by other species, including the
honey bee (Apis mellifera), rat snake {Elaphe ob-
soleta), red-bellied woodpecker (Melanerpes caro-
inus), red-headed woodpecker {Melanerpes ery-
throcephalus), white-breasted nuthatch {Sitta
car o line nsis), tufted titmouse (Parus bicolor),
eastern bluebird (Sialia sialis), common flicker
(Colaptes auratus), pileated woodpecker {Dryo-
copus pileatus), great crested flycatcher (Myiar-
chus crinitus), screech owl (Otus asio), wood
duck (Aix sponsa), starling (Sturnus vulgaris),
gray squirrel (Sciurus carolinensis), fox squirrel
{Sciurus niger), and flying squirrel {Glaucomys
volans) (Baker 1971b, Jackson et al. 1976a).
The fragmentation of nesting habitat, and
thus of the woodpecker's distribution, may be
leading to reduced gene flow between populations
and a loss of ability to colonize new habitat that
may develop.
Subjects needing further investigation include
seasonal variation of food habits and home range
requirements, causes of high nestling mortality
(relative to other woodpeckers), pesticide levels
and their significance, and reintroduction tech-
niques.
LITERATURE CITED/SELECTED
REFERENCES
Affeltranger, C. 1971. The red heart disease of
southern pines. Pages 96-99 in R. L. Thomp-
son, ed. The ecology and management of the
red-cockaded woodpecker. Proc. Symp. Oke-
fenokee Natl. Wildl. Refuge, Folkston, Geor-
gia. May 26-27. U.S. Bur. Sport Fish. Wild,
and Tall Timbers Res. Stn., Tallahassee, Flo-
rida. 188 pp.
Baker, W. W. 1971a. Observations on the food ha-
bits of the red-cockaded woodpecker. Pages
100-107 in R. L. Thompson, ed. The ecology
and management of the red-cockaded wood-
pecker. Proc. Symp. Okefenokee Natl. Wildl.
Refuge, Folkston, Georgia. May 26-27. U.S.
Bur. Sport Fish. Wildl. and Tall Timbers Res.
Stn., Tallahassee, Florida. 188 pp.
1971b. Progress report on life history
studies of the red-cockaded woodpecker at
Tall Timbers Research Station. Pages 44-59 in
R. L. Thompson, ed. The ecology and man-
agement of the red-cockaded woodpecker.
Proc. Symp. Okefenokee Natl. Wildl. Refuge,
Folkston, Georgia. May 26-27. U.S. Bur.
Sport Fish. Wildl. and Tall Timbers Res. Stn.,
Tallahassee, Florida. 188 pp.
Beal, F. E. L. 1911. Food of the woodpeckers in
the United States. U. S. Dep. Agric. Biol. Sur-
vey Bull. 37:1-64.
Beckett, T. A. 1974. Habitat acreage requirements
of the red-cockaded woodpecker. EBBA News
37:3-7.
Crosby, G. T. 1971a. Ecology of the red-cockaded
woodpecker in the nesting season. M.S. Thesis.
Univ. of Florida, Gainesville. 45 pp.
1971b. Home range characteristics of the
red-cockaded woodpecker in north central
Florida. Pages 70-73 in R. L. Thompson, ed.
The ecology and management of the red-cock-
aded woodpecker. Proc. Symp. Okefenokee
Natl. Wildl. Refuge, Folkston, Georgia. May
26-27. U.S. Bur. Sport Fish. Wildl. and Tall
Timbers Res. Stn., Tallahassee, Florida. 188
pp.
Czuhai, E. 1971. Synoptic review of forest
resources and use within the range of the
red-cockaded woodpecker. Pages 108-124 in
R. L. Thompson, ed. The ecology and man-
agement of the red-cockaded woodpecker.
Proc. Symp. Okefenokee Natl. Wildl. Refuge,
Folkston, Georgia. May 26-27; U.S. Bur.
Sport Fish. Wildl. and Tall Timbers Res. Stn.,
Tallahassee, Florida. 188 pp.
Dennis, J. V. 1971. Utilization of pine resin by
the red-cockaded woodpecker and its effec-
tiveness in protecting roosting and nesting
sites. Pages 78-86 in R. L. Thompson, ed.
Proc. Symp. Okefenokee Natl. Wildl. Refuge,
Folkston, Georgia. May 26-27. U.S. Bur.
Sport Fish. Wildl. and Tall Timbers Res. Stn.,
Tallahassee, Florida. 188 pp.
Florida Game and Fresh Water Fish Commission.
1976. Red-cockaded woodpecker. Pages 79-
87 in Cross Florida Barge Canal restudy
report: endangered, threatened, rare, special
concern, status undetermined, and biologi-
cally sensitive species. U.S. Fish Wildl. Serv.
unpublished. 267 pp.
Jackson, J. A. 1971. The evolution, taxonomy,
distribution, past populations and current sta-
tus of the red-cockaded woodpecker. Pages 4-
29 in R. L. Thompson, ed. The ecology and
management of the red-cockaded woodpecker.
Proc. Symp. Okefenokee Natl. Wildl. Refuge,
Folkston, Georgia. May 26-27. U.S. Bur.
Sport Fish. Wildl. and Tall Timbers Res. Stn.,
Tallahassee, Florida. 188 pp.
1974. Gray rat snakes versus red-
cockaded woodpeckers: predator prey adap-
tations. Auk 91(2):342-347.
1976. Rights-of-way management for an
endangered species — the red-cockaded wood-
pecker. Pages 247-252 in Proc. Symp. Envi-
ronmental Concerns in Rights-of-Way Manage-
ment. Mississippi State Univ.,
Jackson, J. A., W. W. Backer, V. Carter, T. Cherry,
and M. L. Hopkins. 1976a. Recovery plan for
the red-cockaded woodpecker. 23 pp.
Jackson, J. A., and R. L. Thompson. 1971. A
glossary of terms used in association with the
red-cockaded woodpecker. Pages 187-188 in
R. L. Thompson, ed. The ecology and man-
agement of the red-cockaded woodpecker.
Proc. Symp. Okefenokee Natl. Wildl. Refuge,
Folkston, Georgia. May 26-27. U.S. Bur.
Sport Fish. Wildl. and Tall Timbers Res. Stn.,
Tallahassee, Florida. 188 pp.
Jackson, J. A., R. Weeks, and P. Shindala. 1976b.
The present status and future of red-cockaded
woodpeckers in Kentucky. Kentucky Warbler
52(4)73-80.
Lay, D. W., and D. N. Russell. 1970. Notes on the
red-cockaded woodpecker in Texas. Auk
87(4):781-786.
Ligon, J. D. 1968. Sexual differences in foraging
behavior in two species of Dendrocopus
woodpeckers. Auk 85(2):203-215.
1970. Behavior and breeding biology of
the red-cockaded woodpecker. Auk 87(2):
255-278.
Murphey, E. E. 1939. Red-cockaded woodpecker.
Pages 75-79 in A. C. Bent, ed. North Ameri-
can woodpeckers. Dover Publication, Inc.
New York.
Nesbitt, S. A., D. T. Gilbert, and D. B. Barbour.
1978. Red-cockaded woodpecker fall move-
ments in a Florida flatwoods community.
Auk 95(1)145-151.
Robbins, C. S., R. Bruun, H. S. Zim, and A.
Singer. 1966. A guide to field identification.
Birds of North America. Golden Press, New
York. 340 pp.
Skorupa, J. A., and M. W. McFarlane. 1979. Sea-
sonal variation in foraging territory of red-
cockaded woodpeckers. Wilson Bull. 88(4):
662-665.
Thompson, R. L. 1976. Changes in status of red-
cockaded woodpecker colonies. Wilson Bull.
88(3):491-492.
Thompson, R. L., and W. W. Baker. 1971. A sur-
vey of red-cockaded woodpecker nesting habi-
tat requirements. Pages 170-186 in R. L.
Thompson, ed. The ecology and management
of the red-cockaded woodpecker. Proc. Symp.
Okefenokee Natl. Wildl. Refuge, Folkston,
Georgia. May 26-27. U.S. Bur. Sport Fish.
Wildl. and Tall Timbers Res. Stn., Tallahassee,
Florida. 188 pp.
U.S. Department of the Interior. 1973. Threa-
tened wildlife of the United States. Com-
piled by Office of Endangered Species and
International Activities, Bur. Sport Fish.
WUdl. Resour. Publ. 114. U.S. Gov. Printing
Office, Washington, D.C. 289 pp.
Wood, D. A. 1975. Status, habitat, home range,
and notes on the behavior of the red-
cockaded woodpecker in Oklahoma. M.S.
Thesis. Oklahoma State Univ. 60 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.8
March 1980
Selected Vertebrate
Endangered Species
Of the Seacoast of the
United States-
IVORYBILLED
WOODPECKER
^r*
W%
h
» W'
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amcntlcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.8
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
IVORY BILLED WOODPECKER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
V
CREDIT: LAURA DIXON, U.S. FISH AND WILDLIFE SERVICE
IVORY BILLED WOODPECKER
Camp ephilus principalis principalis (Linnaeus)
Campephiliis principalis bairdii Cassin
KINGDOM Animalia
CLASS Aves
ORDER Piciformes
FAMILY Picidae
OTHER NAMES American Ivory -billed
Woodpecker, Kent, Ivory-bill, Pearly-
bill, Log-god, Log-cock, White-back
Woodpecker, Poule de Bois, Grand
Pique-bois, Lord -god, Cuban Ivory-
billed Woodpecker, Carpentero Real.
DATE
Entered into SWIS to be determined
Updates to be determined
LEGAL STATUS
Federal: Endangered: (42 FR 36428, 14 July
1977).
States: Endangered: North Carolina; Pro-
tected by Migratory Bird Treaty Act
of 3 July 1918 (40 Stat. 755; 16
U.S.C. 703-711) as amended 3 Decem-
ber 1969. Public Law 91-135. South
Carolina; Georgia; Florida; Alabama;
Mississippi; Texas; Arkansas (black
List, extinct or near extinction).
REASONS FOR CURRENT STATUS
C. p. principalis is probably very close to ex-
tinction because of the scarcity of suitable habitat
(U.S. Fish and Wildlife Service 1973). Ivory-bHls
disappeared when the mature forests they in-
habited were cut over, and the larger timber es-
sential for providing adequate food supply was
removed. In many cases, their disappearance al-
most coincided with logging operations; in other
cases there was no close correlation, but there are
not records of ivory -bills remaining very long after
cutting of forests. Their disappearance in the
Suwannee River region of Florida is believed to
have been due to excessive collecting rather than
logging (Tanner 1942). Herbert Stoddard and
Whitney Eastman shared the conviction that ivory-
bills were reduced to near extinction by gun pres-
sure, but that with large land holdings coming
into possession of ranchers and timber companies,
the bird had a better chance of survivzd (Eastman
1958). Some shooting for sport or curiosity is
known to have occurred, particularly before pas-
sage of protective laws. Although direct killing by
man is not as important as loss of habitat by log-'
ging in reducing their numbers, it could be impor-
tant locally when only a very few are left (Tanner
1942).
C. p. bairdii, the Cuban Ivory-billed Wood-
pecker was once widely distributed in Cuba, but
is now limited to remote areas in Oriente Province.
Land clearing during the speculative sugar boom
of the early 1900's and later lumbering activities
extirpated it from its former range. By far the ma-
jor predator is man, as these woodpeckers are
prized as food by the natives (Lamb 1957).
PRIORITY INDEX
75
DESCRIPTION
C. p. principalis is a very large (crow-sized)
woodpecker, 50 cm long, with 46 cm wingspan.
Its shape is long and slender, with a long tapering
tail (Audubon 1842, Tanner 1942). Both sexes
are mostly glossy blue-black, with tail and pri-
maries duller black. There is a narrow white stripe
on each side of neck, starting below the eye and
continuing down to the folded secondaries, which
are conspicuously white, as are all but 5 of the
outermost primaries and the under-wing coverts.
This makes a large white patch on the rear half of
the wing, narrowing toward the tip. The nasal
plumes and anterior edge of lores are white. The
crest is red in the male, black in the femede. The
iris is pale, clear lemon-yellow. Tarsi and toes are
light gray. The bill is large and ivory-white (Ridg-
way 1914, Tanner 1942). The best field identific-
ation character is the large white patch on the
wing, conspicuous when the bird is perched. Call
notes are a nasal "kent, kent" (Tanner 1942).
Measurements.- Adult male (15): wing, 240-
263 (255.8 mm); tail, 147-160.5 (154.4); culmen,
63-72.5 (68.2); tarsus, 42.5-46 (44.2); outer an-
terior toe, 30-34 (32.1). Aduh female (11): wing,
240-262 (256.4 mm); tail, 151-166 (159.5); cul-
men, 61-67.5 (64.3); tarsus, 40.5-44 (42.6); outer
anterior toe, 30-33.5 (31.7) (Ridgway 1914).
The eggs are pure china-white, exceedingly
glossy, and more pointed than most woodpeckers.
Measurements (13 eggs) 34.5 x 23.6 to 36.8 x
26.9 (av. 34.8 x 25.2 mm) (Bendire 1895).
C. p. bairdii is similar to C. p. principalis but
slightly smaller; the bill is decidedly smaller; nasal
tufts much smaller; and white stripe on side of
head continues nearly to the base of the bill.
Measurements.- Adult male (2): wing, 236-
250 (243 mm); tail, 137-154 (145.2); culmen,
59-61 (60); tarsus, 40-42 (41); outer anterior toe,
30.5-31.5 (31). Adult female (2): wing, 240-255
(247.5 mm); tail, 159.5-165.5 (162.5); culmen,
58-60 (59); tarsus, 41; outer anterior toe, 30.5-
31.5 (31) (Ridgway 1914).
RANGE
C. p. principalis formerly was a resident in the
bottomlands and swampy forests of southeastern
United States from northeastern Texas, south-
eastern Oklahoma, northeastern Arkansas, south-
eastern Missouri, southeastern Illinois, southern
Indiana and southeastern North Carolina, south-
ward to the Brazos River, Texas, the Gulf Coast
and southern Florida (A.O.U. 1957, Bent 1931,
Tanner 1942). There are probably a few still in
southeastern Texas, Louisiana, Florida and South
Carolina (Dennis Unpubl.).
C. p. bairdii was formerly distributed widely
in Cuba, but is now apparently confined to north-
eastern Oriente Province (from the pinares of
Mayari eastward); it was seen at an elevation of ap-
proximately 213 m in 1942 (Bond 1950). Records
in the 20th century come only from northern
Oriente Province in the extreme eastern part of
Cuba. The Sierra del Cristo region apparently had
ivory-bills until at least 1920 (Dennis 1948). A
few pairs were thought by Barbour to be in the
Sierra de Nipe near Mayari (1943); Abelardo
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Moreno in Lamb (1957) found three ivory -bills in
the Sierra de Moa region east of Sierra del Crista].
RANGE MAP
(Adapted from Tanner 1942, Dennis 1967,
Lamb 1957).
STATES/COUNTIES: (from Tanner 1942)
North Ciirolina: New Hanover.
South Carolina: Chesterfield, Clarendon, Berk-
ley, Georgetown, Charlestown,
Barnwell, Allendale, Beaufort.
Georgia: Tattnall, Pierce, Ware.
Florida: Liberty, Gulf, Wakulla, Leon,
Jefferson, Taylor, Dixie, Suwan-
nee, Levy, Baker, Clay, Alachua,
Putnam, Marion, Citrus, Sump-
ter, Volusia, Seminole, Her-
nando, Sumpter, Lake, Polk,
Orange, Osceola, Brevard, Pinel-
las, Clearwater, Hillsborough,
Manatee, Highlands, Okeecho-
bee, Desoto, Charlotte, Collier,
Lee.
Mississippi:
Alabama:
Louisiana:
Monroe, Clay, Hancock, Harri-
son, Jackson, Warren, Bolivar.
Hale, Dallas, Wilcox, Pike,
Marengo.
Morehouse Parish, West Carroll
Parish, East Carroll Parish, Mad-
ison Parish, Franklin Parish,
Tensas Parish, Concordia Parish,
West Feliciana Parish. St. Mar-
tin Parish, Iberville Parish, Iberia
Parish, Lafourch Parish.
White.
Stoddard.
Fulton.
Mississippi, Jackson, Poinsett,
Phillips.
Atoka, Bryan.
Cooke, Jasper, Harris, Brazoria.
Illinois:
Missouri:
Kentucky:
Arkansas:
Oklahoma:
Texas:
HABITAT
C. p. principalis resides in swampy forests, es-
pecially the large bottomland river swamps of the
coastal plain and Mississippi Delta and the cypress
swamps of Florida. It was most abundant in the
lower bottoms of the Mississippi River, the rivers
of South Carolina and Georgia, and in Florida
swamps and swampy hammocks. Habitats were
divided by Tanner (1942) into three main regional
types: (1) Bottomlands of the Mississippi Delta;
(2) River bottoms outside the Mississippi Delta;
(3) the Florida region.
The Mississippi Delta, the alluvial flood plain
of the Mississippi River, stretches from the junc-
tion of the Ohio and Mississippi to the Gulf of
Mexico, and is from 64 to 130 km wide. Ivory-
biU distribution was evidently limited to the higher
parts of the "first bottoms," which were rarely
covered with water more than a few months of
the year. The soil is a moderately well-drained al-
luvial clay. The forest is a sweet gum-oak associa-
tion with dominant trees being sweet gum, bot-
tomland red oak and green ash. Associated with
them are willow oak, water oak, over cup oak,
American elm, cedar elm, hackberry, water
hickory, and pecan. Ivory-bills fed mostly upon
sweet gvim and bottomland red oak.
The river bottoms outside the Mississippi
Delta are the floodplains of the larger rivers of the
southeastern United States, flowing either into
the Atlantic or Gulf of Mexico. The soil is alluvial
clay that is usually flooded annually. Ivory-bills
have been observed in those bottomlands in both
oak -sweet gum and cypress-tupelo forests but were
most common in oat-sweet gum. These habitats
differ from those of the Mississippi Delta primar-
ily in their smaller area, but also in the predomin-
ance of laurel oak and water oak. Dominant for-
est species on dryer sites of these bottoms are
sweet gum, laurel oak and water oak and associated
with their dominants are overcup oak, cow oak,
water hickory, green ash and American elm. West
of the Mississippi Delta in eastern Texas, the
sweet gum-oak association consists of sweet gum
and willow oak as dominants, with overcup oak,
water hickory, green ash, black gum and cedar
elm as important associates. Loblolly and long-
leaf pine woods border all of these river swamps,
but ivory -bills rarely feed in them.
Ivors-bill habitats in the Florida region vary
considerably, although cypress is a dominant tree
in all of them, a condition not found in ivory-bill
habitat outside of the Florida region. Another dif-
ference is that ivory-bills in Florida, unUke else-
where, frequently feed in the pine woods
bordering the swamp. Many ivory-bills
are recorded in swamps along the small rivers of
central and southern Florida where the predomin-
ant trees are baldcypress, red maple, laurel oak,
black gum, with some sweet gum and cabbage
palmetto. Ivory-bills frequently nested in cypress
trees, but only occasionally fed upon them. In the
Mississippi Delta region, they nested only in var-
ious species of hardwood trees (at least in the
Singer Tract), in parts of the woods where the
ground was at least partly covered with water dur-
ing the nesting season (Tanner 1942).
A characteristic of all habitats used by ivory-
bills is that other species of woodpeckers, such as
the pileated and red-bellied woodpeckers, reach
their greatest abundance there also. The most
likely places to look for ivory-bills are bottom-
land forests where big sweet gums and oaks are
abundant, where there are many dead and dying
trees, and where other woodpeckers are abundant.
The many dying and dead trees in old age stands
of timber contain the wood-boring insects eaten
by the woodpeckers. The sweet gum-oak bottom-
land forests supply the best feeding conditions on
the Mississippi Delta. The wood-boring insects are
most abundant in wood 2 or 3 years dead from
storm, fire, logging, or disease. These conditions
occur most often in large, old forests, and their
elimination or isolation has made it increasingly
difficult for ivory-bills to find sufficient food and
to move from one area to another in search of a
variable food supply that has always been more or
less eruptive and undependable (Tanner 1942).
Virgin haidwood is not a necessity and pines are
more important than Tanner thought in the
Neches River Valley (Dennis 1967).
The ivory-bill is a nomadic "disaster species,"
moving into areas where trees have been killed by
fire, storms, insect attack, or flooding (Dennis
1967). Observations in Texas and Florida by
Herbert Stoddard and John Dennis convinced
them that old age or virgin hardwood forests are
not essential as long as there are large numbers
of recently dead trees to supply the type of
wood-boring grubs prefened by ivory-bills. They
thought that dead pine trees, including recently
cut slash, were frequently used (Dennis 1967).
C. p. bairdii. Gunclach (1876) found Cuban
ivory-bills in the high country of Pinar del Rio
and also in low country along river bottoms simi-
lar to the habitat of the American ivory-bill, near
Guantanamo in eastern Cuba. During the last half
of the 19th century, Cuban ivory-bills were found
mainly in high country in pine forests on deep
lateritic soil. They feed in both hardwoods and
pines, but nest and roost almost exclusively in old
pine trees (Pinus cubensis) (Lamb 1957). The
lateritic soil, composed of small, hard nodules of
iron ore, drains very quickly and completely, so
that it can support pine forest up to about 300 m
elevation. Above that, hardwood becomes domin-
ant. Most of the pine land has been lumbered, but
the birds have managed to adapt to changing habi-
tat, living in large dead pines that are still standing
and feeding on dead pines and dead hardwoods,
both of which are infested with wood-boring
beetles. They roost and nest only in pines, and
large enough pines are becoming rare.'
FOOD AND FORAGING
C. p. principalis. Audubon (1842) mentions
grapes, persimmons £ind blackberries as food of
ivory-bills, in addition to beetles and their larvae.
Allen and Kellogg (1937) found ivory-bills digging
trenches in rotten wood, as plicated wood-
peckers do, to get at the large wood-boring beetle
larvae. More often, they scaled off bark from
recently dead trees or from dead branches of liv-
ing trees to get at insects and larvae hidden be-
neath. Most feeding was in dead pines at the
edges of swamps. They sometimes fed on the
ground like flickers.
The most common feeding behavior is to
knock the bark off recently dead trees with side-
wise blows or quick flicks of the bill to uncover
and eat the borers that live between the bark and
the sapwood. When feeding the young, they hold
grubs in the back of the bill while continuing to
scale bark for additional food. Grubs 2.5 to 5 cm
long are used to feed young. Ivory-bill workings
for food show as bare places on recently dead
limbs of trees where the bark has been scaled off
clean for a considerable extent. Pileated wood-
peckers do some scaling, but it is usually confined
to smaller limbs and to those longer dead. They
obtain most of their food by digging in the wood,
while ivory -bills obtain theirs by scaling the bark.
Extensive scaling of bark from a tree so recently
dead that the bark is still tight, with a brownish
or reddish color of the exposed wood showing
that the work is fresh, is one good indication of
the presence of ivory-bills (Tanner 1942). How-
ever, they do also chisel into the wood, making
somewhat conical holes. In the Singer Tract,
Louisiana, most feeding was on sweet gum. Nut-
tail's oak, and hackberry, over 30 cm in diameter.
Wandering and ranges of ivory-bills are prob-
ably controlled by abundance of food. They re-
quire an unusually large supply of certain wood-
boring insects which make up most of their diet
and which is abundant only in occasional localities
for a comparatively short period. Birds remain in
one locality as long as the food lasts, then move,
sometimes for considerable distances, until they
find another area with an adequate food supply
(Tanner 1942).
Contents of three ivory-bUl stomachs from
Louisiana and Texas summarized by Cottam and
Knapper (1939) (including those reported by Beal
1911) were as follows: 46% of combined content
was of animal origin, 45.33% being long-horned
beetles (Cerambicidae), and 0.67% of engraver
beetles (Tomiscus sp.); 54% was of vegetable
origin, 14% being seeds of Magnolia grandiflora,
27% of Carya, 12.67% of seeds of poison ivy
{Rhus radicans), and 0.33% being fragments of
unidentified gall. In three stomachs from a Caro-
lina region, Wilson (1811) found large quantities
of large grubs fitting the description of some
larger larval Cerambycids. In two stomachs from
Louisiana, Goss (1859) found one large Cera m6yx
and the stones of cherries. In debris from an ivory-
bill nest in the Singer Tract, Louisiana, soon after
the young had left, Tanner (1942) found 1 frag-
ment of Elatrid larva; 21 mandibles of Cerambycid
larvae; 1 mandible of a Scarabaeid beetle larva,
and a few fragments of adult insects, probably
Coleoptera. E. A. Mcllhenny [in Bendire 1895),
said they fed on acorns. Alexander Sprunt, Jr.,
in Tanner (1942), saw ivory -bills feeding on black
gum and tupelo berries. Probably, the seasons
when fruits or seeds are eaten are from late sum-
mer to early winter when they are most easily
available; stomachs with large percentages of
vegetable food were taken in November. Ivory -b ills
drink water from hollows in trees (Tanner 1942).
C. p. bairdii. Their feeding habits are about
equally divided between pine and hardwood for-
ests, and they have been seen feeding on both
types of tree. They feed both by scaling bark and
by digging holes in wood, with the majority of
feeding signs of the scahng type. There are no re-
cords of the actual food eaten, but presumably it
consists of larvae of wood-boring beetles like
those preferred by American ivory-bills. Such in-
sects were found in trees used by the Cuban ivory-
bills.
SHELTER REQUIREMENTS
- C. p. principalis nests and roosts in holes in
large dead or living trees, usually hardwoods.
C. p. bairdii requires holes in large dead pine
stubs for both nesting and roosting.
NESTING OR BEDDING
C. p. principalis. Allen and Kellogg (1937)
quote Audubon who believed the nesting hole is
always made in the trunk of a live tree, generally
an ash orhackberry, and at great height. However,
they noted that there were records of nesting in
live cypress, partly dead oaks, a dead royal palm
stub, and an old and nccirly rotten white elm
stump, indicating as great a variety as nest sites
of the pileated woodpecker. Beyer (1900) found
a nest as low as 8 m in a living over-cup oak.
Allen and Kellogg (1937) found a nest in Florida
10 m up in a live cypress and three nests in Louis-
iana in oak and one in a red maple. Nest trees
were very large. One nest was in a dead pin oak
stub about 16 m high; the entrance hole was 13
cm high and 11.4 cm wide; depth of nest cavity,
47 cm, and diameter 20 cm at egg level (Allen
and Kellogg 1937). In Florida, height of nests
from the ground averaged 15 m (Tanner 1942)
with extremes of at least 8 m (Hoyt 1905) and 20
m (Ridgway 1898). In Louisiana, nests averaged
15.5 m and ranged from 12 to 21 m from ground,
all in dead trees or dead parts of living trees where
wood was a bit punky but still quite hard. The
average depth of all reliably measured nest cavities
is 48 cm (Tanner 1942). Nesting and roosting
holes of ivory -bills have oval or irregular entrances
measuring about 13 cm vertically and 10 cm across,
or about 2.5 cm larger than pileated woodpecker
entrance holes (Tanner 1942).
Ivory-bills do not use old nesting holes, but
excavate new ones usually in different trees. One
roosting hole has never been seen occupied by
more than one bird even by a young one still in
the company of its parents. For at least 2 weeks
after leaving the nest, young roost in the open in
trees while the parents roost in holes. The birds
emerge from their roosting holes much later in
the morning than other woodpeckers. Then pairs
and young join together for their daily feeding
flights (Tanner 1942).
C. p. bairdii nest and roost almost exclusively
in holes in old pines. Only one instance of a roost
hole in a hardwood has been reported. Nesting
and roosting sites were found in 16 pine trees
which contained 33 holes dug by ivory-bills
(Lamb 1957). Dennis (1948) found a nest about
10 m from the ground in a dead pine stub. The
opening was approximately 10.2 by 10.2 cm,
forming a rough square. Old holes found by Lamb
(1957) were all considerably lower than 10 m,
averaging about 6 m above ground. The pine stubs
used were never over 8 m tall. In the virgin forest
area, holes were closer to 9 m from ground and
one was nearly 18 m. Measurements of a female's
roost entrance were 15.6 cm (width) by 24.1 cm
(height); inside diameter of the cavity was 25.4
cm and depth of cavity from entrance to bottom
was 32 cm.
RITUAL REQUIREMENTS
A pair of C. p. principalis was observed by
Allen and Kellogg (1937) to clasp bills, evidently
as part of courtship behavior. Tanner (1942) ob-
served a similar event. He also described soft con-
versational notes by both members of a pair when
they exchanged places on the nest. The drumming
display consists of a double tap instead of the
multiple taps or drum-roll of other North Ameri-
can woodpeckers (Tanner 1942).
POPULATION NUMBERS AND TRENDS
C. p. principalis. Earlier accounts gave no ac-
curate or definite statements of abundance, but
indicated that it never was common. An excep-
tion was Audubon (1842) who said it was "very
abundant along the Buffalo Bayou (near Hous-
ton?), Texas." Audubon's "very abundant" may
have meant compared with ivory-bills in other
localities, where Audubon usually described the
species as quite rare (Tanner 1942). Arthur T.
Wayne and his hunters collected 5 ivory-bills
in California Swamp, 97 km (58 mi) south of
Old Town, Florida and saw 4 more several days
after the fifth one was collected. The following
year, 1893, five more were collected from that
swamp. Later observations indicate that those
10 birds were practically all that were present in
that area of 154 km^ . In vicinity of the Wacissa
River, Florida, in 1894 Wayne collected 19 ivory-
bills and some persisted there until about 1937.
On that basis. Tanner (1942) estimated that there
were probably about 12 pairs in that 188 km^
swampy area, or 1 pair per 16 km^ . Wayne (1910)
stated that he saw 200 ivory-bills in Florida dur-
ing the years 1892 to 1894. In the Singer Tract of
about 300 km^ of virgin forest in Louisiana, in
1934, there were about 7 pairs, or 1 pair per 43
sq km. Total population in all areas in 1939 was
estimated at about 24 individuals (Tanner 1942).
The greatest distance a pair traveled from the
roosting area in Singer Tract in breeding season
was 2 km (Tanner 1942). Birds of the Singer
Tract appeared to be sedentary, with ranges up to
6 km or more across. However, birds in other
areas appear to wander considerable distances,
probably in response to the availability of food
(Tanner 1942) near the nesting site.
Bark-stripping from recently dead pines, pos-
sibly the work of ivory-bills, was seen along Men-
ard, Big Sandy, and Village Creeks; along the
Neches River north and northwest of Beaumont;
and near Votaw and Silsbee, Texas, in December
1973 (Orie L. Loucks, Prof, of Botany, U. of
Wisconsin in lit. 17 March 1975).
On 22 May 1976, one ivory-bill was reported
near the mouth of Wolf Creek, flying across
Magnolia Ridge Road north of Beaumont, Texas
(William B. Mounsey, University of the Wilderness
ms. report, 4 Sept. 1976).
There were repeated reports of sightings of
ivory-bills in swamps along the Congaree and
Wataree Rivers, South Carolona, during the per-
iod 1966-67; John V. Dennis (Unpubl.) believed
these to be valid in part.
Herbert Stoddard saw a pair in beetle-killed
pines near Thomasville, Georgia, probably in
1958 Q. V. Dennis ms. 1976). On 3 and 4 March
1950, Eastman reported seeing a male and female
ivory-bill on the Chipola River in northwestern
Florida, and in April 1950 Dennis heard one near
the same place (Dennis 1967, J. V. Dennis ms.
1976).
On 28 Aug. 1966, Bedford P. Brown J. and
Jeffrey R. Sanders, Chicago bird watchers, heard
the distinctive call notes and saw 2 female ivory-
bills on Boiling Creek, a tributary of the Yellow
River at the Elgin Air Force Base in northwestern
Florida;they reported the sighting to J. V. Dennis,
who tried unsuccessfully to find the birds again
(Dennis 1967, Dennis 1976).
A feather from a cavity of a wind-blown tree
northwest of Lake Okeechobee, Florida, around
1965, was identified by Alexander Wetmore and
John Aldrich as that of an ivory-bill. However,
the white patch on the feather was stained brown,
indicating it had been in the tree hole a long time
Q. W. Aldrich pers. comm., Agey and Heinzman
1971).
Near the Neches River north of Beaumont,
Texas, above Dam B Reservoir, in April 1966,
one was seen by Mrs. Olga Hooks Lloyd, bird
watcher of Beaumont, and again in the same area
on 10 December 1966, one was seen by John
Dennis; there are also several other probable re-
cords from theNechesRiver Valley (Dennis 1967).
A sound recording of ivory-bill call notes was
made at Stanford Preserve on Village Creek, north
of Beaumont, Texas, 25 February 1968 by Helen
and Peter Isleib of Cordova, Alaska, and John V.
Dennis. The recording is now in possession of the
National Geo.graphic Society. Isleib found bill
marks on bark -striped trees nearby which measured
the same as the tip of ivory-bill's bill rather than
that of a pileated woodpecker, according to Alex-
ander Wetmore (Peter Isleib pers. comm.). The
sound recording was analyzed by Peter Paul
Kellogg {in lit. 14 February 1969) at Cornell
University Library of Natural Sounds and by
John W. Hardy, Bioacoustics Laboratory, Florida
State University, and both thought it sounded
very much like the call of an ivory-bill, although
Hardy (1975) believed that one of the call notes
of a blue jay was possible.
On 22 May 1971, 2 ivory -bills were sighted
and one was photographed in the Atchafalaya
River swamp west of Baton Rouge, La. by an un-
identified dog trainer. The beginnings of a hole
and fresh chips on the ground under the place the
bird was photographed were found (George H.
Lowery,Jr. pers. comm., Stewart 1971).
James Tanner and PaulSykesin 1968 searched
the same areas in the Neches River valley, Texas,
where John Dennis and others reported evidence
of ivory-bills in 1966, with negative results with
respect to sightings, sound or indication of feed-
ing (Sykes 1968, Tanner in lit. 6 March 1968).
C. p. bairdii. Demiis (1948) found 3 Cuban
ivory-bills, 2 of them a mated pair that were in-
cubating, in April 1948, in the Sierra de Moa,
Oriente Province, Cuba, and heard of another
group of 6 being seen there in 1941.
Lamb (1957) found 4 pairs in the Moa region
and found another area supporting 2 pairs farther
inland at a higher elevation, between the Moa
and Punta Gorda Rivers, but with identical habi-
tat (pine forest). In summary. Lamb (1957) esti-
mated the number of Cuban ivory-bills extant
in July of 1956 as 6 pairs or 12 individuals, all
between the watersheds of the Moa-Cubanas
Rivers to the west and the Punta Gorda River to
the east, along the north coast of Oriente Pro-
vince, and in the extreme headwaters of the Cale-
tura River, a large tributary of the Moa River, and
in the headwaters of the JaquEuii River, a tribu-
Xaxy of the Toa River, and on the divide between
the Toa and Moa Rivers. The population density
was computed as about 40.2 km^ per pair, and
the smallest area estimated for a single pair was
19.3 km^ . Observations were made during March,
June, and early July.
More recent sightings include 2 south of
Cupeyal in late 1967, a female northwest of
Cupeyalin February 1968, and a female at Yateras
in May 1972. All are in the Moa-Guantanamo
region, Oriente Province. Certainly no more than
8 pairs still exist and probably fewer than 6 (L. S.
Varona 1974, and O. H. Garrido 1974 in King
1978).
REPRODUCTION
C. p. principalis. In the Florida region, most
nests have been in living or dead cypress, but also
in bay and cabbage palm (Tanner 1942). Thomp-
son (1896) said ivory-bills nested in long-dead
pine trees in the Okefenokee Swamp region. In
northern Louisiana, Tanner (1942) found nests in
red maple, sweet gum, and Nuttall's oak. In the
same region, Beyer (1900) found nests in Ameri-
can elm and overcup oak. In southern Louisiana,
Mcllhenny in Bendire (1895) said birds built in
cypress or tupelo, preferably partly dead. Audu-
bon (1842) thought they preferred hackberry
trees for nesting.
The presence of eggs has been recorded on
the following dates: Florida (4 records), 4 March
to 19 April; Louisiana (5 records), 6 March to 19
May; Georgia (2 records) 6 and 10 April; Texas
(2 records) 11 April and 3 May (Arthur Allen in
Bent 1939). Time of nesting is evidently irregular.
Scott (1881) collected an incubating female in
Florida 20 January 1880. Scott (1888) found a
nest containing a one-third grown young 17
March 1887. Ridgway (1898) collected a male
that left the nest hole on 15 February 1898. Hoyt
(1905) noted that nesting began in the latter part
of January and eggs were laid by 10 February.
Tanner in Bent (1939) noted young leaving the
nest on 30 March 1937 in Louisiana, and appar-
ently the same pair had young in the last week in
February 1938. In contrast, there are 10 records
of April nesting, 5 for May and 1 (Beyer 1900) of
a young just out of nest in July. The later records
might represent second attempts at nesting.
Florida birds, in general, start nesting earlier than
those in Louisiana (Bent 1939). There is no cor-
relation between the erratic timing of nesting and
weather, except possibly the amount of sunshine.
They have tended to nest earher in cloudy winters.
No correlation with food supply has been found,
but the possible effects are not discounted (Tan-
ner 1942).
According to Mcllhenny in Bendire (1895),
the female does all work of excavating the nesting
cavity, but Audubon (1842) states that both birds
work at excavating. Thompson (1896) also re-
ports that both sexes excavate.
Although Bendire (1895) said there were 5
eggs per clutch and only 1 brood per season,
Arthur Allen in Bent (1939) found the number of
eggs normally not more than 3, and 1 or 2 of
those often infertile. Frequently, if nesting is suc-
cessful at all, it results in a single young rather
than 2 or3. Tanner (1942) figures that the average
number of eggs per set is 2.9 with a range of 1 to
4, and that early nests had fewer eggs than later
ones. He said the period of incubation and length
of time young stay in the nest is unknown. In the
Singer Tract, the male incubated every night and
the female most of the time during the day. They
exchanged places about eight times a day, with
no regularity in the setting time except when the
female relieved the male in the morning and just
before he took his place for the night. Both sexes
share about equally in feeding the young in the
nest and for a while after they leave the nest. Young
follow parents on feeding trips and gradually
learn to obtain food for themselves after about 4
weeks, but still receive food from adults after 2 to
2.5 months (Tanner 1942). Hoyt (1905) states
that after young leave the nest in April, they re-
main with their parents until the mating season in
December. Some young stay with parents even
after another family is fledged the following nest-
ing season. All aggregations of ivory-bills are
probably family groups of one or more years pro-
duction. Tanner (1942) says the average number
of young per brood leaving the nest is 2.11 with
a survival success of 50%. He says there is no data
on survival of young to breeding age nor length of
time an adult can live and successfully breed. The
important difference between nesting of ivory-
bills and other species is that it lays fewer eggs. A
pair may go through a season without any attempt
at nesting. It probably takes at least 2 years to
reach breeding age. Possibly the quantity of food
or lack of synchronization of the reproductive
cycles of a mated pair may determine whether
nesting will occur. The lower rate of reproduction
accentuates the danger inherent in the small size
of the population. The most likely serious preda-
tors on nesting ivory-bills are barred owls, rac-
coons, and nest mites.
Ivory-bill pairs usually nest well separated
from each other (about 4.8 km apart in the Singer
Tract), and, at least occasionally, nest in the same
area year after year. There are no records of birds
protecting territory from trespass by another
ivory-bill (Tanner 1942).
C. p. bairdii. Incubation is shared by both
sexes (Dennis 1948). Feeding and brooding are
thought to be shared by both sexes (Lamb 1957).
Causes of nesting failure are always traceable to
human interference, although crows are thought
by natives to be predators on nests (Lamb 1957).
MANAGEMENT AND CONSERVATION
C. p. principalis. The only way of preserving
the ivory-billed woodpecker is to preserve in their
entirety any areas that can be found where the
birds still exist. A suitable area would be about
6 to 8 km^ of good habitat for each pair. Pos-
sibly some timber may be selectively cut, leaving
dead and dying trees. The quantity of food may
be artificially increased by progressively killing
enough trees at a time to supply a large number
of wood-boring insect larvae (Tanner 1942,
Paul Sykes in lit. 1968). Dennis (1967) found
that trees killed by girdling do not attract as
many wood-boring insects as those killed by
flooding and suggested limited inundation and
cutting of pine slash as a means of increasing
ivory -bill food supply.
In October 1974, a bill was passed by Congress
creating a preserve of 34,000 ha of potential
ivory-bill habitat in the Big Thicket area of
southeastern Texas (Dennis ms. 1976).
Public information is an important part of
ivory-bill management as with other endangered
species. Only an interested and sympathetic public
can save the bird from such threats as big dams,
wholesale clearing of bottomland timber, and
drainage of favorable habitat (Dennis 1967).
In 1967, the U.S. Corps of Engineers halted
the timber management plan at Dam B Reservoir
on Neches River, Texas, in deference to ivory-
bills. Federal and state wardens in area were alerted
and public appeal received positive and gratifying
response (Harry Goodwin in lit. to Roland Cle-
ment 19 December 1967).
The U.S. Fish and Wildlife Service adopted a
policy of informing the public of locations of re-
ported ivory-bill sightings as opposed to restric-
tion of such information advocated by Herbert
Stoddard and some other knowledgeable people
(Letter from Director, Fish and Wildlife Service
to E. V. Komarek of 12 September 1967, mem-
orandum to Secretary of Interior from Director,
Fish and Wildhfe Service of 20 July 1967; Letter
of 1 July 1967 from John Dennis to George M.
Sutton).
C p. bairdii. Establishment of refuges in areas
where birds are still known to exist in the most
important step in management, for without pre-
servation of habitat, fast disappearing due to lum-
bering, the bird cannot survive. Further effort to
locate existing birds is needed to pinpoint pros-
pective refuge areas (Lamb 1957). Plans were
made to erect nesting boxes as an experiment to
substitute for suitable dead pine stubs which were
disappearing. Whether this was done, and if so
what happened, is unrecorded (Lamb 1957). For-
est reservations have been established at Cuepyal
and Jaguane under auspices of the Academy of
Sciences since 1963. Each reservation is watched
and no further exploitation of timber is being per-
mitted, although much of the larger timber has
already been removed (Lamb 1957; Fisher et al.
1969).
The ivory-billed woodpecker is protected un-
der the Protected Migratory Bird Treaty Act, 40
Stat. 755: 16 WSC 703-74, 3 July 1918; as
amended 3 December 1969. PL 91-135.
AUTHORITIES
James T. Tanner
Dept. of Zoology
University of Tennessee
Knoxville , Tennessee 37916
John V. Dennis
3 Joy Street
Nantuckett, Massachusetts 02554
Paul W. Sykes
P.O. Box 2077
Delray Beach, Florida 33444
PREPARER'S COMMENTS
From the evidence presented, I believe that a
few ivory-bills still exist in the United States, but
they are so nomadic that it will continue to be
difficult to verify the occasional sighting. Never-
theless, every effort should be made to locate
nesting pairs and to take measures to preserve
nesting habitat in those locations. Then additional
feeding areas should be artificially created near
such locations. -J. W. Aldrich.
LITERATURE CITED/SELECTED
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10
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Dennis, J. V. 1967a. The ivory-bill flies still.
Audubon Mag. 69:38-45.
Eastman, W. 1958. Ten-year search for the Ivory-
billed Woodpecker. Atlantic Naturalist 13:
216-228.
Fisher, J., N. Simon and J. Vincent. 1969. Wild-
life in danger. New York, Viking Press.
Gosse, P. H. 1859. Letters from Alabama. Lon-
don pp. 91-93 (in Tanner 1942).
Greenway, J. C. 1958. Extinct and vanishing birds
of the World. Amer. Comm. for Intemat.
WildUfe Protection, New York, N.Y. Special
Pub. 13:357-360.
Gundlach, J. 1876. Contribucion a la Ornitologia
Cubana. Habana.
Hardy, J. W. 1975. A tape recording of a possible
Ivory -billed Woodpecker. Amer. Birds 29:
647-651.
Hasbrouck, E. M. 1891. The present status of the
Ivory-billed Woodpecker {Campephilus prin-
cipalis) Auk 8:174-186.
Howe, R. H. and L. King. 1902. Notes on various
Florida birds. Contrib. N. A. Ornith. 1:30.
Howell, A. H. 1907. Birds of Alabama. U.S. Bur.
Biol. Surv. pp. 159-162.
Howell, A. H. 1911. Birds of Arkansas. Bull. Bur.
Biol. Surv. 38:45-46.
Howell, A. H. 1932. Florida bird hfe. New York
pp. 313-315.
Hoyt, R. D. 1905. Nesting of the Ivory-billed
Woodpecker in Florida. Warbler (ser. 2) 1:52-
55.
King, W. B. 1978 (in press). (Revised) Red data
book (Aves). I.C.B.P., Survival Service,
Morges, Switzerland.
Kline, H. A. 1886. Ivory-billed Woodpecker. For-
est and Stream 26:163.
Lamb, G. R. 1957. The Ivory-billed Woodpecker
in Cuba. Research Rept. No. 1, Pan-American
Sect. Intnat. Comm. Bird Preservation.
Mcllhenny, E. A. 1941. The passing of the Ivory-
billed Woodpecker. Auk 58:582-584.
Merriam, C. H. 1874. Ornithological notes from
the south. Amer. Nat. 8:88.
Murphy, R. C. 1929. A second topotype of
Campephilus principalis. Auk 46:376.
Nice, M. M. 1931. The birds of Oklahoma. Pub.
Univ. Okla. 3:116.
Oberholser, H. C. 1938. The bird life of Louisiana.
Bull La. Dept. Cons. 28:38-382.
Pindar, L. O. 1925. Birds of Fuhon County,
Kentucky. Wilson BuU. 37:86.
Ridgway, R. 1898. The home of the ivory-bill.
Osprey 3:35-36.
Ridgway, R. 1914. The birds of North and Middle
America. U.S. Nat. Mus. Bull. 50, pt. 6.
Ridgway, R. 1915. Bird-life in southern Illinois.
Bird-Lore 17:194.
Scott, W. E. C. 1881. On birds observed in Sump-
ter. Levy and Hillsboro Counties, Florida.
Bull. Nuttall Ornith. Club 6:14-21.
Scott, W. E. C. 1888. Supplementary notes from
the Gulf coast of Florida, with a description
of a new species of marsh wren. Auk 5:183-
188.
Scott, W. E. C. 1889. A summary' of observations
on the birds of the Gulf coast of Florida. Auk
6:251.
11
Stewart, J. R., Jr. 1971. Central Southern Region.
American Birds 25:868.
Sykes, P. W., Jr. 1968. Report on the search for
the Ivory-billed Woodpecker [Campephilus
principalis principalis) in eastern Texas, Jan-
uary 1968. Administrative report, Fish and
Wildlife Service.
Tanner, J. T. 1942. The Ivory -billed Woodpecker.
Nat. Audubon Soc. Research Rep. No. 1, New
York.
Thompson, J. M. 1896. An archer's sojourn in the
Okefinoke. Atlantic Monthly 77:486-491.
Thompson, M. 1889. A red-headed family.
Oologist 6:23-29.
U.S. Fish and Wildlife Service. 1973. Threatened
wildlife of the United States. U.S. Dept. of
Interior, Resource Pub. 114, G.P.O. Washing-
ton, D.C.
Wayne, A. T. 1910. Birds of South Carolina. Con-
trib. Charleston Mus.:87-88.
Wetmore, A. 1943. Evidence for the former oc-
currence of the Ivory-billed Woodpecker in
Ohio. Wilson Bull. 55:127.
Williams, R. W., Jr. 1904. A preliminary list of
the birds of Leon County, Florida. Auk 21:
455.
Wilson, A. 1811. American Ornithology. Phil-
adelphia Vol. 4:20-26.
Yell. 1885. The big woodpecker. Forest and
Stream 24:107.
12
Biological Services Program
FWS/OBS-80/01.9
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE OCELOT
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species /\ct of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of f^ndangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704'')8
u
FWS/OBS-80/01.9
March 1980
SKLKCi i:d vi:r 1 kbra 1 i: kndangered spkcies
OF THE SEACOAS 1 OF 1 HE UNFFED STA TES-
THE OCELOT
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National ("oastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald \V. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish antl Wildlife Scr\ice
U.S. Department of the Interior
OCELOT
Felis pardalis Linnaeus
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivora
FAMILY Felidae
OTHER COMMON
NAMES leopard cat, ocelote, tiger cat,
tigrUIo
DATE
Entered into SWIS to be determined
Updates 31 May 1978
LEGAL STATUS
Federal: Endangered (non-U. S. populations
only) (44FR43705, 25 July 1979).
States: Endangered: Massachusetts, Texas.
REASONS FOR CURRENT STATUS
Habitat destruction and degradation from
brush-clearing operations are primarily responsible
for the ocelot's population status (Culbertson
and Schmidly 1974, Davis 1974). Population de-
clines are also affected by predator control activi-
ties and persecution. Serious population declines in
Central and South America are due to pet and fur
trade exploitation (lUCN 1972, Paradiso 1972).
PRIORITY INDEX
Not assigned.
DESCRIPTION
Medium-sized (0.8 to 1.2 m long) spotted cat
wdth moderately long tail. Females average smaller
than males. Ground color is grayish to cinnamon
dorsally, paler on sides; underparts and inside of
limbs are whitish; dark markings form streaks that
run obliquely down sides; areas enclosed by
black-bordered spots are more intense than
ground color. There are two black stripes on
the cheeks; the tail is spotted and ringed with black.
Young have coloration similar to adults, but
darker. Photographs appear in Davis (1974) and
Guggisberg (1975).
RANGE
Extremely rare to nonexistent in Arizona
(Cockrum 1960). In Texas, the ocelot occupies
the lower Rio Grande Valley south of a line from
Eagle Pass to just north of Corpus Christi (Hock
1955, Culbertson and Schmidly 1974, Davis
1974). There has been one record each from the
Trans-Pecos (Schmidly 1977) and northern Texas
(Davis 1951), but these are thought to be released
or escaped captives (W. C. Brownlee and R.
McBride personal communication). In Mexico,
the species occurs in eastern Coahuila, along the
Rio Sabines vicinity of the Sierra de San Marcos,
and south and east of Saltillo (Baker 1956, lUCN
1972). It is also present in suitable habitat in Cen-
tral and South America (lUCN 1972).
The ocelot's former distribution included cen-
tral and eastern Texas, into southern Arkansas
and western Louisiana, and south into Mexico
east of the highlands. Ocelots were also found
from central Arizona south into the Sonoran and
west slopes of the Sierra Madre. There are no
records from Mexico City (C. Conway personal
communication). The Mexican highlands form a
barrier between the race of ocelot that occurs in
Texas {F. p. albescens) and that which occurs (or
occurred) in Arizona (F. p. sonoriensis) Goldman
1925). The ocelot was also distributed transcon-
tinentally throughout Central and South America
as far south as Uruguay and northern Argentina
(Goldman 1943, Hock 1955, Hall and Kelson
1959, lUCN 1972, Guggisberg 1975).
RANGE MAP
Shading refers to present range (Cockrum
1960, Davis 1974, Brownlee 1978, R. McBride
personal communication); dots indicate sightings
and/or kills within the last century.
STATES/COUNTIES
Arizona Cochise, Pima, Santa Cruz.
Texas Cameron, Hidalgo, Kenedy, Willacy.
HABITAT
Ocelots inhabit tropical and subtropical for-
ests ranging from low swamp to upland oak
forests (Hall and Kelson 1959, lUCN 1972). In
the northern part of their range they inhabit
dense, nearly impenetrable chaparral thickets
(Davis 1974); they also use second-growth forests
and partially cleared lands (lUCN 1972).
FOOD AND FORAGING BEHAVIOR
Ocelots hunt mostly at night, but it is not un-
usual for them to be seen in the daytime (Cala-
hane 1947, Hall and Kelson 1959, Denis 1964).
Their food habits are not well known; they are re-
ported to feed on small and moderate-sized mam-
mals, birds, monkeys, tree lizards, and, occasion-
ally, domestic fowl (Hall and Kelson 1959, Denis
1964, Davis 1974).
SHELTER REQUIREMENTS
Ocelots take refuge in caves in rock bluffs,
hollow trees, or dense parts of thickets (Davis
1974).
NESTING AND BEDDING
The den site is usually a cave in a rocky bluff,
hollow tree, or the dense part of a thicket (Davis
1974).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Home range area is not known. Pairs share the
same territory but do not hunt together (Guggis-
berg 1975). They often rest in trees (Davis 1974);
they are excellent tree climbers (Hall and Kelson
1959).
POPULATION NUMBERS AND TRENDS
There are no firm estimates on populations in
the United States or Latin America. lUCN is con-
ducting a status survey in Latin America, and the
Texas Parks and Wildlife Department is making a
field survey in Texas. Preliminary findings of the
Texas survey indicate around 35 at Santa Ana Na-
tional Wildlife Refuge and 24 at Laguna Atascosa
National Wildlife Refuge (Brownlee 1978). How-
ever, others claim that there are only 12 to 15
ocelots on Laguna Atascosa and none are found
a
o
U
o
u
o
Q
d
u
■f-t
X
I
e
ll
o
c
c
.(«
CO
D
■M
.s >.
■t-J
:^
g so
^ o
-a c
^ top
at Santa Ana, due to its small size (688 ha) (R.
McBride, personal communication).
Another survey in South Texas in 1976 esti-
mated the ocelot population at 30 to 40 in Wil-
lacy County, 0 to 2 in Hidalgo County, 12 to 20
in Cameron County, and an unknown number in
Kenedy County (R. McBride personal communi-
cation).
The species rarely occurs in southern Arizona
(Cockrum 1960). Populations are declining in
Texas due to predator control activities and parti-
cularly to brush-clearing operations in the lower
Rio Grande Valley (Culbertson and Schmidly
1974, Davis 1974). Latin American populations
are reduced and in some areas are seriously de-
pleted (lUCN 1972, Paradiso 1972).
REPRODUCTION
There is no fixed breeding season in the tropics
(Denis 1964). In Texas, breeding takes place in
late summer, with young bom in September,
October, and November (Davis 1974). Gestation
period is believed to be about 70 days (Guggis-
berg 1975). Litter size ranges from two to four,
with two being more common (Leopold 1959,
Denis 1964, lUCN 1972). In Texas, average litter
size appears to be one (R. McBride personal com-
munication). Growth and development of young
are not well known.
MANAGEMENT AND CONSERVATION
No recovery team has been appointed at this
time.
Ocelots are protected in Texas and some
Latin American countries, but U.S. populations
are not listed on the official U.S. Endangered Spe-
cies list.
Brush is no longer cleared on the National
Wildlife Refuges in the lower Rio Grande Valley
(lUCN 1972), in order to maintain natural habi-
tat. Laguna Atascosa National Wildlife Refuge is
thought to contain habitat most similar to that
required by ocelots.
AUTHORITIES
W. C. Brownlee
Texas Parks and Wildlife Department
4200 Smith School Road
Austin, TX 78744
W. B. Davis
Department of Wildlife and Fisheries Science
Texas A&M University
College Station, TX 77843
Roy McBride
Department of Biology
Sul Ross State University
Box 725
Alpine, TX 78930
PREPARER'S COMMENTS
The use of questionnaire surveys may lead to
overestimates of populations and inaccurate dis-
tributional patterns when escaped or released cap-
tives are sighted or even when the same individual
is sighted repeatedly. Intensive field surveys are
needed to accurately define the current status of
the ocelot. Its ecology and life history also require
further investigation.
LITERATURE CITED/SELECTED
REFERENCES
Bailey, V. 1905 Biological survey of Texas,
Am. Fauna 25:1-222.
N.
1931. Mammals of New Mexico. N. Am.
Fauna 53:283-285.
Baker, R. H. 1956. Mammals of Coahuila, Mexico.
Univ. Kansas Publ. Mus. Natur. Hist. 9:125-
335.
Bangs, 6. 1898. The land mammals of peninsular
Florida and the coast region of Georgia. Proc.
Boston Soc. Natur. Hist. 28:157-235.
Brownlee, W. C. 1978. Feline status survey. Per-
formance Report, Proj. W-103-R-7. Texas
Parks Wildl. Dep. Austin. 4 pp.
Calahane, V. H. 1947. Mammals of North Ameri-
ca. MacMillan Co., New York. 682 pp.
Chapman, F. M. 1894. Remarks on certain land
mammals from Florida, with a list of species
known to occur in the state. Bull. Am. Mus.
Natur. Hist. 6:333-346.
Cockrum, W. L. 1960. The recent mammals of
Arizona: their taxonomy and distribution.
Univ. Arizona Press, Tucson. 276 pp.
Culbertson, K., and D. J. Schmidly. 1974. Sum-
mary of statements on the status of the rare,
endangered, and peripheral mammals in Texas.
Texas Organization for Endangered Species,
Temple. 8 pp.
Davis, W. B. 1951. Unusual occurrence of the oce-
lot in Texas. J. Mammal. 32:363-364.
1974. The mammals of Texas. Texas
Parks Wildl. Dep. Bull. 41:1-294.
Denis, A. 1964. Cats of the world. Houghton
Mifflin Co., Boston. 144 pp.
Goldman, E. A. 1925. Two new ocelots from
Mexico. J. Mammal. 6:122-124.
1943. The races of ocelot and margay in
Middle America. J. Mammal. 24:372-385.
Goodwyn, F. 1970. Behavior, life history and pre-
sent status of the jaguarundi, Felis yagour-
oundi {Ldicepede) in South Texas. M.A. Thesis.
Texas A&I University, Kingsville. 63 pp.
Guggisberg, C. A. W. 1975. Wildcats of the world.
Taplinger Publ. Co., New York. 382 pp.
Hall, E. R., and K. R. Kelson. 1959. Mammals of
North America. 2 vols. Ronald Press, New
York. 1,083 pp.
Hock, R.J. 1955. Southeastern exotic felids. Am.
Midi. Natur. 53:324-328.
Harlan, R. 1825. Fauna Americana: being a des-
cription of the mammiferous animals in-
habiting North America. Anthony Finely,
Philadelphia. 320 pp.
lUCN. 1972. Red data book. Vol. I, Mamnalia.
Compiled by H. A. Goodwin and C. W. Hollo-
way. lUCN, Morges, Switzerland.
Layne, J. N. 1974. The land mammals of Florida.
In P. J. Gleason, ed. Environments of south
Florida; past and present. Mem. 2 Miami
Geol. Surv. 452 pp.
Leopold, A. S. 1959. Wildlife of Mexico; the
game birds and mammals. Univ. California
Press, Berkeley. 568 pp.
Lowery, G. H., Jr. 1974. The mammals of Louisi-
ana and its adjacent waters. Louisiana State
Univ. Press, Baton Rouge. 565 pp.
Lowman, G. E. 1975. A survey of endangered,
threatened, rare, status undetermined, peri-
pheral, and unique mammals of the south-
eastern national forests and grasslands. USDA
For. Serv., Atlanta, Ga. 121 pp.
Maynard, C.J. 1883. The mammals of Florida. Q.
J. Boston Zool. Soc. 2: 1-50.
Paradiso, J. L. 1972. Status report on cats (Feli-
dae) of the world, 1971. U.S. Fish Wildl. Serv.
Spec. Sci. Rep. Wildl. 157.
Schmidly, D. J. 1977. The mammals of Trans-
Pecos Texas. Texas A&M Press, College Sta-
tion. 225 pp.
Sealander, J. A., and P. S. Gipson. 1974. Threat-
ened Arkansas mammals. Unedited draft re-
report to State of Arkansas Planning Depart-
ment, Little Rock.
Seton, E. T. 1937. Lives of game animals. Vol. I.
Literary Guild, New York.
Silveira, E. K. 1972. A case of cannabalism among
ocelots, Felts parda lis nitis, at Brasilia Zoo.
Int. Zoo. Yearb. 12:182-183.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
GainesvUle, FL 32601
Biological Services Program
FWS/OBS-80/01.10
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE ATTWATER'S GREATER PRAIRIE CHICKEN
''l:^^^
Fish and Wildlife Service
U.S. Department of the Interior
prefacp:
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as air.iiulcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should he directed lo:
Office of Endangered Species
U.S. Fish and W ildlife Senice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be dirccicvl to:
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.10
March 1980
SELECTKD Vl.R 1 KBRA 1 K ENDANGERED SPECIES
OF THE SEACt)AS T OF IHE UNFFED STA lES-
THE ATTWATER'S GREATER PRAIRIE CHICKEN
A Cooperative Effort
by the
National Fish and Wildhfc Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. VVoodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
ATTWATER'S GREATER PRAIRIE CHICKEN
Tympanuchus cupido attwateri Bendire
KINGDOM Animalia
CLASS Aves
ORDER Galliformes
FAMILY Tetraonidae
OTHER COMMON
NAMES Attwater's prairie chicken
greater prairie chicken, heath hen
DATE:
Entered into SWIS to be determined
Update 1 August 1978
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 March
1967)
States: Endangered: Texas
REASONS FOR CURRENT STATUS
Conversion of natural tall grass prairie to agri-
cultural lands for soybeans, grain, sorghum, rice,
and cotton has eliminated extensive amounts of
habitat. Commercial development and urban
sprawl have contributed to the loss of habitat
while overgrazing and oil development have re-
duced habitat quality (Lehmann and Mauermann
1963, W. Shifflett personal communication).
The Attwater's prairie chicken was formerly
killed in great numbers for target practice and
sport (Lehmann 1941).
PRIORITY INDEX
Not assigned.
DESCRIPTION
Prairie chickens are large, hen-like birds; males
weigh almost 1 kg, and females 0.75 kg. They are
brovm barred with black above, buffy barred with
black below, and have a short, rounded, dark tail.
Males have an orange comb above the eye, and,
on each side of the neck, an area of orange skin
that inflates during the courtship display. Tufts
of feathers (pinnae) extend down each side of the
neck, and point forward during courtship.
The Attwater's prairie chicken is distinguished
from the lesser prairie chicken (T. pallidicinctus),
which still occurs in the Texas panhandle, by
orange instead of reddish throat pouches. From
the northern race {T. cupido pinnatus), formerly
found in Texas, it differs in having the back of
the tarsus unfeathered. From the extinct heath
hen {T. cupido cupido), it differs in having pure
white axillaries and rounded pinnae.
Color plates appear in Lehmann (1941) and
Peterson (1947); black-and-white illustrations
appear in Lehmann and Mauermann (1963).
RANGE
This species' range formerly extended over
the entire Gulf coastal prairie of southwestern
Louisiana and Texas, and south to the Rio
Grande (Lehmann and Mauermann 1963). At
present, it is confined to small, disjunct popula-
tions scattered over about 12 Texas counties,
mainly Refugio, Austin, and Colorado. It is extir-
pated in Louisiana.
RANGE MAP
Present range (W. Shifflett personal com-
munication) is shaded on the following page.
STATES/COUNTIES
Texas Aransas, Austin, Brazoria, Cham-
bers,' Colorado, Dewitt, Fort Bend,
Galveston, Goliad, Harris, Jefferson,'
Refugio, Victoria, Waller,' Wharton.
HABITAT
Coastal grassland prairie approaches ideal
conditions when (1) vegetation is diversified and
native grasses, sedges, legumes, brush, and dwarfed
trees provide a variety of cover; (2) knolls, ridges,
and hog wallows are frequent and soils vary from
loose sand to tight clay or silt; and (3) permanent
sources of water are not more than 0.2 km apart
(Lehmann 1941).
Cultivation causes immediate abandonment,
but the birds will usually return after the field lies
' Indicates counties in which V. Lehmann (personal com-
munication) believes the species may no longer occur.
fallow for a year or two (V. Lehmann personal
communication).
FOOD AND FORAGING BEHAVIOR
Food of adults is 85% vegetable and 15%
animal; this ratio is approximately reversed in the
young. Favorite plants are ruellia {Ruellia ciliosa),
perennial ragweed {Ambrosia psilostachya), black-
berry {Rubus sp.), doveweed {Croton sp.), and
sensitive briar {Neptunia lutea); favorite animals
are grasshoppers and beetles.
Except during breeding season, adults feed
twice daily (dawn to 08:00 and 16:00 to dark).
Gizzards collected at noon are usually empty.
Feeding is slow and dehberate. Their capacity is
large; 20 cc in gullet and 30 cc in gizzard. The
variety of foods found in a stomach is im-
mense—up to 29 kinds of food and more than
1,300 items (Lehmann 1941).
SHELTER REQUIREMENTS
These birds require light to medium cover for
roosting, especially on gentle slopes, and medium
heavy to heavy cover for nesting and escape.
Light cover is necessary for booming (see Repro-
duction) and feeding. Hence, considerable habi-
tat diversity is required (Lehmann 1941). Fallow
rice fields and weedy, sandy slopes are used for
brood habitat in Austin and Colorado counties
(W. Shifflett personal communication).
NESTING OR BEDDING
Prairie chickens usually nest on well-drained
mounds or ridges near trails and within 1.3 km of
a booming ground. The nest is a shallow depres-
sion about 18 cm in diameter, more or less roofed
over by a medium to heavy grass cover, and lined
with bits of dead grass and a few feathers (Leh-
mann 1941).
RITUAL REQUIREMENTS
Preferred booming sites are short-grass flats,
often not elevated, 0.4 ha or so in area, and sur-
rounded by moderate grass cover suitable for
nesting.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Permanent sources of surface water no more
than 1.6 km apart are a must. Heavy rainfall in
late April or May is a hazard to nesting (Lehmann
1941).
POPULATION NUMBERS AND TRENDS
The species formerly numbered nearly
1,000,000 birds. In 1937, the population was esti-
mated at 8,700 (Lehmann 1941). In 1962, Leh-
mann and Mauermann estimated the total popula-
tion at 1,335. Since then, fluctuation has been
only slight, and the 1978 estimate was 1,500 indi-
viduals (W. Shifflett personal communication).
REPRODUCTION
Elaborate displays by males on the booming
grounds commence in February, peak in March,
and end in May. Females visit the booming
ground briefly for copulation and then select sites
and construct nests. Peak laying occurs in late
March and early April, at which time an average
of 12 eggs are laid. Hens take two feeding forays
daily, incubating the eggs the remainder of the
day. The incubation period is 23 days. Hatchlings
are precocious and immediately follow the hen
away from the nest. Some young leave the hen at
6 to 8 weeks of age, while others remain into the
fall (Lehmann 1941).
An annual increase of 100% is considered
good, despite the potential for 600% (Lehmann
1941).
MANAGEMENT AND CONSERVATION
Prairie chickens have been protected from
hunting since 1937. Lehmann (1941) conducted a
thorough study of Attwater's prairie chicken.
A 3,200-ha acquisition in the middle of prime
prairie chicken range in Colorado County was de-
signated the Attwater's Prairie Chicken National
Wildlife Refuge. Management techniques used to
improve the habitat include a closely regulated
grazing program, prescribed burning, mowing,
shrub eradication, and experimental crop plantings
(W. Shifflett personal communication).
The Texas Parks and Wildlife Department and
the U.S. Fish and Wildlife Service monitor the
Texas population annually (W. Shifflett personal
communication).
Lehmann and Mauermann (1963) have made
several management recommendations. Sugges-
tions include posting signs on roads to alert
motorists that they are in prairie chicken habitat;
transplanting of chickens from inhospitable areas
east of the Brazos River to better ranges (e.g.,
Victoria County); and organizing and supervising
a predator control program.
A Recovery Team has been appointed.
AUTHORITIES
Bill Brownlee (Recovery Team)
Texas Parks and Wildlife Dept.
4200 Smith School Road
Austin, TX 78744
Royce Jurries
Texas Parks and Wildlife Dept.
1131 Travis Street
Columbus, TX 78934
Val. W. Lehmann
P.O.Box 185
Carrizo Springs, TX 78834
Wayne Shifflett (Recovery Team)
Attwater's Prairie Chicken NWR
P.O.Box 518
Eagle Lake, TX 77434
Nova Silvy (Recovery Team Leader)
Dept. Wildlife and Fisheries Science
Texas A&M University
College Station, TX 77843
James Teer (Recovery Team)
Welder Wildlife Foundation
P.O. Box 1400
Sinton.TX 78387
PREPARER'S COMMENTS
The prairie chicken population is rather stable
due in part to the slowing down of conversion of
prairie lands to croplands. If several large ranches
should alter existing ranching operations, the prai-
rie chicken could be pushed close to extinction.
However, the Attwater's prairie chicken NWR
should prevent the species from becoming extinct.
LITERATURE CITED/ SELECTED
REFERENCES
Brownlee, W. C. 1970-1977. Attwater's prairie
chicken population census. Texas Parks Wildl.
Dep., Austin. Progress Rep.
. 1973. Vegetative requirements of Att-
water's prairie chickens. Texas Parks Wildl.
Dep., Austin. Progress Rep.
Lehmann, V. W. 1941. Attwater's prairie chicken,
its life history and management. N. Am. Fauna
57:65 pp.
Lehmann, V. W. and R. G. Mauermann. 1963
Status of Attwater's prairie chicken. J. Wildl.
Manage. 27:713-725.
Peterson, R. T. 1947. A field guide to the birds.
Houghton Mifflin Co., Boston. 230 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and WildHfe Laboratory
412 N.E. 16th Avenue, Room 250
GainesvUle,FL 32601
Biological Services Program
FWS/OBS-80/01.11
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE CAPE SABLE SPARROW
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data are not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as aiiuridcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Knginccrs in coordina-
tion with the Offices of Kndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Sen'ice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems I cam
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704')8
II
FWS/OBS-80/01.11
March 1980
si:lp:cii:d vi;ri kbraii: kndangered spkcies
OF rin: skacoast oi i hi: united siai es-
THE CAPE SABLE SPARROW
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
ant! the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. VVoodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal F.cosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CAPE SABLE SPARROW
Ammospiza marititna mirabilis Howell
KINGDOM Animalia
CLASS Aves
ORDER Passeriformes
FAMILY Fringillidae
OTHER COMMON
NAMES Cape Sable se2iside sparrow
DATE
Entered into SWIS to be determined.
Updates 17 August 1978, 10 October 1978.
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 March
1967)
States: Endangered: Florida
REASONS FOR CURRENT STATUS
The Cape Sable sparrow inhabits only un-
Credit: Harold Werner
stable interior marshes within a limited area of
southern Florida. There are only three known dis-
junct populations, two of them nearly extir-
pated. Progressive invasion of marshlands by
fast-growing exotic trees (Casuarina, Melaleuca,
Schinus) poses a major threat to the remaining
habitat, which is also under pressure from hurri-
canes, housing development, and inland encroach-
ment of mangroves. Frequent man-induced wild-
fires and predation by feral cats and dogs may
have caused the declines of some previously
extensive colonies (U.S. Department of the
Interior 1973; Werner 1975,1976, 1979; Mac-
Kenzie 1977).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The Cape Sable sparrow is about 13 cm long,
and colored olive-gray with olive-brown tail and
wings. It is marked with a yellow spot before the
eye and a white streak and black whisker along
the jaw. The ventral coloration is light gray to
nearly white, with darker streaking on breast and
sides. It tends to be greener above and whiter
below than other races of seaside sparrows. The
sexes are similar.
Immature birds are streaked dorsally with
brown to light buff, with less streaking on Hght
underparts. The yellow eye spot and dark whisker
are reduced or absent.
Color illustrations appear in Holt and Sutton
(1926), Howell (1932), Peterson (1947), and
Sprunt (1954); Werner (1975) presents black-and-
white photographs.
RANGE
This sparrow is a nonmigratory resident of
fresh to slightly brackish marshes in extreme
southern and southwestern peninsular Florida. It
ranges from 8 to 34 km inland, and is isolated
from other races of seaside sparrow by at least
260 km along the Gulf and 280 km along the At-
lantic coast of the State. Three disjunct, low-
density populations occur in marshes at Cape Sa-
ble, the Big Cypress Swamp, and the Taylor
Slough area of Everglades National Park, the last
being the location of the bulk of the population
(Werner 1975, 1976,1979).
The species was once common landward of
the mangrove zone from Carnestown to Shark
Valley Slough, including the coastal marl prairie
on Cape Sable (Stimson 1956). It is now only
rarely seen in this area (Werner 1979).
Emigration of fledglings is the chief dispersal
mechanism for an otherwise sedentary popula-
tions (Werner 1975, 1976).
RANGE MAP
Known localities for Cape Sable sparrows are
based on surveys conducted in 1970-1975 (re-
drawn from Werner 1976). Critical Habitat is re-
drawn from 42 FR 49685, 11 August 1977.
STATES/COUNTIES
Florida Collier, Dade, Monroe.
HABITAT
The species inhabits seasonally flooded brush-
less subtropical marshes (prairies) of interior sou-
thern Florida. Habitats vary from entirely fresh to
slightly brackish, generally remaining dry most of
the year. Brushy or rocky marshlands are avoided,
as are extremely dense stands of cordgrass {Spar-
tina bakeri) (Werner 1975, 1976, 1979).
Werner (1975, 1976) described Cape Sable
sparrow habitat in terms of four major graminoid
communities: muhly grass {Muhlenbergia filipes)
prairie; short sawgrass [Cladium jamaicensis) prai-
rie; prairies of tall, clumped cordgrass; and prairies
of low cordgrass growing with an irregular spatial
distribution.
Photographs of habitat appear in Holt and
Sutton (1926), Stimson (1968), and Werner
(1975).
FOOD AND FORAGING BEHAVIOR
The Cape Sable sparrow is primarily insecti-
vorous, with beetles and spiders comprising the
bulk of the diet. Amphipods, mollusks, and vege-
table matter are of minor importance (Howell
(1932). They feed almost entirely on or near the
ground, but are occasionally observed in sawgrass
flowers during the breeding season (Werner 1975).
SHELTER REQUIREMENTS
Sparrows generally remain hidden in marsh
vegetation during the hottest part of the day. The
microclimate may be moderated by moisture
from heavy morning dew (Werner 1975, 1979).
NESTING OR BEDDING
A woven grass nest is generally concealed
within a tussock of grass 6 to 37 cm above ground.
Nest construction may be either cupped or
domed (Werner 1975, 1979).
Werner (1975) reported nests primarily in
Muhly grass and occasionally in sawgrass. Nesting
is also known to take place in cordgrass and glass-
wort {Salicornia) (Nicholson 1928, Stimson
1968).
Photographs of nests and nest sites appear in
Stimson (1968) and Werner (1975).
RITUAL REQUIREMENTS
Males often chase females during the breeding
season. A female occasionally extends and vi-
brates her wings when approached by a male. The
significance of such interactions is not known
(Werner 1975).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Fire is a dangerous but vital component of
Cape Sable sparrow habitat. Periodic burning of
interior marshlands is necessary for perpetuation
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of optimal conditions. Sparrow populations are
closely related to the age of vegetation after fires.
Maximum density is achieved within 3 or 4 years
after a burn, followed by a sharp decline as the
dead component of vegetation increases. Pro-
longed absence of fire permits the elimination
of marsh by hardwood invasion. Colonies may be
enhanced or reduced by fire, depending on several
critical factors, such as burn pattern, percent of
contiguous habitat burned, size and percent of
colony evicted, proximity of and isolation from
adjacent colonies, and frequency and season of
burning (Werner 1975, 1976).
The breeding season appears to be closely re-
lated to the hydroperiod of the marsh. Singing
and nesting decrease abruptly when when the on-
set of heavy rains causes the marsh to flood
(Werner 1979), MacKenzie 1977). Dew may be an
important source of moisture during the dry sea-
son (Werner 1975, 1976).
Cape Sable sparrows interact very little with
other birds in their habitat area. Predation on
adult sparrows appears to be low. Mortality may
increase as summer floods inundate protective
vegetation. Nestlings are subject to attack by ants
(Werner 1975, 1976); 19% of nests surveyed by
Werner (1975) at Taylor Slough are known to
have failed due to predation. Similar failure of an
additional 6% is suspected.
The entire nesting cycle is completed within
the confines of a male's territory. Werner (1975)
reported territories of from 0.3 to 6.8 ha, some of
them overlapping. Territories, generally retained
through successive breeding seasons, are defended
by song and occasional chasing and combat. Ave-
rage territory size decreased with increasing popu-
lation density following fire. New residents usual-
ly occupy suboptimal sites adjacent to established
occupants.
POPULATION NUMBERS AND TRENDS
Werner (1976) estimated the total Cape Sable
sparrow population at 1,900 to 2,800 birds. Over
95% live in 8,800 to 12,800 ha of marshland in
the Taylor Slough area of South Florida. Muhly
grass prairie provides the principal habitat. Werner
(1976) described this final stronghold as 'an area
of widely spaced individuals, broken by various
sized patches of unacceptable habitat, on which
islands of greater density appear and disappear,
following the tracks of fires. . . .' A formerly large
population adjoining Big Cypress Swamp (Stim-
son 1956) was devastated by extensive fires
during the spring breeding season (Stimson 1961,
1968; Werner 1976, MacKenzie 1977). Recent
surveys indicate at least a 95% reduction since
1955. Only two singing males were recorded at
Ochopee site in 1975, compared with 10 in 1970
(Werner 1975, 1976, 1979).
The species was regularly sighted on the coastal
marl prairie of Cape Sable between 1918 and
1935 (Howell 1919, 1932; Holt and Sutton 1926;
Nicholson 1928; Semple 1936; Stimson 1956).
Believed to have been extirpated by a severe hur-
ricane on 2 September 1935 (Stimson 1956,
1968), they were rediscovered on Cape Sable near
Little Fox Lake in 1970 (Werner 1971). Altera-
tion of the habitat by the storm of 1935 apparent-
ly is responsible for the population decline. Only
a few widely spaced individuals remain (Werner
1975, 1976).
REPRODUCTION
The potential breeding season slightly exceeds
5 months, extending from February to August.
Up to three broods are produced in a single sea-
son. The pair bond may change between broods
or continue over two consecutive years (Werner
1975, 1979).
Normally, three or four eggs are laid per nest,
rarely two or five. Eggs are incubated by the fe-
male. Incubation requires more than 11 days.
Both parents feed the young. Flightless young
leave the nest 9 to 11 days after hatching; they
are capable of short flights about 2.5 weeks after
hatching (Werner 1976, 1979). They become
independent at about 45 days (MacKenzie 1977).
Fledglings begin to molt in July; the post-
nuptial molt of adults is completed August to
September (Werner 1975).
Photographs of nestlings appear in Werner
(1975).
MANAGEMENT AND CONSERVATION
Recovery efforts have emphasized determina-
tion of ecological requirements as well as restora-
tion and maintenance of habitat. Primary manage-
ment strategies involve controlling water, fire, and
exotics to maintain historic conditions. Acquisi-
tion and management of private land in Big Cy-
press area has been recommended (Werner 1979,
MacKenzie 1977).
Specific management recommendations pro-
vided by Werner (1975) are: (1) eliminate exotic
plants and animals from areas occupied by Cape
Sable sparrows; (2) employ controlled periodic
burns to retard hardwood invasion of suitable
marshes; (3) restrict burning to August and Sep-
tember, to approximate timing of natural light-
ning strikes that follow the breeding season;
(4) start burns with a single ignition spot to pre-
vent entrapment of birds in junction zones;
(5) to promote optimum population size, bum
15% to 20% of the habitat each year, using small,
evenly distributed fires in the oldest roughs; and
(6) the habitat of small colonies should not be
burned, but the burning of limited adjacent areas
could prove beneficial by expanding the habitat.
Critical Habitat has been designated to
include 'areas of land, water, and airspace in the
Taylor Slough vicinity of Collier, Dade, and Mon-
roe Counties' (42 FR 49685, 11 August 1977; 42
FR 47840, 22 September 1977). Rulemaking in-
cludes areas both within and outside Everglades
National Park.
No recovery team has been appointed.
AUTHORITIES
Sonny Bass
Research Biologist
Everglades National Park
P.O. Box 279
Homestead, FL 33030
J. A. Kushlan
Research Biologist
Everglades National Park
P.O. Box 279
Homestead, FL 33030
J.C.Ogden
Research Department
National Audubon Society
115 Indian Mount Trail
Tavernier, FL 33070
W. B. Robertson, Jr.
Research Biologist
Everglades National Park
P.O. Box 279
Homestead, FL 33030
L. A. Stimson
4339 S.W. 9th Terrace
Miami, FL 33134
H. W. Werner
Research Management Specialist
Carlsbad Caverns and Guadalupe Mountains
National Parks
3225 National Park Highway
Carlsbad, NM 88220
PREPARER'S COMMENTS
Previouslv known as Ammospiza mirabilis
(Howell 1932).
PREPARER'S COMMENTS
Previously known as Ammospiza mirabilis
(Howell 1932; Stimson 1954, 1968; American
Ornithologist's Union 1957), the Cape Sable spar-
row was considered the last avian species identi-
fied in continental United States (Stimson 1968;
Werner 1975, 1976). It has recently been designa-
ted a subspecies of A. maritima (Eisenmann
1973). Morphologically and behaviorally similar
to other subspecies of seaside sparrows (Griscom
1944, Stimson 1968), it is unique in its geographi-
cal isolation and confinement to interior marshes
(Werner 1975, 1976, 1979).
LITERATURE CITED/SELECTED
REFERENCES
American Ornithologist's Union. 1957. Check-
Hst of North American Birds, 5th ed. Port
City Press, Baltimore.
Anderson, W. 1942. Rediscovery of the Cape
Sable seaside sparrow in Collier County. Fla.
Nat. 16:12.
Beecher, W. J. 1955. Late-Pleistocene isolation of
salt-marsh sparrows. Ecology 36:23-28.
Dietrich, A. L. 1938. Observations of birds seen
in South Florida. Fla. Nat. 11:101.
Eisenmann, E. (Chairman). 1973. Thirty-second
supplement to the American Ornithologist's
Union check-list of North American birds.
Auk 40:411-419.
Griscom, L. 1944. A second revision of the sea-
side sparrows. La State Univ. Mus; Zool.
Occas. Paper 19:313-328.
Holt, E. G., and G. M. Sutton. 1926. Notes on
birds observed in southern Florida. Ann. Car-
negie Mus. 16:409-439.
Howell, A. H. 1919. Description oa a new seaside
sparrow from Florida. Auk 36:86-87.
. 1932. Florida bird life. Coward-McCann,
Inc., New York. 579 pp.
MacKenzie, J. P. S. 1977. Birds in peril. Hough-
ton Mifflin Co., Boston. 191 pp.
Nicholson, D. J. 1928. Nesting habits of seaside
sparrows in Florida. Wilson Bull. 40:225-237.
. 1938. An historical trip to Cape Sable.
Fla. Nat. 11:41-44.
Ogden, J. C. 1972. Florida region. Am. Birds 26:
852.
Peterson, R. T. 1947. A field guide to the birds.
Houghton Mifflin Co., Boston. 230 pp.
Robertson, W. B., Jr., and J. A. Kushlan. 1974.
The southern Florida avifauna. Pages 414-452
in P. J. Gleason, ed. Environments of South
Florida: present and past. Miami Geological
Society, Memoir 2.
Sample, J. B. 1936. The Cape Sable sparrow and
hurricanes. Auk 53:341.
Sprunt, A., Jr. 1954. Florida bird life. Coward-
McCann, Inc., New York. 527 pp.
Stimson, L. A. 1944. Rediscovery of the Cape
Sable sparrow confirmed. Fla. Nat. 17:31-32.
. 1948. Cape Sable sparrow still in Collier
County. Fla. Nat. 21 :68-69.
1953. Cape Sable seaside sparrow. Fla.
Nat. 26:57.
. 1954, Cape Sable seaside sparrow: Am-
mospiza mirabilis (Howell). Pages 479-481 in
A. Sprunt, Jr. Florida bird life. Coward-Mc-
Cann, Inc., New York. 527 pp,
. 1956. The Cape Sable seaside sparrow:
its former and present distribution. Auk 73:
489-502.
1961. Cape Sable sparrows: fire and
range extension. Fla. Nat. 34:139-140.
. 1968. Cape Sable sparrow. Pages 859-
868 in O. L. Austin, Jr. Life histories of
North American cardinals, grosbeaks, bun-
tings, towhees, finches, sparrows, and allies.
U.S. Natl. Mus. Bull. 237.
U.S. Department of the Interior. 1973. Threat-
ened wildlife of the United States. Compiled
by Office of Endangered Species and Interna-
tional Activities. Bur. Sport Fish. Wildl.
Resour. Publ. 114. U.S. Gov. Printing Office,
Washington, D.C. 289 pp.
Werner, H. W. 1971. Cape Sable sparrows redisco-
vered on Cape Sable. Auk 88:432.
. 1975. The biology of the Cape Sable
sparrow. U.S. Gov. Rep., Everglades Natl.
Park. 215 pp.
. 1976. Distribution, habitat, and origin of
the Cape Sable seaside sparrow. M.S. Thesi
Univ. of Fla., Tampa. 53 pp.
. 1979. Cape Sable seaside sparrow. Pages
19-20 m H. W. Kale, ed. Rare and endangered
biota of Florida, Vol. 2, Birds. Univ. Presses
of Florida, Gainesville.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.12
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
LEATHERBACK SEA TURTLE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoasl of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of F,ndangercd Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.12
March 1980
SELIXTKD V1:R 1 EBRAl i: KNDANGERED SPECIES
OF THE SEACOAS 1 OF 1 HE UNITED S lATES-
LEATHERBAGK SEA TURTLE
A Cooperative Effort
by the
National Fish and WildHfc Laboratory,
the Office of Endangered Species
and the
National (Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidcll Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wiltllife Service
U.S. Department of the Interior
CREDIT: P. C. H. PRITCHARD
LEATHERBACK SEA TURTLE
Dermochelys coriacea Linnaeus
KINGDOM Animalia
CLASS Reptilia
ORDER Testudinata
FAMILY Dermochelyidae
OTHER COMMON NAMES leathery turtle
DATES
Entered into SWIS to be determined
Updates: . .22 September 1976, 20 January 1977.
LEGAL STATUS
Federal: Endangered (35 FR 8495;2June 1970).
States: Endangered: Georgia, Maryland,
Mississippi, New York, North Carolina,
South Carolina, Texas.
Protected: Alabama
REASONS FOR CURRENT STATUS
General population reduction and overuse by
humans are the primary factors determining the
status of leatherback turtles. Exposure and vul-
nerability while nesting make overharvesting by
man possible.
Sea turtles require relatively undisturbed
beaches for nesting. Increasing development and
human activity on beaches is incompatible with
successful turtle reproduction. Illumination of
beaches at night can distract hatchlings away
from the sea (McFarlane 1963).
Harvesting eggs has put tremendous pressure
on this species. Occasionally adults are slaughtered
on nesting beaches. Meat is used as fish bait and
as human food (lUCN 1968, Pritchard 1979a,
Rebel 1974).
Predation is extensive — particularly on hatch-
lings. Pritchard (1971) identified several species
of crabs, fishes, reptiles, and mammals that prey
on hatchlings. Predation on adults is generally
limited to sharks and killer whales {Orcinus orca)
Caldwell and Caldwell 1969).
Incidental catches by commercial shrimping
and fishing boats result in entanglement and sub-
sequent drowning in the trawler nets.
Littering of the seas is believed to have some
impact on sea turtle populations. Plastic bags have
the appearance of jellyfish, but cause death when
eaten by turtles (Rebel 1974).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The largest of all turtles, it is easily disting-
uished by its leathery skin. The neck and limbs
are thick and feebly retractible. Average carapace
length is 155cm. Adults generally weigh from
290 to 590kg. The carapace is triangular, and is
covered with a layer of rubbery skin rather than
horny shields. Carapace has seven longitudinal
ridges. Head and neck are black or dark brown
with a few white or yellow blotches. Each side of
the gray upper jaw has a tooth-like cusp. The
lower jaw is hooked anteriorly. Paddle-like claw-
less limbs are black with white margins, and may
have white spots.
Hatchlings are dark brown or black with
white or yellow carapacial keels and flipper mar-
gins. Skin is covered with small scales that become
thinner with each molt — starting about 3 weeks
after hatching.
Black-and-white photographs are in Carr
(1952, 1967), Bustard (1973), Rebel (1974),
Riedman and Witham (1974), and LeBuff (1976).
Deraniyagala (1939) presents a complete pictorial
and descriptive anatomical discussion.
RANGE
The leatherback is widely distributed in the
oceans of the world. From tropical Atlantic,
Pacific, and Indian Oceans, they are found as far
north as British Columbia, Newfoundland, and
the British Isles. They are found as far south as
Australia, Cape of Good Hope, and Argentina.
Other bodies of water, such as the Mediterranean
Sea are also inhabited.
In spite of what appears to be a large range, it
represents a reduction as compared to the range
in earlier times (lUCN 1968).
Major nesting beaches are in Malaya, Surinam,
French Guiana, Mexico, Costa Rica, South Africa,
Dominican Republic, and U.S. Virgin Islands.
Annual United States nesting is restricted to
Florida, although one nesting incident in 1966 in
North Carolina was reported by Schwartz (Per-
sonal communication). Atlantic Coast nests have
been recorded from Flagler Beach to Miami, with
the majority of records from Palm Beach and
Martin Counties. There appears to be a small
population that nests regularly on Hutchinson
Island, Martin County. On the Gulf Coast, nesting
is common in March and April (A. F. Carr per-
sonal communication).
Two reported nesting occurrences in Florida
have been on publicly owned beaches: Sebastian
State Park, Brevard County, and St. Vincent
National Wildhfe Refuge, Franklin County. Other
recorded nesting beaches are in private ownership.
They are protected by the Florida Department of
Natural Resources which, according to Witham
(Personal communication), protects all nesting
sea turtles.
RANGE MAP
Distribution off the continental United States
is illustrated by shading adjacent coasthnes.
Recorded nesting localities are depicted by dots.
STATES /COUNTIES
Alabama: Baldwin, Mobile.
California: Del Norte, Humboldt, Los Angeles,
Marin, Mendocino, Monterey, Orange,
San Diego, San Francisco, San Luis
Abispo, San Mateo, Santa Barbara,
Santa Cruz, Sonoma, Ventura.
Delaware: Kent, Sussex.
Florida: Bay, Brevard Broward, Charlotte,
Citrus, Collier, Dade, Dixie, Duval,
Flagler, Franklin, Gulf, Hernando,
Hillsborough, Indian River, Jefferson,
Lee, Levy, Manatee, Martin, Monroe,
Nassau, Okaloosa, Palm Beach, Pasco,
Pinellas, St. Johns, St. Lucie, Santa
Rosa, Sarasota, Volusia, Wakulla,
Walton.
Georgia: Bryan, Camdon, Chatham, Glynn,
Liberty, Mcintosh.
Louisiana: Cameron, Iberia, Jefferson, Lafourche,
(Parishes) Plaquemines, St. Bernard, St. Mary,
Terrebonne, Vermilion.
Maine: Cumberland, Hancock, Knox, Lincoln,
Sagadahoc, Waldo, Washington, York.
Maryland: Calvert, Dorchester, Somerset, St.
Marys, Talbot, Worchester.
pp^
SCALE
<^c ;oo 2»o aoo uiles
200 300 40O ■iLOMC'reBs
CONIC PROJECTION
vest LOMCtTWOC
Eastern distribution of leatherback sea turtle offshore (shading) and nesting beaches (dots)
Massachusetts: Barnstable, Dukes, Essex, Middle-
sex, Nantucket, Norfolk, Plymouth.
Mississippi: Hancock, Harrison, Jackson.
New
Hampshire: Rockingham.
New Jersey: Atlantic, Cape May, Cumberland,
Monmouth, Ocean.
New York: Nassau, Suffolk.
North
Carolina: Brunswick, Carteret, Currituck, Dare,
Hyde, New Hanover, Onslow, Pender.
Oregon: Clatsop, Coos, Curry, Douglas, Lane,
Lincoln, Tillamook.
Rhode
Island: Newport, Washington.
South
Carolina: Beaufort, Charleston, Colleton,
Georgetown, Horry.
Texas: Aransas, Brazoria, Calhoun, Cameron,
Chambers, Galveston, Jefferson,
Kenedy, Kleberg, Matagorda, Nueces,
Willacy.
Virginia: Accomack, Gloucester, Lancaster,
Mathews, Middlesex, Northampton,
Northumberland, Virginia Beach,
York.
Washington: Clallam, Grays Harbor, Jefferson,
Pacific.
HABITAT
Leatherbacks are the most pelagic of the sea
turtles, and are often found near the edge of the
continental shelf. In Northern waters, they some-
times enter shallow estuarine bays (Deranigayala
1939, Pope 1939).
FOOD AND FORAGING BEHAVIOR
Although apparently omniverous, consuming
sea urchins, squid, crustaceans, tunicates, fish,
blue-green algae, and floating seaweed, its princi-
pal food is jellyfish. Deeply notched , sharp edged
jaws appear adapted for holding and cutting soft-
bodied prey. The mouth has fleshy papillae, and
the throat has a valve which probably assist in
swallowing and retaining soft-bodied prey (Prit-
chard 1971).
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
Leatherbacks require a sloping sandy beach
backed up by vegetation for nesting. There must
be sufficient slope so that the crawl to dry sand
is not too far. The depth of the coarse dry sand is
important because the female first excavates a pit
for her body and then must reach moist sand so
that she can make the proper flask-shaped nest
(Pritchard 1969a, Witham 1976). Preferred
beaches are mainland or island areas near deep
water and rough seas (Rebel 1974).
Site specificity such as that displayed by the
green turtle (Chelonia mydas) is not apparent for
the leatherback. Lund (1974) suggests renesting is
generally greater than 7 miles from the initial nest.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
Estimates are based on counts of nesting fe-
males: Trengganu (Malaya), 4,000; French Guiana,
15,000; Costa Rica, 1,000; Trinidad, Surinam,
Tongaland, and Ceylon, 200 to 400; and Jalsico
(Mexico) to northern Peru, 5,000. Pritchard
(1969b) estimates the world female breeding
population to be from 29,000tto 40,000.
Lund (1974) estimates that about 25 leather-
back nest in Florida each year; The number of
females involved is uncertain.
REPRODUCTION
The nesting season varies per locality; Suri-
nam and Guyana, May to July; Costa Rica, April
to July; Silebache (includes French Guiana) Miy
to August; Trinidad, March to August; and the
Danish West Indies, March to May (Rebel 1974).
Florida nesting season is from April to August.
Females nest at night at intervals of 2 to 3
years. As many as six cluches may be laid a
season with an average inter-nesting period of 10
days (Pritchard 1969a; Lund 1976) Clutches av-
erage 80 to 85 eggs with the last layer of eggs
generally abnormal. The white spherical eggs are
approximately 50 to 54 mm in diameter (Pope
1939; Lund 1976). Incubation takes from 55 to
74 days and emergence of the hatchlings occurs at
night.
Animals mature in 6 to 10 years. Mating takes
place in shallow water offshore of the laying
beach. Occasionally males will crawl up the beach
in an attempt to mate with nesting females (Le-
Buff 1976). Photographs of courtship and mating
are in Bustard (1973).
MANAGEMENT AND CONSERVATION
The Malayan Nature Society transplanted eggs
into enclosures to increase numbers of hatchlings.
This project is now under the control of Malayan
Fisheries Department (lUCN 1968).
Marquez M. (1976) recommended formation
of seven natural reserves for the coast of Mexico.
These are the major Mexican sea turtle nesting
sites and include Playa de la Escolbilla, Oaxaca
State, a leatherback nesting beach.
Other protective measures should include ef-
forts to curtail the loss of leatherbacks in fishing
or shrimping trawls, protection of nesting beaches
from turtles, and a limitation of development on
nesting beaches (Pritchard 1971).
Legal protection to varying degrees exists in
Mexico, Costa Rica, Surinam, French Guiana,
Ascension Island, Trust Territory of the Pacific,
Tahiti, Fiji Islands, Kingdom of Tonga, Australia,
Sarawak, British Indian Ocean, Seychelle's Islands,
South Africa and the United States (Pritchard
1969b; U.S. Department of Commerce 1976).
The leatherback is listed in Appendix 1 of
1976 Convention on Internation Trade in Endan-
gered Species of Wild Fauna and Flora. Commer-
cial trade in this species is subject to strict regula-
tion, and both an export and import permit are
necessary for trade by participating countries.
Critical Habitat has been determined to in-
clude a major nesting area on Sandy Point Beach
at the western edge of St. Croix, U.S. Virgin Is-
lands (43 FR 43688-43689; 26 September 1978),
as well as adjacent waters (44 FR 17710-17712;
23 March 1979).
AUTHORITIES
Archie Carr
Department of Zoology
University of Florida
Gainesville, FL 32611
George Hughes
Oceanographic Research Institute
P.O. Box 736
Durban, Natal
South Africa
Frank Lund
P.O. Box 541
Jupiter Island, FL 33458
Nicholas Mrosovsky
Department of Zoology
University of Toronto
Ontario , Canada M5 3 1 A 1
Peter Pritchard
Florida Audubon Society
P.O. Drawar 7
Maitland,FL 32751
J. P. Schults
Surinam Forest Service
P.O. Box 436
Paramaribo, Surinam
PREPARER'S COMMENTS
Nesting records for the United States are not
complete because the entire coast has not been
surveyed.
LITERATURE CITED/SELECTED
REFERENCES
Brongersma, L. D. 1970. Miscellaneous notes on
turtles. 111. Koninkl. Nederl. Akademic Weten-
schappen-Amoterdam. Proc. Serv. C. 73(4):
323-335.
Bustard, R. 1973. Sea turtles, natural history, and
conservation. Taplinger Publ.,New York. 220
pp.
Caldwell, D. K. 1959. On the status of the Atlan-
tic leatherback turtle, Dermochelys coriacea
coriacea, as a visitant to Florida nesting
beaches, with natural history notes. Quart. J.
Florida Acad. Sci. 21(3):285-291.
Caldwell, D. K., and M. C. Caldwell. 1969. Addi-
tion of the leatherback sea turtle to the
known prey of the killer whale, Orcinus orca..
J. Mammal. 50(3):636.
Caldwell, D. K., and W. F. Rathjen. 1969. Unre-
corded West Indian nesting sites for the leath-
erback and hawksbill sea turtles, Dermochelys
coricaea and Ertmochelys i. imbricata. Copeia
1969(3):622-623.
Carr, A. F. 1952. Handbook of turtles. Turtles of
the United States, Canada, and Baja Califor-
nia.Comstock Publ. Assoc, Cornell Univ.
Press, Ithaca, N.Y. 542 pp.
. 1967. So excellent a fishe. The Natural
History Press, Garden City, N.Y. 248 pp.
Carr, A. F., and A. R. Main. 1973. Report on an
inquiry into ecological implications of a turtle
farming project. In Turtle farming project in
northern Australia. Union Offset Co., PTY,
Limited, Canberra, Australia. 80 pp.
Carr, A. F., and L. H. Ogren. 1959. The ecology
and migrations of sea turtles, 3. Dermochelys
in Costa Rica. Ame. Mus. November 1958. 29
pp.
Deraniyagala, P. E. P. 1939. Tetrapod reptiles of
Ceylon. Vol. 1. Columbo Mus. Publ. Ceylon.
412 pp.
Ernst, C. H. and R. W. Barbour. 1972. Turtles of
the United States. Univ. Kentucky Press, Lex-
ington. 347 pp.
Frair, W., R. G. Ackman, and N. Mrosovsky.
1972. Body temperature oi Dermochelys cor-
iacea: Warm turtle from cold water. Science
177:791-0793.
lUCN. 1968. Red data book. Vol. 3. Amphibia
and reptiha. Compiled by R. E. Honegger.
lUCN, Morges, Switzerland.
LeBuff, C. R., Jr. 1976. Tourist turtle. Florida
Wildl. 30(2): 16-17.
Lund, F. 1974. A survey of marine turtle nesting
in the United States. Unpubl. 39 pp.
. 1979. Atlantic leatherback. Pages 54-55
in R. W. McDiarmid ed. Rare and endangered
biota of Florida, Vol. 3, Amphibians and rep-
tiles. Univ. Presses of Florida, Gainesville.
Marquez M., R. 1976. Reserves naturales papa la
conservation de las tortugas marinas en Mexi-
co. INP/SI 83:1-22.
McFarlane, R. W. 1963. Disorientation of logger-
head hatchlings by artificial road lighting.
Copeial963(l):153.
Montoya, A. E. 1969. Programas de investigacion
y conservacion de las tortugas marinas en
Mexico. Pages 34-53 in Marine turtles. lUCN
New Publ. Ser. Suppl. Paper 20.
Neill, W. H., E. R. Stevens, E. G. Carey, K. D.
Lawson, N. Mrosovsky, and W. Frair. 1974.
Thermal inertia versus thermoregulation in
'warm' turtles and tunas. Science 184:1008-
1010.
Pope, C. H. 1939. Turtles of the United States
and Canada. Alfred A. Knopf, Inc., New
York. 343 pp.
Pritchard, P. C. H. 1969a. Sea turtles ot the Gui-
anas, Bull. Fla. St. Mus. 13(2): 120-132.
. 1969b. Summary of world sea turtle su-
vival situation. lUCN Bull. 2(11):90-91.
1971. The leatherback or leathery turtle
Dermochelys coriacea. lUCN Monogr. 1.
Morges, Switzerland. 39 pp.
Rebel, T. P. 1974. Sea turtles and the turtle in-
dustry of the West Indies, Florida, and the
Gulf of Mexico. Rev. Ed. Univ. of Miami
Press, Coral Gables. 250 pp.
Riedman, S. R., and R. Witham. 1974. Turtles
extinction or survival? Abelard-Schuman,
New York. 156 pp.
Survival Service Commission. 1969. Marine tur-
tles. Proc. working meeting of marine turtle
specialists organized by lUCN at Morges,
Switzerland.
Tweedie, M. W. F. 1953. The breeding of the
leathery turtle. Proc. Zool. Soc. London 123
(2):273-275.
U.S. Department of Commerce. 1976. Proposed
listing of the green sea turtle [Chelonia my-
das), loggerhead {Caretta caretta), and pacific
ridley sea turtle (Lepidochelys olivacea) as
threatened species under the Endangered
Species Act of 1973. Draft EIS. Natl. Oceanic
Atm. Admin., Natl. Marine Fish. Serv. Wash-
ington, D.C.
Witham, R. 1976. Dermochelys coriacea. In H. O.
Hillestad, D. B. Means, and W. W. Baker, eds.
Endangered and threatened vertebrates of the
southeastern United States. Tall Timbers Res.
Stn. Misc. Publ. 4.
Worth, R. F., and J. B. Smith. 1976. Marine turtle
nesting on Hutchinson Island, Florida in
1973. Florida Dep. Nat. Resour. Marine Res.
Lab. Florida Mar. Res. Publ. 18.
Yerger, R. W. 1965. The leatherback turtle on the
Gulf coast of Florida. Copeia 1963 (3):365-
366.
Zim, H. S., and H. M. Smith. 1953. Reptiles and
amphibians. A Guide to familiar America spe-
cies. Simon and Schuster, New York. 147 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.13
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
6REEN SEA TURTLE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubHc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of F^ndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Ct)mputer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.13
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF IHE UNFI ED STATES-
GREEN SEA TURTLE
A Cooperative Effort
by the
National Pish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. VVoodard
National Coastal Ecosystems Team
NASA-Slidcll Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
Credit: O. James
GREEN SEA TURTLE
Chelonia mydas Linnaeus
KINGDOM Animalia
CLASS Reptilia
ORDER Testudinata
FAMILY Cheloniidae
OTHER COMMON
NAMES Edible turde, tortuga verde.
DATE
Entered into SWIS to be determined.
Updates. . .22 September 1976, 8 February 1977.
LEGAL STATUS
Federal Endangered : waters of Florida and Pa-
cific Coast of Mexico (43 FR 32800-
32811, 28 July 1978).
Threatened: elswhere throughout its
range (43 FR 32800-32811, 28 July
1978).
States: Endangered: Florida, Maryland, Mis-
sissippi, New Jersey, Texas.
Protected: Alabama, Georgia, North
Carolina, South Carolina.
REASONS FOR CURRENT STATUS
The green turtle's vulnerability while nesting
has led to its overexploitation for food by local
populations seeking a readily available source of
protein food. Recent technological advances such
as freezing and canning have increased its use for
food, and an increase in demand for turtle pro-
ducts (leather, cosmetics, and pharmaceuticals)
has placed considerable stress on the species.
Nesting populations in Bermuda, Florida, the
Greater Antilles, and Jamaica have been extir-
pated or nearly so (U.S. Department of Com-
merce 1976).
In many areas, excessive predation on eggs
and hatchlings has substantially reduced recruit-
ment, causing populations to decrease.
Relatively undisturbed beaches are necessary
for nesting. Increasing development and use of
beaches is incompatible with sea turtle reproduc-
tion. Illumination of beaches at night, for exam-
ple, can distract hatchlings away from the sea
(McFarlane 1963).
Sea turtles are caught incidentally to commer-
cial fishing and shrimping activities. Some turtles
are eaten by fishermen, some are sold in local
markets, and some are mutilated or drowned as
a result of entanglement in trawls (U.S. Depart-
ment of Commerce 1976).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The green turtle is a large sea turtle with a
broad, heart-shaped shell and small head. Flippers
are paddle shaped, each with a single claw. Adults
are 91 to 122 cm long and weigh 100 to 200 kg.
The color varies widely. In general, hatch-
lings have black carapaces, white plastrons, and
white margins on the shell and limbs. Adults have
a smooth, keelless carapace, colored light to dark
brown, with brown mottling. The plastron is
whitish to hght yellow; the upper surface of the
head is light brown with yellow markings; sides of
the head are brown with broad yellow margins;
the neck is dusky above and yellow near the shell
below. The tail and flippers are colored like the
carapace and plastron.
The carapace can be identified by four costal
plates, none of which borders the nuchal shield,
and by the absence of jagged marginals. Large
scutes on the carapace do not overlap. There is
only one pair of prefrontals between the eyes.
This species is illustrated in Carr (1967), Par-
sons (1962), Ernst and Barbour (1972), Rebel
(1974), and Riedman and Witham (1974).
RANGE
The green turtle is distributed world-wide in
waters above 20° C in the coldest month. Juveniles
are sometimes found over a wider temperature
range and thus a greater area. Green turtles live in
waters off the North American coast from Massa-
chusetts to Mexico and from British Columbia to
Baja California (U.S. Department of Commerce
1976). They are uncommon off California.
Major nesting grounds in the Western Hemi-
sphere include the Michoacan Coast, Mexico; Tor-
tuguero, Costa Rica; Shell Beach, Guyana; Bigi
Santi, Surinam; and Aves Island. See Hirth (1971)
for nesting sites in the Eastern Hemisphere.
Known annual nesting in the continental U.S.
is limited to small nesting populations of the east
coast of Florida, from Brevard County to Brow-
ard County. Jupiter and Hutchinson Islands have
the greatest number of nests. For detailed infor-
mation on nesting at Hutchinson Island, see Galla-
ger et al. (1972) and Worth and Smith (1976).
Their former distribution included nesting
beaches on the Dry Tortugas, Cayman Islands,
several other previously undisturbed islands, and
more extensively in Florida (lUCN 1968).
A small population of Pacific green turtles
{Chelonia mydas agassizii) bask and nest on some
of the islands in the Hawaiian Islands National
Wildlife Refuge. These islands are Federally owned
and protected, although until recently, according
to Balazs (1976), turtles had been harvested.
In Florida, green turtles have nested on pub-
licly owned lands in Merritt Island and Hobe
Sound National Wildlife Refuges, and St. Lucie
and Sebastian State Parks, Other nesting beaches
are privately owned and the Florida Department
of Natural Resources, according to R. Witham
(personal communication), protects all nesting on
these beaches.
RANGE MAP
Distribution offshore continental U.S. is illus-
trated on the following maps by shading of the
adjacent coastline; nesting beaches are depicted
by dots.
STATES/COUNTIES
Alabama Baldwin, Mobile. Mobile.
California Del Norte, Humboldt, Los Angeles,
Marin, Mendocino, Monterey, Orange,
San Diego, San Francisco, San Luis
Obispo, San Mateo, Santa Barbara,
Santa Cruz, Sonoma, Ventura.
Delaware Sussex.
Florida Bay, Brevard, Broward, Charlotte, Cit-
rus, Collier, Dade, Dixie, Duval, Flag-
ler, Franklin, Gulf, Hernando, Hillsbo-
rough, Indian River, Jefferson, Lee,
Levy, Manatee, Martin, Monroe,
Nassau, Okaloosa, Palm Beach, Pasco,
Pinellas, St. Johns, St. Lucie, Santa
Rosa, Sarasota, Volusia, Wakulla, Wal-
ton.
Georgia Bryan, Camden, Chatham, Glynn, Li-
berty, Mcintosh.
SCALE
0 to
ioo
io ioo ^^o
_^oo »,
lE
0 -ex
"°
3CMJ 400
K,.0«£T
"
CONIC
PROJECTION
Eastern distribution of green sea turtles offshore (shading) and nesting beaches (dots)
4
Louisiana
(parishes)
Maryland
Massa-
chusetts
Mississippi
New
Jersey
Cameron, Iberia, Jefferson, Lafourche,
Plaquemines, St. Bernard, St. Mary,
Terrebonne, Vermilion.
Worcester.
Barnstable, Dukes, Essex, Middlesex,
Nantucket, Norfolk, Plymouth.
Hancock, Harrison, Jackson.
Atlantic, Cape May, Monmouth,
Ocean.
New York Nassau, Suffolk.
North
Carolina
Oregon
Rhode?
Island
South
Carolina
Texas
Virginia
Wash-
ington
Brunswick, Carteret, Currituck, Dare,
Hyde, New Hanover, Onslow, Pender.
Clatsop, Coos. Curry, Douglas, Lane,
Lincoln, Tillamook.
Newport, Washington.
Beaufort, Charleston, Colleton,
Georgetown, Horry.
Aransas, Brazoria, Calhoun, Cameron,
Chambers, Galveston, Jefferson,
Kenedy, Kleberg, Matagorda, Nueces,
Willacy.
Accomack, Northampton.
Clallam, Grays Harbor, Jefferson, Pa-
cific.
HABITAT
The green turtle inhabits comparatively shal-
low waters inside reefs and in bays and inlets.
Favored habitat appears to be lagoons and shoals
with an abundance of marine grass and algae. This
habitat type occurs in much of the Gulf of Mexi-
co and Caribbean shore waters and around many
oceanic islands. Green turtles are long-distance
migrants and are occasionally seen in the open sea
en route from feeding grounds to nesting beaches
or vice versa (Carr 1952, 1967).
FOOD AND FORAGING BEHAVIOR
Adults are largely herbivorous, feeding on
marine algae and shallow water pastures of marine
grasses (including Thalassia, Zostera, Cymodocea,
and Halophila) (Carr 1952, Randall 1965). Small
mollusks, sponges, crustaceans, and jellyfish are
often consumed (Carr 1952,Hirth 1971).
SHELTER REQUIREMENTS
Pacific green turtles bask on reefs or uninhab-
ited islands.
Some green turtles have been known to rest
with their shells wedged under rocks or reefs on
the bottom of the sea (Carr 1952).
Witham (1976b) states that hatchlings find re-
fuge and food in sargassum [Sargassum sp.). Frick
(1975) observed hatchlings resting on sargassum
clumps.
NESTING OR BEDDING
Successful nesting requires a sloping beach
platform with open ocean exposure and minimal
disturbance. A variety of textures of sand can be
used, but it must be friable and well drained (A.
F. Carr personal communication). The presence
or absence of vegetation does not appear to be
critical; vegetation can be helpful in nesting be-
cause roots may prevent the sand from crumbling;
on the other hand, it can be detrimental if the
roots pierce the eggshells (Caldwell 1959). On
beaches without rooted plants, rain or the wetting
of the sand by the turtle aids in preventing the
sand from crumbling (Bustard 1973, Carr and
Main 1973).
Illustrated descriptions of nesting appear in
Hendrickson (1958), Carr and Ogren (I960), and
Hirth(1977).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
Estimates of the breeding female population
for the Western Hemisphere nesting areas are:
Sarawak Turtle Islands 10,000
Surinam 2,500
Tortuguero 6,000
Aldabra fewer than 1,000
Australia 75,000, but includes
flatbacks {Chelonis depressa) and
loggerheads {Caretta caretta)
Florida 50
(Hirth 1971, Lund 1974).
Ehrenfeld (1974) estimates the world popu-
lation of green turtles at between 100,000 and
400,000.
Numbers fluctuate vvdth losses of nesting
beaches to storms or development. Thus, it is
difficult to predict trends. According to A. F.
Carr (personal communication), persons inves-
tigating green turtles around the world are con-
cerned with the apparent decline in total numbers.
REPRODUCTION
Breeding season varies with locality:
Michoacan Coast, Mexico .... May to September
Tortuguero, Costa Rica June to November
Shell Beach, Guyana March to August
Bigi Santi, Surinam February to August
Aves Island March to December
Florida April to July
(Hirth 1971, Rebel 1974).
Nocturnal nesting occurs at 2-, 3-, or 4-year
intervals (Carr and Ogren 1960, Hirth 1971). As
many as seven clutches are laid in one season,
with renesting occurring at 9- to 13-day intervals.
Carr and Ogren (1960) plotted returns of turdes
at Tortuguero in 1956 through 1959 and observed
that renesting was usually within 1.5 km of the
previous site.
Clutch size varies from 75 to 200 eggs with in-
cubation taking from 48 to 70 days, depending
on beach and water conditions (Carr and Hirth
1962). Hatchlings generally emerge at night with-
in a period of 48 hours (lUCN 1968, Rebel 1974).
Animals are believed to reach maturity in 4 to
6 years in tropical waters and 5 to 13 years in
temperate waters. Hirth (1971) estimates survival
rates to maturity to be 1% to 3% of the hatch-
lings. A. F.Carr (personal communication to L.H.
Ogren 1975) estimates survivorship in Costa Rica
to be about 0.1% of the hatchlings.
Copulation occurs near the nesting beach
(Bustard 1973). Whether the female stores sperm
for successive laying seasons has not been deter-
mined. Photographs of courtship and mating
appear in Booth and Peters (1972).
MANAGEMENT AND CONSERVATION
Carr (1969) has made several suggestions for
protection of the green turtle: Export of turtle
products should be discontinued and catches for
local use should be strictly supervised. No turtle
boats should be permitted to operate off the
northern 20 km of Tortuguero Beach. The sale or
storage of calipee in Costa Rica should also be
prohibited.
The Survival Service Commission (1969) sug-
gests that setting aside Ascension, Astove, Aves,
Aldabra, Jabal Aziz, Sabah Turtle Islands, and the
French Frigate Shoal as turtle islands could help
stabilize turtle populations in those areas.
Marquez M. (1976) recommends formation of
several natural reserves for the coast of Mexico,
which include the major Mexican sea turtle nest-
ing sites. Two reserves, Playa de Rancho Nuevo,
Tamaulipas State, and Isla Contoy, Quintana Roo
State, are green turtle nesting areas.
Cayman Turtle Farm, Ltd., has made exten-
sive efforts at captive breeding. At present, the
farm is dependent on natural stocks of eggs, and
according to Hirth (1971), the project cannot be
considered a complete success until it is indepen-
dent of those sources. Some biologists suggest
that such a project may never be ecologically or
economically efficient. Others are concerned that
turtle production could increase demand and thus
increase the pressure on natural stocks (Ehrenfeld
1974).
Hatcheries operate in Australia, Malaysia,
Mexico, United States, and other countries. These
and similar efforts to curtail natural predation
could have a beneficial impact on numbers.
Stocking has been attempted in many areas, but
there have been no indications of definite success.
Green turtles are legally protected to varying
degrees in the following places: Mexico, Costa
Rica, Panama, Surinam, French Guiana, Ascen-
sion Island, Trust Territory of the Pacific, Tahiti,
Fiji Islands, Kingdom of Tonga, Australia, Sara-
wak, British Indian Ocean and Seychelles Islands,
Israel, South Africa, Europa Islands, and the U.S.
(Pritchard 1969, U.S. Department of Commerce
1976,43 FR 32800-32811).
Most populations are hsted under Appendix I
of the Convention on International Trade in
Endangered Species of Wild Fauna and Flora;
commercial trade is subject to strict regulation,
and both an export and import permit are neces-
sary for trade by participating countries. Austra-
lian populations are listed under Appendix II,
which requires export permits for trade in those
populations.
AUTHORITIES
George H. Balazs
University of Hawaii at Manoa
Hawaii Institute of Marine Biology
P.O. Box 1346, Coconut Island
Kaneohe, HI 96744
Archie Carr
Department of Zoology
University of Florida
Gainesville, FL 32611
Harold Hirth
Department ofBiology
University of Utah
Salt Lake City, UT 84112
George Hughes
Oceanographic Research Institute
P.O. Box 736
Durban, Natal
South Africa
Frank Lund
P.O. Box 541
Jupiter Island, FL 33458
Peter Pritchard
Florida Audubon Society
P.O. Drawer 7
Maitland,FL 32751
J. P. Schulz
Surinam Forest Service
P.O. Box 436
Paramaribo, Surinam
Ross Witham
Florida Department of Natural
Resources
Marine Research Lab
P.O. Box 941
Jensen Beach, FL 33457
PREPARER'S COMMENTS
Nesting records for the U.S. are not complete,
as the entire coast has not been surveyed. Recruit-
ment in Florida is probably very low due to pre-
dation, particularly by raccoons [Procyon lotor).
LITERATURE CITED/SELECTED
REFERENCES
Allen, E. R., and W. T. Neill. 1953. Know your
reptiles: the green turtle. Florida Wildl. 7(4):
19,32.
Banks, E. 1937. The breeding of the edible turtle
(Chelonia mydas). Sarawak Mus. J. 4(15):
523-532.
Balazs, G. H. 1976. Green turtle migrations in the
Hawaiian Archipelago. Biol. Conserv. 9:125-
140;
Booth, J., and J. A. Peters. 1972. Behavioural
studies on the green turtle {Chelonia mydas)
in the sea. Anim. Behav. 20(4):808-812.
Bustard, H. R. 1970. The adaptive significance
of coloration in hatchling green sea turtles.
Herpetologica 26:224-227.
• 1973. Sea turtles, natural history and
conservation. Taplinger Publ., New York
220 pp.
Caldwell, D. K. 1959. The loggerhead turtles of
Cape Romain, South Carolina. Bull. Florida
St. Mus.4(10):340.
. 1963. The sea turtle fishery of Baja
California, Mexico. Calif. Fish and Game
49(3):140-151.
Caldwell, D. K., and A. F. Carr. 1957. Status of
the sea turtle fishery in Florida. Trans. N.
Am. Wildl. Comm. 22:457-463.
Carr, A. F. 1952. Handbook of turtles: The
turtles of the United States, Canada, and Baja
California. Comstock Publ. Assoc, Cornell
Univ. Press, Ithaca, New York. 542 pp.
• 1964. Transoceanic migrations of the
green sea turtle. Bioscience 14(8):49-52.
. 1967. So excellent a fishe. The Natural
History Press, Garden City, N. Y. 248 pp.
. 1969. Sea turtle resources of the Carib-
bean and Gulf of Mexico. lUCN Bull. 2(10):
74-83.
. 1970. Green sea turtles in peril. Natl.
Parks Conserv. Mag. 44(271): 19-24.
1972. Great reptiles, great enigmas.
Audubon Mag. 74(2)24-35.
-. 1975. The Ascension Island green turtle
colony. Copeia 1975(3):547-555.
Carr, A. F., and H. Hirth. 1962, The ecology and
migrations of sea turtles: 5 comparative fea-
tures of isolated green turtle colonies. Am.
Mus. Novit. 2091:1-42.
Carr, R. F., and R. M. Ingle. 1959. The green tur-
tle {Chelonia mydas) in Florida. Bull Mar. Sci.
Gulf-Carib. 9(3):315-320.
Carr, A. F., and A. R. Main. 1973. Report on an
inquiry into ecological implications of a turtle
farming project. In Turtle farming project in
northern Australia. Union Offset Co., Pty.,
Limited, Canberra, Australia. 80 pp.
Carr, A. F., and L. Ogren. 1960. The ecology and
migrations of sea turtle, 4. The green turtle in
the Caribbean Sea. Bull. Am. Mus. Nat. His-
tory 121(l):l-48.
Cox, G. A., and R. G. Mauermann. 1976. Inciden-
tal catch and disposition of sea turtles by the
Brownsville-Port Isabel Gulf shrimp fleet. Un-
publ.
Deraniyagala, P. E. P. 1939. Tetrapod reptiles of
Ceylon. Vol. 1. Columbo Mus. Publ. Ceylon.
412 pp.
Ehrenfeld, D. 1974. Conserving the edible sea tur-
tle: Can mariculture help? Am. Sci. 62(1):23-
31.
Ehrhart, L. M. 1976. Final report to the National
Aeronautics and Space Administration Ken-
nedy Space Center. A study of a diverse
coastal ecosystem of Florida. Office of Gradu-
ate Studies and Research, Florida Technologi-
cal Univ. Orlando. Unpubl.
Ernst, C. H., and R. W. Barbour. 1972. Turtles of
the United States. Univ. Kentucky Press, Lex-
ington. 347 pp.
Frick, J. 1971. Observations on sea turtles at Al-
dabra Atoll. Phil. Trans. Roy. Soc. London
Ser. Bull. 260:273-410.
Gallagher, R. M., M. H. Hollinger, R. M. Ingle, and
C. R. Futch. 1972. Marine turtle nesting on
Hutchinson Island, Florida in 1971. Florida
Dep. Nat. Resour. Spec. Sci. Rep. 37. 11 pp.
Harrisson, T. 1956. The edible turde {Chelonia
mydas) in Borneo. 4. Growing turtles and
growing problems. Sarawak Mus. J. 7(7):233-
239.
Hendrickson, J. R. 1958. The green sea turtle,
Chelonia mydas, (Linn.) in Malaya and Sara-
wak. Proc. Zool. Soc, London. 130(4) :455-
535.
Hendrickson, J. R., and E. Balasingam. 1966.
Nesting beach prefemces of Malayan sea tur-
tles. BuU. Natl. Mus., Singapore. 33(10):69-76.
Hirth, H. F. 1971. Synopsis of biological data on
the green turtle, Chelonis mydas. F.A.O. Fish.
Syn. 85. Firm/585. SAST-Green Turtle-5.31
(07)005.02.
Ingle, R. M., and W. F. G. Smith. 1949. Sea tur-
tles and the turtle industry of the West Indies,
Florida, and the Gulf of Mexico. Univ. Miami.
Press, Miami.
lUCN. 1968. Red data book. Vol. 3. Amphibia
and reptiha. Compiled by R. E. Honegger.
lUCN, Morges, Switzerland.
Lund, F. 1974. A survey of marine turtle nesting
in the United States. Unpubl. 39 pp.
.1979. Atlantic green turtle. Pages 23-24
in R. W. McDiarmid ed. Rare and endangered-
biota of Florida, Vol. 3. Amphibians
and reptiles. Univ. Presses of Florida, Gaines-
ville.
Marquez M., R. 1976. Reservas naturales para la
conservacion de las tortugas marinas en Mexi-
co. INP/SIi83: 1-22.
McFarlane, R. W. 1963. Disorientation of logger-
head hatchlings by artificial road lighting.
Copeia 1963(1): 153.
Montoya, A. E. 1969. Programas de investigacion
y conservacion de las tortugas marinas en
Mexico. Pages 34-53 in Marine turtles. lUCN
New Publ. Ser. Suppl. Paper 20.
Parsons, J. J. 1962. The green turtle and man.
Univ. of Florida Press, Gainesville. 126 pp.
Pope, C. H. 1939. Turtles of the United States
and Canada. Alfred A. Knopf, Inc. New York.
343 pp.
Pritchard, P. C. H. 1969. Summary of world sea
turtle survival. lUCN Bull. 2(11):90-91.
Randall, J. 1965. Grazing effect on sea grasses by
herbivorous reef fishes in the West Indies.
Ecology 46:225.
Rebel, T. P. 1974. Sea turtles and the turtle in-
dustry of the West Indies, Florida, and the
Gulf of Mexico. Univ. of Miami Press, Miami.
250 pp.
Riedman, S. R., and R. Witham. 1974. Turtles ex-
tinction or survival? Abelard-Schuman, New
York. 156 pp.
Schwartz, F.J. 1976. Behavioral and tolerance re-
sponses to natural cold winter water tempera-
tures by three species of sea turtles in North
Carolina. Page 5 in Florida and interregional
conference on sea turtles. Florida Dep. Nat.
Resour. Marine Res. Lab. Unpubl.
Survival Service Commission. 1969. Marine turtles.
Proc. Work. Meet. Marine Turtle Special,
organized by lUCN at Morges, Switzerland.
True, F. W. 1884. The fisheries and fishery indus-
tries of the United States, Sec. 1, Pt. 2. The
useful aquatic reptiles and batrachians. U.S.
Comm. Fish.
U.S. Department of Commerce. 1976. Proposed
listing of the green sea turtle [Chelonia my-
das), loggerhead [Caretta caretta), and Pacific
ridley sea turtle {Leptdochelys olivacea) as
threatened species under the Endangered Spe-
cies Act of 1973. Draft EIS. Natl. Oceanic
Atm. Admin., Natl. Mar. Fish. Serv. Washing-
ton, D.C.
U.S. Department of the Interior. 1973. Threat-
ened wildlife of the United States. Compiled
by Office of Endangered Species and Interna-
tional Activities, Bur. Sport Fish. Wildl. Re-
sour. Publ. 114. U.S. Gov. Printing Office,
Washington, D.C. 289 pp.
Walker, W. F. 1971. Swimming in sea turtles of
the family Cheloniidae. Copeia 1971(2):229-
233.
. 1976a. Chelonia mydas account. In H.
O. Hillestad, D. B. Means, and W. W. Baker,
eds. Endangered and threatened vertebrates of
the southeastern United States. Tall Timbers
Res. Sta. Misc. Publ. 4.
1976b. Evidence for ocean-current me-
diated dispersal in young turtles, Chelonia
mydas (Linnaeus). M.S. Thesis. Univ. of Okla-
homa, Norman. 48 pp.
Worth, D. F., and J. B. Smith. 1976. Marine turtle
nesting on Hutchinson Island, Florida, in
1973. Florida Dep. Nat. Resour. Mar. Res.
Lab. Publ. 18.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.14
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE TEXAS BLIND SALAMANDER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal F^cosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
FWS/OBS-80/01.14
March 1980
SELECTED VERTEBRA IE ENDANGERED SPECIES
OF THE SEACOAST OF FHE UNITED STATES-
THE TEXAS BLIND SALAMANDER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and W'iltllife Service
U.S. Department of the Interior
CREDIT: J.JOHNSON
TEXAS BLIND SALAMANDER
Typhlomolge rathbuni Stejneger
KINGDOM Animalia
CLASS Amphibia
ORDER Caudata
FAMILY Plethodontidae
OTHER COMMON NAMES None
DATE
Entered into SWIS to be determined.
Updates 21 November 1977, 31 March 1978,
22 August 1978.
LEGAL STATUS
Federal Endangered (32 FR 4000, 11 March
1978.
States Endangered: Texas
REASONS FOR CURRENT STATUS
The Texas blind salamander is endangered be-
cause of its extremely restricted distribution in a
fragile subterranean ecosystem. At accessible lo-
cations (especially Ezell's Cave), sightings de-
clined sharply in the 1960's, due probably to
overcollecting by scientific, commercial, and
hobbyist collectors. When the cave was sealed to
prevent human entrance, the bat colony that
supplied a presumably important energy resource
in the form of guano was eliminated.
Recent research (Longley 1978) has shown
that the major part of the population is located in
inaccessible parts of the Edwards Aquifer and is
probably stable at present. However, there is a po-
tential for contamination of the aquifer, as well as
evidence that extensive groundwater withdrawal
is causing the head in that aquifer to decrease.
This could ultimately lead to intrusion of poor
quality water from adjacent aquifers (Longley
1978).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The Texas blind salamander is all white or
pinkish with blood-red external gills and tooth-
pick-like legs. Head and snout are strongly flat-
tened with two small black dots representing
vestigial eyes beneath the skin. It reaches a total
length of 13 cm. Color photographs appear in
Mohr and Poulson (1966), Zahl (1972), and
Conant (1975).
RANGE
The species occurs only in subterranean
waters of the Edwards Aquifer near San Marcos,
Hays County, Texas. It can be seen only in caves,
sinkholes, and fissures, including Ezell's Cave,
Primer's Well, and Rattlesnake Cave. Two addi-
tional sites, Johnson's Well and Wonder (Beaver)
Cave, have been recorded, but no Texas blind
salamanders have been seen in either location for
many years (Russell 1976). Longley (1978) has
placed nylon nets over Pipe Spring at San Marcos
Springs and the type locality artesian well, Aqua-
tic Station, Southwest Texas State University in
San Marcos, and collected Texas blind salaman-
ders as they were washed out of the aquifer.
Their total distribution is limited to about
104 km? mostly beneath the city of San Marcos
(Russell 1876). However, based on known distri-
bution (see range map), the figure could be less
than this, perhaps as little as 10 km^ (F. E. Potter
personal communication).
RANGE MAP
Current localities (observed within the past 5
years) are indicated by dots. Former localities
(not observed within the past 50 years) are shown
by triangles (F. E. Potter personal communica-
tion).
STATES/COUNTIES
Texas Hays.
HABITAT
The blind salamander inhabits water-filled
caverns of the San Marcos Pool of the Edwards
Aquifer. It is knovm only from incidental speci-
mens washed out of the aquifer or found near the
water surface in caves. Water quality is considered
very good with average temperatures in the vicini-
ty of 21° C (Longley 1978).
FOOD AND FORAGING BEHAVIOR
These salamanders feed on any living inverte-
brates they can handle, including a tiny snail
species, copepods, amphipods, and a shrimp. Cap-
tive specimens have been maintained for up to 2
years on epigeal forms of daphnia and other small
crustaceans, suggesting use of any such forms as
may happen to wash into the aquifer from the
surface.
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
Not known.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
The blind salamander is completely adapted
to the cave environment.
POPULATION NUMBERS AND TRENDS
Information is lacking on population size, but
it is believed to be stable because of the large
number of specimens and high percentage of ju-
veniles washed out of the aquifer by springs and
artesian wells. In the 2-year period 1975-1977,
Longley (1978) netted 133 specimens, 32 of
which washed from the type locality artesian well
and 101 of which washed from Pipe Spring at San
Marcos Springs, a previously unreported locality
for the species.
REPRODUCTION
Information on reproduction is scanty. One
gravid female contained 39 eggs. Tiny specimens
less than 2 cm long have been found throughout
the year (Longley 1978). Dunn (1926) reported
eggs laid March 15, 1886, and spermatheca
packed with spermatozoa in early fall of 1916.
Gravid females have been observed each month of
the year (F. E. Potter personal communication).
Brandon (1971) discusses tesdcular lobes ranging
from zero to four in the seven specimens he ex-
amined. There appears to be a correlation be-
tween size (age class), number of lobes, and num-
ber of times sperm has been produced.
lin
^ 116
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in
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o
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UJ
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o
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o
-4-)
J2
MANAGEMENT AND CONSERVATION
Following the original fear that the species
was being depleted, the Nature Conservancy pur-
chased Ezell's Cave in 1967. The bat colony that
had roosted there and contributed much of the
basic energy for the cave and local aquifer com-
munity had been almost eliminated when the cave
entrance was sealed some years before. Although
efforts to reestablish the bat colony have thus far
been unsuccessful, a few cave-dwellers, including
Typhlomolge rathbuni, are still infrequently ob-
served (F. E. Potter personal communication).
The continued protection afforded by offi-
cial Hsting should minimize the potential impact
of collectors on the few cave habitats accessible
to them. The major part of the blind salamander's
habitat is inaccessible, but adverse impact is pro-
jected as the aquifer level declines with increased
ground water usage (Longley 1978). Concurrent
vkdth this decline in water quantity is the increased
potential for urban pollution as more and more
urbanization takes place along the aquifer re-
charge zone. General management plans, de-
signed by action agencies to maximize recharge
and minimize introduced contaminants, mostly to
benefit human consumption, should also benefit
the diverse assemblage of species occupying the
aquifer, including Typhlomolge rathbuni. A study
is currently underway at Ezell's Cave to deter-
mine existing environmental conditions (water
quality) and present trends in faunal abundance
and diversity. The results should serve as a basis
on which to monitor conditions in the cave.
AUTHORITIES
Glenn Longley
Biology Department
Southwest Texas State University
San Marcos, TX 78666
Floyd E. Potter, Jr.
Texas Parks and Wildlife Department
4200 Smith School Road
Austin, TX 78744
Samuel S. Sweet
Department of Biological Sciences
University of California, Santa Barbara
Santa Barbara, CA 93106
PREPARER'S COMMENTS
Mitchell and Reddell (1965) placed the Texas
blind salamander in the genus Eurycea; however,
Brandon (1971) retained Typhlomolge. New data
on skull morphology support the continued
recognition of the genus Typhlomolge (Potter and
Sweet 1979).
LITERATURE CITED/SELECTED
REFERENCES
Brandon, R. A. 1971. North American troglobi-
tic salamanders: Some aspects of modification
in cave habitats with special reference to
Gyrinophilus palleucus. Bull. Natl. Speleol.
Soc. 33:1-2L
Conant, R. 1975. A field guide to reptiles and am-
phibians of eastern and central North America,
2nd ed. Houghton Mifflin Co., Boston.
429 pp.
Dunn, E. R. 1926. The salamanders of the family
Plethodontidae. Smith College Publications,
Northampton, Mass. 441 pp.
Longley, G. 1975. Environmental assessment,
Upper San Marcos River watershed. Environ-
mental Sciences of San Marcos. Soil Conserv.
Serv. Contract AG-48-SCS 01256. 367 pp.
. 1978. Status of the Texas blind salaman-
der. Endangered Species Report 2. U.S. Fish
Wildl. Serv., Albuquerque. 45 pp.
Mitchell, R. W., and J. R. Reddell. 1965. Eurycea
tridentifera , a new species of troglobitic sala-
mander from Texas and a reclassification of
Typhlomolge rathbuni. TexasJ.Sci. 17:12-27.
Mohr, C. E., and T. L. Poulson. 1966. The Hfe of
the cave. McGraw-Hill Book Co., New York.
232 pp.
Potter, F. E., Jr. 1963. Gross morphological varia-
tion in the genus Typhlomolge with descrip-
tion of a new species. Unpubl. M.S. Thesis.
Univ. of Texas, Austin. 66 pp.
Potter, F. E., Jr. and S. S. Sweet. 1979. Generic
boundaries in Texas cave salamanders and a
redescription of Typhlomolge robusta (Am-
phibia: Plethodontidae). Copeia. In press.
Russell, W. H. 1976. Distribution of troglobitic
salamanders in the San Marcos area. Hays
County, Texas. Unpubl. Rep. 7601, Texas
Assoc. Biol. Invest. Trogl. Eurycea, Austin.
35 pp.
Zahl, P. A. 1972. The shadowy world of salaman-
ders. Natl. Geog. Mag. 142(1):104-117.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.15
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
YUMA CLAPPER RAIL
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the Fndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.15
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
YUMA CLAPPER RAIL
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
YUMA CLAPPER RAIL
Rallus longirostris yumanensis Dickey (1923)
KINGDOM Animalia
CLASS Aves
ORDER Gruiformes
FAMILY Rallidae
OTHER COMMON NAMES None
DATE
Entered into SWIS to be determined
Update to be determined
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 Mar
1967). Migratory Bird Treaty Act of
3 July 1918 (40 Stat. 755; 16 U.S.C.
703-711) as amended 3 December
1969. Public Law 91-135.
State: California: Rare
Arizona: Protected
REASONS FOR CURRENT STATUS
Grinnell (1914) did not report any clapper
rails and noted that marshes were few and small
in size during his 3-month vertebrate survey along
the Lower Colorado River from 15 February to
15 May 1910, suggesting a considerable increase
in both habitat and rails as a result of damming
of river since then (Ohmart and Smith 1973).
Since clappers normally do not return to the
Colorado River habitat until about April 22-25, it
would have been easy for Grinnell to miss the
influx of birds.
Reclamation projects along the Colorado
River have both created and destroyed marsh
habitat. Dam construction may have generally
increased habitat by creating marshes. Therefore,
birds may now be at the northernmost point of
their historic range.
Channelization has eliminated large areas of
habitat near Yuma since 1963 (Tomlinson and
Todd 1973).
PRIORITY INDEX
8
DESCRIPTION
R. I. yumanensis is a chicken-size bird, gray-
brown above with cinnamon brown breast, flanks
barred gray and white, and a white patch under
the upturned tail. Bill, legs, and toes are long.
Similar to R. I. levipes but paler underparts,
duller and more olivaceous upper parts. More
slender tarsus and bill (Dickey 1923). Very simi-
lar to levipes of southern coastal California,
differing only in more slender proportions and
paler coloration (Van Rossem 1929). Similar to
rhizophorae (of Sonora) in paleness but, in gen-
eral, more brownish (less grayish) and has more
pointed wings because of difference in length of
primaries. Compared with levipes, it is paler on
breast and throat, has grayer flanks, duller wing
coverts and more extensive brown in the crown
(Banks and Tomlinson 1974). Ripley (1977),
although noting the above differences, is dubious
about the validity of yumanensis as a subspecies.
Measurements — Male: Wing 143.1-160.1 mm
(av. 149.8 mm); tail 60.3-69.0 (av. 64.2); exposed
culmen 55.4-61.8 (av. 59.2); tarsus 47.9-55.0
(av. 50.3); middle toe without claw 50.3-54.6
(av. 52.7); weight 222-307 g (av. 256g). Female:
wing 135.6-148.5 mm (av. 141.8mm); tail 57.8-
62.6 (av. 59.9); exposed culmen 51.9-58.2 (55.5);
tarsus 43.0-49.5 (45.4); middle toe without claw
46.5-51.1 (49.0); weight 192-268g (219g). Banks
and Tomlinson 1974).
Eggs are oval, glossy, pale pinkish buff and
cartridge buff, with sparse spots of varying brown
shades;41.8 by 28.8mm (Bent 1926).
RANGE
This rail breeds, at present, in marshes along
the Colorado River in California and Arizona,
from Needles to the Topock marsh south to the
Colorado River delta in Sonora; west to marshes
along the southeastern Salton Sea, California;
east, locally, to the Gila River near Tacna, Ari-
zona, and possibly the Salt River near Phoenix
(Tomlinson and Todd 1973, Ohmart and Smith
1973). It is thought to winter, at least in part, in
coastal and interior marshes and coastal mangrove
swamps in Mexico, including Estero Mescales,
8 km north of Teacapan, and Castillo, 11km east
of Mazatlan, Sinaloa; also at Laguna San Felipe,
Puebla (Banks and Tomlinson 1974).
The former more restricted range is documen-
ted by Dickey (1923), Bent (1926, Van Rossem
(1929), Moffitt (1932), Abbott (1940), and
Grinnell and Miller (1944).
Lack of authentic winter records in the north-
em breeding areas is noted by Phillips et al.
(1964), Todd (1971), and Tomlinson and Todd
(1973). Winter records of clapper rails (presum-
ably yumanesis but possibly wanderers of other
subspecies) at Salton Sea appearing on two 1976
Christmas bird counts published in American
Birds 31 (4):880, 1977 and confirmed by R. Guy
McCaskie (pers. comm.) ; and also at Topock Marsh
in January 1974 (Smith 1974), suggest that some
individuals do not migrate.
Breeding populations along the Colorado Riv-
er, the Colorado River delta and at Salton Sea have
been identified from specimens as yumanensis
(Banks and Tomlinson 1974). No specimens rep-
resentative of the small, isolated populations
along the Gila and Salt Rivers in Arizona, nor any
winter specimens from the Colorado River or
Salton Sea have been critically examined.
RANGE MAP:
The following map depicts breeding range
(from Tomlinson and Todd 1973) and winter re-
cords (from Banks and TomHnson, 1974).
STATES/COUNTIES:
California: Imperial, Riverside, San Bernardino.
Arizona: Maricopa, Mohave, Yuma.
HABITAT
The Yuma clapper rail requires freshwater or
NEV.
ARIZONA
SAN BERNARDINO CO.
Needles '
MOHAVE CO.
I Topock Marsh
Topock
Gorge
LakeHavasu "*>s^ Bill Williams
Delta
Parker Dam
CC
o
<
Headgate Rock Dam_^
-jC
Parker
Lost Lake »\
Wl)(/ar
Lake IVIodvalya
RIVERSIDE CO.
BIythe
Palo Verde Diversion Dan
U.S. Highway 60-70
Palo Verde Lagoon
Davis L.(
Three Fingers L.
Draper L.Jl]
Taylor Ferry
/)£> Cibola L.
YUMA CO.
IMPERIAL CO.
/Xii Aniencan
Morelos
Martinez I
Laguna Dam i
» Dobe L.
Imperial Dam
r.^^'y
oi
n^
'MIttry L.
Yuma
Tacna
BAJA CALIFORNIA
Colorado River Delta
• Riito
MARICOPA CO.
RWef
WINTER
RECORDS
SONORA
Known range of the Yuma clapper rail.
3
brackish stream sides and marshes, associated
with heavy riparian and swamp vegetation (Grin-
nel and Miller 1944), such as alkaline cattail mar-
shes (Phillips et al. 1964). In general, habitat con-
sists of shallow-water marshes containing dense
stands of cattail (Typha latifolia) and big bulrush
or tule (Scirpus acutus), in both brackish and
freshwater situations. Shallow water with mud
flats available for feeding are selected over areas
where water is deep and steep banks prevalent.
Stands of cattail and tules dissected by narrow
channels of water 1.6 to 7 m wide had densest
populations, according to Tomlinson and Todd
(1973). Preferrred breeding habitat is light cattail
or tule stands with downed vegetation, with adja-
cent dry land a must (Ohmart and Smith 1973).
Water of breeding habitat on the Colorado River
Delta in Mexico is salty and growths of cattails
and tules appear to be limited to small fresh or
brackish sloughs. The vegetation is characterized
by an overstory of saltcedar {Tamarix sp.) and an
understory of iodine bush {Allenrolfia occident-
alis), all quite different from the habitat above
the delta (Tomlinson and Todd 1973).
The rail seems to require wet mudflat or sand-
bar sites, covered rather densely with mostly ma-
ture vegetation exceeding 0.5m. Optimum condi-
tions are created by open ponds or channels as
opposed to unbroken stands of vegetation. Salt-
cedar {Tamarix pentandra) stands are rarely
utilized (Todd 1971).
The following habitat characteristics appeared
in locations where clapper rail densities were
higher than average:
1. Water, flowing through many small
channels from 0.5 to 3 m wide, either covered
with vegetation or appearing as small bodies
of open water 0.02 to 0.2 hectares in size.
2. Extensive areas of water of depth less
than 0.3 m little or no daily fluctuation in
water level.
3. High ground in strips or, less importantly,
as small islands.
4. Emergent vegetation, cattail or bulrush,
with little or no Phragmites sp., which are too
high and have few down stems (Gould 1975).
At Topock Marsh, the highest densities were
found in light cattail stands — lowest, in heavy
stands. Dense cattail had 0.9 rails per 10 ha; light
light cattail 1.9 per 10 ha; dense buUrush 1.7 per
10 ha; light bullrush 1.8 per 10 ha. The majority
of rails were in the ecotone between emergent
vegetation and higher ground, either at the shore-
line or on hummocks in the marsh (Smith 1974).
Winter habitat probably includes mangrove
swamps on the Pacific coast of Sinaloa and fresh-
water marshes of Puebla (Banks and Tomlinson
1974).
FOOD AND FORAGING BEHAVIOR
The major food is invertebrates, with little
vegetable matter. Crayfish [Procambarus and
Oropectes) were the dominant food in 9 of 10
stomachs from Topock Marsh south to Imperial
Reservoir. Of two stomachs from the confluence
of the Gila and Colorado Rivers, one contained
primarily (98%) freshwater clams {Corbicula sp.),
and the other, 97% isopods. Colorado River delta
specimens contained a greater variety of food, but
the majorcomponents were water beetles and fish.
Of 16 stomachs, 9 had crayfish, 1 1 had insect frag-
ments, 4 had water beetles, 4 had fish, and 3
contained clams. Other insects in small amounts
were weevils, damselfly nymphs, dragonfly
nymphs, grasshoppers, and insect eggs. Spiders,
leeches, prawns, and a small mammal bone were
also found. Plant material consisted of twigs (10%
in one stomach), 2 legume seeds (1 stomach) and
18 unidentified black seeds (3 stomachs). Like
other subspecies of clapper rail, yumanensis seems
to be a selective opportunist whose variety of
food is limited by its availability in the particular
habitat (Ohmart and Tomlinson 1977). In To-
pock Marsh, crayfish are an important part of the
diet. Crayfish are found in all areas where rails
occurred, with the greatest number of crayfish in
water 8 to 15 cm deep with abundent stems and
leaves lying in the water. Floating and recumbent
vegetation is important in foraging areas, as they
provide habitat for crayfish and a platform to
walk on (Smith 1974).
SHELTER REQUIREMENTS
None other than mentioned in other sections.
NESTING OR BEDDING
In three nests, material consisted of black
sticks with dry leaves on them, two nests were
made of fine stems with dry blossoms attached.
Nests were located on hummocks and in the
crotches of small shrubs just above water in dense
cattail and tamarisk associations (Abbott 1940).
RITUAL REQUIREMENTS
The rail appears to defend its territory (Tom-
linson in Ohmart and Smith 1973). The 'clatter
call' is given in unison by male and female and
with adjacent birds (Tomlinson in Ohmart and
Smith 1973). Territory size averages 1.44 ha
(Smith 1974).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None other than specified in other sections.
POPULATIONS NUMBERS AND TRENDS
A multiagency Yuma clapper rail census re-
corded 889 along the Colorado River in 1973 and
787 in 1974. In the Imperial Valley, 134 were
recorded in 1974 (no count in 1973). Total count
for the Colorado River and Imperial Valley com-
bined in 1974 was 921 rails (Gould 1975). Esti-
mated local populations in 1973 were: Topock
Marsh 109-136, Topock Gorge-52-65, Bill Wil-
liams River delta-21-35, Colorado River delta-
145 (Cornelius 1972, Ohmart and Smith 1973).
Total population was estimated at over 1,000
(Tomlinson in Ohmart and Smith 1973).
Factors regulating populations include preda-
tion by raccoon and coyote (Abbot 1940), bob-
cat, feral house cat, dog, and Cooper's hawk
(Todd in Ohmart and Smith 1973); and habitat
destruction (Tomlinson and Todd 1973, Corne-
lius 1972).
REPRODUCTION
A nest with 7 slightly incubated eggs was
found near the Salton Sea on 12 May 1940. Obser-
vations along the lower Colorado River in summer
(8 May to 16 September) include an adult with
3 young 2 weeks old on 17 July (Phillips et al.
1964); and a nest with one egg, another with two
eggs, and two empty found on 5 May. A nest with
six fresh eggs, one with seven fresh eggs, and one
with seven slightly incubated eggs were found on
12 May. Abbot (1940) reports a clutch of six
slightly incubated eggs on 26 May, and five
clutches ranging from six to seven eggs (average
6.5).
The incubation period is unknown, but is
probably similar to other clapper rails, 21 to 23
days (Ohmart and Smith 1973).
Hatching success is suggested by three quar-
ter- to half-grown downy chicks found on 23
June (Tomlinson in Ohmart and Smith 1973) and
three 2-week-old young (Phillips et al. 1964).
A breeding period in May and June was indi-
cated by responses to taped calls (Tomlinson and
Todd 1973).
MANAGEMENT AND CONSERVATION
An annual or periodic index to abundance of
rails will be important in view of constant change
in the rivers through reclamation projects. Taped
call notes could be used to obtain such an index,
but would be costly to maintain. Since cattails
and tules are so important as habitat, estimates of
rail population size can be determined by inspec-
tion of aerial photographs taken periodically. Prior
research could determine average density of rails
for specific habitat types and sizes. Spot checks
on the ground would help to determine accuracy
(Tomlinson and Todd 1973).
State, national, and international uses of
Colorado River water are all involved in manage-
ment of Yuma clapper rails. Occasionally, such
management, even that for fish and wildlife, if it
involves dredging out cattail growth, is detrimental
to rail survival. The main requirement in manage-
ment for Yuma clapper rails is that extensive
growths of cattails and tules must be preserved
throughout its range.
AUTHORITIES
Richard L. Todd
Arizona Department of Game and Fish
2222 West Greenway Road
Phoenix, Arizona 85023
Roy E. Tomlinson
U.S. Fish and Wildlife Service
P.O. Box 1306
Albuquerque, New Mexico 87103
Philip M. Smith
1613 W.Peoria Ave. No. 2
Phoenix, Arizona 85029
Robert D. Ohmart
Department of Zoology
Arizona State University
Tempe, Arizona 85281
Gale W. Monson
8831 N. Riviera Drive
Tucson, Arizona 85704
Gordon I. Gould, Jr.
1080 E.Nevada
Blythe, California 92225
PREPARER'S COMMENTS
Research attention focused on the Yuma
clapper rail because of its endangered status has
cleared up two important matters. Despite doubts
expressed by Ripley (1977), Banks and Tomlin-
son (1974) have shown, on the basis of critical
study of adequate specimens, that it is a taxa-
nomically valid subspecies, examples of which can
be recognized as migrants when away from their
breeding areas.
Also, it has been found that the populations
of this rail will respond to changing distribution
of its preferred type of habitat, which has now
been described in detail, by extending its range
(Tomlinson and Todd 1973, Ohmart and Smith
1973, Smith 1974, Gould 1974). This informa-
tion should make possible the continued exis-
tence of this subspecies by means of specific habi-
tat management for it, in addition to the several
other competing land uses in the limited sites
available along the lower Colorado River and
adjoining areas.— John W. Aldrich.
LITERATURE CITED/SELECTED
REFERENCES
Abott, C. G. 1940. Notes from the Salton Sea,
California. Condor 42:264.
Banks, R. C, and R. E. Tomlinson. 1974. Taxo-
nomic status of certain Clapper Rails of south-
western United States and northwestern Mexi-
co. Wilson Bull. 86:325-335.
Bent, A. C. 1926. Life histories of North Ameri-
can marsh birds. U.S. Nat. Mus. Bull. 135:275.
California Department of Fish and Game. 1978.
At the crossroads: a report on California's
endangered and rare fish and wildlife. Bian-
nual Rep. 103 pp.
Cornelius, S. S. 1972. Yuma Clapper Rail census,
Havasu National Wildlife Refuge. Unpubl.
Dickey, D. R. 1923. Description of a new Clapper
Rail from the Colorado River valley. Auk 40:
90-94.
Gould, G. I., Jr. 1975. Yuma Clapper Rail study-
censuses and habitat distribution 1973-74.
California Dep. Fish Game Admin. Rep. 75-2,
April 1975.
Grinnell, J. 1914. An account of the mammals
and birds of the lower Colorado Valley. Univ.
Cahf. Publ. Zool. 12:51-294.
Grinnell, J., and A. H. Miller. 1944. The distribu-
tion of the birds of California. Pac. Coast Avi-
fauna 27:128.
Moffitt, J. 1932. Clapper Rails occur on marshes
of Salton Sea, California. Condor 34: 137.
Oberholser, H. C. 1937. A revision of the Clapper
Rails [Rallus longirostris Boddaert). Proc.
U.S.Nat. Mus. 84:313-354.
Ohmart, R. D., and R. E. Tomlinson. 1974. Food
of western Clapper Rails. Wilson Bull. 89:332-
336.
Ohmart, R. D., and R. W. Smith. 1973. North
American Clapper Rails [Rallus longirostris),
literature survey with special consideration
being given to the past and current status of
yumanensis. Bur. Reclam. Rep.
PhilUps, A., J. Marshall, and G. Monson. 1964.
The birds of Arizona. Univ. Arizona Press,
Tucson. 31pp.
Ridgway, R., and H. Friedmann. 1941. The birds
of North and Middle America, Part 9. U.S.
Nat. Mus. Bull. 50.
Ripley, S. D. 1977. Rails of the world. David R.
Godine, Boston.
Smith, P. M. 1974. Yun.a Clapper Rail study,
Mohave County, Arizona, 1973. California
Dep. Fish Game, Spec. Wildl. Investig. Prog.
Rep. Job 11-5.9. June 1974.
Todd, R. C. 1971. Report on the study of the
Yuma Clapper Rail along the Colorado River.
Prepared for Colorado River WildUfe Council
Meeting,, April 5-6, 1971, Las Vegas, Nevada.
Tomhnson, R. D., and R. L. Todd. 1973. Distri-
bution of two western Clapper Rail races as
determined by responses to taped calls. Con-
dor 75:177-183.
Van Rossem, A.J. 1929. The status of some Paci-
fic Coast Clapper Rails. Condor 31:213-215.
Biological Services Program
FWS/OBS-80/01.16
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
SANTA BARBARA SONG SPARROW
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Knginccrs in coordina-
tion with the Offices of F.ndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should bo directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.16
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
SANTA BARBARA SONG SPARROW
A Cooperative Effort
by the
National Fish and Wildhfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coeistal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
PHOTO OF CUOSELY REUATED SUBSPECIES
CREDIT: HERBERT CLARKE
SANTA BARBARA SONG SPARROW
Melospiza melodia graminea Townsend (1890)
KINGDOM Animalia
CLASS Aves
ORDER Passeriformes
FAMILY Fringillidae
OTHER COMMON
NAMES Bell Finch, Coast Song
Sparrow, California Song Sparrow.
DATE
Entered into SWIS To be determined
Updates To be determined
LEGAL STATUS
Federal: Endangered. (42 FR 36427), 14 July
1977.
States: Protected: California.
REASONS FOR CURRENT STATUS
There is general agreement among ornitho-
logists who have searched for song sparrows on
Santa Barbara Island during the breeding season
that the subspecies M. m. graminea is extinct (G.
L. Hunt, Jr. pers. comm. to J. W. Aldrich, 18
Aug. 1972, and Warren King, 26 March 1974;
Small and Henderson 1974). The main reason for
its decline and extinction was elimination of
dense vegetation over the entire island by feral
domestic rabbits whose population exploded
from 1953-1959; and, finally, extensive fire in
1959 that destroyed the remaining vegetation and
litter down to the mineral soil. Feral cats, which
were numerous in earlier times, along with bam
owls {Tyto alba) and American kestrels {Falco
sparverius) may have contributed to the decline,
particularly after the 1959 fire destroyed the
concealing vegetation (Small and Henderson
1974).
PRIORITY INDEX
None assigned.
DESCRIPTION
M. m. graminea is small, brownish gray above,
white streaked with black below, with a black
spot on chest. It is distinguished from related
races of song sparrow by shorter wings and tail,
and, except for tnicronyx of San Miguel Island,
by more grayish (less brownish) coloration. It
differs from M. m. clementae of the Other Chan-
nel Islands in smaller bill, tarsus, and feet; from
M. m. cooperi, of the adjoining mainland; in
smaller bill; from coronatorum of Los Coronados
Islands in larger tarsi and feet. Nineteen adults
measured: length, 5.50 - 6.12 in (140 - 155 mm)
av. 5.80 in (147 mm); wing, 2.25 - 2.50 in (57.2
- 63.5 mm) av. 2.35 in (59.7 mm); bill, 0.40 -
0.46 in (10.2 - 11.7 mm) av. 0.43 in (10.9 mm)
(Townsend 1896, Grinnell 1897, 1928, Van
Rossen 1924).
RANGE
Formerly a permanent resident on Santa Bar-
bara Island off southwestern California, it was
confined to that island (Willett 1933; Grinnell
and Miller 1944, American Ornithologist Union
1957).
RANGE MAP
See range map on following page.
STATES/COUNTIES
C^difomia: Los Angeles.
HABITAT
Santa Barbara Island, with an area of only 2.6
km^ , is 61 km from the mainland and 37 km
from the nearest other island (Santa Catalina). No
permanent fresh water is present on the island. A
low ridge extends along the western edge with
high points 171 and 193 m above sea level. The
central portion is a graded slope, almost level in
some areas, then falling steeply to the ocean on
the eastern edge, which is cut by a number of
canyons. It has an equable climate, and practically
never frosts; rainfall is only 12 in (30.5 cm), but
wet fogs are frequent. Vegetation was formerly
long, coarse grass growing thick and tangled every-
where, making walking difficult (Townsend 1890).
Sparse brush covered slopes and ravines (Grirmell
1897). The song sparrows were found where the
brush afforded protection (Wright and Snyder
1913). Song sparrows used bushes for nesting,
and fog supplied the moisture essential to birds of
this type (Grinnell and Miller 1944). In more
recent times, thickets of giant tree-sunflower
{Coreopsis gigantea) were abundant (Philbrick
1972; Sumner 1958). Gross changes in vegetation
have taken place in the 20th century as a result of
agriculture, the effect of overgrazing by rabbits,
and more recently, a severe fire ( 1959) that burned
over almost the entire island and eliminated most
of the remaining vegetation (Small and Henderson
1974).
FOOD AND FORAGING BEHAVIOR
No information is available, but the diet pre-
sumably consisted of insects and small seeds, the
same as mainland populations.
SHELTER REQUIREMENTS
Dense growths of grass (Townsend 1890),
sparse brush cover on slopes and ravines used for
nesting (GrinneU 1897), and thickets of giant
tree-sunflower (Small and Henderson 1974) were
all important cover vegetation for song sparrows.
NESTING AND BEDDING
Five nests were all practically the same size
and composition. A typical one was supported by
obliquely growing twigs of a bush and lined with
yellow grasses, in marked contrast to the larger
brown grass and weed stems of which the nest
structure was built (Grinnell 1897).
RITUAL REQUIREMENTS
Song, notes and actions are the same as those
of mainland races of song sparrow, a strongly
territorial species.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None is known other than dense vegetative
cover for nesting and concealment.
POPULATION NUMBERS AND TRENDS
Once extremely abundant - in fact, the most
abundant bird - on the small island of Santa Bar-
bara (Townsend 1890). They were numerous
SAN FRANCISCXD
CALIFORNIA
t>t^'
LOS ANGELES
t
SANTA BARBARA ISLAND
Total range of the Santa Barbara song sparrow.
everywhere, especially on the brush-covered fields
on the southern part of the mesa (Grinnell 1897).
They were abundant residents on Santa Barbara on
first of May 1908, and at times were "fairly
swarming" in short scrub (Howell 1917), abun-
dant residents on the island in 1911 (Willett
1933); permanent residents, abundant, but no
year given (Grinnell and Miller 1944). By 1967,
the subspecies was extinct according to Kenneth
Baker, National Park Service (pers. comm. to
Chnton Lostetter, FWS 1972). Monitoring of all
land birds by means of periodic searches of all
parts of Santa Barbara Island from 15 May to 23
July 1972 failed to produce sight or sound of a
song sparrow (George L. Hunt, Jr. pers. comm.
1972). No trace of song sparrows was found on
the isleind by diligent search by Small and Hender-
son from 13 to 17 May 1974, despite open condi-
tion of land that made it easy to observe any bird
present. A song sparrow sighting on the island by
Robert DeLong on 19 August 1967 may be the
last record of the subspecies (Small and Hender-
son 1974). Records of single song sparrows in
1972 and 1973 by George Hunt, Jr. were thought
by him to be migrants of other races from the
mainland, since they were associated with waves
of migrant landbirds (Small and Henderson 1974).
REPRODUCTION
Nest containing 2 eggs, advanced in incuba-
tion, were found 16 June 1911. At that date most
of the young were already full grown (Willett
1912 and 1933). In mid-May 1897, fuU-grovm
juveniles were numerous, more so than adults,
which were all apparently engaged in nest build-
ing or raising second broods. From 3 to 5 eggs
were laid per set. Five sets averaging 3.8 eggs each
were secured on May 14 and 15. Eggs averaged
0.61 X 0.78 in (15.5 x 19.8 mm), with extremes
of 0.70 to 0.82 in (17.8 - 20.8 mm) in length and
0.57 to 0.64 in (14.5 - 16.3 mm) in diameter
(GrinneU 1897).
MANAGEMENT AND CONSERVATION
It is now too late to manage for this species,
but the example may be useful in planning habi-
tat restoration and management to prevent ex-
tinction of other wildlife species on Santa Bar-
bara Island. The Santa Barbara song sparrow is
protected by the Migratory Bird Treaty Act and
by the National Monument status of the island,
under the National Park Service.
AUTHORITIES
George Hunt, Jr.
U. of Calif.
Irvine, Calif.
Kenneth Baker
Nat. Park Service
John Small
Point Reyes Bird Observatory
R. P. Henderson
Point Reyes Bird Observatory
PREPARER'S COMMENTS
The evidence seems conclusive that the subspe-
cies of song sparrow that bred on Santa Barbara Is-
land (M. m. graminea) is now extinct and that
song sparrows observed on the island from time
to time are probably migrants from other breed-
ing areas.
LITERATURE CITED
American Ornithologists Union. 1957. Check-list
of North American Birds, 5th Edition. Balti-
more, Md., Amer. Omith. Union.
Grinnell, J. 1897. Report on the birds recorded
during a visit to the islands of Santa Barbara,
San Nicholas and San Clemente in spring of
1897. Pasadena Academy of Sciences Pub.
1:1-21(6).
Grinnell, J. 1928. The Song Sparrow of San
Miguel Island, California. Proc. Biol. Soc.
Wash. 41:37-38.
Grinnell, J., and A. H. Miller. 1944. The distribu-
tion of the birds of California. Pac. Coast
Avifauna 27:608 pp. (554).
Howell, A. B. 1917. Birds of the islands off the
coast of southern California. Pac. Coast Avi-
fauna 12:1-127(80).
Van Rossem, A. J. 1924. A survey of the Song
Sparrows of Santa Barbara Islands. Condor
26:217-220.
Philbrick, P. N. 1972. The plants of Santa Barbara
Island, California. Madrono 21:329-393.
Small, J., and R. P. Henderson. 1974. Part II.
Santa Barbara Island, hi "The status of the
Song Sparrow and Bewick's Wren on San
Clemente Island and Santa Barbara Island,
California" by R. M. Stewart, John Smail,
William Clow and R. P. Henderson. Report
to Endangered Species Office U. S. Fish and
Wildlife Service by Point Reyes Bird Observa-
tory, Bolinas, Calif. 94924. Oct 1974.
Sumner, L. 1958. The rabbits of Santa Barbara
Island, a progress report. In Smail and Hen-
derson 1974. (hsted above).
Townsend, C. H. 1890. Scientific results of ex-
ploration by the U. S. Fish Commission Al-
batross. No. XIV - Birds from the coasts of
western North America and adjacent islands,
collected in 1888-89 with descriptions of new
species. Proc. U. S. Nat. Mus. 13:131-142
(139).
Willett, G. 1912. Birds of the Pacific slope of
southern California. Pac. Coast Avifauna 7:1-
122(84).
Willett, G. 1933. A revised list of the birds of
southwestern California. Pac. Coast Avifavma
21:1-204(185).
Wright, H., and G.K.Snyder. 1913. Birds observed
in the summer of 1912 among the Santa Bar-
bara Islands. Condor 15:86-92(91).
Biological Services Program
FWS/OBS-80/01.17
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
ESKIMO CURLEW
^^^^^£^^^
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc ncH necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.17
March 1980
SELECTED VERTEBRA IE ENDANGERED SPECIES
OF THE SEACOAST OF IHE UNITED STATES-
ESKIMO CURLEW
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
lOIO Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
M-
i»^-
•^-iV:.^
•.-^
►>. ."f ^.r VT' • -^i .. . c ■* *
ESKIMO CURLEW
(Numenius borealis Forster)
KINGDOM Animalia
CLASS Aves
ORDER. Charadriiformes
FAMILY Charadriidae
OTHER COMMON
NAMES Doughbird, fute, prairie
pigeon, pipi-pi-uk or tura-tura
courlis du nord or Corbigeau
des Esquimaux, chittering
curlew, zarapito, Chorlo
campino and Chorlo grande.
DATE
Entered into SWIS To be determined
Updates To be determined
LEGAL STATUS
Federal: Endangered: (32 FR 4001, 11 March
1967; 35 FR 12122, 29 July 1970).
States: Endangered: South Carolina, Texas,
Alaska.
CREDIT: PES, WASHINGTON, D.C.
REASONS FOR CURRENT STATUS
The evidence is overwhelming that unrestricted
hunting for the market, particularly during north-
ward spring migrations through the midwestem \
prairies of the United States, and to a lesser
degree in the fall migration in southeastern Labra-
dor and (after severe storms) on the coast of Mas-
sachusetts, drastically and rapidly reduced this cur-
lew's population between 1870 and 1890 (Mackay
1892, Forbush 1912, p. 427, Swenk 1915, Bent
1929, p. 126, Greenway 1958, Vincent 1966, Fish
and Wildlife Service 1973). Other factors that
may have contributed to its rapid decline are
severe storms during long overocean migrations
(Townsend and Allen 1907, Forbush 1912);habi-
tat altered by cultivation and grazing on winter-
ing grounds and the spring migration route
(Cooke 1910, Dement'ev and Gladkov 1951); and
a succession of unsuccessful breeding seasons
caused by unfavorable weather (Banks 1977). A
characteristic of the Eskimo curlew that may have
contributed to its rapid decline was its tame nature
and extreme gregariousness, making it easy to
shoot (Swenk 1916, Bent 1929 ,p. 127,Coues 1861
MacKay 1892). Its continued failure to recover
after hunting in the United States was banned by
the Migratory Bird Treaty Act in 1916 must be
attributable to some unique characteristic that
makes it more vulnerable to environmental condi-
tions than other shorebirds with similar migration,
breeding, and wintering ranges. That characteris-
tic may be a greater concentration of all members
of the population at all times, making it more
vulnerable to short but critical periods of unfavor-
able weather and habitat conditions (Banks 1977).
The conversion of native grasslands to cultivated
fields in both the main wintering area in southern
South America and the principle migration route
through the tall grass prairie of the United States,
which coincided with the population decline
(Cooke 1910, Dement'ev and Gladkov 1969) and
has continued to the present is one likely reason
for its failure to recover.
It may be that only the natural grasslands of
the southern South American pampas and the tall
grass prairies of the United States could produce
enough easily available food, in the form of grass-
hopper egg pods, to supply the energy for both
the curlew's exceptionally long migration flights
and the initiation of breeding in the spring. The
Arctic tundra presumably provides enough crow-
berries and blueberries to support the fall migra-
tion, but the South American and North Ameri-
can grasslands, whose more productive areas are
now largely cultivated, may not provide enough
suitable insect life in winter and early spriiig to
enable the curlews to travel their long traditional
migration routes.
PRIORITY INDEX
55
DESCRIPTION
Eskimo curlews are medium-sized shorebirds
(about 30 cm long), smaller than whimbrels with
shorter (about 5 cm), more slender, slightly down-
curved bills; uniformly dark (rather than barred)
primaries; greenish (rather than gray) legs; more
blackish above with unstriped dark crowns. They
may be distinguished from very similar little cur-
lews [Numenius minutus) which breed in north-
east Asia and migrate through western Asia to
Austraha, by their generally darker and more
buffy coloration with v-shaped black marks, in-
stead of streaks, below and darker cinnamon buff
coloration under wings (Forrand 1977). The two
forms are considered races of the same species by
Dement'ev and Gladkov (1951) but as two distinct
species by the American Ornithologists' Union
(1957).
RANGE
N. borealis formerly nested in the Arctic
tundra of northwestern Mackenzie between the
Mackenzie and Coppermine Rivers (MacFarlane
1891, Swainson and Richardson 1881). There
have been several probable sightings by Canadian
Wildlife Service personnel east of the Mackenzie
River delta in a general area where nesting is
known to have occurred formerly. They probably
nested in Alaskan tundra west to the Bering Sea
(Nelson 1887, Murdock 1885); they wintered in
grasslands of southern South America from
southern Brazil and Uruguay, with a few probably
north of Buenos Aires, Argentina (Cooke 1910),
south to middle-eastern Argentina, chiefly north
of the Chubut River; casually to Chile and Tierra
del Fuego (Greenway 1958, Barrows 1884, Bent
1929, Cooke 1910, Swenk 1926, Sclater and
Hudson 1889,Wetmore 1926).
Fall migration (adults preceeding young), be-
ginning in July, was southeasterly from the breed-
ing grounds to a feeding and staging area on the
coast of southern Labrador (Audubon 1835,
Townsend 1907,Coues 1861, Austin 1932, Todd
1963); thence via Newfoundland and Nova Scotia
(Tufts 1961, Peters and Burleigh 1951) over the
Atlantic Ocean directly to eastern South America,
and ending on the wintering grounds in early Sep-
tember (Sclater and Hudson 1889 , Barrows 1884).
Severe storms occassionally forced the birds to
land on the north Atlantic coast of the United
States (Bent 1929,Forbush 19 12, Sage and Bishop
1913, Palmer 1949, Griscom and Snyder 1955),
Bermuda, and the eastern islands of the West
Indies (Bond 1956).
There was a much smaller flight down the
west side of Hudson Bay with a few individuals
reaching points on the Great Lakes and even Cin-
cinnati, Ohio, and Cooke Co. Texas (Cooke 1910,
Hagar and Anderson 1977). By what route those
birds reached the wintering grounds (if they did)
is unknown.
Spring migration began in late February (Bar-
rows 1884), heading northwest from the winter-
ing area, probably across the Andes in Chile, the
Pacific Ocean, northern Middle America (Guate-
mala - Salvin 1861), and the Gulf of Mexico, to
the coasts of Texas and Louisiana, arriving there
in early March (Greenway 1958); thence gradually
northward, primarily through the prairies of mid-
dle United States, to eastern South Dakota; thence
rather quickly to the breeding grounds, arriving
before the end of May (Bent 1929). There was
some accidental in-migration in Greenland, Ice-
land, Britain, the Falkland Islands, and north-
eastern Siberia (American Ornithologists' Union
1957).
RANGE MAP
A map of breeding and wintering areas and
migration routes is shown on the following page.
STATES/COUNTIES
Alaska:
Cape Lisboume, Kotzebue
Sound, Nulato, Point Barrow, St.
Michael, St. Paul Id.
Arkansas:
Washington.
Colorado:
Denver.
Connecticut:
Middlesex, New Haven, Toland.
Illinois:
Cooke.
Indiana:
Knox, White.
Iowa:
Des Moines, Jackson, Johnson,
Polk, Poweshiek.
Kansas:
Douglas, Ellis, Lyon, Russell,
Sedgwick, Woodson.
Louisiana:
Acadia, Jefferson Davis, Orleans,
Plaquemines.
Maine :
Cumberland, Hancock, Knox,
Lincoln, Penobscot.
Massachusetts:
Barnstable, Dukes, Essex, Nan-
tucket, Suffolk.
Michigan :
Kalamazoo, St. Clair.
Missouri:
Jasper, St. Louis, Vernon.
Nebraska:
Adams, Buffalo, Douglas, Fill-
more, Hall, Hamilton, Lincoln,
Madison, York.
New York:
Monroe, Nassau, Niagara, Suffolk,
Queens.
Ohio:
Hamilton.
Oklahoma:
Osage.
Pennsylvania: Erie.
South Dakota: Brown, Clay, Douglas, Hanlin,
Pennington, Yankton.
Texas: Aransas, Bexar, Cameron, Cal-
houn, Cooke, Galveston, Kendall,
Lampassas, Nueces, Pecos, San
Patricio, Victoria, Washington,
Wise, Young.
Wisconsin: Dodge.
HABITAT
The Eskimo curlew nested on treeless Arctic
tundra, fed in open natural grassland and tundra,
burned prairies, meadows, pastures, plowed lands,
and intertidal zones during migration and on win-
tering grounds. Most of the time was spent in the
North American tundra and tall-grass prairie, and
the South American pampas (MacFarlane 1881;
Coues 1861, 1874; Cooke 1910; Swenk 1916;
Bent 1929; Greenway 1958).
FOOD AND FORAGING BEHAVIOR
The chief food in late summer on Arctic tun-
dra, including the migration staging area in Labra-
dor, was crowberry (Empetrum nigrum and blue-
berry Vaccinium sp.). (Audubon 1835; Coues
1861, 1874; Townsend 1907; Greenway 1958.)
A small species of snail abounding on rocks in
intertidal areas in southern Labrador was also
eaten extensively (Coues 1861). Ants were men-
tioned as food on tundra breeding areas (Swainson
and Richardson 1881, Bent 1929, Coues 1861).
Grasshoppers [Melanoplus sp.) and their egg cap-
sules or pods, obtained by probing in unplowed
prairie land, were important food on the spring
migration (Swenk 1916). On plowed land, they
fed on white grubs and cutworms (Swenk 1916).
Adult grasshoppers would not be available on the
prairies as early in spring as curlews were moving
through successive temperature zones on way
north (U.S. Entomological Commission 1877), so
only egg pods and emerging young grasshoppers
were present at that time.
SHELTER REQUIREMENTS
None known.
NESTING OR BEDDING
Nests are shallow depressions in the surface of
KNOWN BREEDING RANGE
Dots Indicate Published Records
D
cC^^'
<s>
Known range of the Eskimo curlew
KNOWN
WINTER RANGE
4
open tundra, lined sparsely with decayed leaves
and dried grasses (MacFarlane 1891).
There is no information on any special roost-
ing behavior or requirements. Roosting is evidently
in the open in the same habitat as nesting and
winter and migration feeding.
RITUAL REQUIREMENTS
None known. Presumably, there is an aerial
territory flight song like other shorebirds have.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Open Arctic tundra for nesting and summer
feeding. Extensive natural upland grassland for
winter and migration feeding.
POPULATION NUMBERS AND TRENDS
No exact population counts or even reliable
estimates have ever been made. Comments of
early observers were that the curlews were for-
merly present in tremendous numbers on the
Labrador and prairie migration stopovers. They
arrived in Texas in "immense flocks" in spring
from 1856-1875 (Bent 1929).
In Kansas, they were abundant as late as 1878,
but were much reduced in 1879 and decreased
rapidly after that (Bent 1929). From 1866 to
1888, they reached Omaha, Neb., in late April,
remaining in force for a week to 10 days. Enorm-
ous flocks consisting of thousands of individuals
formed dense masses extending a quarter to a half
mile (0.4-0.8 km) in length and a hundred or
more yards (91 m) in width which, when alighting,
would cover 40 to 50 acres (16 to 24 hectares) of
ground. At that time, they were slaughtered liter-
ally by the cart-load. Their numbers in the prairies
began diminishing rapidly in the early 1880's
(Swenk 1916). On the Labrador migration staging
area in 1833, Audubon (1835) described great
flocks that reminded him of passenger pigeon
abundance. A "cloud" of curlews in fall migration
was seen on the Magdalen Islands in 1890— perhaps
the last big flocks seen in the east (Forbush 1912).
During the last 50 years, very few have been seen
during migration at any one time. The most recent
records are Galveston Island, Texas, 22 March
1959 (1), 3 April 1960 (1), 31 March 1961 (1),
24 March 1962 (1), and 31 March 1962 (2)
(Emanuel 1962); Barbados, West Indies, fall migra-
tion 1963 (1 specimen); North Point, West side of
James Bay, Ontario, 15 August 1976 (2).
REPRODUCTION
Nests are very difficult to find. Incubating
birds flush long before observers approach.
Eggs, usually 4 to a clutch, resemble the grass in
color, being dark brownish green to blue blotched
with brown, more heavily on the larger end
(MacFarlane 1891). Eggs were present in nests at
Fort Anderson, Mackenzie from May 27 through
June 13 (MacFarlane 1891). Eggs were found at
Point Lake, Mackenzie, on 13 June 1822 (Swain-
son and Rich2u:dson 1881). Time of hatching and
length of time young duce dependent on adults is
unknown, but by the end of July the breeding
season is over for the most part and the adults
head south, soon to be followed by the young.
MANAGEMENT AND CONSERVATION
With the present state of our knowledge of
the distribution of the remaining Eskimo curlew
population, particularly during the breeding and
wintering seasons, nothing can be done to manage
it except to rigidly protect the occasional indivi-
duals that are discovered; to keep a lookout for
them in areas where they formerly concentrated,
such as the southern tip of the Labrador Peninsula,
the coast of New England and Long Island, the
tail-grass prairie belt from the coast of Texas and
Louisiana north to South Dakota, and the Argen-
tina grasslands from Buenos Aires south to the
Chubut River. Also, special effort should be made
to locate the present breeding grounds of the
small remaining population, with special attention
given to the former breeding area along the Arctic
coast between the Anderson and Coppermine
Rivers and south to Great Bear Lake, Mackenzie.
It might be beneficial to increase the area of un-
plowed grassland along the migration routes and/
or in wintering grounds.
AUTHORITIES
No living person has had enough personal ex-
perience with Eskimo curlews to be considered an
authority on the species.
PREPARER'S COMMENTS
The preparer has been most impressed by the
failure of the Eskimo curlew population to re-
cover from its rapid decline in the 1880's after
hunting was stopped, in contrast with the golden
plover, a shorebird with a similar migration pat-
tern with which it associated in migration and on
its wintering grounds. The decrease in availability
of grasshopper egg pods, which were much more
numerous before most of the natural grassland of
the tall grass prairie belt was plowed up to plant
crops (U.S. Entomological Commission 1877)
may have been the main reason for failure to re-
cover. Grasshoppers avoid cultivated land for egg
laying and the great destructive flights of grass-
hoppers in the American prairies declined at
about the same time as the Eskimo curlew; both
coincided with the extensive breaking of the
prairie sod in the American Midlands.
LITERATURE CITED/SELECTED
REFERENCES
American Ornithologists 's Union. 1957. Checklist
of North American Birds, 5th ed. Port City
Press, Baltimore, Md.
Audubon, J. J. 1835. Ornithological Biography,
Vol. 3:69.
Austin, O. L., Jr. 1932. The birds of Newfound-
land Labrador. Memoirs Nuttall Ornith. Club.
Cambridge, Mass.
Baerg, W. J. 1931. Birds of Arkansas. Agric. Exp.
Sta. U. of Arkansas, Fayetteville.
Banks, R. C. 1977. The dechne and fall of the
Eskimo Curlew, or why did the curlew go ex-
taille? Amer. Birds 31:127-134.
Barrows, W. B. 1884. Birds of lower Uruguay.
Auk 1:313-319.
Beardslee, C. S. and H. D. Mitchell. 1965. Birds of
the Niagara Frontier Region. Bull. Buffalo
Soc. Nat. Sci. 22.
Bent, A. C. 1929. Life histories of North Ameri-
can shorebirds. Part IL U.S. Nat. Mus. Bull.
146.
Bond, J. 1956. Checklist of birds of the West
Indies. Acad. Nat. Sci. Philadelphia.
Bull, J. 1964. Birds of the New York Area. New
York, Harper and Row Publishers.
Butler, A. W. 1898. The Birds of Indiana. Indiana
Dept. Geol. & Nat. Resources 22nd Ann. Re-
port. Indianapolis.
Coues, E. 1861. Notes on the ornithology of
Labrador. Proc. Acad. Nat. Sci. Philadelphia
13:215-257.
. 1874. Birds of the Northwest; a hand-
book of the ornithology of the region drained
by the Missouri River and its tributaries. U.S.
Geol. Surv. Terr. Wise. Publ. No. 3, 791 pp.
Cooke, W. W. 1910. Distribution and migration of
North American shorebirds. Bur. Biol. Surv.
Bull. 35:1-100.
Dement'ev, L. P., and N. A. Gladkov, ed. 1969.
Birds of the Soviet Union Vol. III. Translated
from Russian by Israel Program for Scientific
Translators, Jerusalem pp. 356-360.
DuMont, P. A. 1934. A revised Hst of the birds of
Iowa. U. of Iowa Studies 15(5):1-171.
Emanuel, V. L. 1961. Another probable record of
an Eskimo Curlew on Galveston, Island, Texas.
Auk 78:259-260.
. 1962. Texans rediscover the near extinct
Eskimo Curlew. Audubon Magazine 64:162-
165.
Farrand, J., Jr. 1977. What to look for: Eskimo
and Little Curlews compared. Amer. Birds 31 :
137-138a.
Fish and Wildlife Service. 1973. Threatened Wild-
life of the United States. Resource Publ. 114,
289 pp.
Forbush, E. H. 1912. A history of the game birds,
wildfowl and shorebirds of Massachusetts and
adjacent states. Mass. State Board of Agric.
Boston 622 pp. (p. 416)
. 1925. Birds of Massachusetts and other
New England states. Part I. Mass. Dept. Agric.
Boston.
Greenway, J. C, Jr. 1958. Extinct and vanishing
birds of the world. Spec. Publ. No. 13 Amer.
Comm. Intemat. Wildlife Protection. New
York 518 pp.
Griscom, L. and D. E. Snyder. 1955. The Birds of
Massachusetts, and annotated and revised list.
Salem, Peabody Museum.
Griscom, L. and E. V. Folger. 1948. The Birds of
Nantucket. Harvard Univ. Press, Cambridge.
Hagar, J. A. and K. S. Anderson. 1977. Sight re-
cord of Eskimo Curlew. Amer. Birds 31:135-
136.
Housse, P. R. 1945. Las Aves de Chile. Ediciones
de la Universidad de Chile.
Jones, Lynds. 1903. Birds of Ohio. Ohio State
Acad. Sci. Spec. Papers No. 6.
Johnson, A. W. 1965. The birds of Chile. Buenos
Aires, Piatt Establecimientos Graficos S.A.
Johnston, R. F. 1960. Directory to the bird-
life of Kansas. U. of Kansas, Mus. Nat. Hist.
Misc. Pub. 23:172.
MacFarlane, R. 1891. Notes on and list of birds
collected in Arctic America, 1861-1866.
{Numeniiis p. 429) Proc. U.S. Nat. Mus. 14:
413-446.
MacKay, G. H. 1892. Habits of the Eskimo Cur-
lew {Nurnenius borealis) in New England.
Auk 9:16-21.
Merriam, C. H. 1877. A review of the birds of
Connecticut. New Haven, Tuttle, Morehouse
and Taylor Printers.
Murdock,J. 1885. Bird Migration at Point Barrow,
Arctic Alaska. Auk 2:63.
Murphye, R. C. 1933. Probable record of Eskimo
Curlew {Nurnenius borealis) at Montauk Point,
N.Y. Auk 50:101-102.
Nelson, E. W. 1887. Report on Natural history
collections made in Alaska between years
1877 and 1881. Arctic series of publications
issued in connection with the Signal Service,
U.S. Army. Wash. D.C. G.P.O.
Oberholser, H. C. 1938. The bird Hfe of Louisiana.
Louisiana Dept. of Conservation. Bull. 28.
. 1974. The bird life of Texas. Vol. I. U.
of Texas Press, Austin.
Palmer, R. S. 1949. Maine birds. Bull. Mus. Comp.
ZooL, Harvard, 102:656 pp.
Peters, H. S. and T. D. Burleigh. 1951. The Birds
of Newfoundland. Dept. Nat. Resources,
Newfoundland, St. Johns.
Ridgway, R. 1919. The birds of North and Middle
America. Part VIII. Bull. 50. U.S. Nat. Mus. p.
413.
Roberts, T. S. 1932. The birds of Minnesota. Vol.
I. Minneapolis, Univ. of Minnesota Press.
Sage, J. H. and L. B. Bishop. 1913. The birds of
Connecticut. Conn. Geol. and Nat. Hist. Surv.
Bull. 20.
Salvin, O. 1861. A list of species to be added to
the ornithology of Central America. Ibis 39:
351-356.
Sclater, P. L. and W. H. Hudson. 1889. Argentine
Ornithology. Vol. 2. R. H. Porter London, p.
192.
Swainson, W. and J. Richardson. 1881. Birds
(part second) Fauna Boreali-Americana. Lon-
don, John Murray.
Swenk, M. H. 1916. The Eskimo Curlew and its
disappearance. Smithsonian Rep. for 1915,
pp. 325-340.
. 1926. The Eskimo Curlew in Nebraska.
Wilson Bull. 38:117-118.
Todd, W. E. C. 1940. Birds of Western Pennsyl-
vania. Univ. Pittsburg Press.
. 1963. Birds of the Labrador Peninsula
and adjacent areas. Toronto, Univ. Toronto
press.
Townsend, C. W. and G. M. Allen. 1907. Birds of
Labrador. Proc. Boston Soc. Nat. Hist. 33:
277-428.
Tufts, Robie W. 1961. The birds of Nova Scotia.
Nova Scotia Museum, HjJifax.
U.S. Entomological Commission. 1877. First An-
nual Report of the U.S. Entomological Com-
mission for the year 1877 relating to the
Rocky Mountain Locust.
Vincent, J. (Compiler). 1966. Red data book.
Vol. 2. Aves. Intematl. Union for Conserva-
tion of Nature and Natural Resources. Morges.
Switzerland.
Wetmore, A. 1926. Observations on the birds of
Argentina, Paraguay, Uruguay and Chile. U.S.
Nat. Mus. Bull. 133:448 pp.
. 1939. Recent observations on the Eski-
mo curlew in Argentina. Auk 56:475.
Widmann, O. 1907. Birds of Missouri. St. Louis.
Williams, G. G. 1959. Probable Eskimo Curlew on
Galveston Island, Texas. Auk 76:539-541.
Wood, N. A. 1951. The birds of Michigan. Univ.
of Mich Press, Ann Arbor.
Biological Services Program
FWS/OBS-80/01.18
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
SOUTHERN SEA OTTER
ill a/ a
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoasl of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.58
11
FWS/OBS-80/01.18
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
SOUTHERN SEA OTTER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
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CREDIT: W. C. LOY, USFWS
SOUTHERN SEA OTTER
Enhydra lutris nereis [Merriam]
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivora
FAMILY Mustelidae
OTHER COMMON
NAMES California sea otter
DATE
Entered into SWIS To be determined
Updates 1 November 1978
LEGAL STATUS
Federal Threatened (42 FR 2965, 14 Jan 1977)
States Fully protected: California
REASONS FOR CURRENT STATUS
The southern sea otter is the southernmost
form of a marine mammal that originally ex-
tended along the rim of the Pacific Ocean from
the northern islands of Japan to Baja California.
Originally estimated to number from 16,000 to
10,000 off the California coast, the species was
heavily hunted for its valuable fur by Russian,
Indian, Spanish, British, and American traders. By
1911, the California population was reduced to
an estimated 50 animals. Following protection
afforded by the International Fur Seal Treaty of
1911 and by the State of California in 1913, the
population began to recover. A survey conducted
by the California Department of Fish and Game
in 1976 estimated the population at about 1,860
individuals located along a 257-km span of coast-
line (Woodhouse et al. 1977). The population
now (1978) occurs 2dong about 320 km of coast-
line. Considering that there are oil depots at both
ends of the current range (Moss Landing and
Morro Bay) and increased tanker traffic offshore.
and that sea otters may die if their fur is contami-
nated by oil, the current population size and dis-
tribution is marginal to insure survival.
PRIORITY INDEX
Not assigned.
DESCRIPTION
Although the smallest of marine mammals,
the sea otter is the largest member of the weasel
family. Adult otters in California weigh from 20
to 39 kg and are from 1 to 1.6 m long. The dense,
dark-browTi fur may become grizzled with age,
especially in older males, and the forelimbs are
modified for feeding, while the flipper-like hind
limbs are used in swimming (Merriam 1904,
Kenyon 1969). The skull is flattened dorsally,
with large nasal openings; two rear upper cheek
teeth are wider than long; there are three upper
premolars and two lower incisors on each side of
the jaw (Hall and Kelson 1959).
RANGE
Originally found from Morro Hermoso, Baja
California, north along the Pacific Coast to Cali-
fornia, Oregon, and Washington, the sea otter is
currently restricted to coastal waters of central
California. Established populations now occur
from Ano Nuevo Island in the north to Avila
Beach to the south (Wild and Ames 1974). Occa-
sional individuals are seen in California as far
north as Humboldt County and as far south as
Los Angeles County.
RANGE MAP
The extent of coastline supporting established
populations of the southern sea otter is shown
by shading on the accompanying map (Wild and
Ames 1974, R. J. Jameson, pers. observ.).
STATES/COUNTIES
California
HABITAT
Santa Cruz, Monterey, San Luis
Obispo
Otter habitat is the neritic zone within 3 miles
of shore; the animals are usually found within
one-half mile of shore and occasionally on shore.
Associated with a wide variety of sublittoral
communities, the presence of an adequate food
supply is an important factor in determining the
presence of sea otters (Woodhouse et al. 1977).
Although found over sandy substrates, sea otter
population centers seem to be associated with
hard substrates and stands of kelp [Macrocystis
pyrifera and Nereocystis lutkeana). Water depth
ranges from 0 to 36 m, in California, sea otters
are usually found in depths of less than 25 m.
Shelter from storm waves seems to be a require-
ment, either in the form of kelp beds or sheltered
coves (Kenyon 1969).
FOOD AND FORAGING BEHAVIOR
Food supply represents a critical environ-
mental factor for sea otters. A sea otter consumes
about 20% to 25% of its body weight in food
each day (Kenyon 1969), and a meal passes
through the digestive tract in about 2.8 hr (Stull-
ken and Kirkpatrick 1955). This large volume of
food is apparently needed to maintain body tem-
perature in cold waters (Morrison et al. 1974). To
meet these requirements, Loughlin (1977) found
that radio-tagged otters spent 34% of their total
time (or 73% of their active time) in foraging. Sea
otters are active both day and night, and 45% of
their foraging is nocturnal (Loughlin 1977,Shimek
and Monk 1977). Feeding is usually in water 1 to
25 m deep. Occasionally, individuals will leave the
water at low tide to forage for mussels {Mytilus
calif ornianus) , but they return to the water to eat
them (R. J. Jameson, unpubl.). Foraging dives
range from 10 to 120 sec, but are usually less than
60 sec long. Depth of water and availability and
type of prey seem to be important factors in de-
termining the length of these foraging dives. Food
is usually gathered from the bottom, but Califor-
nia sea otters spend considerable time foraging in
the kelp canopy where snails {Tegula spp) and
kelp crabs {Pugettia producta) are abundant. A
stone 'tool' may be used to break hard-shelled
molluscs, such as abalone, from the bottom
(Houk and Geibel 1974) and at the surface, this
tool may be used as an anvil to break mollusc
shells (Hall and Schaller 1964). Woodhouse et al.
(1977) conclude that the type of food eaten cor-
responds to availability more than to preference,
although foraging energetics certainly favors the
collecting of food items with high caloric rewards.
Foods commonly taken by sea otters in Call-
't. Ano Nuevo
Monterey
Bay
Pt. Lobos
Pt. Sur
MONTEREY
O
O
O
O
o
Morro Bay
Avila
-.1 L
miles
Pt. Conception
The range of the southern sea otter
3
fornia include the following: abalone [Haliotis
sp.), Turban snails {Tegula sp.), sea urchins
(Strongylocentrotus sp.), rock crabs {Cancer
sp.), kelp crabs {Pugettis. sp.), mussels {Mytilus
sp), Pismo clams (Tivela stultorum), and octopus
[Octopus sp.). (Hall and Schaller 1964, Ebert
1968, Wild and Ames 1974, Miller et al. 1975,
Woodhouse et al. 1977).
In recently reoccupied habitat, the diet may
consist almost exclusively of readily available
large food items such as abalone, sea urchins,
and Pismo clams, but with continued occupancy
and consequent depletion of large, easily obtained
food items, the otters' diets become more diverse
(WUd and Ames 1974).
SHELTER REQUIREMENTS:
Sea otters in California do not 'haul out' as
frequently as they do in Alaska; however, in re-
cent years several regular hauling areas have been
discovered. These areas are used primarily during
the winter and spring months when kelp beds are
reduced in size, and air and water temperatures
are lower (R. J. Jameson, unpubl.). Most of the
year, otters seek shelter in the extensive beds of
g\3int kelp that occur throughout most of the
present range.
area for extended periods (Loughlin 1977). Dur-
ing the breeding season, a male-female pair may
remain together for only one copulation, or may
form a bond that can last several days (Kenyon
1969, Vandevere 1970). It is not known if a spe-
cial set of habitat characteristics are required for
courtship and breeding, but available information
suggests there are none.
OTHER ENVIRONMENTAL REQUIREMENTS
Lacking a layer of blubber, sea otters depend
on air trapped in their dense fur for insulation
against heat loss to their environment (Kenyon
1969). With captives, Kenyon discovered that
soiled fur rapidly loses its insulating ability, ren-
dering the animal a sure victim to hypothermia.
Recent experiments in Alaska have shown that
sea otters exposed to crude oil floating on the
water can die (Siniff et al. 1977). This finding is
of critical importance, since it relates to the po-
tential damage to otters from offshore oil pollu-
tion. Although no wild sea otter deaths have yet
been reported from oil pollution in California,
a major spill could be very damaging (California
Department of Fish and Game 1976).
NESTING OR BEDDING:
Sea otters may roll themselves in kelp fronds
while resting or sleeping, presumably to avoid
transport by wind and currents. Mothers wdll
similarly leave pups in kelp while diving for food
(Fisher 1939). Typical haul-out areas are rela-
tively low-lying intertidal rocks covered with a
lush growth of a variety of marine algae.
RITUAL REQUIREMENTS:
Sea otters are most frequently members of
aggregations known as 'rafts' (LoughUn 1977,
Schneider 1978). Territoriality may be expressed
by some males who defend areas near female
rafts, but the evidence for territoriality is circum-
stantial and needs to be better documented.
Otters are known to remain in the same limited
POPULATION NUMBERS AND TRENDS
The rate of increase of the California sea otter
population has been 5.4% per year since 1940
(Woodhouse et al. 1977). The present population
is far below the 16,000 estimated as the potential
for California waters (California Department of
Fish and Game 1976), or the estimate of 47,800
given by Miller (1974) for all waters from the
Oregon-California border to Morro Hermoso, Baja
California. While the currently occupied range
may be nearing carrying capacity, population
growth has occurred in recent years by expansion
of the range both up and down the coast (Wild
and Ames 1974). Observations indicate that high
densities of otters occur at the limits of the range.
These groups, consisting primarily of males, move
into unoccupied territory as the food supply
behind them is depleted (California Department
of Fish and Game 1976). If expansion and popu-
lation growth are allowed to continue at current
rates, doubling of population size could be
expected every 15 years.
Resident sea otters are relatively sedentary;
the average linear dimension of the home range is
about 2.5 km (Loughlin 1977). Females have
larger home ranges (averaging 80 ha) than males
(38.5 ha) (Loughlin 1977); however, recent tag-
ging studies indicate that some otters, particularly
males, use large segments of coastline up to 160
km (R. J. Jameson Unpubl.). These individuals
spend the winter with large rafts of males near the
ends of the range, returning to the central part
during the peak breeding season (summer). They
have few natural enemies. White sharks appar-
ently prey on sea otters (Morejohn et al. 1975).
Most mortality seems to occur in young of the
year and older individuals (Kenyon 1969, More-
John et al. 1975). Sea otters have moderate to
heavy loads of internal parasites (Morejohn et al.
1975), and these may cause higher mortality
during times of stress. Due to the high daily
caloric requirement, severe winter storms pre-
venting feeding may be responsible for additional
mortality (Wild and Ames 1974),
REPRODUCTION
Some female sea otters begin reproductive
activity at 3 years of age, and all over 6 years are
reproducing. Males do not mature until 5 or 6
years of age (Schneider 1978). Breeding season
peaks from July to September (R.J.Jameson Un-
publ.), and pupping is most frequent from Novem-
ber through March (Vandevere 1970). Implanta-
tion is delayed 4 to 4'/2 months, and development
then proceeds for 4 to 4V2 months, making the
gestation period 8 to 9 months overall (Sinha et
al. 1966, Schneider 1978). Twinning is rare
among sea otters, and it is unlikely that a female
could support two pups. Two years has been ac-
cepted as the interval between births (Kenyon
1969), but recent studies in Alaska and California
indicate that some females pup in consecutive
years.
MANAGEMENT AND CONSERVATION
The California sea otter population currently
represents a slowly recovering population of ma-
rine mammals occupying only part of its original
range. With few natural enemies, continued growth
in numbers and range may be expected. Because
of its relatively low reproductive potential, con-
tinued protection from human predation and
habitat degradation will be necessary if this
recovery is to continue. The most recent sea-otter
survey, conducted by the California Department
of Fish and Game in 1979 resulted in an estimate
of less than 1,500, a decrease of about 300 from
the 1976 census. Since the weather conditions
during the 1979 survey were unfavorable, it is
considered quite low.
Kelp forests growing along rocky coasts ap-
pear to constitute optimum otter habitat in Cali-
fornia. Estes and Palmisano (1974) indicated that
sea otter predation on herbivores may be impor-
tant in the maintenance of large stands of kelp. If
increased growth of macrophytes is correlated
with this predation by sea otters, their presence in
an area may increase primary productivity. The
trophic consequences of the reestablishment of
the sea otter population remain to be seen. Mean-
while, protecting the kelp bed habitat from physi-
cal and chemical degradation will continue to be
important for the entire littoral ecosystem.
The southern sea otter now occupies only a
small fraction of its original range. The continuity
of the sea otters' current territory makes the
species quite vulnerable to catastrophic events
and augments the importance of establishing
satellite populations in other parts of its former
range to insure its survival. Previous translocation
programs have demonstrated the feasibility of es-
tablishing new populations of the sea otter (Ken-
yon 1969, Jameson et al. 1978). Estabhshment
of satellite populations will greatly decrease the
probability of the entire population's being des-
troyed by an epizootic or an oil spill.
In the past 10 years, questions have arisen
concerning the compatabiHty of the sea otter
with sport and commercial shellfishing interests.
The abundance of large abalones, sea urchins, and
Pismo clams diminishes following reoccupancy of
an area by sea otters (Miller 1974, Woodhouse et
al. 1977). However, other studies indicated that
after the initial period of reinvasion, the diet of
the sea otter becomes more diverse, and the com-
munity food web may enjoy a broader base
resulting from reduction of the numbers of large
herbivores (Palmisano and Estes 1977, Rosenthal
and Barilotti 1973), and a consequent increase of
macrophytic algae. Under such a regime of preda-
tion, large herbivores are usually restricted to
refugia protected from carnivores (Lowry and
Pearse 1973, Cooper et al. 1977). This was appar-
ently the undisturbed condition of California sub-
littoral communities (Rashkin 1972). The super-
abundance of large herbivores upon which the
commercial shellfish industry is based is appar-
ently an artifact of the historic reduction in sea
otter numbers. Herbivore-carnivore numbers can
be expected to return to equihbrium following
the sea otter's reoccupancy of its former range.
AUTHORITIES
Ronald J . Jameson
National Fish and Wildlife Laboratory
Piedras Blancas Field Station
P.O. Box 67
San Simeon, California 93452
Karl W. Kenyon
U.S. Fish and Wildlife
11990 Lakeside Place N.W.
Seattle, Washington 98125
Service (retired)
A. M.Johnson
National Fish and Wildlife Laboratory
Anchorage Field Station
4454 Business Park Boulevard
Anchorage, Alaska 99503
James E. Estes
National Fish and Wildlife Laboratory
CCMS; Applied Science Building
University of California
Santa Cruz, California 95064
J. E. Vandevere
93 Via Ventura
Monterey, California 93940
Tom Loughlin
National Marine Fisheries Service
Marine Mammals and Endangered Species Div.
F-33 NMFS
Washington, D.C. 20235
D. Miller and J. Ames
California Department of Fish and Game
2201 Garden Road
Monterey, California 93940
PREPARER'S COMMENTS
Recently there has been some controversy
over the systematic status of the southern popula-
tions of sea otter (Roest 1973, 1976; Davis and
Lidicker 1975). Pointing to an apparent latitudi-
nal cline in some skull measurements, Roest
(1973) contended that the southern sea otter is not
subspecifically distinct, but represents one end of
a size continuum. Davis and Lidicker (1975)
argue that available evidence is best interpreted to
suggest that a genetically distinct group of sea ot-
ters exists off the California coast.
Subspecies are often recognized primarily on
morphological criteria, which usually are the
result of genetic divergence due to selection with-
in different sets of environmental parameters. To
date, most attention has been paid to relatively
few cranial measurements in sea otters. The north-
ern and southern populations of the sea otter ap-
pear to display some differences in diet and ana-
tomy as well as in cranial morphology (Wood-
house et al. 1977, Miller 1974), which may or
may not be genetically based. It is certainly pos-
sible that selection has resulted in some genetic
divergence between these populations. However,
we suggest the application of modern systematic
techniques aimed at assessing this genetic distance
between populations (such as karyology, protein
electrophoresis, immunology, and perhaps DNA
annealing) to adequately resolve this controversy.
Meanwhile, it would be inappropriate to sacrifice
the protection afforded the recovering southern
populations to a disagreement over an as-yet
unresolved taxonomic issue. This issue was ad-
dressed by the USFWS in the Federal Register (14
January 1977): "This question actually is not
relevant to the matter at hand, because sections 3
and 4 of the Act allows [sic] the listing of popu-
lations of species in portions of their range, as
well as entire species and subspecies. Since the
southern sea otter does form a significant popu-
lation, it can be treated independently under the
Act, regardless of its taxonomic status. The Ser-
vice decided, however, to utilize the subspecific
designation Enhydra lutris nereis in this rule-
making, although this decision had no connection
with the decision to list as threatened."
LITERATURE CITED/SELECTED
REFERENCES
Barabash-Nikiforov, I. I., V. V. Reshetkin, and N.
K. Shidlovskaya. 1947. The sea otter (kalan).
Transl. from Russian by A. Birron and Z. S.
Cole. 1962. Nat. Sci. Found, and U.S. Dept.
Int., Washington, D.C. (Israel Program for Sci.
Transl.).
Bolin, R. L. 1938. Reappearance of the southern
sea otter along the California coast. J. Mamm.
19:301-303.
Boolootian, R. A. 1961.
California sea otter.
47:287-292.
The distribution of the
Calif. Fish and Game
California Dept. of Fish and Game. 1976. A
porposal for sea otter protection and research,
and request for the return of management to
the state of California., 2 vol., unpubl.
Cooper, J. M. and A. Hines. 1977. Subtidal aba-
lone populations in an area inhabited by sea
otters. Veliger 20:163-167.
Davis, J. and W. Z. Lidicker, Jr. 1975. The taxon-
omic status of the southern sea otter. Proc.
Calif. Acad. Sci. 40:429-437.
Hall, K. R. L. and G. B. Schaller. 1964. Tool-
using behavior of the California sea otter. J.
Mammal 45:287-298.
Houk, J. L. and J. J. Geibel. 1974. Observation
of underwater tool use by the sea otter,
Enhydra lutris Linnaeus. Calif. Fish and Game
60:207-208.
Jameson, R. J., A. M.Johnson and K. W. Kenyon.
1978. The status of translocated sea otter
populations in the eastern Pacific Ocejm.
Proc. 2nd Conf. Biol. Marine Mamm. p. 8.
Kenyon, K.W. 1969. The sea otter in the eastern
Pacific ocean. No. Amer. Fauna. 68 pp.
Lensink, C. J. 1962. The history and status of
sea otters in Alaska. Unpubl. Ph.D. disserta-
tion, Purdue Univ. pp.
Loughlin, T. R. 1977. Activity patterns, habitat
partitioning, and grooming behavior of the sea
otter, Enhydra lutris, in California. Unpubl.
Ph.D. dissertation, Univ. of Calif., Los
Angeles. pp.
Lowry, L. F. and J. S. Pearse. 1973. Abalones
and sea urchins in an area inhabited by sea
otters. Mar. Biol. 23:213-219.
Ebert, E.E. 1968. A food habits study of the
hem sea otter, Enhydra lutris nereis. Calif.
Fish and Game 54:33-42.
Estes, J.A. 1977. Population estimates and feed-
ing behavior of sea otters, pp 51 1-525 in M.L.
Merritt and R. G. Fuller, eds., The Environ-
ment of Amchitka Island, Alaska. U.S. ERDA
TID-26712.
Estes, J.A. and J. F. Palmisano. 1974. Sea otters:
their role in structuring nearshore communi-
ties. Science, 185:1058-1060.
Fisher, E. M. 1939. Habits of the southern sea
otter. J. Mammal 20:21-36.
Hall, E. R. and K. R. Kelson. 1959. The mammals
of North America. Ronald Press, New York.
Merriam, C. H. 1904. A new sea otter from
southern California. Proc. Biol. Soc. Washing-
ton 17:159-160.
Miller, D.J. 1974. The sea otter Enhydra lutris.
Calif. Dept. Fish and Game, Marine Resources
Leaflet 7. pp.
Miller, D. J., J. E. Hardwick and W. A. Dahlstrom.
1975. Pismo Clams and sea otters. Calif. Dept.
Fish and Game, Marine Resources Tech. Rep.
31. pp.
Morejohn, G. V., J. A. Ames, and D. B. Lewis.
1975. Post mortem studies of sea otters,
Enhydra lutris L., in California. Calif. Dept.
Fish and Game, Marine Resource Tech. Rep.
30. pp.
Morrison, P., M. Rosenmann, and J. A. Estes.
1974. Metabolism and thermoregulation in
the sea otter. Physiol. Zool., 47:218-229.
Palmisano, J. F. and J. A. Estes. 1977. Ecological
interactions involving the sea otter, pp. 527-
567, in M. L. Merritt and R. G. Fullers, eds.,
The Environment of Amchitka Island, Alaska.
U.S. ERDATID- 26712.
Rashkin, P. 1972. Monterey Peninsula shell
mounds — some general remarks. Monterey
Co. Archael. Soc. Quart. 1:5.
Roest, A. I. 1973. Subspecies of the sea otter,
Enhydra lutris. Los Angeles Co. Mus. Contrib.
Sci. 252:1-17.
. 1976. Systematics and the status of sea
otters, Enhydra lutris. Bull. So. Calif. Acad.
Sci. 75:267-270.
Rosenthal, R.J. and D. C. Barilotti. 1973. Feed-
ing behavior of transplanted sea otters and
community interactions off Chichagof Island,
southeast Alaska, pp. 74-88, in W.J. North,
ed., Calif Inst Tech., Kelp Habitat Improv.
Proj.,Ann. Rep. 1 July 1972-30 June 1973.
Sandegren, F. E., E. W. Chu, and J. E. Vandevere.
1973. Maternal behavior in the California sea
otter. J. Mammal 54:668-679.
Scheffer, V. B., and F, Wilke. 1950. Validity
of the subspecies Enhydra lutris nereis, the
southern sea otter. J. Washington Acad, Sci.
40:269-272.
Schneider, K. B. 1978. Sex and age segregation
of sea otters. Alaska Dept of Fish and game,
Final Rep. , Job 8.9R 45 pp.
Shaw, S. B. 1971. Chlorinated hydrocarbon
pesticides in California sea otters and harbor
seals. Calif. Fish and Game 57:290-294.
Siniff, D. B., A. M. Johnson, and T. W. Williams.
1977. Observations on responses of sea otters
[Enhydra lutris) to oil contamination. Proc.
2nd Conf. Biol. Mar. Mammal, p. 32.
Stephenson, M. D. 1977. Sea otter predation on
pismo clams in Monterey Bay. Calif. Fish and
Game 63:117-120.
Stullken, D. E., and C. M. Kirkpatrick. 1955.
Physiological investigation of captivity mor-
tality in the sea otter {Enhydra lutris). Trans.
20th N. Amer. Wildl. Conf. pp. 476-494.
Vandevere, J. E. 1970. Reproduction in the sea
otter, pp. 221-227, in Proc. 7th Ann. Conf.
on Biol. Sonar and diving Mammals, Stanford
Res. Inst., Menlo Park, Calif.
Wild, P. W. and J. A. Ames. 1974. A report on
the sea otter, Enhydra lutris L., in California.
Calif. Dept Fish and Game, Marine Resources
Tech. Rep. 20. PP-
Woodhouse, C. D., R. K. Cowen, and L. R.
Wilcoxon. 1977. A summary of knowledge of
the sea otter, Enhydra lutris, L., in California
and an appraisal of the completeness of bio-
logical understanding of the species. U.S. Mar.
Mammal Comm. Rep. No. MMC-76/02 (PB
270 374).
ACCOUNT PREPARED BY
National Fish and Wildlife Laboratory
Piedras Blancas Field Station
P.O. Box 67
San Simeon, California 93452
National Fish and Wildlife Laboratory
1300 Blue Spruce Drive
Fort Collins, Colorado 80524
Shimek, S. J. and A. Monk. 1977. Daily activity
of sea otter off the Monterey peninsula,
California. J. Wildl. Mgmt. 41(2):277-283.
Sinha, A. A., C. H. Conaway, and K. W. Kenyon.
1966. Reproduction in the female sea otter.
J. Wildl. Mgmt. 30(1):121-130.
Biological Services Program
FWS/OBS-80/01.19
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
MORRO BAY KANGAROO RAT
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Knginecrs in coordina-
tion with the Offices of P^ndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.19
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
MORRO BAY KANGAROO RAT
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Shdell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT: GLENN R. STEWART
MORRO BAY KANGAROO RAT
Dipodomys heermanni morroensis (Merriam)
KINGDOM Animalia
CLASS Mammalia
ORDER Rodentia
FAMILY Heteromyidae
OTHER COMMON NAMES Heermann's
kangaroo rat
DATE
Entered into SWIS to be determined
Updates 16 October 1978
LEGAL STATUS
Federal Endangered (35 FR 16047, 13 Oct
1970).
States Endangered: California
REASONS FOR CURRENT STATUS
Historically, this isolated subspecies has occu-
pied a range of less than 10 km^ (Grinnell 1922).
Recent appraisals have documented a continually
shrinking range and, concomitantly, decreased
population size. Three factors have contributed to
this decline (Congdon and Roest 1975):
1. Direct loss of habitat due to growth of resi-
dential areas.
2. Successional changes of sparsely vegetated
areas into more thickly vegetated chaparral
communities.
3. Increased predation by domestic cats hunting
in fields adjacent to human dwellings.
PRIORITY INDEX
Not assigned
DESCRIPTION
The darkest colored of all kangaroo rats, this
population was originally described as a distinct
species, partly on the basis of its dark pelage and
markings (Merriam 1907, Grinnell 1922). The hip
stripe, characteristic of the genus, is often incom-
pletely formed, thus failing to completely sepa-
rate the thigh patch from the rest of the back
(Grinnell 1922, Stewart and Roest 1960). The
white side tail stripes are narrow; less than one-
half the width of the dark dorsal and ventral
stripes. The tail is often completely black just be-
hind the white basal collar (Grinnell 1922). Audi-
tory bullae are less highly inflated than neighbor-
ing subspecies of D. heermanni (Grinnell 1922,
Boulware 1943).
RANGE
This rat is found in six disjunct patches of
habitat just south and southeast of Morro Bay,
California. The total area currently occupied is
estimated at 1.3 km^ (Roest 1977). All current
populations occupy remnants of the historical
range, and can be enclosed by a circle 6.5 km in
diameter.
RANGE MAP
Distribution is shown on the accompanying
map (after Roest 1977). Occupied areas are indi-
cated by shading.
STATES /COUNTIES
California: San Luis Obispo.
HABITAT
Habitat consists of early serai stages of the
chaparral community, where vegetation is low
and sparse and shrubs are widely scattered (Cong-
don and Roest 1975), on medium-textured sandy
loam (Stewart and Roest 1960). Plants typical of
the habitat include Lotus scoparius, Erigonum
parvifolium. Salvia mellifera, and scattered annual
grasses (Stewart and Roest 1960).
FOOD AND FORAGING BEHAVIOR
Food in the wild not known. In the related
Tulare subspecies (D. h. tularensis) , food consists
primarily of seeds of grasses and shrubs during the
dry season, and grass and herb cuttings during the
rainy season. Also, small amounts of insect mater-
ial are taken seasonally (Tappe 1941, Fitch 1948).
The Morro Bay subspecies probably stores small
amounts of food material in its burrows, as
reported for the Tulare subspecies (Fitch 1948).
Captive Morro Bay kangaroo rats eat seeds and
leaves of native plants found within their range,
including those of Lotus, Dudleya, and Bromus
(Stewart and Roest 1960).
SHELTER REQUIREMENTS
Like other heteromyid rodents, the Morro
Bay kangaroo rat constructs its own burrow in
sandy soil. Burrows are often located along low
ridges near open space, particularly in areas of
thick brush (Congdon and Roest 1975).
The burrow of D. h. morroensis consists of
shallow tunnels (15 to 30cm below the surface)
measuring 1.8 to 3 m long, with two or three
chambers and a similar number of escape tunnels,
the latter terminating about 2.5 cm below ground
level (Stewart and Roest 1960).
NESTING OR BEDDING
Nests are located in one of the chambers of
the burrow systems (Stewart and Roest 1960).
No description of the nest of this subspecies is
available, but Tappe (1941) described the nest of
the Tulare kangaroo rat as an ovoid chamber 12.7
by 15 cm and 10 cm high, which was lined with
fine grass stems, fine grass roots, and husks of
grass seeds. The nests were used only during the
breeding season. A similar nest may be made by
the Morro Bay kangaroo rat.
RITUAL REQUIREMENTS
The behavior of this free-living nocturnal
rodent is not known. Information for the related
great basin kangaroo rat (Z). microps) indicates a
need by members of the genus Dipodomys for
open ground around the burrow (Kenagy 1976).
In D. microps, courtship is limited to drumming
miles
Current distribution of the Morro Bay kangaroo rats.
by the male around the burrow of the female.
Receptive females emerge and copulation quickly
commences in the vicinity of her burrow. Com-
peting males interact, including bouts of locked
fighting, near the female's burrow (Kenagy 1976).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
The population density of other species of
kangaroo rats has been shown to be inversely
related to the density of shrubby vegetation
(Rosenzweig 1973). This relationship reflects the
requirements of saltatory locomotion and the
erratic-leaping type of escape behavior that is
highly developed in kangaroo rats. Kenagy (1976)
suggests the leaping behavior may also function in
reducing male-to-male physical contact during
conflicts over access to females' burrows. Stewart
and Roest (1960) report the invasion of a burned
area by the Morro Bay kangaroo rat. Evidence
indicates that open, sparsely vegetated habitat is a
critical requirement for populations of this
kangaroo rat.
POPULATION NUMBERS AND TRENDS
The population of this subspecies is directly
related to both the quality and quantity of avail-
able habitat. The range was originally described
by Grinnell (1922) as "less than 4 miles square"
(41 km^ ). Stewart and Roest (1960) reported the
area of remaining habitat to be less than 6.5 km^ .
In 1960, the population numbered about 8,000
individuals (Congdon and Roest 1975). A 1971
study indicated a range of 3.6 km^ and a popu-
lation of 3,000 kangaroo rats Congdon 1971, in
Congdon and Roest 1975). Roest (1977) indi-
cated that the range is now reduced to 1.3 km^
and the population numbers are between 1,500
and 2,000 individuals.
The density varies from 42 kangaroo rats per
hectare in the most favorable habitat to an esti-
mated 5 per hectare in marginal habitat. Territory
size is estimated to be about 0.07 ha (Roest
1977). Roest (1977) indicates considerable sea-
sonality in their activity, with little or no activity
in January and February, and increasing activity
through early summer. A decline in activity was
noted in mid-summer, and a fall resurgence is
suggested.
REPRODUCTION
Most young are probably born from February
through August, but breeding may continue
throughout the year with a possible second peak
of breeding in the fall (Roest 1977). A similar
concentration of breeding in the spring and
summer months occurs in D. h. tularensis, which
also may have multiple litters per year (Fitch
1946). Average litter size is probably just over
three. Four recorded litters for the Morro Bay
subspecies were 4, 3, 3, and 3 (Stewart and Roest
1960). Other subspecies of D. heermanni are
reported to have average litter sizes of 2.6 to 3.7
(Fitch 1946).
MANAGEMENT AND CONSERVATION
A summary of management problems for this
subspecies is presented in Congdon and Roest
(1975). The most significant adverse factor is the
loss of habitat, caused by conversion of parts of
the former range into suburban housing develop-
ments. While kangaroo rats may be tolerant of the
proximity of human dwellings (Stewart and
Roest 1960), they are eliminated from developed
land. A concomitant problem is predation by do-
mestic cats near houses. A more serious problem
in remaining prime habitat is the absence of wild
fires, leading to higher shrub density and the
development of a mature chaparral community.
This subspecies may invade habitat created by
burning (Stewart and Roest 1960). It disappears
from fields as the shrub height and density in-
crease (Congdon and Roest 1975). Management
should thus be directed towards prevention of
further loss of habitat by a moratorium on devel-
opment of any remaining Morro Bay kangaroo rat
habitat and maintenance of existing habitat in the
early stages of succession, which constitutes the
favored condition for this subspecies. Both these
management programs could be most easily
achieved within the confines of a publicly owned
preserve. Recently, a 20-ha refuge for this animal
was established by the California Department of
Fish and Game for the westernmost ('Dunes')
population (see range map)(Gustafson 1978). But
the habitat in this refuge contains the lowest
population density of Morro Bay kangaroo rats
measured by Roest (1977). Other areas of prime
habitat should be acquired. Areas of favorable
habitat which are used by off-road vehicles are
likely to suffer severe deterioration (U.S. Depart-
ment of the Interior 1973), and such use should
be curtailed.
Fitch, H. S. 1948. Habits and economic relation-
ships of the Tulare kangaroo rat. J. Mammal.
29:5-35.
AUTHORITY
Aryan Roest
Biological Sciences Department
California Polytechnic State University
San Luis Obispo, California 93407
PREPARER'S COMMENTS
One reason for preserving this distinctive iso-
lated population of kangaroo rats is its scientific
status. From its first description in 1907 to the
time of Boulware's publication (1943), it was
recognized as a separate species. While no avail-
able evidence indicates that it should not be con-
sidered a subspecies of D. heermanni, Boulware
(1943:393) found the characters of the Morro
Bay kangaroo rat to be most different from its
nearest neighbor (Z). h. jolonensis) and most simi-
lar to a more removed southern relative (Z). h.
arenae). Further study on the taxonomic position
of the Morro Bay kangaroo rat is indicated. The
species has a high reproductive potential and col-
lection of a few specimens for critical examina-
tion would present no danger to the population.
The population of this subspecies is continu-
ously declining and without protective measures,
'the Morro Bay kangaroo rat is almost certainly
doomed to extinction.' (Congdon and Roest
1970).
LITERATURE CITED/SELECTED
REFERENCES
Boulware, J. T. 1943. Two new subspecies of
kangaroo rats (genus Dipodomys) from south-
ern California. Univ. California Publ. Zool.
46:391-396.
Congdon, J., and A. Roest. 1975. Status of the
endangered Morro Bay kangaroo rat. J.
Mammal. 56:679-683.
Csuti, B. A. 1971. Karyotypes of kangaroo rats
from southern California. J. Mammal. 52:
202-206.
Grinnell, J. 1922. A geographical study of the
kangaroo rats of California. Univ. California
Publ. Zool. 24:1-124.
Gustafson, J. 1978. Morro Bay kangaroo rat land
acquisition. California Dep. Fish Game Non-
game Wildl. Invest. Final rep. E-1-1. 3 pp.
Kenagy, G. J. 1976. Field observations of male
fighting, drumming, and copulation in the
Great Basin kangaroo rat {Dipodomys mi-
crops).]. Mammal. 57:781-785.
Merriam, C. H. 1907. Descriptions of ten new
kangaroo rats. Proc. Biol. Soc. Washington
20:75-80.
Roest, A. 1977. Distribution and population esti-
mate of the Morro Bay kangaroo rat. Califor-
nia Dep. Fish Game Nongame Wildl. Invest
Final Rep. E-1-1. 19 pp.
Rosenzweig, M. L. 1973. Habitat selection experi-
ments with a pair of coexisting heteromyid
rodent species. Ecology 54: 111-117.
Stewart, G. R. and A. I. Roest. 1960. Distribution
and habits of kangaroo rats at Morro Bay. J.
Mammal. 41:126-129.
Tappe, D. T. 1941. Natural history of the Tulare
kangaroo rat. J. Mammal. 22:117-148.
U.S. Department of the Interior. 1973. Threatened
wildlife of the United States. Resource Publ.
114.289 pp.
PREPARED/UPDATED BY
National Fish and Wildlife Laboratory
1300 Blue Spruce Drive
Fort ColHns, Colorado 80524
Biological Services Program
FWS/OBS-80/01.20
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
CALIFORNIA LEAST TERN
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services vi the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Flndangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.20
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
CALIFORNIA LEAST TERN
A Cooperative Effort
by the
National Fish and WUdlife Laboratory,
the Office of Endjingered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CALIFORNIA LEAST TERN
Sterna albifrons browni
KINGDOM Animalia
CLASS Aves
ORDER Charadriiformes
FAMILY Laridae
OTHER COMMON
NAMES Least tern, brown tern,
brown least tern
DATE
Entered into SWIS To be determined
Updates 22 Nov 1978
LEGAL STATUS
Federal
States
Endangered (35 FR 16047, 13 Oct
1970; 35 FR 18320, 2 Dec 1970)
Endangered: California
REASONS FOR CURRENT STATUS
The least tern nests on sandy ocean beaches
and salt flats near lagoons and estuaries. Coastal
highway construction, recreational development,
and beach cottages have reduced habitat and per-
mitted increased access to tern nesting beaches.
Feeding areas have been developed, filled in, and
polluted (Wilbur 1974). Continuing loss of shore-
side nesting and feeding habitat, along with high
levels of human disturbance and domestic preda-
tors at the remaining colonies, have been respon-
sible for a decline in numbers up to the present
times (Craig 1971, Wilbur 1974).
DESCRIPTION
Least terns (subfamily Sterninae) are the
smallest of the terns, measing 23 cm long with a
wingspread of about 51 cm (Davis 1968). They
are characterized by a black cap and white fore-
head, grey wings with black tips and leading edges,
yellow legs, and a black-tipped yellow bill. The
sexes are similar. Immatures have darker plumage
and dark bills; the contrast between their white
heads and dark eye stripes is often distinctive (K.
Bender in Wilbur 1974). It has faster wingbeats
than other terns, and can hover longer than the
larger terns.
The California least tern cannot be reliably
differentiated from other races of the least tern in
the field on the basis of plumage characteristics
alone (Burleigh and Lowery 1942).
Illustrations and descriptions appear in Ridg-
way (1919) and Bent (1921).
PRIORITY INDEX
Not assigned.
RANGE
The historical breeding range of the California
least tern extended along the California coast
from Moss Landing, Monterey County, to San
Jose del Cabo in southern Baja California (Daw-
son 1924, Grinnell 1928, Grinnell and Miller
1944, A.O.U. 1957). Some least terns have nested
in a number of locations farther north; between
1939 and 1954, they nested at the mouth of the
Pajaro River, Santa Cruz County (Pray 1954,
Wilbur 1974), and nesting near San Francisco Bay
in Alameda, Alameda County, was confirmed in
1967 (Chandik and Baldridge 1967). There are
numerous spring and summer records for this area
(Grinnell and Wythe 1927, Allen 1934, Sibley
1952, DeBenedictis and Chase 1963, Chase and
Paxton 1965). Wilbur (1974) suggests that
nestmg may have occurred here previously. In
south San Francisco Bay, 60 were counted on 1
Sept 1968, and a breeding colony of 30 pairs was
established on Bay Farm Island, Alameda County,
in 1969 (Gill 1977). A colony of 15 pairs was also
reported in 1969 from Bair Island, San Mateo
County (Anderson 1970). The nesting range has
apparently always been widely discontinuous,
with the majority of birds nesting in southern
California from southern Santa Barbara County
south through San Diego County (Wilbur 1974).
Known nesting sites in Baja California are Scam-
mons Lagoon (Bancroft 1927, Grinnell 1928),
San Jose del Cabo (Lamb 1927, Grinnell 1928),
Ensenada (Bahia Todos Santos), and Baliia de San
Quintin (Massey 1977).
The California least tern is migratory, usually
arriving at its breeding area during the last week
of April and departing again in August (Davis
1968, 1974; Massey 1971; Swickard 1971, 1973),
although terns have been recorded as early as 1 3
March and as late as 31 October (Sibley 1952,
Wilbur 1974).
There are six records north of San Francisco
Bay and four records of single birds seen at Hum-
boldt Bay, Humboldt County; two specimens
were collected at Fort Steven, Clatsop County,
Oregon (DeSante et al. 1972, CLTRT, in prep.)
Migrating least terns have also been recorded from
the Colorado River valley (Grater 1939, Phillips
et al. 1964) and at the Salton Sea, Imperial Coun-
ty (McCaskie and Cardiff 1965, McCaskie 1971).
The winter distribution of the California least
tern is unknown. Least terns in Guatemala
(Griscom 1932) and Veracruz (Warner and Mengel
1951) have been identified as members of the
subspecies browni. California least terns are
thought to migrate to Peru, but Murphy (1936)
doubts this. Recent efforts to find them in Peru
have been unsuccessful.
RANGE MAP
Known nesting locations for the California
least tern are shown on the accompanying map
(from Wilbur 1974).
STATES/COUNTIES:
California: Alameda, Imperial, Los Angeles,
Monterey, Orange, San Diego, San
Mateo, Santa Barbara, Santa Cruz,
Ventura.
Mexico: Locations of breeding colonies in Baja
California south of Bahia de San
Quintin are unknown.
HABITAT
Nesting colonies require undisturbed flat areas
with loose substrate, such as sandy upper sea
beaches or open expanses of fill-dirt or dried mud
characterized by little or no vegetation. For forag-
ing, they need adjacent open ocean, surf-line,
estuary or lagoon with a good supply of small fish
(Grinnell and Miller 1944; Craig 1971; Massey
1971; Swickard 1971, 1973; Wilbur 1974).
FOOD AND FORAGING BEHAVIOR
Least terns often feed in small loose groups of
2 to 10 birds. While fishing, they hover above the
water. If prey is sighted, the bird quickly plunges
to the surface, usually breaking its fall just before
reaching the water so it is only partially sub-
merged instead of diving beneath the surface.
Often, a series of short plunges and repeated hov-
erings will occur before the actual dive, which is
then usually made from only a few centimeters
above the water (Bent 1921, Hardy 1957).
They have not been seen eating anything but
fish, including northern anchovy {Engraulis mor-
dax), shiner perch {Cymanogaster aggregata),
anchovy [Anchoa compressa), topsmelt (Athertn-
ops affinis), killifish {Fundulus parvtpinnts) , ']a.ck-
smelt [Atherinopsis calif orniensis) , California
grunion [Leuresthes tenuis) and mosquito fish
(_) Knovm occupied since 1970
ELDORADO
V
S CRUZ
5'
25//281 AJ30)^'"' °'^'^°
I25/ 29
22- ^^ - y
'32'
27
39
41
'ao^
43
(34)
California least tem nesting locations.
{Gambusia affinis) (Swickard 1971;Massey 1972,
1974; CLTRT in prep). Other subspecies occa-
sionally feed on Crustacea, molluscs, sand eels
(Ammodytes), annelid worms, and insects
(Tompkins 1959).
SHELTER REQUIREMENTS
At first the young are weak and helpless.
Adults brood chicks continuously the first day.
By the second day, chicks make short walking
trips from the nest. From the third day on, they
become more active and are increasingly mobile
(CLTRT in prep.)
NESTING OR BEDDING:
Nests are simple, shallow depressions in sand,
dirt or gravel, usually without twigs or other
materials. Bent (1921) states that on beaches
where there are shells, the birds may encircle their
scrapes with them. Likewise, Swickard (1971,
1973) reported finding one nest, located on flat
ground, that was completely lined with small
twigs; Massey (1974) reported that nearly 20% of
the nests she studied were partly or completely
shell-lined.
In sand, the bird scoops out the nest depres-
sion (Davis 1968, 1974; Massey 1971, 1974;Swi-
ckad 1971). But in hard soil, such as dried mud or
fill, the nest can be any kind of natural or artifi-
cial depression, including a dried boot track (Swi-
ckard 1971).
Scrapes are circular, about 20 mm deep and
120 mm in diameter (Hardy 1957). Temporary
scrapes are used for brooding when chicks begin
to wander (Hardy 1957).
Least terns are colonial, but usually do not
form dense concentrations (Wilbur 1974). The
distance between nests is 3 to 5 m or more (Wolk
1954; Hardy 1957; Massey 1971, 1974). Nesting
densities may be as low as 3 to 7 per ha (Swickard
1971, 1973), although they are often greater; 145
pairs nested on about 0.75 ha at FAA Island in
Mission Bay, San Diego County, California.
RITUAL REQUIREMENTS
Nesting beaches are usually used as a parading
ground during courtship (Hardy 1957, Wilbur
1974).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Open expanses free from human disturbance
are necessary for courtship, resting, and nesting;
adjacent unpolluted lagoons or estuaries are
needed for feeding (Longhurst 1969, Craig 1971,
Wilbur 1974).
POPULATION NUMBERS AND TRENDS
Thousands of terns once nested on the beaches
and near the estuaries of the Pacific Coast of
North America from Monterey Bay south to cen-
tral Baja California. These flocks have dwindled
rapidly in the last century. In 1970, the popula-
tion in California was estimated at less than 300
pairs, scattered over 15 nesting colonies (Craig
1971), but this study was not made in the nesting
season and did not attempt to obtain precise
population data. The California Department of
Fish and Game has since conducted annual cen-
suses. In 1973, initial surveys estimated a popula-
tion size of 624 pairs, with nesting activity at 19
sites (Bender 1973). The 1974 surveys estimated
the breeding population at 582 pairs. Breeding
activity was recorded at 20 sites, but nesting
occurred at only 16 of them (Bender 1974). The
estimated breeding population was 600 pairs in
1975 and 664 pairs in 1976 (Atwood et al. 1977).
In 1977, 775 breeding pairs were located at
29 colony sites, but 9 colonies were abandoned or
had poor nesting success because of human distur-
bance, predation, or flooding (Atwood et al.
1977). This was the largest breeding population
since the surveys began, but the greater numbers
found are due in part to increased survey staff
and more accurate survey methods (Atwood et al.
1977).
The breeding population of the California
least tern is difficult to assess for several reasons.
Birds may shift breeding sites from year to year,
breeding dates may not be uniform, and age at
first reproduction and longevity are unknown.
Less standardized and concentrated censusing in
the past no doubt missed nesting birds at some
locations, and some censuses were too late or too
early to find breeding birds.
Local perturbations compound the difficul-
ty in estimating population trends. A colony at
Huntington Beach (Sunset Aquatic Regional
Park), Orange County, had 51 nests in 1968
(Davis 1968), 25 in 1969 (Massey 1971), 12 in
1970 (Massev 1971) and only 1 or 2 in 1972
(Bender 1973, 1974). At this site, fencing was in-
stalled and vegetation removed to improve nesting
success, but the birds gradually stopped nesting
there for unknown reasons, and had not returned
as of 1978. In 1969, a colony of 15 pairs was
reported from Bair Island, San Mateo County
(Anderson 1970). The birds did not use this area
in 1971, but were present again from 1972 to
1975; 14 active scrapes were located in July 1975
(Gill 1977).
REPRODUCTION
California least terns are sociable and colonial
nesters (Davis 1974). They are easily disturbed on
the nest and will take flight for aerial territorial
defense. If repeatedly disturbed, they abandon
their nests (Hardy 1957, Davis 1974, Wilbur
1974). Eggs are small, measuring about 31 by 24
mm, and are buff-colored with various brown and
purple streaks and specks. One to four are laid,
with two to three the most common clutch size
(Davis 1968, 1974; Anderson 1970; Swickard
1971, 1973; Massey 1972). Eggs are laid late in
the morning on consecutive days (Davis 1968,
Massey 1971, 1974).
Nesting starts in mid-May, with most of the
nests completed by mid-June (Bent 1921, Davis
1968, Swickard 1971, Massey 1974). Late season
(July and August) nests may be renests (Wilbur
1974). Chambers (1908) believed that terns often
reared more than one brood per season, but other
authorities do not consider the least tern a multi-
ple-nesting species (Wilbur 1974).
Incubation, which begins with the laying of
the first egg, is irregular at first, but become regu-
lar once the clutch is completed (Davis 1968,
Swickard 1971, Massey 1972). The female does
most of the incubating, but both parents partici-
pate (Davis 1968, 1974).
The incubation period varies from 17 to 28
days; the normal length is 20 to 25 days (Hagar
1937; Hardy 1957; Davis 1968, 1974; Swickard
1971; Massey 1972). An incubation period of 14
to 16 days given by Bent (1921) apparently is in
error. Eggs hatch on consecutive days. Chicks are
initially weak, but become strong and mobile by
the third day (Davis 1968, 1974).
In two colonies, hatching success was 80% to
90%, and fledging rates varied betwen 1 1% and
50% (Swickard 1971, Massey 1972). Infertility
appears to be a minor cause of egg failure (Wilbur
1974). Predation pressures appear to be high on
both eggs and fledgings. Domestic predators
(house cats and dogs) and Norway rats {Rattus
norvegicus) are often implicated (Chambers 1908;
Edward 1919; Craig 1971; Swickard 1971, 1973;
Pentis 1972; Atwood et al. 1977).
In the past, high tides caused heavy losses of
California least tern eggs (Shepardson 1909,
Sechrist 1915). Now few colonies are found in
areas where tides are a problem. However, in areas
with soil less permeable than beach sands, sum-
mer rains can cause serious nest losses. Swickard
(1971) noted that birds nesting on salt flats had
only 43% hatching success after flooding from
heavy rains, while on adjacent beach sands, rain
water percolated through the nests and there was
a 90% hatching success.
MANAGEMENT AND CONSERVATION
The least tern can tolerate some kinds of
human activities; they have nested successfully
between runways on military air fields (Atwood
et al. 1977), in a small fenced enclosure on a pub-
lic beach (Anon 1970, Massey 1972), beside a
shopping center parking lot (Atwood et al. 1977),
and on man-made fill alongside coastal lagoons
and estuaries (Massey 1972, Pentis 1972). How-
ever, use of any site depends on a nearby body of
water containing abundant small fishes and on
protection from harassment and predation (Wil-
bur 1974). Fencing, posting and educational
displays, and limited predator control provide
protection for nesting birds (Craig 1971; Massey
1972; Bender 1973, 1974; Atwood et al. 1977).
Suitable feeding areas appear to be in critically
short supply. Protecting those few that remain
and restoring others that no longer support ade-
quate fish populations because of reduced tidal
flow or pollution may be the most significant
management plan for the tern (Wilbur 1974).
Since least terns require open nesting areas,
natural succession tends to eliminate suitable
nesting areas, particularly on man-made sites.
Vegetation removal is advocated to encourage the
continued use of those nesting beaches under-
going succession.
At Bair Island, the substrate is sandy loam
which, when wet, sticks to tern eggs. In 1977,
artificial dry sand mounds about 1 m in diameter
were distributed in the colony as an alternative
nesting substrate (Atwood et al. 1977). Elsewhere,
clay pipes, clay roofing tiles, and special concrete
blocks strategically placed in nesting areas have
been used successfully by chicks for shade and
protection (Atwood et al. 1977).
Continued banding projects will be useful to
determine postbreeding movements to wintering
areas, to understand site-faithfulness, intercolony
aggregations, and age at sexual maturity (Rypka
1977).
A recovery plan has been developed by the
California Least Tern Recovery Team and is
currently being reviewed by the Fish and Wildlife
Service. The plan lists proposed actions for each
colony.
AUTHORITIES
Sanford R. Wilbur
Patuxent Wildlife Research Center
California Field Station
Ojai, California 93023
Alan Craig
California Dept. of Fish and Game
1416 Ninth Street
Sacramento, California 95814
PREPARER'S COMMENTS
It is unclear whether the number of California
least terns is actually increasing or if the apparent
increase is a function of the increased level of
research on this subspecies. Overall trends are
difficult to elucidate from the cursory survey data
that is currently available. Better coordination is
needed to standardize the censusing of colonies.
Least terns are adaptable and even small
colonies could form the basis of substantial new
colonies. Thus all colonies, regardless of their size,
should be protected as much as possible from
human harassment and predation. At present, the
concentration of birds in a few breeding locations
(33% of the total population is at two locations in
San Diego County) makes the future of this
species precarious.
LITERATURE CITED / SELECTED
REFERENCES
Allen, A. 1934. The season: San Francisco region.
Bird Lore 36(4):316.
American Ornithologists. Union. 1957. Check Ust
of North American birds. 5th edition. 691 pp.
Anderson, W. 1970. The California least tern
breeding in Alameda and San Mateo Counties.
California Fish and Game 56(2): 136- 137.
Anonymous. 1970. Least tern. Wandering Tattler
17(9):4.
Atwood, J. L., P. D. Jorgenson, R. M. Jurek, and
T. D. Manolis. 1977. Cahfornia least tern
census and nesting survey, 1977. California
Department of Fish and Game Nongame
Wildlife. Investigations, E-1-1, Job V-2.11,
Job Final Report. 35 pp.
Bancroft, G. 1927. Breeding birds of Scammons
Lagoon, Lower California. Condor 29(1):29-
57.
Bender, K. 1973. California least tern census and
nesting survey, 1974. California Department
of Fish and Game Nongame Wildlife Investi-
gation W-54-R, Progress Report. 47 pp.
. 1974. California least tern population
and nesting survey, 1974. California Depart-
ment of Fish and Game Nongame Wildlife
Investigation W-54-R, Job Final Report. 17
pp.
Bent, A. C. 1921. Life history of North American
gulls and terns. Nat. Mus. Bull. 13.
Burleigh, T. D. and G. H. Lowery, Jr. 1942. An
inland race of Sterna albifrons. Occas. Pap.
Mus. Zool. Louisiana State Univ. 10:173-177.
Chambers, W. L. 1908. The present status of the
least tern in southern California. Condor 10
(6):237.
Chandik, T. and A. Baldridge. 1967. Nesting,
middle Pacific coast region. Audubon Field
Notes 21(5):600-603.
Chase, T. and R. O. Paxton. 1965. Middle Pacific
coast region. Audubon Field Notes 19(5):
574-576.
Craig, A. M. 1971. Survey of California least tern
nesting sites. California Department of Fish
and Game Project W-54-R-4, Job II-5.1. Job
Completion Report. 55 pp.
Davis, M. E. 1968. Nesting Behavior of the least
tern {Sterna albifrons). M. Sc. thesis, Univ.
Cahf., Los Angeles. 72 pp.
1974. Experiments in the nesting behav-
ior of the least tern Sterna albifrons browni.
Proc. Linnaean Soc. N.Y. 72:25-43.
Dawson, W. L. 1924. The birds of California.
South Moulton County, San Diego. 2,162 pp.
De Benedictis, P. and T. Chase, Jr. 1963. Middle
Pacific Coast Region. Audubon Field Notes
17(5):480-483.
DeSante, D., R. Le Valley, and R. Stallcup. 1972.
Middle Pacific coast region. American Birds
26(1):112-118.
Edwards, H. A. 1919. Losses suffered by breeding
birds in southern California. Condor 21(2):
65-68.
Gill, R. Jr. 1977. Breeding avifauna of the south
San Francisco Bay estuary. Western Birds
8:1-12
Grater, R. K. 1939. New bird records for Nevada.
Condor 41(1):30.
Grinnell,J. 1928. A distributional summation of
the ornithology of Lower California. Univ.
Cahf., Publ. Zool. 32(l):l-300.
Grinnell, J. and A. Miller. 1944. The distribution
of the birds of California. Pacific Coast Avi-
fauna 27:1 -608.
Grinnell, J. and M. W. Wythe. 1927. Directory of
the bird-life of the San Francisco Bay region.
Pacific Coast Avifauna 18: 1 - 160.
Griscom, L. 1932. The distribution of bird-life
in Guatemala. Bull. Amer. Mus. Nat. Hist.
64:1-439.
Hagar, J. A. 1937. Least tern studies — 1935 and
1936. Bull. Mass. Audubon Soc. 21(4):5-8.
Hardy, J. W. 1957. The least tern in the Missis-
sippi valley. Publ. of Museum, Mich. State
Univ., Biol. Series l(l):l-60.
Lamb C. C. 1927. Notes on some birds of the
southern extremity of Lower California.
Condor 29(3):155-157.
Leach, H. R. and L. O. Fisk. 1972. At the cross-
roads: a report on California's endangered and
rare fish and wildlife. Calif Dept. of Fish and
Game. 99 pp.
Longhurst, A. R. 1969. The status of an endang-
ered bird {Sterna albifrons) in San Diego
County, 1969. U.S. Bur. Commer. Fish.,
Lajolla. Unpbl. Rep. 7 pp.
McCaskie, G. 1971. Southern Pacific Coast
Region. American Birds 25(5):905-908.
McCaskie, G. and E. A. Cardiff. 1965. Notes on
the distribution of the parasitic jaeger and
some members of the Laridae in California.
Condor 67(6):542-544.
Massey, B. W. 1971. A breeding study of the
California tern. California Dept. Fish and
Game Wildlife Manage. Branch. Admin. Rep.
71-9. 22 pp.
. 1972. Breeding biology of the Califor-
nia least tern. M.Sc. thesis, California State
Univ. at Long Beach. 101 pp.
. 1974. Breeding biology of the California
least tern. Proc. Linnaean Soc. N.Y. 72:1-24.
. 1977. Occurrence and nesting of the
least tern and other endangered species in
Baja California, Mexico. Western Birds 8:
67-71.
Murphy, R. C. 1936. Oceanic Birds of South
America. Macmillan, N.Y. 1,245 pp.
Pentis, A. 1972. Who will save the least tern?
Environment Southwest 446:6-8
Phillips, A., J. Marshall, and G. Monson. 1964.
The birds of Arizona. Univ. Arizona Press,
Tucson. 212 pp.
Pray, R. H. 1954. Middle Pacific coast region.
Audubon Field Notes 8(5):326-327.
Ridgway, R. 1919. The birds of North and Middle
America, part VIII. Bull. U.S. Nat Mus. 50:
1-793.
Rypka, D. D. 1977. California least tern banding
project, 1977. Calif Dept. Fish and Game,
Nongame Wildlife Investigations Project
E-l-l,Job 2. 12, Job Final Report. 7 pp.
Sechrist, E. E. 1915. Least Tern. Oologist 32
(1):18.
Shepardson, D. I. 1909. Notes on the least tern.
Oologist 26(9): 152.
Sibley, C. G. 1952. The birds of the south San
Francisco Bay region. Unpubl. Rep. 42 pp.
(Copy at Oakland Public Museum).
Swickard, D. 1971. The status of the California
least tern at Camp Pendleton, 1971. Camp
Pendleton Marine Corps base. Unpubl. Rep.
30 pp.
. 1973. Status of the least tern at Camp
Pendleton, California. California Birds 3(3):
49-58.
Tompkins, I. R. 1959. Life history notes on the
least tern. Wilson Bulletin 71(4):313-322.
Warner, D. W. and R. M. Mengel. 1951. Notes on
birds from the Veracruz coastal plain. Wilson
Bulletin 63(4):288-295.
Wilbur, S. R. 1974. The literature of the Califor-
nia least tern. Bur. Sport Fish Wildl. Spec. Sc.
Report-Wildlife 175. 18 pp.
Wolk, R. G. 1954. Some Preliminary observations
on the reproductive behavior of the least tern
(Sterna albifrons antillarum Lesson). M.Sc.
thesis, Cornell Univ., Ithaca, N.Y. 42 pp.
ACCOUNT PREPARED/ UPDATED BY
National Fish and Wildlife Laboratory
1300 Spruce Drive
Fort Collins, Colorado 80524
Biological Services Program
FWS/OBS-80/01.21
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
KIRTLAND'S WARBLER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of P'.ndangercd Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.21
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
KIRTLAND'S WARBLER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
KIRTLAND'S WARBLER
Dendroica kirtlandii (Baird)
KINGDOM Animalia
CLASS Aves
ORDER Passeriformes
FAMILY Parulidae
OTHER COMMON
NAMES Jack pine warbler
DATE
Entered into SWIS To be determined
Updates 24 Oct 1977, 10 Jan. 1978
LEGAL STATUS
Federal Endangered (32 FR 4001, 11 Mar
1967; 35 FR 18320, 2 Dec 1970).
States Endangered: Florida, Georgia, Michi-
gjui, North Carolina, South Carolina.
REASONS FOR CURRENT STATUS
Small population size, restricted geographic
distribution, and extremely narrow habitat re-
quirements are the major causes of this species'
precarious position. Prevention of fire and other
forest management practices have eliminated much
suitable breeding habitat. Nest parasitism by the
brown-headed cowbird [Molothrus ater) has also
contributed to past population declines. A pos-
sible, but as yet unknown, factor may be opera-
ting against Kirtland's warbler on the wintering
grounds in the Bahama Islands (KWRT 1976).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The adult male has upper parts of bluish slate
gray, streaked with black, a black mask and white
eye ring, underparts dull yellow, sides streaked
with black, and fuscous wings without bars. The
adult female has upper parts bluish gray with fus-
cous streaks, underparts of pale yellow, and
fuscous speckling on the breast. It is large for a
warbler— 14 to 15 cm long, with a short, stout bill.
Color illustrations appear in Mayfield (1960)
and the various field guides.
RANGE
Kirtland's warbler breeds in the northern
Lower Peninsula of Michigan and winters in the
Bahama Islands (Mayfield 1960). Strays have re-
cently (1977, 1978) been observed near Peta-
wawa, Ontario, Kazabazua, Quebec, and Black
River Falls, Wisconsin (Ryel 1978b). Lane (1975)
observed a male and female near Veracruz, Mexi-
co in 1974. The strays are not believed to be evi-
dence of additional populations (H. Mayfield,
pers. comm.).
RANGE MAP
The known distribution of the Kirtland's
warbler is depicted on the following map (KWRT
1976).
STATES/COUNTIES
Breeding
Michigan Alcona*, Alpena*, Clare*, Crawford,
Iosco, Kalkaska, Montmorency*,
Ogemaw, Oscoda, Otsego*, Presque
Isle*, Roscommon, Wexford.
*historical breeding localities in which the warb-
ler has not been recorded since the 1972 breeding
survey (Ryel 1978a).
HABITAT
This warbler has very specific breeding habitat
requirements, including stands of jack pine {Pinus
banksiana) 2 to 6 m high (8 to 21 years old) inter-
spersed with many small openings, mmimal
ground cover, and little or no hardwoods.The
stands are usually on Grayling sands which drain
very rapidly. Jack pine stands supporting breeding
Kirtland's warblers are usually 30 ha or more in
area. Historically, fire maintained the habitat by
killing mature jack pines and opening up the
forest for natural regeneration. The warbler then
used these immature stands until they beccmie too
old.
The preferred wintering habitat on the Baha-
ma Islands is not known, but Radabaugh (1974)
suggests use of low, broad-leafed scrub, the pre-
vailing form of vegetation.
FOOD AND FORAGING BEHAVIOR
A variety of insects are taken from the ground,
air, or pine foliage. Kirtland's warblers often
hover at the ends of branches as they pluck in-
sects out of the pine needle clusters. They also
eat berries.
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
The ground nest is built of sedges and grasses,
rounded in shape with an inside diameter of 50 to
60 mm. Although a cover is not built, the place-
ment of the nest under a grass tussock provides a
canopy and often a tunnel entrance.
RITUAL REQUIREMENTS
Males call from perches or from the ground.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
A breeding pair requires about 12 ha of young
jack pine habitat for their territory (KWRT 1976).
POPULATION NUMBERS AND TRENDS
In 1951, there were an estimated 432 singing
males; in 1961, 502; and in 1971, 201. In the 7
years since then (1972-78), there have been 200,
216, 167, 179, 200, 218, and 196 singing males
counted on the breeding ground (Mayfield 1975,
Ryel 1978b). In 1974 and 1978, there were unex-
plained declines in singing males (Ryel 1978b).
Mayfield (1953) stated that females and males are
about equal in numbers; thus, the 1978 total for
the species could be estimated at 400 individuals.
REPRODUCTION
They generally lay five eggs, which hatch in
mid-June. Prior to cowbird control, nesting suc-
cess averaged 1.4 fledglings per pair. After the
cowbirds were trapped, the success rate rose to
4 fledglings per pair.
Known distribution of the Kirtland's warbler.
MANAGEMENT AND CONSERVATION
The State of Michigan in 1957 set aside three
tracts of 1,040 ha each (Mayfieid 1963, Radtke
and Byelich 1963). Two were planted with open
stands of jack pine, and all have attracted Kirt-
land's warblers (KWRT 1976).
The Kirtland's Warbler Management Area, es-
tablished by the U.S. Forest Service in 1961,
consists of some 1,620 ha of jack pine forest
which is being managed for the warblers. Manage-
ment activities on these refuges have included
selective cutting, burning, and replanting to
achieve the desired habitat type. The warblers
have nested successfully in each of these areas
(KWRT 1976).
An intensive program to eliminate nest para-
sitism by the brown-headed cowbird was begun in
1972 and has proven immensely successful.
The Kirtland's Warbler Recovery Team
drafted a Recovery Plan (1976) whose primary
objective is to reestablish a wild population
throughout the former range at a minimum level
of 1,000 pairs. Five steps are necessary to reach
the primary objective:
1. Maintain and develop suitable nesting
habitat throughout the former range
2. Protect the species on its wintering
grounds and along the migration route
3. Reduce key factors adversely affecting re-
production and survival
4. Monitor breeding populations to evaluate
responses to management practices and envi-
ronmental changes
5. Reintroduce the species into the Upper
Peninsula of Michigan or in other States to
establish independent, self-sufficient popula-
tions.
AUTHORITIES
John Byelich (Recovery team leader)
P.O.Box 306
Mio, Michigan 48647
George W. Irvine (Recovery Team)
Huron-Manistee National Forest
421 South Mitchell Street
Cadillac, Michigan 49601
Nels Johnson (Recovery Team)
Regional Biologist
Michigan Department of Natural Resources
P.O.Box 128
Roscommon, Michigan 48653
Wesley R.Jones (Recovery Team)
U.S. Fish and Wildlife Service
Federal Building, Fort Snelling
Twin Cities, Minnesota 55111
Harold Mayfieid (Recovery Team)
9235 River Road
Waterville, Ohio 43566
Robert Radtke (Recovery Team)
U.S. Forest Service
633 West Wisconsin Avenue
Milwaukee, Wisconsin 53203
L.H. Walkinshaw
4691 Timberlane Road
Lake Wales, Florida 33853
Richard Winters (Recovery Team)
U.S. Fish and Wildlife Service
301 Manley Miles Building
1405 South Harrison Road
East Lansing, Michigan 48823
PREPARER'S COMMENTS
None.
LITERATURE CITED/SELECTED
REFERENCES
KWRT (Kirtland's Warbler Recovery Team). 1976.
Kirtland's warbler recovery plan. pp.
Lane, J. 1975. Kirtland's warbler in Mexico. Am.
Birds 29(1):144.
Mayfieid, H. F. 1953. A census of the Kirtland's
warbler. Auk 70:17-20.
. 1960. The Kirtland's warbler. Cranbrook
Inst. Sci., Bloomfield Hills, Mich. pp;
. 1962. 1961 decennial census of the Kirt-
land's warbler. Auk 79:173-182,263-268.
-. 1963. Establishment of preserves for the
Kirtland's warbler in the state and national
forests of Michigan. Wilson Bull. 75:216-220.
. 1972. Third decennial census of Kirt-
land's warbler. Auk 89:263-268.
. 1973a. Census of Kirtland's warbler in
1972. Auk 90:684-685.
Statistical Serv. Rep. 167.
. 1973b. Kirtland's warbler census. Am.
Birds 27:950-952.
. 1975. Numbers of Kirtland's warblers.
Jack-Pine Warbler 53:39-47.
Orr, C. D. 1975. 1974 breeding success of the
Kirtland's warbler. Jack-Pine Warbler 53:
59-66.
Radabaugh, B. E. 1974. Kirtland's warbler and its
Bahama wintering grounds. Wilson Bull. 96:
374-383.
Radtke, R., and John Byelich. 1963. Kirtland's
warbler management. Wilson Bull. 75:
208-215.
Ryel, L. A. 1976. 1975 census of Kirtland's warb-
ler. Jack-Pine Warbler 54:2-6.
. 1978a. How to see a Kirtland's warbler.
Birding 10:53-58.
1978b. Kirtland's warbler status, June
Shake, W. F., and J. P. Mattsson. 1975. Three
years of cowbird control: an effort to save the
Kirtland's warbler. Jack-Pine Warbler 53:
48-53.
Walkinshaw, L. H. 1972. Kirtland's warbler— en-
dangered. Am. Birds 26:3-9.
Walkinshaw, L. H., and W. R. Faust. 1974. Some
aspects of Kirtland's warbler breeding biology.
Jack-Pine Warbler 52:64-75.
1975, 1974 Kirtland's warbler nesting
1978. Michigan Dep. Nat. Resour. Surveys
success in northern Crawford County, Michi-
gan. Jack-Pine Warbler 53:54-58.
Zimmerman, D. A. 1956. Jack pine association in
the Lower Peninsula of Michigan: its structure
and composition. Ph.D. thesis, Univ. Mich.,
Ann Arbor.
PREPARED/UPDATED BY
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Rm. 250
Gainesville, Florida 32601
Biological Services Program
FWS/OBS-80/01.22
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
HAWKSBILL TURTLE
.^*v
C*.r.iiiJ
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Knginecrs in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.22
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
HAWKSBILL TURTLE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
SlidelL Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
HAWKSBILL TURTLE
Eretmochelys imbricata Linnaeus
KINGDOM Animalia
CLASS Reptilia
ORDER Testudinata
FAMILY Cheloniidae
OTHER COMMON NAMES carey
DATES
Entered into system To be determined
Updates. . 22 September 1976; 14 February 1977
22 May 1979
LEGAL STATUS
Federal: Endangered (35 FR 8491 ;2 June 1970).
States: Endangered: Florida, Georgia, Mary-
land, Mississippi, New York, Texas.
Protected: Alabama, North Carolina
REASONS FOR CURRENT STATUS
The hawksbill turtle is clearly being extir-
pated from Atlantic and Carribean waters (Carr
and Stancyk 1975). The major cause is exploita-
tion for the shell, but meat and skin are also used.
Eggs are harvested by man and other predators.
Relatively undisturbed beaches are required
for nesting. Increasing development and modifi-
cation of beaches are incompatible with sea turtle
reproduction. For example, illumination of
beaches at night can result in distraction of hatch-
lings away from the sea (McFarlane 1963).
Other causes contribute to low total numbers.
These include increased numbers of snorkelers
and scuba divers who spear turtles for sport and
prize (Carr and Stancyk 1975). Littering of seas,
according to Rebel (1974), may cause fatalities
(e.g. if a plastic bag is mistaken for a jellyfish and
consumed). Also, tremendous predation of hatch-
lings and eggs by camiverous ants, crabs, fishes,
reptiles, birds, and mammals limits recruitment
(Rebel 1974).
PRIORITY INDEX
Not assigned
DESCRIPTION
The hawksbill is a small sea turtle with an
elongated, oval shell with overlapping scutes on
the carapace. The head is relatively small; flippers
have two claws. Barnacles are often found on the
carapace and plastron. General coloration is
brovwi with numerous splashes of yellow, orange,
or reddish-brown on the carapace. The plastron is
yellowish with black spots on intergular and post-
anal scales. Juveniles are black or very dark brown
with light brown or yellow on edge of shell, limbs,
and raised ridges on carapace. Adults are 76 to 89
cm long, and weigh 43 to 75 kg.
It is the only sea turtle with two pairs of pre-
frontal scales on the head and four costal plates
on each side of the carapace.
Illustrated in Carr (1952, 1967), Ernst and
Barbour (1972), Bustard (1973), Rebel (1974),
and Riedman and Witham (1974).
RANGE
The hawksbill is scattered throughout the
world's tropical waters. Distribution in the Atlan-
tic Ocean extends from southern Brazil to Massa-
chusetts.
They nest on scattered islands and shores
generally between 25° latitude north and south.
Some western hemisphere nesting sites include:
the tropical Gulf Coast of Mexico, West Indies,
Bahamas, and scattered beaches off Central and
South America. Continental United States nesting
is limited to infrequent Florida nestings. Maps of
prominent nesting beaches are compiled in Sur-
vival Service Commission (1969).
The sea turtle's range probablyhas not changed
significantly, but numbers have declined consider-
ably. Many nesting beaches have been abandoned
either due to natural disaster (hurricanes, erosion,
etc.), alteration of habitat, or commercial use by
man.
RANGE MAP
Distribution in waters off the continental
United States is illustrated by shading of adjacent
coastal States and counties. Nesting records are
depicted by dots.
STATES /COUNTIES
Alabama: Baldwin, Mobile.
Delaware: Sussex.
Florida: Bay, Brevard, Broward, Charlotte,
Citrus, Collier, Dade, Dixie, Duval,
Flagler, Franklin, Gulf, Hernando,
Hillsborough, Indian River, Jefferson,
Lee, Levy, Manatee, Martin, Monroe,
Nassau, Okaloosa, Palm Beach, Pasco,
Pinellas, St. James, St. Lucie, Santa
Rosa, Sarasota, Volusia, Wakulla,
Walton.
Georgia: Bryan, Camden, Chatham, Glynn,
Liberty, Mcintosh.
Louisiana: Cameron, Iberia, Jefferson, Lafourche,
(Parishes) Plaquemines, St. Bernard, St. Mary,
Terrebonne, Vermilion.
Maryland: Worchester.
Massachusetts: Barnstable, Dukes, Essex, Middle-
sex, Nantucket, Norfolk, Plymouth.
Mississippi: Hancock, Harrison, Jackson.
New Jersey: Atlantic, Cape May, Monmouth,
Ocean.
New York: Nassau, Suffolk.
North
Carolina: Brunswick, Carteret, Currituck, Dare,
Hyde, New Hanover, Onslow, Pender.
Rhode
Island: Newport, Washington.
South
Carolina: Beaufort, Charleston, Colleton,
Georgetown, Horry.
Texas: Aransas, Brazoria, Calhoun, Cameron,
Chambers, Galveston, Jefferson,
Kenedy, Kleberg, Matagorda, Nueces,
Willacy.
Virginia: Accomack, Northampton.
HABITAT
Hawkbills frequent rocky areas, reefs, shallow
coastal areas, lagoons of oceanic islands, and
narrow creeks and passes (Carr 1952). They are
generally found in water less than 20m deep.
Hatchlings are often found floating in masses of
sea plants (Pope 1939).
FOOD AND FORAGING BEHAVIOR
Work by Carr et al. (1966) demonstrates that
United States distribution (shading) and nesting records (dots) for the hawksbill turtle.
although turtles are omnivorous, these prefer
invertebrates. Stomach samples include ectoprocts
of the genera Amthia and Steganoporella, a
hydroid probably of the genus Sertularia, evidence
of remains of sea urchins, and the major food
item appears to be the sponge, Geodia gibberosa
(Carr et al. 1966). Rebel (1974) lists mangrove,
algae, fish, barnacles, mollusks, and jellyfish as
hawksbill food.
They feed on the bottom and forage close to
shores and reefs. Divers have observed them scrap-
ing and chewing at reef faces. They have also been
seen swimming around refuse dumped in shore
waters at Ascension Island (Carr and Stanyck
1975).
SHELTER REQUIREMENTS
Not Known.
Surinam June to July
Seychelles September to November
Ceylon — northwest coast April to May
—southwest coast November to February
(Rebel 1974).
Females nest nocturnally every 2 to 3 years
but several times a season at 2-week intervals.
Clutch size varies, but Carr et al. (1966) reported
an average at Tortuguero to be 160 eggs. Carr and
Main (1973) estimate that incubation lasts 60 days
and believe that one in 1,000 or 10,000 survives
to maturity.
According to Ernst and Barbour (1972)
maturity takes from 3 to 4 years.
Mating takes place in shallow water near the
nesting beaches (Pope 1939). Photographs of
courtship and mating are in Bustard (1973).
NESTING OR BEDDING
Hawkbills use a variety of beach types for
nesting. Carr et al (1966) stated that nearly every
undisturbed deep-sand beach in the tropics may
be visited by nesting females.
Lund (1979) states that nesting occurs ran-
domly. Carr and Stanyck (1975) believe there is
some degree of site preparation, and Deraniyagala
(1939) reports incidents of females returning to
previous nesting sites.
RITUAL REQUIREMENTS
Not Known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not Known.
POPULATION NUMBERS AND TRENDS
Population estimates are not available. Carr
and Stanyck (1975) point out that evidence from
work at Tortuguero indicates a decline in popu-
lation from 1956 to 1970.
REPRODUCTION
The nesting season varies with locality:
Bermuda April to June
British West Indies June to October
Costa Rica May to November
Venezuela May to August
Guyana August
MANAGEMENT AND CONSERVATION
Laws protect the hawksbill to varying degrees
in the following areas: Mexico, Costa Rica, Puerto
Rico, Surinam, French Guiana, Ascension Island,
Trust Territory of the Pacific, Tahiti, Fiji Islands,
Kingdom of Tonp^, Australia, Sarawak, British
Indian Ocean and Seychelle's Islands, Israel
South Africa, and Europa Island (Pritchard 1969;
U.S. Department of Commerce 1976).
Hawksbills are listed under Appendix 1 in the
Convention on International Trade in Endangered
Species of Wild Fauna and Flora. Commercial
trade in the species is subject to strict regulation,
and both an export and import permit are neces-
sary for trade by participating countries.
Marquez (1976) recommends preservation of
several natural reserves along the coast of Mexico.
Among these is Isla Contoy in Quintana Roo
State, a nesting beach for hawksbills.
Carr (personal communication) recommends
that a ban be placed on international traffic and
tourist sale of tortoise shells and protection of all
nesting beaches.
Pritchard (1969) suggested the following con-
servation activites: more research devoted to
studying the missing year (1st year) of the hatch-
lings and developing a practical means of tagging
hatchlings; investigating population dynamics and
the possibility of sustained yield management;
and promoting publicity concerning protection of
sea turtles after population stability is deter-
mined.
Critical Habitat has been proposed for beach-
es on Culebra and Mona Islands, Puerto Rico (43
FR 22224-22225, 24 May 1978).
The species adapts well to captivity and the
chances for captive breeding are are good, but
the cost may be prohibitive (Witham 1976).
AUTHORITIES
Archie Carr
Department of Zoology
University of Florida
Gainesville, FL 32611
George Hughes
Oceanographic Research Institute
P.O. Box 736
Durban,
Natal, South Africa
Frank Lund
P.O. Box 541
Jupiter Island, FL 33458
Bernard Nietschmann
Department of Geology
University of Michigan
Ann Harbor, MI 48104
Peter Pritchard
Florida Audubon Society
P.O. Drawer 7
Maitland,FL 32751
PREPARER'S COMMENTS
None.
LITERATURE CITED/SELECTED
REFERENCES
Bustard, R. 1973. Sea turtles, natural history and
conservation. Taplinger Publ., New York. 220
pp.
Carr, A. F. 1952. Handbook of turtles. Turtles of
the United States, Canada, and Baja Califor-
nia. Comstock Publ. Assoc, Univ. Cornell
Press, Ithaca, N.Y. 542 pp.
. 1967. So excellent a fish. The Natural
History Press, Garden City, N.Y. 248 pp.
. 1972. Great reptiles, great enigmas.
Audubon March: 24-35.
Carr, A. F., H. F. Hirth, and L. Ogren. 1966. The
ecology and migrations of sea turtles, 6. The
hawksbill turtle in the Caribbean Sea. Amer.
Mus. Nov. 2248. 29 pp.
Carr, A. F., and A. R. Main. 1973. Report on an
inquiry into ecological implications of a turtle
farming project. In Turtle farming project in
northern Australia. Union Offset Co., Pty,
Limited, Canberra, Australia. 80 pp.
Carr, A. F., and S. Stancyk. 1975. Observations
on the ecology and survival outlook of the
hawksbill turtle. Biol. Conserv. (8): 161-172.
Deraniyagala, P. E. P. 1939. Tetrapod reptiles of
Ceylon. Vol. 1. Columbo Mus. Publ. Ceylon.
412 pp.
Ernst, C. H., and R. W. Barbour. 1972. Turtles of
the United States. Univ. of Kentucky Press,
Lexington. 347 pp.
lUCN. 1968. Red data book. Vol. 3. Amphibia
and reptilia. Compiled by R. E. Honegger.
lUCN. Morges, Switzerland.
Lund, F. 1979. Atlantic hawksbill. Pages 24-25 in
R. W. McDiarmid, ed., Rare and endangered
biota of Florida, Vol. 3, Amphibians and rep-
tiles. Univ. Presses of Florida, Gainesville.
Marquez M., R. 1976. Reserves naturales para la
conservacion de las tortugas marinas en Mexi-
co. INP/SIi83: 1-22.
McFarlane, R.W. 1963. Disorientation of logger-
head hatchlings by artificial road lighting.
Copeia 1963(1):153.
Montoya, A.E. 1969. Programs de investigacion y
conservacion de las tortugas marinas en Mexi-
co. Pages 34 to 53 in Marine turtles. lUCN
New Publ. Ser. Suppl., Paper 20.
Pope, C. H. 1939. Turtles of the United States
and Canada. Alfred A. Knopf, Inc., New
York. 343 pp.
Pritchard P. C. H. 1969. Summary of world sea
turtle survival situation. lUCN Bull. 2(11):90-
91.
Rainey, W. E., and P. C. H. Pritchard. 1972. Dis-
tribution and management of Ceiribbean sea
turtles. Trans. N. Am. Wildl. Nat. Resour.
Conf. 37:1-17.
Rebel, T. P. 1974. Sea turtles and the turtle in-
dustry of the West Indies, Florida, and the
Gulf of Mexico. Rev. ed. Univ. of Miami
Press, Coral Gables. 250 pp.
Riedman, S. R., and R. Witham. 1974. Turtles: ex-
tinction or survival? Abelard-Schumann New
York. 156 pp.
Schmidt, S., and R. P. Witham. 1961. In defense
of the turtle. Sea Frontiers 7(4):21 1-219.
Survival Serivce Commission. 1969. Marine tur-
tles. Proc. working meeting of marine turtle
specialists organized by lUCN at Morges,
Switzerland.
U.S. Department of Commerce. 1976. Proposed
listing of the green sea turtle {Chelonia my-
das). Loggerhead (Caretta caretta), and pacific
ridley sea turtle {Lepidochelys olivacia) as
threatened species under the Endangered Spe-
cies Act of 1973. Draft EIS. Natl. Oceanic
Atm. Admin., Natl. Marine Fish. Ser., Wash-
ington, D.C.
Walker, W. F. 1971. Swimming in sea turtles of
the family Cheloniidae. Copeia 1971(2):229-
233.
Witham, R. P. 1976. Eretmochelys imbricata ac-
count. In H. O. Hillestad, D. B. Means, and W.
W. Baker, eds. Endangered and threated ver-
tebrates of the southeastern United States.
Tall timbers Res. Stn. Misc. Publ. 4.
Zim, H. S., and H. M. Smith. 1953. Reptiles and
amphibians: A guide to familiar American
species. Simon and Schuster, New York. 147
pp.
Biological Services Program
FWS/OBS-80/01.23
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE INDIANA BAT
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the F.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Kngincers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.23
March 1980
SELFXTKD VER 1 EBRAl K ENDANGERED SPECIES
OF IHE SEACOAST OE 1 HE UNITED STATES-
THE INDIANA BAT
A C<)opcrati\c Effort
by the
National Fish and WiklHfe Laboratory,
the Office of Endangered Species
ant! the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Scr\ ices
Fish and Wildlife Ser\ice
U.S. Department of the Interior
CREDIT: STEVEN R. HUMPHREY
INDIANA BAT
Myotis sodalis Miller and Allen
KINGDOM Animalia
CLASS Mammalia
ORDER Chiroptera
FAMILY Vespertilionidae
OTHER COMMON
NAMES Indiana myotis, social bat,
Kentucky brown bat
DATE
Entered into SWIS to be determined
Updates 28 April 1978
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 March
1977)
States: Endangered: Florida, Georgia, Indiana,
Illinois, Michigan, Mississippi, Mis-
souri, New Jersey, New York, North
Carolina, Ohio, South Carolina, Ten-
nessee, Vermont.
REASONS FOR CURRENT STATUS
Approximately 87% of the entire species
population winters in only seven caves (Humph-
rey 1978). Disturbance or vandalism by man dis-
rupts hibernation and results in deaths and abor-
tions of young (Hall 1902, Humphrey and Scud-
der 1978). Intrusion by vandals, spelunkers, and
biologists have accounted for losses (IBRT 1975).
Hall (1962) discusses clustering behavior of Indi-
ana bats in caves and states that frequent disrup-
tion of congregations could lead to extinction.
Loss of habitat has caused 50% of the decline
over recent years (Humphrey 1978). Some hiber-
nation sites have been rendered unsuitable as a
result of blocking or impeding air flow into the
caves, thus changing the cave's climate (Humphrey
1978). Urbanization and deforestation have con-
tributed to the species decline (Mohr 1972).
Humphrey et al. (1974) found that females breed
in scattered nurseries in trees and suggests that
losses of large amounts of summer habitat could
be catastrophic.
Natural factors such as a low population
growth rate (Humphrey and Cope 1977), floods
(Griffin 1953, DeBlase et al. 1965), freezes
(Humphrey 1978), and cave-ins (Hall 1962) tend
to inhibit recovery.
PRIORITY INDEX
None designated.
DESCRIPTION
Adults are 41 to 49 mm long and weigh 6 to 9
g. The fur is huffy-brown on shoulders and sepia
brown dorsally; underparts are pinkish white.
Wing membrane and ears are blackish brown.
Dark, fuscous brown hairs on and between toes
do not extend beyond the tips of the claws. The
calcar is keeled. FHght-age young lack distin-
guishing markings. Barbour and Davis (1969),
Humphrey (1975, 1977), and IBRT (1975) con-
tain illustrations.
RANGE
Indiana bats range throughout most of the
eastern U.S. from New Hampshire to Iowa to
eastern Oklahoma, and southeast to the pan-
handle of Florida. Their winter range is much
smaller, being restricted primarily to Alabama,
Tennessee, Kentucky, Indiana, Missouri, and West
Virginia.
RANGE MAP
Dots on the following map represent winter
caves; cross-hatched area is summer range (Hum-
phrey 1978).
STATES/COUNTIES
Alabama: Blount, Calhoun, Cherokee, Cullman,
DeKalb,Etowah, Jackson, Limestone,
Madison, Marshall, Morgan.
Arkansas: Baxter, Benton, Carroll, Clark, Clay,
Cleburne, Conway, Craighead, Craw-
ford, Faulkner, Franklin, Fulton, Gar-
land, Greene, Hot Spring, Howard,
Independence, Izard, Jackson, John-
son, Lawrence, Logan, Lonoke, Madi-
son, Marion, Montgomery, Newton,
Perry, Pike, Polk, Pope, Prairie, Pulas-
ki, Randolph, Saline, Scott, Searcy,
Sebastian, Sevier, Sharp, Stone, Van
Buren, Washington, White, Woodruff,
Yell.
Con-
necticut: Fairfield, Hartford, Litchfield, Middle-
sex, New Haven, Tolland, Windham.
Florida: Jackson.
Georgia: Bartow, Catoosa, Chattooga, Chero-
kee, Cobb, Dade, Dawson, Fannin,
Floyd, GUmer, Bordon, Haralson,
Lumpkin, Murray, Paulding, Pickens,
Polk, Towns, Union, Walker, White,
Whitfield.
Illinois: Adams, Bond, Brown, Bureau, Cal-
houn, Carroll, Cass, Champaign, Chris-
tian, Clark, Clay, Clinton, Coles,
Cook, Crawford, Cumberland, DeKalb,
DeWitt, Douglas, DuPage, Edgar, Ed-
wards, Effingham, Fayette, Ford,
Franklin, Fulton, Gallatin, Greene,
Grundy, Hamilton, Hancock, Hardin,
Henderson, Henry, Iroquois, Jackson,
Jasper, Jefferson, Jersey, Jo Daviess,
Johnson, Kane, Kankakee, Kendall,
Knox, Lake, LaSalle, Lawrence, Lee,
Livingston, Logan, McDonough,
McLean, Macon, Macoupin, Madison,
Marion, Marshall, Mason, Menard,
Mercer, Monroe, Montgomery, Mor-
gan, Moultrie, Ogle, Peoria, Perry,
Piatt, Pike, Pope, Putnam, Randolph,
Richland, Rock Island, St. Clare, Sa-
line, Sangamon, Schuyler, Scott,
Shelby, Stark, Stephenson, Tazewell,
Union, VermiHon, Wabash, Warren,
Washington, Wayne, White, Whiteside,
Will, Williamson, Woodford.
Indiana: All counties.
Iowa: Adair, Adams, Appanoose, Cedar,
Clarke, Clayton, Chnton, Dallas, Davis,
Decatur, Des Moines, Dubuque, Fre-
mont, Henry, Iowa, Jackson, Jasper,
Jefferson, Johnson, Jones, Keokuk,
Lee, Louisa, Lucas, Madison, Mahaska,
Marion, Monroe, Montgomery, Musca-
tine, Page, Polk, Poweshiek, Ringgold,
Scott, Taylor, Union, Van Buren, Wa-
pello, Warren, Washington, Wayne.
Kansas: Atchison, Bourbon, Brovm, Cherokee,
Crawford, Doniphan, Jefferson, John-
son, Leavenworth, Linn, Miami,
Wyandotte.
Kentucky: Adair, Allen, Anderson, Barren,
Bath, Bell, Bourbon, Boyd, Boyle,
_
..I
-^ I
1 '
4
^^^^^^^;^^^3'~\
b
-4-~
x \^.y^A^
^
V
Y",
^
^1
Tvs
■ 1
^1
'
\ •■-<y
^-v^VC
^
K /
\
\
■/ INDIANA BAT
1 1 SUMMER
' \
\ /
\ __--A^—
— ,r\
•
WINTER CAVES
)
,^>^^
^
V ■
WINTER CAVES
(CRITICAL
HABITAT)
)
0
500
\
km
Map showing the range, winter caves and critical habitat of the Indiana bat.
Bracken, Breathitt, Breckenridge, Bul-
litt, Butler, Caldwell, Campbell, Car-
roll, Carter, Casey, Christian, Clark,
Clay, Clinton, Crittenden, Cumber-
land, Daviess, Edmonson, Elliott, Es-
till, Fayette, Fleming, Floyd, Franklin.
Gallatin, Garrard, Grant, Grayson,
Green, Greenup, Hancock, Hardin,
Harlan, Harrison, Hart, Henderson,
Henry, Hopkins, Jackson, Jefferson,
Jessamine, Johnson, Kenton, Knott,
Knox, Larue, Laurel, Lawrence, Lee,
Leslie, Letcher, Lewis, Lincoln,
Livingston, Logan, Lyon, McCreary,
McLean, Madison, Magoffin, Marion,
Martin, Mason, Meade, Menifee, Mer-
cer, Metcalfe, Monroe, Montgomery,
Morgan, Muhlenberg, Nelson, Nicholas,
Ohio, Oldham, Owen, Owsley, Pendle-
ton, Perry, Pike, Powell, Pulaski,
Robertson, Rockcastle, Rowan, Rus-
sell, Scott, Shelby, Simpson, Spencer,
Taylor, Todd, Trigg, Trimble, Union,
Warren, Washington, Wayne, Webster,
Whitley, Wolfe, Woddford.
Maryland: Allegany, Carroll, Frederick, Garrett,
Montgomery, Washington.
Massa-
chusetts:
Berkshire, Franklin, Hampshire,
Hampden, Middlesex, Worcester,
Michigan: Allegany, Barry, Berrien, Branch, Cal-
houn, Cass, Clinton, Eaton, Genesee,
Hillsdale, Ingham, Iron, Jackson, Kala-
mazoo, Kent, Lenawee, Livingston,
Ottowa, St. Joseph, Shiawassee, Van
Buren, Washtenaw, Wayne.
Missouri: Adair, Andrew, Atchison, Audrain,
Barry, Barton, Bates, Benton, Bol-
linger, Boone, Buchanan, Butler, Cald-
well, Callaway, Camden, Cape Girar-
deau, Carroll, Carter, Cass, Cedar,
Chariton, Christian, Clark, Clay, Clin-
ton, Cole, Cooper, Crawford, Dade,
Dallas, Daviess, DeKalb, Dent, Doug-
las, Franklin, Gasconade, Gentry,
Greene, Grundy, Harrison, Henry,
Hickory, Holt, Howard, Howell, Iron,
Jackson, Jasper, Jefferson, Johnson,
Knox, Laclede, Lafayette, Lawrence,
Lewis, Lincoln, Linn, Livingston,
McDonald, Macon, Madison, Maries,
Mercer, Miller, Moniteau, Monroe,
Montgomery, Morgan, Newton, Nada-
way, Oregon, Osage, Ozark, Perry,
Pettis, Phelps, Pike, Platte, Polk,
Pulaski, Putnam, Ralls, Randolph,
Ray, Renolds, Ripley, St. Charles,
St. Claire, St. Francois, Ste. Gene-
vieve, St. Louis, St. Louis City, Saline,
Schuyler, Scotland, Scott, Shannon,
Shelby, Stoddard, Stone, Sullivan,
Taney, Texas, Vernon,Warren, Wash-
ington, Wayne, Webster, Worth,
Wright.
Nebraska: Nemaha, Pavmee, Richardson.
New
Hampshire: Cheshire, Grafton, Hillsborough,
Merrimack, Sullivan.
New Jersey: Bergen, Passaic, Sussex, Warren.
New York: Albany, Broome, Cayuga, Chemung,
Chenango, Columbia, Cortland, Dela-
ware, Dutchess, Essex, Franklin, Ful-
ton, Genessee, Hamilton, Herkimer,
Jefferson, Lewis, Madison, Montgo-
mery, Oneida, Onondaga, Orange, Os-
wego, Otsego, Putnam, Rensselaer, St.
Lawrence, Saratoga, Schenectady,
Schoharie, Sullivan, Tioga, Tompkins,
Ulster, Warren, Washington.
North
Carolina: Alleghany, Ashe, Avery, Buncombe,
Burke, Caldwell, Cherokee, Graham,
Haywood, Jackson, McDowell,
Macon, Madison, Mitchell, Surrey,
Swain, Tatauga, Wilkes, Yancey.
Ohio: Adams, Allen, Ashland, Athens, Aug-
laize, Belmont, Brown, Butler, Car-
roll, Champaign, Clark, Clermont,
Clinton, Columbiana, Coshocton,
Crawford, Darke, Defiance, Delaware,
Erie, Fairfield, Fayette, Frankhn, Ful-
ton, Gallia, Greene, Guernsey, Hamil-
ton, Hancock, Hardin, Harrison,
Henry, Highland, Hocking, Holmes,
Huron, Jackson, Jefferson, Knox,
Lawrence, Licking, Logan, Lorain,
Lucas, Madison, Marion, Medina,
Meigs, Mercer, Miami, Monroe, Mont-
gomery , Morgan, Morrow, Muskingum,
Noble, Ottawa, Paulding, Perry, Pick-
away, Pike, Preble, Putnam, Richland,
Ross, Sandusky, Scioto, Seneca, Shel-
by, Stark, Summit, Tuscarawas, Union,
Van Wert, Vinton, Warren, Washing-
ton, Wayne, Williams, Wood, Wyandot.
Oklahoma: Adair, Cherokee, Craig, Delaware,
Haskell, Latimer, Le Flore, Mayes,
McCurtain, Mcintosh, Muskogee, Ot-
tawa, Pittsburg, Pushmataha, Sequoy-
ah, Wagone.
Penn-
sylvania:
Adams, Allegheny, Armstrong, Beaver,
Bedford, Berks, Blair, Bradford, But-
ler, Cambria, Carbon, Centre, Clear-
field, Clinton, Columbia, Cumber-
land, Dauphin, Fayette, Franklin,
Fulton, Greene, Huntingdon, Indi-
ana, Jefferson, Juniata, Lackawanna,
Lancaster, Lawnrence, Lebanon, Lu-
zerne, Lycoming, Mifflin, Monroe,
Montour, Northumberland, Perry,
Pike, Schuylkill, Snyder, Somerset,
Sullivan, Susquehanna, Tioga, Union,
Washington, Wayne, Westmoreland,
Wyoming, York.
Tennessee: Anderson, Bedford, Bledsoe, Blount,
Bradley, Campbell, Cannon, Carter,
Cheatham, Claiborne, Clay, Cocke,
Coffee, Cumberland, Davidson, De-
Kalb, Dickson, Fentress, Franklin,
Grainger, Greene, Grundy, Hamblen,
Hamilton, Hancock, Hickman, Jack-
son, Jefferson, Johnson, Knox, Lin-
coln, Loudon, McMinn, Macon,
Marion, Marshall, Maury, Meigs, Mon-
roe, Montgomery, Moore, Morgem,
Overton, Pickett, Polk, Putnam, Rhea,
Roane, Robertson, Rutherford, Scott,
Sequatchie, Sevier, Smith, Stewart,
Sullivan, Summer, Trousdale, Unicoi,
Union, Van Buren, Warren, Washing-
ton, White, Williamson, Wildon.
Virginia: Albemarle, Alleghany, Amherst, Ap-
pomattox, Augusta, Bath, Bedford,
Bland, Botetourt, Buchanan, Bucking-
ham Campbell, Carroll, Clarke, Craig,
Culpeper, Cumberland, Dickenson,
Fauquier, Floyd, Fluvanna, Franklin,
Frederick, Giles, Goochland, Grayson,
Greene, Henry, Highland, Lee, Lou-
doun, Louisa, Madison, Montgomery,
Nelson, Orange, Page, Patrick, Pittsyl-
vania, Prince William, Pulaski, Rappa-
hannock, Roanoke, Rockbridge, Rock-
ingham, Russell, Scott, Shenandoah,
Smyth, Spotsylvania, Tazewell, War-
ren, Washington, Wise, Wythe.
West
Virginia: All counties.
Wisconsin: Crawford, Grant, Green, Iowa, Lafay-
ette, Richland.
HABITAT
Winter hibernation is restricted to caves with
specific climatic conditions, namely, temperature
4° to 8° C and relative humidity from 66% to 95%
(Barbour and Davis 1969, Humphrey 1978).
These requirements are met only in a narrow zone
close to a cave entrance (Hall 1962).
Sexes and age classes diverge in summer.
Females and juveniles live in stream bank and
floodplain areas, roosting under the bark of trees
(Humphrey et al. 1977). Males use floodplain
ridges and hillside forests and usually roost in
caves (LaVal et al. 1976, 1977).
FOOD AND FORAGING BEHAVIOR
Females and juveniles eat mainly small, soft-
bodied flying insects, primarily Diptera (J.J. Bel-
wood and S. R. Humphrey personal communica-
tion). In early summer, femcdes and juveniles for-
age along stream banks most of the time, and
shift later to include trees and the edges of flood-
plain forests (Humphrey et al. 1977). Foraging
areas average 1.47 ha per animal in early summer,
and 4.54 ha per animal by mid-summer (Humph-
rey et al. 1977). Males forage in densely wooded
areas at tree-top height (LaVal et al. 1976, 1977).
SHELTER REQUIREMENTS
Winter caves have stable, cool temperatures
(usually 4° to 8° C, but sometimes as low as 1° C),
and high humidity (Humphrey and Scudder 1979,
LaVal etal. 1976).
NESTING OR BEDDING
Summer nursery roosts are located under the
bark of live or dead hardwoods, which serves as a
buffer against sudden temperature changes, shel-
ters the bats from weather and predators, and acts
as a solar heat collector (Humphrey et al. 1977).
RITUAL REQUIREMENTS
The bats swarm near caves from August to
November in Missouri and from mid-August to
the end of October in Indiana and Kentucky
(Cope and Humphrey 1977). Swarming is des-
cribed as a phenomenon in which large numbers
of bats fly in and out of a cave entrance from
dusk to dawn, while relatively few roost in the
cave during the day (Humphrey and Cope 1976).
The significance of this activity is not known; it
has been suggested that it relates to mate selec-
tion and copulation, adjustment to and familiari-
zation with seasonal ranges, and/or rest stops
during migration (IBRT 1975, M. D. Tuttle per-
sonal communication).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
It is possible that in some instances, the gray
bat (My Otis grisescens), another Federally hsted
endangered species, may compete for roosting
space in winter caves (Hall 1962) and could pos-
sibly displace the Indiana bat from them.
POPULATION NUMBERS AND TRENDS
The recent discovery of two new winter caves
has brought estimates of total population to
509,000 (Richter et al. 1978). The bat once
numbered 1 million, but major catastrophies,
primarily flooding, killed many in wintering caves
(Hall 1962, IBRT 1975).
Humphrey (1978) recorded a 28% decline in
total numbers from 1960 to 1975 (Table 1).
Table 1. Population estimates and trends for the
Indiana bat, by state
1960
1975
Trend
State
population
population
percentage
Missouri
311,433
285,983
-8.2
Kentucky
209,796
55,782
-73.4
Tennessee
7,554
Arkansas
1,700
—
Indiana
102,823
104,824
+1.9
Illinois
339
194
-42.8
Virginia
580
—
West Virginia
1,757
—
Pennsylvania
1,002
—
New York
.....
500
—
Vermont
,
. .not censused. .
Totals
625,393
458,874
-26.6
The decline in Kentucky was due to flooding,
disturbance, and man-caused alteration of cave
climate. Humphrey (1976) believs the Indiana bat
is headed for extirpation in Kentucky. Missouri
reductions are a result of winter weather variation.
Incidents of vandalism and the collapse of a mine
contributed to the decline in Illinois (Mohr 1962,
Humphrey 1978). The overall trend at present
will lead to the species' extinction in 50 years,
but most losses of winter habitat are reversible
and this eventuality can be avoided (Humphrey
1978).
REPRODUCTION
Copulation occurs in the fall with the females
storing the sperm over the winter (Barbour and
Davis 1969). The females form nursery colonies
under the bark of trees in late spring (usually
May) and give birth to one young in late June or
early July (Humphrey et al. 1977). Only females
and young occupy the nursery roost (Humphrey
et al. 1977). The longevity record for males is
13.5 years; for females, 14.8 years. The survival
rate for females is high for the first 10 years,
and for males, for the first 6 years only (Hum-
phrey and Cope 1977).
MANAGEMENT AND CONSERVATION
The primary conservation effort to date has
been to control human access to caves by the
use of gates. The National Park Service, the U.S.
Forest Service, and some State agencies have
installed gates on several caves. The Recovery
Team suggests the gating of several more caves
and mines, which will give protection to 99% of
the bats (IBRT 1975). In one instance (Colossal
Cave, Kentucky), a gate modified cave micro-
climate and made it unsuitable for bats (Hum-
phrey 1978). Gates or fences have also increased
the bats' susceptibility to predation at the en-
trances (Tuttle 1977). Other conservation efforts
include a moratorium on issuance of bat bands by
the U.S. Fish and Wildlife Service and restrictions
on use of pesticides such as DDT. The National
Speleological Society has appointed a Bat Con-
servation Task Force and has taken some specific
measures to protect bats.
The original Recovery Team (since disbanded)
outlined a Recovery Plan (IBRT 1975) listing
three objectives critical to preservation of the
species: (1) protecting winter habitat by securing
caves and mines and restricting entry; (2) initi-
ating a public information and education pro-
gram; and (3) monitoring population levels and
habitat quality. A new Recovery Team is being
formed (J. M. Engel personal communication).
Other management needs include protection
of the summer riparian environment (Martin
1973, Humphrey 1978).
The majority of the species winters in public-
ly owned caves, but many private caves are also
used (R. K. LaVal personal communication).
Public aquisition of certain of the latter, including
four in Missouri and one in West Virginia, is being
sought at present (R. LaVal personal communica-
tion, J. Rawson personal communication).
Table 2 lists pertinent information on pub-
licly owned wintering caves.
Dr. Tom Kunz (Recovery Team)
Department of Biology
Boston University
2 Cummington
Boston, Massachusetts 02215
Dr. Don Wilson (Recovery Team)
National Fish and Wildlife Laboratory
National Museum of Natural History
Washington, D.C. 20560
Dr. Merlin Tuttle (Recovery Team)
Vertebrate Division
Milwaukee Public Museum
Milwaukee, Wisconsin 53233
AUTHORITIES
J. T. Brady (Recovery Team)
Biologist
U.S. Army Corps of Engineers
210 N. 12th Street, Room 853
St. Louis, MO 63101
J. B. Cope
Joseph Moore Museum
Earlham College
Richmond, IN 47374
J. M. Engel
U.S. Fish and Wildlife Service
Federal Building, Fort Snelling
Twin Cities, MN 55111
S. R. Humphrey
Florida State Museum
Museum Road
Gainesville, FL 32611
R. K. LaVal (Recovery Team Leader)
Missouri Department of Conservation
Fish and Wildlife Research Center
1110 College Avenue Columbia,
Columbia, MO 65201
R. Z. Mumford
Dept. of Forestry and Conservation
Purdue University
Lafayette, IN 47904
A. Richter
Joseph Moore Museum
Earlham College
Richmond, IN 47374
PREPARER'S COMMENTS
None.
Table 2. Ownership of caves, Critical Habitat, and
recent (1975) population estimates for Indiana bat
caves (Humphrey 1978, Richter et al. 1978).
[F=Federal, P=Private, S=State, CH=Critical Habitat.]
State
County
Cave
Owner-
ship
Population
estimate
Alabama
?
Santa
F
?
Several caves F
700
in Sylamore
Forestry Dist.
Madison
Denney
?
1,000
Illinois
LaSaUe
Blackball
Mine
(CH), S
192
?
No. Ill
S
50,000
?
No. 175
S
1,250
?
No. 376
S
100,000
Crawford
Wyandotte
(CH), S
1,460
Crawford
Saltpeter
S
95
Greene
Ray's
(CH),P
2,700
Kentucky Carter
Bat
(CH), S
40,000
Edmonson
Coach
(CH), P
4,500
Edmonson
Dixon
F
3,600
Edmonson
Long's
F
7,600
Edmonson
Colossal
F
14
Edmonson
Wilson
F
0
Edmonson
Bat
F
68
?
Carter
S
Missouri
Iron
PUot Knob
(CH), P
Shannon
No. 047
(CH),)
46,000
Washington No. 029
(CH), S
81,800
Crawford
No. 021
(CH). F
12,850
Franklin
No. 009
(CH), S
21,000
Franklin
No. 017
S
Camden
No. 053
S
Tennessee Blount White Oak (CH), F
Blowhole
6,050
? Nicajack F
Anderson Norris Dam F
West
Virginia Pendelton Hellhole (CH), P
1,500
Tucker Big Springs F
(Blowing Cave)
150
Tucker Cave Hollow F
23
lucifugus, in Indiana and north-central Ken-
tucky. Am. Soc. Mammal. Spec. Publ. 4:1-81.
1977. Survival rates of the endangered
Per 42 FR 40687, 11 August 1977.
Tuttie, personal communication, 1978.
UTERATURE CITED/SELECTED
REFERENCES
Barbour, R. W., and W. H. Davis. 1969. Bats of
America. Univ. Press, Kentucky, Lexington.
286 pp.
Cope, J. B., and S. R. Humphrey. 1977. Spring
and autumn swarming behavior in the Indiana
hdLt,Myotis sodalis. J. Mammal. 58:93-95.
DeBlase, A. F., S. R. Humphrey, and K. S. Drury.
1965. Cave flooding and mortality in bats in
Wind Cave, Kentucky. J. Mammal. 46:96.
Engel,J.M. 1976. The Indiana bat, Afyofw sodalis,
a bibliography. U.S. Fish Wildl. Serv., Spec.
Sci. Rep. 196. 11 pp.
Griffin, D. G. 1953. Deluge underground. Bull.
Natl. Speleol. Soc. 15:34-37.
Hall, J. S. 1962. A life history and taxonomic
study of the Indiana bat, Myotis sodalis.
Reading Pub. Mus. and Art Gallery Sci. Publ.
12:3-68.
1972. The status oi Myotis sodalis, the
West Virginia Dep. Natur. Resour. Elkins.
5 pp.
Hall, E. R., and K. R. Kelson. 1959. The mam-
mals of North America. Vol. 1. Ronald Press,
New York. 546 pp.
Humphrey, S. R. 1975. Cover photo. Bioscience
25(9).
1977. Keep 'em flying. Animal Kingdom
89(5).
1978. Status, winter habitat, and manage-
ment of the endangered Indiana bat,
Myotis sodalis. Q. J. Florida Acad. Sci.
41(2):65-76.
Humphrey, S. R., and J. B. Cope. 1976. Popula-
tion ecology of the little brown bat, Myotis
Indiana bat, Myotis sodalis. J. Mammal. 58:
32-36.
Humphrey, S. R., J. B. Cope, A. R. Richter, and
P. P. Humphrey. 1974. Summer ecology of
the endangered Indiana bat. Fifth Am. Symp.
Bat Res., Texas Tech. Univ., Lubbock.
Humphrey, S. R., A. R. Richter, and J. B. Cope.
1977. Summer habitat and ecology of the en-
dangered Indiana bat, Myotis sodalis. J. Mam-
mal. 58:334-346.
Humphrey, S. R., and S. J. Scudder. 1979. Indi-
ana bat. Pages 3-4 in J. N. Layne, ed. Rare
and endangered biota of Florida, Vol. 1,
Mammals. Univ. Presses of Florida, Gaines-
ville.
IBRT (Indiana Bat Recovery Team). 1975. Reco-
very plan for the Indiana bat. Draft. U.S. Dep.
Interior, Fish Wildl. Serv. 34 pp.
LaVal, R. K., R. L. Clawson, W. Caire. L. R. Win-
gate, and M. L. LaVal. 1976. An evaluation of
the status of Myotine bats in the propsed
Meromec Park and Union Lake project areas,
Missouri. School of Forestry, Fisheries, and
Wildlife, Univ. of Missouri, Columbia. 136 pp.
LaVal, R. K., R. L. Clawson, M. L. LaVal, and W.
Caire. 1977. Foraging behavior and nocturnal
activity patterns of Missouri bats, with em-
phasis on the endangered species Myotis grises-
cens and Myotis sodalis. J. Mammal. 58:592-
599.
Martin, R. L. 1973. The current status of bat pro-
tection in the United States of America.
Period. Biol. 75:153-54.
Mohr, C. E. 1972. The status of threatened spe-
cies of cave-dwelling bats. Bull. Natl. Speleol.
Soc. 34:33-37.
Mumford, R. E. 1974. The status of the Indiana
bat and the eastern wood rat on the Wayne-
Hoosier National Forest, Indiana. Purdue
Univ. Dep. Forestry and Conserv. Spec. Study.
Richter, A. R., D. A. Seerley, J. B. Cope, and J.
H. Keith. 1978. A newly discovered concen-
tration of hibernating Indiana bat, Myotis so-
dalis, in southern Indiana. J. Mammal. 59: 191.
Tuttie, M. D. 1977. Gating as a means of protec-
ting cave dwelling bats. Pages 77-82 in Na-
tional Cave Management Symposium Proc.
1976. Speleobooks, Albuquerque. 106 pp.
Biological Services Program
FWS/OBS-80/01.24
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE ATLANTIC SALT MARSH SNAKE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.24
March 1980
SELFCl KD VKR TEBRAl i: ENDANGERED SPECIES
OF IHE SEACOAS T OF 1 HE UNITED STATES-
THE ATLANTIC SALT MARSH SNAKE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Ser\'ices
Fish and Wildlife Service
U.S. Department of the Interior
ATLANTIC SALT MARSH SNAKE
Nerodia fasciata taeniata Cope
KINGDOM Animalia
CLASS ReptUia
ORDER Squamata
FAMILY Colubridae
OTHER COMMON
NAMES salt water snake
eastern Florida water snake
east coast striped water snake
DATE
Entered into SWIS to be determined
Updates 3 January 1978, 11 October 1978,
21 May 1979
LEGAL STATUS
Federal: Threatened
States:
(42 FR
29 November 1977)
Endangered: Florida
60743-60745,
REASONS FOR CURRENT STATUS
This species has a Umited geographical range
and is restricted to habitat that is subject to pro-
gressive disturbances. Development on coastal
barrier islands threatens the snake's remaining
habitat. Draining and diking operations promote
hybridization and genetic swamping by an adja-
cent freshwater race, the Florida water snake
Nerodia fasciata pictiventris (Kochman and
Christman 1979a; 42 FR 60743-60745; 29
November 1977).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The Atlantic salt marsh snake is a small water
snake approximately 70cm long. Its dorsal pat-
tern is variable, combining elements of blotching
and longitudinal striping. It has a median pair of
dark brown stripes on pale olive ground color,
often fragmented posteriorly into longitudinal
series of blotches. Its sides have a row of dark
blotches that may merge to form short stripes in
the neck region. The belly is reddish-brown to
black with a median row of yellowish spots.
Color photograph is in Dodd (1978); black
and white photographs in Carr and Goin (1942),
Wright and Wright (1957) and Kochman and
Christman (1979a).
RANGE
This species is endemic to the following
localities on the Atlantic coast of Florida:
Volusia County - vicinity of National Gardens,
Daytona Beach, and New Smyrna Beach
Brevard County — Merritt Island National
Wildlife Refuge at Playalinda Beach (R.
Demmer personal communication) and vici-
nity of Micco (Kochman and Christman
1979a)
Indian River County - Gifford (H. Kochman
unpublished data) and vicinity of Vero Beach
(Kochman and Christman 1979a).
Snakes exhibiting various degrees of intermed-
iacy between the Atlantic salt marsh snake, the
Florida water snake, and the mangrove water
snake (Nerodia fasciata compressicauda) occur at
various localities throughout the known range of
N.f. taeniata (Neill 1958,Kockman 1977).
There is no evidence to suggest that the range
of this species has changed significantly (42 FR
60743-60745; 29 November 1977).
RANGE MAP
Dots on the following map represent specific
localities or general areas from which one or more
specimens have been taken (adapted from Koch-
man and Christman 1979a).
STATES/COUNTIES
Florida: Brevard, Indian River, Volusia.
HABITAT
The Atlantic salt marsh snake inhabits coastal
salt marshes and mangrove swaimps. It has been
specifically reported along shallow tidal creeks
and pools in association with glasswort {Salicornia
perennis) (Carr and Goin 1942), blackrush {/un-
cus roemerianus) (Niell 1958), and black man-
grove (Avicennia germinans) (H. Kale personal
communication, H. Kochman unpublished data).
It occurs in saline environments ranging from
brackish to full strength seawater.
FOOD AND FORAGING BEHAVIOR
The species is ecologically and behaviorally
similar to the Gulf salt marsh snake (N. f. clarki).
It forages in shallow water during low tidal stages
(Carr and Goin 1942, Neill 1958, Kochman and
Christman 1979a), apparently feeding upon dense
congregations of small fishes that become entrap-
ped during the falling tide (Neill 1958). A similar
feeding strategy has also been reported for the
Gulf salt marsh snake (Pettus 1956).
Although primarily regarded as nocturnal
(Carr and Goin 1942, Kochman and Christman
1979a), Atlantic salt marsh snakes may be strong-
ly influenced by ddal rhythms and have been
reported to forage in daylight during favorable
tides (Neill 1958). They are also known to enter
minnow traps in shallow water (H. Kale personal
communication) .
SHELTER REQUIREMENTS
Little specific information is available. One
specimen reportedly withdrew into a fiddler crab
(Uca) burrow when approached during daylight
(Carr and Goin 1942).
NESTING OR BEDDING
Not known.
RITUAL REQUIREMENTS
Not knowTi.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Coastal races of N. fasciata are reproductively
compatible with adjacent freshwater populations
and require ecological isolation to maintain
genetic integrity (Pettus 1956, 1963; Conant
1975, Kochman 1977). The Atlantic salt marsh
snake appears to hybridize freely with the Florida
water snake in ecotonal areas of distributional
overlap (Kochman 1977). It can remain genetical-
ly distinct only if hybridization is limited to
narrow zones of ecological transition between
freshwater and saline habitats. Alteration of
coastal wetlands through drainage and diking
enhances hybridization and promotes genetic
introgression, i.e. swamping, by the Florida water
snake (Kochman and Christman 1979a; 42 FR
60743-60745; 29 November 1977).
POPULATION NUMBERS AND TRENDS
No population estimates are available. Ap-
proximately 50 specimens are known to science
(H. Kochman unpublished data).
Map indicating the range of the Atlantic salt marsh snake along the eastern coast of Florida. Dots
represent sites where specimens have been collected.
REPRODUCTION
Very little information is available. One cap-
tive female gave birth to nine young (eight alive
and one stillborn) in late August (H. Kochman
unpublished data). Their reproductive biology is
probably similar to the Gulf salt marsh snake:
2 to 14 live young bom during midsummer
(Kochman and Christman 1979a, 1979b), with an
average litter size of 6 to 7 (H. Kochman unpub-
lished data).
MANAGEMENT AND CONSERVATION
Portions of their known range include State
and Federal lands along the Atlantic coast of
Florida: Tomoka State Park (Volusia County),
Merritt Island National Wildlife Refuge (Volusia
and Brevard Counties), Cape Kennedy Air Force
Station (Brevard County), Patrick Air Force Base
(Brevard County), and the Pelican Island
National Wildlife Refuge (Indian River County).
In a letter to the U.S. Fish and Wildlife Ser-
vice, Office of Endangered Species (December
1976), H. Kochman emphasized that habitat alter-
ation and subsequent hybridization pose the
chief threats to the Atlantic salt marsh snake. In
view ofM /a5aa to 's high variability along Florida's
Atlantic coast, it was concluded that conservation
measures must be oriented toward safeguarding
habitats and populations rather than a specific
phenotype.
The designation of Critical Habitat in coastal
areas of Volusia, Brevard and Indian River coun-
ties has been recommended by representatives of
various Federal, State and private organizations
(42 FR 60743-60745; 29 November 1977).
No formal advisory committee or recovery
team has been established.
AUTHORITIES
Archie F. Can-
Department of Zoology
University of Florida
Gainesville, FL 32611
Richard Demmer
District V Naturalist
Division of Recreation and Parks
Florida Department of Natural Resources
Route l,Box 107AA
Clermont, FL 32711
Howard I. Kochman
National Fish and Wildlife Laboratory
412 N. E. 16di Avenue, Room 250
GainesvUle, FL 32612
PREPARER'S COMMENTS
The distribution and status of the Atlantic
salt marsh snake remain essentially unknown.
Further study may extend its knovsm home range
northward and southward along the Atlantic
coast of Florida. Potentially suitable habitat
should be surveyed for undiscovered populations,
followed by designation of Critical Habitat in ap-
propriate areas. A special interagency advisory
committee with recovery team functions should
be established to evaluate alternatives for effec-
tive conservation and management.
Water snakes from scattered localities along
the Gulf coast may exhibit a pattern of striping
and spotting similar to the Atlantic salt marsh
snake (Conant 1975). In many instances, this is
the result of hybridization between the longi-
tudinally striped Gulf salt marsh snake and ad-
jacent cross-banded races (Pettus 1956, 1963). It
has been suggested that the Atlantic salt marsh
snake may likewise be the product of hybridi-
zation dating back to Pleistocene contact of
striped and banded populations (Kochman 1977,
Kochman and Christman 1979a). Dunson (in
preparation) does not consider the Atlantic salt
marsh snake sufficiently distinct from the man-
grove water snake to warrant subspecific status.
UTERATURE CITED/SELECTED
REFERENCES
Carr, A. F., and C. J. Coin. 1942. Rehabilitation
of Matrix sipedon taeniata Cope. Proc. New
England Zool. Club 21:47-54.
Conant, R. 1975. A field guide to reptiles and am-
phibians of eastern and central North America.
Houghton Mifflin Co., Boston 429 pp.
Dodd, C. K., Jr. 1978. Amphibians & reptiles, the
declining species. Water Spectrum 10(1):24-
32.
Dunson, W. A. Occurrence of partially striped
forms of the mangrove snake Nerodia fasciata
compressicauda Kennicott and comments on
the status of N. f. taeniata Cope. Unpubl. MS.
Kochman, H. I. 1977. Differentiation and hybri-
dization in the Matrix fasciata complex (Rep-
tilia:Serpentes): A nonmorphological ap-
proach. M.S. Thesis, University of Florida,
Gainesville. 105 pp.
Kochman, H. I., and S. P. Christman. 1979a. At-
lantic salt marsh snake. Pages 27-28 in R. W.
McDiarmid, ed. Rare and endangered biota of
Florida, Vol. 3, amphibians and reptiles. Univ.
Presses of Florida, Gainesville.
. 1979b. Gulf salt marsh snake. Pages 62-
63 in R. W. McDiarmid, ed. Rare and endan-
gered biota of Florida, Vol. 3, amphibians and
reptiles. Univ. Presses of Florida, Gainesville.
Neill, W. T. 1958. The occurrence of amphibians
and reptiles in saltwater areas, and a bibliog-
raphy. Bull. Mar. Sci. Gulf and Caribb. 8(1):
1-97.
Pettus, D. 1956. Ecological barriers to gene ex-
change in the common water snake {Matrix
sipedon). Ph. D. Dissert. Univ. of Texas, Aus-
tin. 87 pp.
. 1963. Salinity and subspeciation in Nat-
trix sipedon. Copeia 1963(3) :499-504.
Wright, A. H., and A. A. Wright, 1957. Handbook
of snakes of the United States and Canada.
Vol. 1. Comstock PubUshing Associates, Ith-
aca. 564 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.25
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE DUSKY SEASIDE SPARROW
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.'38
u
FWS/OBS-80/01.25
March 1980
SELtXIKD VI:R I EBRA 1 1: ENDANGERED SPECIES
OF I HE SEAC:OAS 1 OF 1 HE UNITED STA TES-
THE DUSKY SEASIDE SPARROW
A Cooperative P^ffort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
DUSKY SEASIDE SPARROW
Ammospiza maritima nigrescens Ridgway
KINGDOM Animalia
CLASS Aves
ORDER Passeriformes
FAMILY FringUIidae
OTHER COMMON
NAMES black and white shore finch,
black shore finch.
DATE
Entered into SWIS To be determined.
Updates 10 October 1978.
LEGAL STATUS
Federal: Endangered (32 FR 4001; 11 March
1967).
States: Endangered: Florida.
REASONS FOR CURRENT STATUS
The dusky seaside sparrow is distributed in an
extremely limited area in northern Brevard
County, Florida. It is adapted to narrow, un-
stable zones of vegetation within salt marshes.
Diking of the marshes for mosquito control has
altered the vegetation and the species has disap-
peared on northern Merritt Island. Wildfires and
marsh drainage for housing, roads and pasture
have reduced available habitat in the St. Johns
River Basin (Sharp 1968, 1970; Baker 1973,
1976, in press; USDI 1973; DSSRT draft). Aerial
spraying in coastal marshes with DDT and other
insecticides to control mosquitoes from 1942 to
1953 may have been responsible for a 70% reduc-
tion in the population (Trost 1968).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The dusky seaside sparrow is about 15 cm
long. Adults have black or blackish brown upper
parts, edged with grayish olive, white venter with
heavy black streaking, bright yellow on lores and
at bend of wing. The wings and tail are fuscous to
fuscous black, edged with light yellowish olive.
The sexes are similar.
Juvenal plumage is lighter in color with nar-
rov/er tan streaking, and reduced yellow on lores
and wings.
Color plates appear in Howell (1932), Peter-
son (1947), Sprunt (1954), and Trost (1968).
A black and white photograph appears in Eber-
hart(1968).
RANGE
The dusky seaside sparrow is a sedentary
inhabitant of brackish marshes and savannahs in
northern Brevard County, Florida. The 1978
survey of singing males documented their pre-
sence in the St. Johns River Basin at the following
three locations: St. Johns National Wildlife
Refuge, approximately 1 km south of the refuge,
and between the north and south forks of the
BeeHne Highway (Baker in press). The sub-
species is isolated from other populations of sea-
side sparrow by 120 km to the north, 200 km to
the west, and 400 km to the south (Trost 1968).
Dusky seaside sparrows were formerly abun-
dant in Indian River salt marshes on Merritt Is-
land, from Dummitt Creek south to Banana Creek
(Trost 1968). Sharp (1968) determined the main-
land distribution to be from Salt Lake south near-
ly to Florida Highway 520 and between 1-95 and
the St. Johns River.
RANGE MAP
Critical habitat and 1977 range are indicated
on the following page. The dusky seaside sparrow
apparently no longer occurs on Merritt Island (J.
Baker, personal communication).
STATES/COUNTIES
Florida: Brevard.
HABITAT
The species inhabits salt marshes with highly
fluctuating water levels and salinities. It prefers
moist zones dominated by cordgrass {Spartina
bakerii), 3 to 5 m above mean sea level. The St.
Johns River marshes present a savannah-like
aspect, being dotted with small ponds, salt pans,
cabbage palms (Sabal palmetto), and hammocks
(Sharp 1968, 1970; Baker 1976;DSSRT draft).
Sharp (1968, 1969a, 1970) believes zones of
short and tall interdigitating plants form the pre-
ferred microhabitat. The Merritt Island habitat
was composed of a heterogeneous mosaic of the
tall cordgrass and black rush (Jiincus roemerianus)
with short saltgrass {Distichlis sptcata), saltwort
{Batis maritima) and glasswort {Salicornia peren-
nis). In the St. Johns River savannahs, fires and
varying salinities and water levels produce the tall-
short pattern with cordgrass.
Black and white habitat photographs may be
found in the following: diked salt marsh-Eberhart
(1968); former Merritt Island habitat— Baynard
(1914) and Nicholson (1928); St. Johns River sa-
vannah—Sharp (1969b).
FOOD AND FORAGING BEHAVIOR
The subspecies is largely insectivorous. Six
stomachs contained 37% grasshoppers and crick-
ets and 25% spiders. Other items included miscel-
laneous insects, seeds, and tubers (Howell 1932).
Trost (1968) observed duskies feeding on small
snails, a dragonfly larva, a butterfly, and possibly
ants.
They forage at or near the ground (Baker
(1976, in press). Feeding has been observed on
dikes and in tidal zone (Trost 1968).
SHELTER REQUIREMENTS
The dusky sparrow remains concealed in
dense vegetation during the nonbreeding season
(Trost 1968). Glasswort on Merritt Island (Bay-
nard 1914) and cordgrass in the St. Johns flood-
plain (Sharp 1968, Baker 1973) form the pre-
ferred shelter.
NESTING OR BEDDING
The dusky seaside sparrow constructs a cupped
nest from grasses. Nests are placed in tussocks
2 to 35 cm above ground and may be covered
with arched vegetation (Nicholson 1928).
Nest sites include tussocks of glasswort, black
rush, cordgrass, salt grass, and wire grass (Sporo-
bolus virginicus) (Baynard 1914; Nicholson 1928,
1929; Trost 1968). Baker (in press) reported a
nest in a salt bush (Bacharis angusti folia).
Baynard (1914) and Nicholson (1928) present
black and white photographs of nests and nest
sites on Merritt Island.
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RITUAL REQUIREMENTS
During the breeding season males sing from
prominent perches atop the glasswort, grasses, or
rushes. A male occasionally will flutter slowly
upward 7 to 12 m and then descend to its perch
while singing. Males chase females in low erratic
flights over the tops of vegetation (Nicholson
1928,Trost 1968).
OTHER CRITICAL ENVIRONMENTAL *
REQUIREMENTS
Nesting is confined to a male's territory,
which is defended by song and occasional chasing.
Occupied nests were found within 12 m of one
another in the 1930's, but territories appear to
have increased to 100 m in diameter in recent
years, due to thinning of marsh vegetation and re-
duction in bird densities. Banding has shown that
males defend the same territory year after year.
Banded juveniles have returned a year later and
established territories 300 m from where they
were fledged. Birds may fly 200 m or more from
nest sites to feeding areas (Trost 1968).
Predation appears to occur mainly on eggs
and young mostly by rats (Oryzomys palustris,
Sigmodon hispidus), raccoons [Procyon lotor), and
many snakes (Lampropeltis getulus, Agkistrodon
piscivorous). Fish crows (Corvus ossifragus) and
boat-tailed grackles [Casstdix mexicanus) may rob
nests. Ants may force adults to abandon nests,
especially during high water (Nicholson 1928;
Trost 1968). Nesting red-winged blackbirds
[Agelaius phoeniceus) constantly harass duskies
which nest nearby (DSSRT draft).
The dusky occupies a median position on the
marsh moisture gradient between the 3 and 5 m
contour lines. Above this elevation, the drier
marsh has woody growth and is vulnerable to
wildfire, and the eastern meadowlark [Sturnella
magna) replaces the dusky. In lower areas with
standing water and/or dense cordgrass, least
bitterns (Ixobrychus exilis), king rails (Rallus
elegans) and probably black rails [Laterallus
jamaicensis) occur. The dusky prefers patches of
cordgrass with heights of 0.5 to 1.5 m and a
density of approximately 1,700 stems/m^ (Sharp
1968, 1970).
Natural fires seem to play a beneficial role in
maintaining dusky habitat along the St. Johns
River. Most lightning fires occur in the rainy
season when the marsh is wet and humidity high;
they move slowly and burn small, discontinuous
areas, removing woody plants and allowing the
tall and short cordgrass savannah required by the
duskies to regenerate (Sharp 1968, Baker in
press). Birds in the immediate vicinity are dis-
placed, but presumably recolonize afterward.
POPULATION NUMBERS AND TRENDS
Dusky sparrow populations were probably
stable until the 1940's, when Nicholson estimated
that aerial insecticide spraying from 1942 to 1953
reduced numbers 70% by 1957 (Trost 1968). De-
creases may have resulted from reproductive
failure and disruption of the food chain. Sharp
(1968) estimated that there were 2,000 pairs on
Merritt Island prior to spraying, based on his ob-
served densities and the reports of Baynard
(1914) and Nicholson (1928). Impoundment of
the island marshes in the mid-1950's and resulting
changes in salt marsh vegetation reduced popula-
tions to four localities having 70 pairs in Trost's
1961-1962 study (Sharp 1968). Sharp (1970)
found 33 or 34 males in his 1968 survey. Baker
(in press) located two males at the north end of
Merritt Island in 1977 and none in the 1978 sur-
vey.
Sharp (1970) found 372 singing males in the
St. Johns River marshes in the spring of 1968 and
estimated the probable total of the mainland
population to be 894 pairs. The St. Johns Nation-
al WUdlife Refuge surveys for 1970 and 1972-
1978 documented 143, 110, 54, 37, 47, 11, 12,
and 9 singing males, respectively. Winter wildfires
apparently caused the most drastic reductions.
Singing males in the entire St. Johns Basin num-
bered 28 in 1977 and 24 in 1978; no females
were seen in 1978, although some were seen in
1977 (Baker, in press).
REPRODUCTION
Dusky sparrows breed from March to August,
with two egg-laying peaks— one from late April to
early May and the other from late June to early
July, indicating production of two broods during
an average season. Pair formation and copulation
have not been observed (Trost 1968). One to five
eggs have been found in nests; four is the most
frequent clutch size (Baynard 1914, Nicholson
1928). The female incubates the eggs for 12 to 13
days and broods the young for 9 days more. Ju-
veniles stay in the territory about 20 days more,
after which the male may drive them away. Both
parents feed the young (Trost 1968).
The fledglings start to molt in late August and
are almost identical to adults by November. The
postnuptial molt of adults begins in August and
concludes by October. The light edges of the con-
tour feathers wear off during the winter and
produce the dark nuptial plumage (Trost 1968).
The average longevity is unknown, but an
adult banded in 1972 was seen tv^dce in 1978 (Ba-
ker in press).
A color plate of a fledgling appears in Trost
(1968), and black and white photographs of eggs
and nestlings, in Baynard (1914) and Nicholson
(1928).
MANAGEMENT AND CONSERVATION
The conflict between mosquito control and
the dusky 's stringent habitat requirements has
made conservation extremely difficult. Attempts
to recover the dusky on Merritt Island have in-
cluded lowering impoundment water levels, con-
necting an impoundment to the Indian River by a
culvert, and removal of a 3,000-m dike. The high
water levels and consequent vegetation changes,
such as shrub encroachment, have prevented these
measures from succeeding so far (DSSRT draft,
Baker, in press).
Management of the mainland dusky popula-
tion involves primarily land acquisition and con-
trolled burning. On the St. Johns National Wild-
life Refuge, destructive winter wildfires now are
checked by firebreaks maintained by refuge per-
sonnel. Experiments to control brush encroach-
ment include small prescribed bums, summer
burning, and herbicide use. A refuge addition of
1,320 ha in the vicinity of the Beeline Highway is
under negotiation (Baker, in press).
In addition, the Recovery Team calls for the
determination of habitat requirements, refining of
habitat manipulation techniques, population
monitoring by annual survey, and restoration of
habitat such as that on Merritt Island (DSSRT
draft).
The 1978 survey documented 9 singing males
on the St. Johns National Wildlife Refuge and 15
in the rest of the river basin. Most of those out-
side the refuge inhabit the area of a proposed
refuge addition between the forks of the Beeline
Highway (Baker, in press).
Critical Habitat is designated as the mainland
area bounded by 1-95, the St. Johns River, and
Florida Highways 45, 528, and 529, and as mos-
quito-control impoundments T-IO-J and T-IO-K
on Merritt Island National Wildlife Refuge (41 FR
53074, 3 December 1976; 42 FR 40685, 11 Au-
gust 1977; 42 FR 47849, 22 September 1977).
AUTHORITIES
J. L. Baker (Recovery Team)
U.S. Fish and Wildlife Service
Merritt Island National Wildlife Refuge
P.O. Box 6504
Titusville, FL 32780
H. W. Kale (Recovery Team)
Ornithological Research Division
Florida Audubon Society
35 1st Court S.W.
Vero Beach FL 32960
B. F. Sharp
U.S. Fish and WUdlife Service
Lloyd 500 Building
500 N.W. Multnomah Street
Portland, OR 97232
P. W. Sykes (Recovery Team)
Fish and Wildlife Service
P.O. Box 2077
Del Ray Beach. FL 33440
L. E. Williams (Recovery Team)
Wildlife Research Office
Florida Game and Fresh Water Fish
Commission
4005 South Main Street
Gainesville, FL 32601
PREPARER'S COMMENTS
The bird was named Ammodromus marttimus
var. nigrescens by Ridgway in 1873. It was desig-
nated Ammospiza nigrescens by Howell (1932),
American Ornithologists' Union (1957), and Trost
(1968). It was redesignated as a race of ^mmoipzza
maritima (Eisenmann 1973). It is geographically
isolated from, but morphologically similar to,
other races of seaside sparrow (Trost 1968). It is
unique in its extremely limited distribution (Chap-
man 1912).
LITERATURE CITED/SELECTED
REFERENCES
American Ornithologists' Union. 1957. Check-list
of North American birds. 5th ed. Port City
Press, Baltimore. 641 pp.
Baker, J. L. 1973. Preliminary studies of the
dusky seaside sparrow on the St. Johns Na-
tional Wildlife Refuge. Proc. Annu. Conf.
Southeast Assoc. Game Fish Comm. 27:207-
214.
. 1979. Dusky seaside sparrow. Pages 16-
19 in H. W. Kale, ed. Rare and endangered
biota of Florida, Vol. 2, Birds. Univ. Presses
of Florida, Gainesville.
. In press. Status of dusky seaside spar-
ro-w. In Rare and Endangered Wildlife Sym-
posium. Georgia Dep. Natur. Resour. and
Georgia Chapter Wildl. Soc. Aug. 3-4, 1978.
Athens.
Baynard, O. E. 1914. The dusky seaside sparrow
{Passerherbulus nigrescens). Oologist 32(7):
130-134.
Beecher, W. J. 1955. Late -Pleistocene isolation of
salt-marsh sparrows. Ecology 36:23-28.
Chapman, F. M. 1912. Handbook of birds of east-
ern North America (rev. ed.). D. Appleton
and Co., New York.
DSSRT (Dusky Seaside Sparrow Recovery Team).
Dusky seaside sparrow recovery plan. U.S.
Fish Wildl. Serv. 15 pp.
Eberhart, J. 1968. The dwindling dusky. Sci.
News. 93:501-501.
Eisenmann, E. (Chairman). 1973. Thirty-second
supplement to the American Ornithologist's
Union check-list of North American birds.
Auk 40:411-419.
Howell, A. H. 1932. Florida bird life. Coward-
McCann, Inc. New York. 579 pp.
Maynard, C. J. 1875. A new species of finch from
Florida. Am. Sportsman 5:248.
Nicholson, D. J. 1928. Nesting habits of seaside
sparrows in Florida. Wilson Bull. 40:225-237.
. 1929. Breeding of the dusky seaside
sparrow on the mainland of Florida. Auk 46:
391.
Peterson, R. T. 1947. A field guide to the birds,
2nd ed. Houghton Mifflin Co., Boston. 230
pp.
Sharp. B. E. 1968. Numbers, distribution, and
management of the dusky seaside sparrow.
M.S. Thesis. Univ. Wisconsin, Madison. 76 pp.
. 1969a. Conservation of the dusky sea-
side sparrow on Merritt Island, Florida. Biol.
Conser. 1(2):175-176.
. 1969b. Let's save the dusky seaside spar-
row. Fla. Natur. (April 1969):68-70.
-. 1970. A population estimate of the
Sincock, J. L. 1958. Waterfowl ecology of the
St. Johns River Valley as related to proposed
conservation areas and changes in the hydro-
logy from Lake Harney to Ft. Pierce, Florida.
Florida Game Fresh Water Fish Comm, Fed,
Aid Project W-19-R.
Sprunt, A., Jr. 1954. Florida bird life. Coward-
McCann, Inc. New York. 527 pp.
Trost, C. H. 1968. Dusky seaside sparrow. Pages
849-859 in O. L. Austin, Jr., ed. Life histories
of North American cardinals, grosbeaks, bunt-
ings, towhees, finches, sparrows, and allies.
U.S. Natl. Mus. Bull. 237. Part 2.
USDI (U.S. Department of the Interior). 1973.
Threatened wildlife of the United States.
Compiled by Office of Endangered Species
and International Activities. Bur. Sport Fish
Wildl. Resour. Publ. 114. U.S. Gov. Printing
Office, Washington, D.C. 289 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
dusky seaside sparrow. Wilson Bull. 8(2): 158-
166.
Biological Services Program
FWS/OBS-80/01.26
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
HAWAIIAN GOOSE (NENE)
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed lo:
Office of Endangered Species
U.S. Fish and Wildlife Senice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.26
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
HAWAIIAN GOOSE (NENE)
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
SlideU, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and WUdhfe Service
U.S. Department of the Interior
HAWAIIAN GOOSE (Nene)
Branta (Nesochen) sandvicensis (Vigors)
KINGDOM Animalia
CLASS Aves
ORDER Anseriformes
FAMILY Anatidae
OTHER COMMON
NAMES Nene; Lava Goose (Elder 1958)
DATE
Entered into SWIS To be determined
Update To be determined
LEGAL STATUS
FEDERAL
Endangered (42 FR 36426, 14 July
1977). Listed: Appendix I, 1973
Convention on International Trade
in Endangered Species of Wild Fauna
and Flora.
States:
Endangered: Hawaii.
REASONS FOR CURRENT STATUS
This highly specialized goose is adapted to life
in an environment that was originally predator-
free and waterless in the Hawaiian lava flows.
Their weak wings, exceptionally long flightless
period during molt, and young that are unable to
fly because of their exceptionally long develop-
ment period made them vulnerabfe to introduced
predators (Elder 1958;Elder and Woodside 1958).
There is evidence that overgrazing by sheep
and goats may have adverse effects on the nene
range, and the introduced mongoose is definitely
a predator on the young. The spread of the mon-
goose corresponded to the period of nene decline.
Mongooses and dogs are probably the most sig-
nificant predators, and the tameness and
curiosity of the nene makes them extremely vul-
nerable to predation as well as to poaching (Elder
1958; Elder and Woodside 1958). Zimmerman
(1974) believes that the information necessary
to identify and correct lethal dangers is still lack-
ing.
Hunting is believed to have contributed heavily
to their original decline, especially since ignorance
of the winter breeding season permitted hunting
at the time when geese were either incubating,
followed by broods, or molting and flightless
(Henshaw 1902 in Elder and Woodside 1958).
Hunting of this species was made illegal in 1911
(Ripley 1965).
A gradual increase in the nene population in
recent years has resulted from introduction of
artificaUy propagated stock on Hawaii and Maui,
the creation of sanctuaries in cooperation with
land owners, and control of predators at release
and breeding sites (Hawaii Div. Fish and Game
1972). However, it is not known if the present
population can maintain itself or increase in the
wild without additional introductions. The intro-
duced population on Maui is clearly not main-
taining itself. The future of the species in the wild
will remain in doubt until it can be shown that re-
production is high enough to offset losses from
all sources (King in press).
PRIORITY INDEX
39
DESCRIPTION
The nene is a medium-sized goose, with a bill
comparatively long, broad, and high near its base
and depressed at the tip. The crown, face, area
around eyes, throat, and a wide band on hind
neck are deep black; sides of head and sides and
front of neck are tawny buff; the feathers of the
neck have dark bases forming deep furrows and
producing a striped appearance; there is a blackish
ring around the base of the neck, broadening on
the back; the breast is pale brown; and the rest of
the body plumage is grayish brown barred with
whitish buff, the feathers having pale borders;
the rump is black; coverts are white with gray
bases; rectrices and primaries are black. The iris
is brown. Bill and legs are black. The tarsus and
toes are long and strong, with much reduced
webbing. The female is smaller than the male,
with shorter neck and slightly duller, darker plu-
mage (Delacour 1954).
Measurements. - male: wing 372 to 378 mm;
the tail is 147 to 158 mm; culmen, 40 to 47 mm;
tarsus, 81 to 90 mm. In the female: wing 350 to
368 mm; tail 144 to 151 mm; culmen 40 to 42
mm; and tarsus 73 to 78 mm (Delacour 1954).
Reported weights of wild specimens were as
follows: Males - summer, 2,074 g; winter (breed-
ing season), 2,370 g. Females - summer 1,762 g,
winter, 2,095 g (Kier et al. 1962).
Immatures.- Duller and more mottled.
Downy Chicks.- Grayish brown; forehead,
sides of head, throat and center of underparts
whitish buff; a dark spot on ear coverts and
whitish spots on base of wing.
Eggs.- Creamy white; five to eight in clutch;
80 X 50 mm (Delacour 1954). The clutch size for
captive birds at Pohakuloa and those recorded in
the wild on Hawaii is three to five eggs.
This species has become highly specialized
anatomically for its original environment, with
legs and feet best adapted to a strictly terrestrial
environment and wings poorly developed for
flight compared to other related geese. It is so
different morphologically from Branta that it
should be accorded a separate genus (MiUer 1937).
RANGE
The Hawaiian goose or nene is native to and
resident on the Island of Hawaii; it has been intro-
duced and is still surviving on Maui, Hawaiian Is-
lands (Delacour 1954). It occurs at several local-
ities on the slopes of Mauna Loa and Hualalai
volcanoes, chiefly between the 1,600- and 2,400-
m elevations. Its former range was reported to
have been much more extensive , even reaching to
the seashore, although records of earlier writers
are dubious (Baldwin 1945; Elder 1958). Nene
bones considered of Pleistocene or early recent
age are found on all the main Hawaiian Islands;
those of most recent age, probably dating to the
Polynesian occupancy, are on Molokai (Storrs
Olson pers. comm. 1977).
At present, breeding is recorded primarily in
three areas: Keauhou Sanctuary, on the east slope
of Mauna Loa ; Kahuku Sanctuary , on the southern
slope of Mauna Loa; and Keauhou 2 Sanctuary,
on the southeast slope of Haulalai (Hawaii Div.
Fish & Game 1972).
The only known major seasonal movement of
the populations is from the winter breeding areas
on the higher mountain slopes to a summering
area on the saddle between Mauna Loa and Mauna
Kea. The summering flock makes daily flights
from their nighttime range in the vicinity of Puu
Oo Ranch southeast of Mauna Kea and north of
the Saddle Road to the daytime range on lava
flows on the slopes of Mauna Loa, at least 15
miles away (Woodside 1956; Ripley 1965;Hawaii
Div. Fish & Game 1972).
RANGE MAP
The following map is adapted from Elder and
Woodside (1958); National Park Service, Hawaii
17004, 1963; Zimmerman 1975.
STATES/COUNTIES
Hawaii: Hawaii, Maui
HABITAT
Nene are usually seen on mountain slopes be-
tween 1525 and 2440 m elevation. Nesting and
feeding habitat is supplied by kipukas, or islands
of vegetation on lava flows, ranging in size from
less than 1 ha to several thousand hectares.
Vegetation grows on decomposing lava in various
stages of succession from lichens on bare rock to
such pioneering plants as ohelo [Vaccinium sp.),
kukainene [Coprosma emodioides), gosmore {Hy-
pochaeris radicata), pukeawe {Styphelia tameia-
meiae), and various grasses. Older kipukas with
deeper soils are vegetated with aalii [Dodonaea
viscosa), mamane (Sophora chrysophylla), ohia
[Metrosideros collina), and koa [Acacia koa) in
association with other shrubs and grasses. Rain-
fall is one of the most important factors in de-
composition of lava and the progress of ecological
succession. Average annual rainfall from 1960
through 1969 in the nene range on Mauna Loa at
2,042 m was 154 cm (Hawaii Div. Fish and Game
1972).
FOOD AND FORAGING BEHAVIOR
Nene are vegetarians. They consume a variety
of available foods, but show a marked preference
for gosmore [Hypochaeris radicata), eating the
leaves, buds, and flowers. Ohelo {Vaccinium sp.)
and kukainene [Coprosma emodioides) are the
most important food berries in nene sanctuaries.
The geese feed most heavily during morning and
late afternoon. Goats and pigs are not numerous
enough to be serious competitors to nene, even
though they graze on the same plants. In some
instances pigs may be beneficial by stimulating
growth of gosmore and other succulents. At the
present time, food resources are not considered a
limiting factor to nene in the wild (Baldwin 1947;
Hawau Div. Fish and Game 1972).
Although nene do eat berries, their preference
is for greens, especially the succulent leaves, stems
and buds of pusdele [Sonchus oleraceus) and gos-
more [Hypochaeris radicata). Also, the nene strips
seeds from the heads of grasses and sedges. At
more frequented altitudes, 1525 to 2440 m, there
is no evidence of food shortage at any season.
Grasses and greens are never scarce in that zone of
high moisture and infrequent frosts (Elder 1958).
In captivity, nene prefer green feed over com-
mercial mixtures (Hawaii Div. Fish & Game 1972).
Nene in the wild do not appear to be attracted
to water. At waterholes, they seem to be con-
cerned only with succulent green vegetation for
food. Drinking water is obtained from fog and
dew condensed on vegetation (Elder 1958; Hawaii
Div. Fish & Game 1972).
SHELTER REQUIREMENTS
Low bushes for concealing nests are the only
known shelter requirements (Hawaii Div. Fish and
Game 1972).
NESTING OR BEDDING
Most nests have been found under pukeawe
bushes or scrub ohia trees, where the goose scoops
out a shallow depression in the litter or duff.
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Leaves, twigs, and down are used to cover eggs
when the incubating bird leaves to forage (Hawaii
Div. Fish and Game 1972).
RITUAL REQUIREMENTS
Nene are more jealous and quarrelsome than
is common among geese. The gander defends his
territory, mate, and brood savagely, but some-
times exhibits the unusual behavior of attacking
his own mate (Delacour 1954).
POPULATION NUMBERS AND TRENDS
The wild population may have totaled 25,000
or more in the 18th century. Reduction began
between 1778 and 1850, progressed rapidly until
1900, and tapered off slowly between 1900 and
1930. In 1953, the estimated population of wild
birds was only 33 (Baldwin 1945). Smith (1952)
estimated the total population in the wild at less
than 30. Since 1940, coincident with conserva-
tion efforts, the population has obviously increased
if the earlier estimates were reasonably correct.
Counts on the summer roosting area near Puu Oo
Ranch north of the Saddle Road on the southeast
slope of Mauna Kea have ranged from 42 in 1966
to 114 in 1969. An increase during the 17 years
of counting in that area (since 1955) was in-
dicated, although inconsistent. About 90% of
birds in the summer roosting area are unhanded
and presumed to be wild rather than pen-reared
(Hawaii Div. Fish and Game 1972).
The number of individual birds observed with
eggs or young in the three nene sanctuaries have
been: 1966, 2; 1967, 32; 1968, 36; 1969, 12;
1970, 16; 1971, 12. Of these parent birds, 40%
were banded, indicating they were released pen-
reared birds; and 60% were unhanded, indicating
they were either wild or offspring of released
birds. There is no indication of a new population
developing from releases on the new sanctuary at
Kipuka Ainahou, northeast of Mauna Loa be-
tween Access Road and Saddle Road (Hawaii Div.
Fish and Game 1972).
Counts of introduced population on Maui re-
corded 43, with 2 produced in the wild in August
1973; 45 were seen in October 1974, with un-
handed birds shovwng up increasingly. Nests or
young were found in 1972 and 1974 (Monthly re-
ports of Hawaii Div. Fish and Game).
The actual number of nene in the wild is un-
known!. The "educated guess" is at least 600 wild
and released birds and their progeny on Hawaii
and a third as many more (200) on Maui (Wood-
side in Zimmerman 1974).
REPRODUCTION
Clutch size has been reported as 5 to 8 eggs
(Delacour 1954), although 3 to 5 eggs per clutch
were recorded for the wild population on Hawaii
and the captive birds at Pohakuloa. The incuba-
tion period is 28 to 31 days (Ripley 1965).
The nesting season may run from October
through March. February to late May is a flight-
less period for young and a molting period for
adults (Ripley 1965). Wild pigs, dogs, cats and
mongooses may threaten nene eggs and goslings
and even adult birds during the 4- to 6-week
flightless period (N. Santos m Zimmerman 1975).
Young grow slowly, requiring 10 to 12 weeks to
reach the flying stage; this is nearly twice the time
required by Canada geese. Adults are completely
grounded for 4 to 6 weeks by the wing molt;
thus, part or all of the nene family may be vulner-
able to ground predators for 3 months or more
each year (Elder and Woodside 1958). Breeding
potential is low because they rarely reach sexual
maturity or lay fertile eggs in captivity until 3
years of age or more. Six pairs observed in the
wild produced an average of only 2 young annually
(Elder and Woodside 1958).
In the captive flock at Shmbridge, England,
the sex ratio is equal. Sixty-two percent laid eggs
at the end of their second year. Ganders between
their fourth and eighth years have fertilized the
most eggs. Females laid the most fertile eggs in
their fourth year. Clutch size at Shmbridge aver-
aged 3.95 eggs, compared to 3.7 at Pohakuloa,
Hawaii. In Hawaii, nene lay their first eggs in
November; in England, about 9 February. Day
length, rather than temperature, is thought to
induce breeding. Long days inhibit breeding and
induce molting (Kier et al. 1967).
In the captive flock at Pohakuloa, inbreeding
of original stock was determined to be the cause
of low fertility; infusion of a new wild bird strain
and selection of birds for productivity greatly in-
creased fertility. Fertility increased with age of
breeders to 75% at 15 years in the Shipman strain
and to 100% at 8 years in the wild strain. Dates of
first egg laying at Pohakuloa ranged from 21 Sep-
tember (1969) to 5 December (1953), and was
most frequent during the first 2 weeks in Novem-
ber. Time of nesting in the wild is generally simi-
lar to that at Pohakuloa. Nene in the wild have
nested as early as October and as late as April,
depending on weather conditions.
Nest sites are usually located in kipukas,
which tends to isolate nesting pairs. Nests are left
uncovered until the last egg is laid, then thoroughly
covered with down, leaves, and twigs before the
female leaves. The male guards the nest from an
elevated lookout point a short distance away and
gives warning at the approach of danger. The fe-
male leaves the nest when approached during the
early stages of incubation, but sits tight during
late stages. Males have never been found incubat-
ing. Most pairs return to the same kipuka each
year.
Released birds will pair with wild mates as well
as with other released birds, although of 56 pairs
in the wild, only a few cases of captive reared
birds mated with other released individuals were
observed. No evidence has been seen of renesting
if the nest is deserted; in contrast, renesting is
common in captivity when the first clutch has
hatched and young have been removed (Hawaii
Div. Fish and Game 1972).
In 1956-57, the breeding area at Keauhou on
Mauna Loa had at least 6 adult pairs and 1 "un-
employed" bird; 12 young were produced, for an
average of 2 young per pair. The one unmated
bird indicates either that reproduction was poor
the preceding year or that most nonbreeders
spend their time elsewhere (Elder and Woodside
1958).
MANAGEMENT AND CONSERVATION
In 1949, lUCN placed the Hawaiian Goose on
its list of the 13 most threatened bird species in
the world, stimulating the beginning of a restora-
tion program with $6,000 appropriated by the
Territory of Hawaii. Captive rearing was started at
a former Civilian Conservation Corps Camp at
Pohakuloa, on the saddle between Mauna Loa and
Mauna Kea, with 4 birds from Herbert C. Ship-
man's aviary in Hilo, 1 from the Honolulu Zoo,
and 1 wild bird caught by a hunter's dog. In
1950, 2 birds were reared at Pohakuloa. In 1951,
3 were sent from Shipman's aviary to the wild-
fowl Trust at Slimbridge, England, to start a rear-
ing project there. In 1958, the U.S. Fish and Wild-
life Service provided a grant of $15,000 per year,
later increased to $25,000, for a nene restoration
project. The nene was officially designated the
territorial— now State— bird of Hawaii (Ripley
1965).
In early attempts at propagation, only 1 in 5
eggs hatched at either Pohakuloa or Slimbridge.
After about 10 years of frustratingly poor pro-
duction, inbreeding was identified as the cause, so
several wild birds were added to the captive flock.
The result was to almost triple the yield of fertile
eggs. Careful selection of goslings for quality
eliminated a "hairy dowTi" mutant. Old and un-
productive adults were also culled from stock
(Zimmerman 1975).
Through the 1973-74 breeding season, 1,306
goslings were raised at Pohakuloa and about one-
third as many at Slimbridge and other sites in
Europe and North America. Most of these birds
have been released to the wild— 934 on Hawaii
and 391 on Maui. The non-Pohakuloa reared
birds were released on Maui, all before 1971.
Since 1971, only Pohakuloa-reared birds have
been released on both islands. Production at
Pohakuloa has been between 100 and 150 gosHngs
per year at an average cost of $250.00 per gosling
(D. Woodside in Zimmerman 1975).
Birds are released into the wild in a flightless
stage, mostly as young between 2 and 4 months
old, but some as molting adults, into predator-
proof enclosures of habitat having natural food
available in addition to artificial food and water.
From there, birds fly over the fence into the wild
after their flight feathers grow. This is known as
the "gentle release" method (Hawaii Div. Fish
and Game 1972). The "nene park" method, pro-
posed by Peter Scott, where young are hatched
under incubating nene in large pens in natural
habitat, was tried, but the Hawaii Div. of Fish and
Game considered it too costly to build large en-
closures and assure predator-proof conditions.
Nene park methods would seem to avoid the pos-
sibility of imprinting goslings to unnatural condi-
tions of artificial brooders and pens, and has
other possible advantages (Pratt 1972). No ade-
quate testing to show the relative merits of the
two methods has been done (Zimmerman 1975).
Propagated stock has been liberated on the
island of Hawaii in three areas known to be fre-
quented by nene, starting on 17 March 1960.
These areas were established as "sanctuaries" by
cooperative agreement with the land owners.
They are: Keauhou Sanctuary- 1,2 7 8 ha on the
eastern flank of Mauna Loa; Keauhou 2 Sanctu-
ary-514 ha on southeastern slope of Haulalai in
North Kona; and Kahuku Sanctuary, on the
southern flank of Mauna Loa. A nene park has
been built in Hawaii Volcanoes National Park on
the Island of Hawaii with Pohakuloa stock. All
released birds are marked with bright, color-coded
plastic leg bands that identify where and when
liberated (Hawaii Div. Fish and Game 1972).
Roughly half of the 100 geese released each year
survive their first year, and at least 25% survive at
least 7 years. Data are still too sparse to construct
complete life tables (D. Woodside in Zimmerman
1975). Older birds, when liberated, appear to
leave flocks of younger birds and range over a
wider area. Data are insufficient to determine if
age at the time of release affects adaptability or
survival. Some released birds disperse to unknown
areas, then reappear after 4 or 5 years (Hawaii
Div. Fish and Game 1972).
A total of 391 birds have been released in
Haleakala Crater on Maui within National Park
boundaries, some every year from 1962 through
1970 except 1967. The first nests were found
there in 1968. A total of 38 nests have been
found and 1 1 young are known to have hatched.
Five dead goslings were attributed to heavy rains.
Mature young were found with parents in May
1971. In January 1972, nene produced the pre-
vious year were observed paired with Slimbridge-
reared birds. The tendency of Maui nene is to re-
turn to the same locality for nesting and some-
times, to the same site. An exceptional case of
dispersal was the one bird that flew back to
Hawaii and appeared at Pohokuloa (Hawaii Div.
of Fish and Game 1972). No thorough study of
the Maui population has ever been conducted
(Berger 1972).
Predator control has involved the use of
poison, injected into chunks of meat scattered in
crevasses frequented by predators. This bait was
placed throughout the sanctuaries for control of
dogs, cats, pigs, and mongooses. Additional bait
was placed in the vicinity of nene nests. Poisoning
is believed to have been effective, at least against
rats and dogs. Bait was placed so as to prevent its
being eaten by Hawaiian hawks, which species
was observed to harass nene on two occasions,
but is not believed to be a serious predator because
of its small number. Since the inception of the
predator control program on sanctuaries, only
two known incidents of predation have been ob-
served: a partly consumed carcass of a gosling,
presumed to have been the work of a rat, and 3
adult nene killed by dogs (Hawaii Div. Fish and
Game 1972).
Among measures proposed for the manage-
ment of nene are: preservation of the natural en-
vironment, including the establishment of perman-
ent refuges; control of predators and feral grazing
and browsing animals in breeding areas; continua-
tion of the captive and "nene park" propagation
program, if necessary to reinforce propagation or
extend the population to new range; conduct of
field studies to follow up on the fate of released
captive-reared birds and appraise total popula-
tions (U.S. Fish and Wildlife Service 1973; King
in press); expansion of education program. More
specific recommendations of the Hawaii Dept. of
Land and Natural Resources (1976) were to con-
tinue the propagation project in 1976 to furnish
nene for release in South Kona; reduce propaga-
tion effort to 24 breeders and produce only 1
brood per pair; continue use of same propagation
methods; discontinue plucking primaries because
deformed primaries can result when this is done;
expand the information and education program.
At least five nene have been illegally killed within
the last 2 yeais. In only one instance was the kill-
ing believed to be malicous; the remainder ap-
peared to have been through ignorance.
In addition to 3 nene sanctuaries cooperatively
managed on private lands, the Division of Fish
and Game announced the establishment of a State
sanctucuy, Kipuka Ainahou Nene Sanctuary con-
sisting of 15,540 ha on the northeast slope of
Mauna Loa between the Mauna Loa access road
and Saddle Road, approved 13 March 1974.
AUTHORITIES
David H. Woodside
Hawaii Division of Fish and Game
1151 Punchbowl Street
Honolulu, Hawaii 96813
Eugene Kridler
U.S. Fish and Wildlife Service
835 Akumu Street
Kailua, Hawaii 96734
Paul H. Baldwin
Department of Zoology
Colorado State University
Fort Collins, Colorado 80521
William Elder
108 Stephens Hall
Columbia, Missouri 65201
Ernest Kosaka
Division of Fish and Game
1179 Punchbowl Street
Honolulu, Hawaii 96813
PREPARER'S COMMENTS
Of the 2 to 36 individual nene observed in
Hawaii sanctuaries with eggs or young each year
from 1966 to 1971, 40% were banded, indicating
they were released pen-reared birds, and 60%
were unhanded, indicating they were either wild
or the offspring of released birds. Since 90% of
the birds observed on the summering grounds
were unhanded, it appears that this group draws
from a wider breeding population than that re-
corded in the sanctuaries, where only 60% were
unhanded. On the other hand, in 2 years with
high counts in the breeding area (32 in 1967 and
36 in 1968), 46 and 45 appeared in the summer-
ing area in 1967 and 1968, respectively, showing
only a small difference in the two annual counts.
This would seem to indicate that most of the
breeding population of the three sanctuaries
moves to the Puu Oo summer area. On the other
hand, with over 100 birds observed in the sum-
mering area in 2 different years, it would appear
that probably less than half of the total popula-
tion is observed in either the breeding season or
the summer counts in most years. In any case,
with data so far presented for Hawaii and Maui,
the "educated guess' of 600 for the two islands
(Zimmerman 1974) appears far too optimistic
and points up the need to establish much more
detailed investigation of the population as a pri-
mary objective of the nene restoration project in
the future.
It is hoped that a census technique, using
transects and census blocks, will be developed by
1980. If this approach proves valid in the sanctu-
aries, a method for censusing the entire nene
range will be devised.
LITERATURE CITED/SELECTED
REFERENCES
Baldwin, P. H. 1945. The Hawaiian Goose, its
distributiou and reduction in numbers. Con-
dor 47:27-37.
Baldwin, P. H. 1947. Food of the Hawaiian Goose.
Condor 49:108-120.
Berger, A. 1972. Hawaiian birdlife. Honolulu,
Univ. of Hawaii Press, p. 75.
Berger, A. 1972a. Hawaiian Birds. 1972. Wilson
Bull. 84:212-222.
Delacour, J. 1954. The waterfowl of the world.
Vol. 1. London, Country Life Limited.
Elder, W. H. 1958. Nene in Hawaii. Preliminary
report on the Nene in Hawaii. Wildfowl Trust.
Ninth Annual report. 112-117.
Elder, W. and D. Woodside. 1958. Biology and
management of the Hawaiian Goose. Trans.
23rd N. Amer. Wildlife Conf. p. 206.
Hawaii Department of Land and Natural Re-
sources (contributed by Ronald L. Walker).
1976. Nene restoration project, 1 July 1972-
30 June 1975. Elepaio 36(9):104-108.
Hawaii Division of Fish and Game. 1972. A re-
port of the nene restoration program. Unpub-
lished administrative report of Hawaii Div.
Fish amd Game.
Kier, J. 1975. Returning the Hawaiian Goose to
the wild, pp. 115-123, in Breeding endangered
species in captivity, R. D. Martin ed. London,
New York, San Francisco, Academic Press.
Kier, J., H. S. Roberts and R. Warren. 1967. The
Hawaiian Goose in Captivity, unpub. ms.
King, W. In press. (Revised) Red data book Vol.
2 - Aves. Internationa] Union for the Conserv-
ation of Nature and Natural Resources, Sur-
vival Service, Morges, Switzerland.
Miller, A. H. 1937. Structural modifications in
the Hawaiian Goose [Nesochen sandvicensis).
A study in adaptive evolution. Univ. Cali-
fornia Press 42:1-80.
Pratt, J. 1972. Research study proposal for inves-
tigation of behavior of the Hawaiian Goose
under the "Nene park" plan. Elepaio 33:33-
34.
Ripley, S. D. 1965. Saving the Nene, World's rarest
goose. Nat. Geog. Nov. 1965:745-754.
Schwartz, C. W. and E. Schwartz. 1948. An eco-
logical survey of the game birds in the
Hawiian Islands with recommendations for
management. Board of Commissioners of
Agric. and Forestry, Territory of Hawaii, pp.
273-275.
Scott, P. A. 1962. A project for a Nene park in
Hawaii. Elepaio 22:80-81.
Smith, J. D. 1952. The Hawaiian Goose (Nene)
restoration program. J. Wildl. Mgmt. 16:1-9.
State of Hawaii Department of Game and Natural
Resources. 1976. Nene restoration project. 1
July 1972-30 Jan. 1975. Elepaio 36:104-108.
U.S. Fish and Wildlife Service. 1973. Threatened
Wildlife of the United States. Resource Publi-
cation 114, U.S. Department of Interior,
G.P.O.
Woodside, D. H. 1956. Wild Nene on Hawaii.
Elepaio 16:67-68.
Zimmerman, D. R. 1974. Return of the nene.
Animal Kingdom. May, June, July 1974.
Zimmerman, D. R. 1975. To save a bird in peril,
Chap. 5. New York Coward, McCann &
Geoghegan.
Biological Services Program
FWS/OBS-80/01.27
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
COLUMBIAN WHITE TAILED DEER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubhc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of PLndangercd Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.27
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
COLUMBIAN WHITE TAILED DEER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-SIidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
COLUMBIAN WHITE-TAILED DEER
Odocoileus virginianus leucurus [Douglas]
KINGDOM Animalia
CLASS Mammalia
ORDER Artiodactyla
FAMILY Cervidae
OTHER COMMON
NAMES Pacific white-tailed deer,
tideland deer, cotton-tail deer
DATE
Entered into SWIS To be determined
Updates 8 Nov. 1978
LEGAL STATUS
Federal . . .Endangered (32 FR 4001, 11 March
1967).
States Endangered: Oregon, Washington.
REASONS FOR CURRENT STATUS
The Columbian white-tailed deer was once
abundant in the low and moist prairie habitat of
the Willamette River Valley of Oregon and north-
ward across the Columbia River into the river
valleys of southern Washington (Cowan 1936,
Gavin 1979). Supression of burning by Indians
and conversion of land for agricultural uses have
eliminated the native grass-herb association upon
which the deer depended. Drainage of beaver
ponds and their accompanying microenvironment
may also have eliminated essential habitat (Gavin
1978). These deer were also shot for food and
sport until early in this century, by which time
they were extirpated from most of their former
range.
PRIORITY INDEX
Not assigned.
DESCRIPTION
This subspecies is a small-sized white-tailed
deer, with a greyish-brown tail distinguishing it
from the race of yellow-tailed deer (O. v. ochrou-
rus) of eastern Washington (Cowan 1936). The
skull is smaller in all measurements, and there is
little possibility of confusion with neighboring
subspecies when adult specimens are compared
(Cowan 1936). The posterior margin of the palate
is even with or extending little farther (2 mm)
than the posterior end of the third upper molar.
Antlers are exceptionally small for northern races
of the white-tailed deer.
RANGE
It occurs in three widely separated groups of
relict populations. The first is found primarily
within the boundaries of the Columbian White-
Tailed Deer National Wildlife Refuge, which con-
sists of several islands and the adjacent mainland
near the mouth of the Columbia River. These
deer are also found on private land in the lower
Columbia River area: Puget Island, Washington,
Wallace Island, Oregon, and near Westport, Ore-
gon.
A search conducted by the Washington De-
partment of Game did not locate any Columbian
white-tailed deer between these populations and
Portland, Oregon, to the east. Recently, Colum-
bian white-tailed deer were found at a second
Columbia River area near Camas and Washougel,
Clark County, Washington (T. A. Gavin, pers.
comm.). A third group of populations occur in
the foothilUs near Roseburg, Oregon, some 320
km (200 mi) to the south.
RANGE MAP
Location of known populations is indicated
on the accompanying map (after Gavin in press).
STATES/COUNTIES
Washington . . .Wahkiakum, Clark
Oregon Clatsop, Columbia, Douglas,
Multnomah
HABITAT
The Columbia River herd prefers low-lying
islands and bottomlands. Much habitat has been
converted to pasture and is enclosed within dikes
or levees. Native trees and shrubs, occuring in
patches, are composed mostly of Sitka spruce
{Picea sitchensis), dogwood (Cornus stolonifera) ,
Cottonwood {Populus trichocarpa), red alder
(Alnus rubra), and willow (Salix) (Gavin 1979).
Common pasture grasses and forbs include
Festuca, Dactylus, Trifolium, Ranunculus, and
Lolium. Carex and /uncus are found in wet areas.
Common shrubs are Rubus, Rosa, Sambucus, and
Symphoricarpos (Gavin 1979). Forest cover in
the refuge areas has been reduced from 70% in
1939 to 17% in 1972 and the pasture lands are
heavily used by the Columbian white-tailed deer
(Suring 1974). The deer are less abundant on
islands (e.g.. Price and Hunting Islands) that are
primarily brushy and heavily wooded (Gavin
1978).
The Roseburg herd is found from river bot-
toms into rolling hills covered by oak woodland.
Major plants include Quercus garryana. Arbutus
menziesii, Acer macrophyllum, Fraxinus latifolia,
and the shrubs Rosa and Symphoricarpos.
Annucd grasses are interspersed among trees
(Gavin in press). Much habitat is privately owned
and used for sheep ranching.
FOOD AND FORAGING BEHAVIOR
The Columbian white-tailed deer is unusual in
its tendency to feed almost exclusively on herba-
ceous vegetation (grasses and forbs). Scheffer
(1940) reported finding only grass in a sample of
four stomachs, and Gavin (in press) found only
herbaceous material in the contents of 33 stom-
achs. Food plants include Ranunculus, Trifolium,
Alopecurus, and Phalris. There is some browsing
in the spring on twigs of Cornus, Rubus, Sambu-
cus, Lonicera, Symphoricarpos, and Fraxinus. Of
all the feeding deer observed by Suring (1974)
99% were grazing. These deer appear to prefer
feeding on pasture where grasses and forbs are
kept short and in a palatable stage of growth by
cattle grazing (Gavin in press). However, they
avoid such areas when cattle are actually present
(Suring 1974). Suring (1974) suggests there is
more feeding during the night than during day-
light hours, and that the percentage of time spent
feeding varies seasonally, with the greatest per-
centage of active time (90%) in summer devoted
to feeding.
MILES
The locations of known populations of Columbian white-tailed deer.
The University of Washington is conducting a
detailed study of the food habits of this deer,
including food availability and plant community
characteristics on the refuge.
SHELTER REQUIREMENTS
Resting deer seek cover in woodlands or
brushy areas adjacent to feeding pastures (Suring
1974). The physiognomy of cover seems more
important than the particular species composi-
tion.
NESTING OR BEDDING
Scheffer (1940) reports that these deer are
seen to emerge from v^^illow thickets about day-
break to begin feeding activities. They are also
reported to bed down in mint or hay fields
(Scheffer 1940). Fawns are sometimes discovered
resting in high grass of unmowed pastures in mid-
summer (U.S. Fish and Wildlife Service 1977).
RITUAL REQUIREMENTS
A relatively solitary animal, the average group
size is two to three individuals (Suring 1974).
The largest aggregations seen by Suring (1974)
occurred in winter and usually did not exceed 10
animals. Intraspecific interactions are relatively
infrequent in this deer, and then they are of low
intensity (only 17% of all male-male conflicts
result in physical contact, as opposed to 42% in
Michigan whitetails) (Suring 1974). Most of the
high-intensity threat actions observed were during
the rut.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Juxtaposition of grazing and cover areas
seems to be critical for this deer. Large expanses
of grazing land, providing ample forage, are
unused unless there is cover nearby. Tenasillahee
Island supports a low density of whitetails, pri-
marily because the available cover is concentrated
around the perimeter of the island (Gavin in
press). Diking of Columbia River islands is essen-
tial to prevent mass mortality, especially to
fawns, during high water (Scheffer 1940, Gavin
1978) and to provide grazing habitat.
POPULATION NUMBERS AND TRENDS
The entire lower Columbia River population
was estimated to number 300 to 400 individuals,
with the refuge population exhibiting a very
stable trend from 1972 to 1977 (Gavin 1979).
The refuge mainland population ranged from a
total of 164 to 215 deer during those years, or
3.67 to 4.82 deer per ha (Gavin 1979). Densities
on adjacent islands, both refuge-owned and pri-
vate, were much lower (e.g., about 1 deer per 100
ha on Puget Island, a private island). The sex
ratio of yearling and adult deer on the refuge was
quite stable from 1972 to 1977 at about three fe-
males per male (Suring 1974, Gavin 1979). Mean
home range size was 103.6 ha (N=18) and 108.6
ha (N=7) for adult females and adult males, re-
spectively (Gavin 1979).
All sources agreed that this population has
probably been at carrying capacity for some time.
Stability of population size has been enforced by
low fawn recruitment and moderate adult mortal-
ity. Fawn mortality is very high (69% to 80%)
during the summer, and adult mortality is often
associated with bacterial infections and parasites
(Gavin 1979).
Population density varies greatly among parts
of the refuge, with some refuge islands notably
underpopulated (e.g., Tenasillahee Island). Small
populations of whitetails are in danger of extinc-
tion by chance catastrophic events, such as
flooding of the Columbia River. Little informa-
tion is available for the Roseburg herd, since most
of the habitat is on private land and research was
only recently begun on these deer. Rough esti-
mates indicated there might be 1,900 whitetails
in Douglas County (Gavin in press). There is no
indication that recent hunting has been detrimen-
tal to the deer. On the other hand, sheep ranchers'
clearing of brushy cover used by deer is contin-
uing at a high rate, and this could be very dam-
aging.
REPRODUCTION
Rutting begins during the first week of No-
vember and reaches its peak later in the month
(U.S. Fish and Wildlife Service 1977). Circum-
stantial evidence indicates that some deer are re-
productively active through March (U.S. Fish and
Wildlife Service 1977). No specific information is
available on gestation period, but researchers have
assumed it is similar to that of the eastern white-
tail (210 days). In November of 1972, 1974, and
1975, the fawn/doe ratio was 35, 60, and 37 per
100, respectively Many does are observed without
fawns, and there are few reports of twins, both of
which further indicate a low survival rate for
fawns in the Columbia River herd, presumably
due to poor conditions of individuals in a popula-
tion at or near carrying capacity (Suring 1974).
Most births occur around the second week of
June. Female fawns are not known to breed their
first year. There is no information available on
the population dynamics of the Roseburg herd.
MANAGEMENT AND CONSERVATION
Habitat within the Columbian White-Tailed
Deer National Wildlife Refuge is currently pro-
tected, and efforts should be directed toward its
improvement. Originally an animal of wet prairies,
riparian habitat, and river bottoms, the deer on
the refuge feed on grazed and mowed pasture.
Maintaining pasture land in favorable condition
for deer is a primary task of refuge personnel.
Habitat management on Tenasillahee Island to
produce more dispersed cover would increase the
carrying capacity of this part of the refuge (Gavin
in press). Reducing the time spent by cattle in
favorable areas might increase their utilization by
deer (Suring 1974). But Gavin (pers. comm.) and
Suring and Vohs (1979) pointed out that cattle
grazing is a benefit in maintaining herbaceous
vegetation on the pastures in a short, actively
growing state. Timing of mowing should be tied
to the reproductive cycle, so as to avoid fawn
mortality (U.S. Fish and Wildlife Service 1977).
Although the present density of deer appears to
have no adverse effect on the habitat (T. A. Gavin
pers. comm.), as Suring (1974) concludes, 'a
healthy population (one able to weather times of
stress) is to be desired, not one of maximum den-
sity.' Surplus animals should be used to reestab-
lish the species in other areas of its former range
that can be identified as favorable habitat (U.S.
Fish and Wildlife Service 1977).
These deer are responsible for some agricul-
tural damage (Scheffer 1940, U.S. Fish and Wild-
life Service 1977). Their current protected status
engenders reluctance on the part of local manage-
ment and agricultural agencies to establish satel-
lite herds, due to the difficulty in controlling
damage to crops and orchards (U.S. Fish and
Wildlife Service 1977). The policy concerning
management of surplus animals should be reeval-
uated for species demonstrated to have a high
intrinsic rate of increase. Loss of habitat, rather
than mortality, seems to be the primary reason
for the decline of such species. For example, it
has been demonstrated that reestablished herds of
Tule elk can grow rapidly (McCullough 1969).
Since conservation of the Columbian white-tailed
deer required maintenance of a minimum of five
viable subpopulations (U.S. Fish and Wildlife Ser-
vice 1977), that goal should not be subverted by
resistance stemming from technical restrictions
placed on control of new populations.
The Roseburg herd poses special management
problems, initially because so few data are avail-
able on its population size and dynamics. Studies
are urgently needed to determine these data prior
to developing a management program. Habitat
preservation is needed for this herd. Urbanization
and clearing are ongoing problems. The primary
range of this herd is closed to hunting. Deer dam-
age to ornamentals and gardens is an increasingly
serious problem.
There have been repeated allegations that the
white-tailed deer of the Roseburg herd interbreed
occasionally with the sympatric black-tailed deer
(Gavin in press, U.S. Fish and Wildlife Service
1977). But in this area deer can usually be easily
assigned to one species or the other by field
observations (Gavin, personal communication).
Evidence indicates that hybridization between
these two species is possible in captive animals
(Gavin in press) and probably occurs in the wild.
The Washington Department of Game is con-
ducting a study of the Columbia River islands as
potential release sites for this species. A recovery
team has been established and has drafted a
recovery plan.
AUTHORITIES
Thomas A. Gavin
Department of Ecology, Fisheries and Wildlife
Oklahoma State University
Stillwater, Oklahoma 74074
E. Charles Meslow
Oregon Cooperative Research Unit
Oregon State University
CorvaHis, Oregon 97331
Winston Smith
Department of Fisheries and Wildlife
Oregon State University
Corvallis, Oregon 97331
Fred Martinsen
Columbia White-Tailed Deer Recovery
Team (Leader)
Washington Department of Game
600 North Capitol Way
Olympia, Washington 98501
PREPARER'S COMMENTS
Most of our information about this deer is
from studies made on the Columbia River herd.
Since the Roseburg herd occupies different habi-
tat, generalization about the biology and manage-
ment of this deer should be extrapolated to the
Roseburg herd with caution.
LITERATURE CITED /SELECTED
REFERENCES
Bailey, V. 1936. The mammals and life zones of
Oregon. North Am. Fauna 55:1-416.
Columbia White-Tailed Deer Recovery Team.
1977. Columbian white-tailed deer ref overy
plan. Draft Rep.
Cowen, I. McTaggert. 1936. Distribution and
variation in deer (genus Odocoileus) of the
Pacific coastal region of North America. Calif.
Fish and Game 22:155-246.
Dalquist, W. W. 1948. Mammals of Washington.
Univ. Kans. Pubis. Mus. Nat. Hist. 2:1 -444.
Gavin, T. A. 1978. Status of Columbian white-
tailed deer [Odocoileus Virginianus leucurus):
some quantitative uses of biogeographic data,
pp. 185-202 in Threatened Deer. lUCN,
Merges, Switzerland. 434 pp.
. 1979. Population ecology of the Colum-
bian white-tailed deer. Ph.D. dissertation.
Oregon State Univ. Corvallis, Oregon 149 pp.
In press. The Columbian white-tailed
in Halls, L. K. (ed). The Ecology and
deer.
Management of White-Tailed Deer. The Stack-
pole Co., Harrisburg, Pa.
McCullough, D. R. 1969. The tule elk. Its history,
behavior, and ecology. Univ Calif. Publ. Zool.
88:1-209.
Scheffer, V. B. 1940. A newly located herd of
Pacific white-tailed deer. J. Mamm. 21:
271-282.
Suring, L. H. 1974. Habitat use and activity
patterns of the Columbian white-tailed deer
along the lower Columbia River. Unpubl.
M. S. thesis, Oregon State Univ.
Suring, L. H. and P. A. Vohs, Jr. 1979. Habitat
use by Columbian white-tailed deer. J. Wildl.
Manage. 43:610-619.
ACCOUNT PREPARED /UPDATED BY
National Fish and Wildlife Laboratory
1300 Blue Spruce Drive
Fort Collins, Colorado 80524
Biological Services Program
FWS/OBS-80/01.28
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
SALT MARSH HARVEST MOUSE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the F.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal F,cosystems Team
U.S. Fish and Wildlife Scr\ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.28
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
SALT MARSH HARVEST MOUSE
A Cooperative Effort
by the
National Fish and Wildlife Lziboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
SALT MARSH HARVEST MOUSE
Reithrodontomys raviventris Dixon
KINGDOM Animalia
CLASS Mammalia
ORDER Rodentia
FAMILY Cricetidae
OTHER COMMON
NAMES Red-bellied harvest mouse,
Petaluma marsh harvest mouse
DATE
Entered into SWIS To be determined.
LEGAL STATUS
Federal Endangered (35 FR 16047, 13 Oct
1970).
States Endangered: California
REASONS FOR CURRENT STATUS
Distribution of the salt-marsh harvest mouse
is Hmited to native salt marsh habitat bordering
bays, estuaries, and rivers of the San Francisco
Bay region. Destruction of salt marsh habitat by
land filling and diking has greatly reduced and
fragmented the habitat of this species (Leach
1976,Shellhammer 1977).
PRIORITY INDEX
Not assigned.
DESCRIPTION
This species is distinguished from the western
harvest mouse {Reithrodontomys megaiotis) by
darker coloration on the back (especially in the
southern subspecies), many hairs being tipped
with black, especially mid-dorsally. Underparts
are white to fulvous; hairs at the anterior base of
the ear often form an ochraceous tuft. Hind feet
and tail are usually very dark. The tip of the tail is
blunt as opposed to pointed in R. megalotis, and
the thickness of the tail is greater (2.1 mm) 20
mm distal from the base (Fisler 1965:14). The
skull is longer than in R. megalotis (Dixon 1908,
1909; Howell 1914; Hall and Kelson 1959).
RANGE
R. raviventris is restricted to salt and brackish
marshes bordering south San Francisco, San
Pablo, and Suisun Bays. It is found on both sides
of the Golden Gate and the Sacramento River.
Two subspecies have been described, which differ
in some aspects of morphology, coloration, and
behavior (Fisler 1965). The easternmost occur-
rence of the species is in the vicinity of Antioch,
eastern Contra Costa County, and the extreme
western occurrence is in Sacramento County. A
marginal record from Grand Island, 2 miles north
of Knight's Landing, Solano County (Hall and
Kelson 1959) is actually a capture of/?, megalotis
(Fisler 1965).
RANGE MAP
Distribution is shown on the accompanying
map (Fisler 1965, Shellhammer, 1977). To make
the locations of populations visible on the map,
the width of the coastal strip occupied by the
mice has been exaggerated. Often the distribution
is limited to a linear strip of marsh within a few
hundred feet of the coast.
STATES/COUNTIES
California
HABITAT
Sonoma, Solano, Marin, Contra
Costa, San Mateo, Alameda, Santa
Clara, Sacramento, Napa.
This mouse is restricted to salt and brackish
marshes where plants provide a dense mat of
cover, ideally 0.2 to 1.0 m high, and a network of
spaces on the ground (Hooper 1944, Wondolleck
et al. 1976). Pickleweed {Salicornia) is the most
important indicator species; other plants typical
of R. raviventris habitat are Atriplex semibaccata,
A. patula, Grindelia cunifolia, Spartina, and
Distichlis.
FOOD AND FORAGING BEHAVIOR
The salt marsh harvest mouse's diet is domi-
nated by green vegetation. It can eat salt grass
(Distichlis) and pickleweed (Salicornia), as well
as some seeds. Seasonal shifts in diet are influ-
enced by available food plants: much more green
vegetation is eaten in winter (Fisler 1965).
SHELTER REQUIREMENTS
It is restricted to areas with considerable
cover, mostly in the form of halophytic vegeta-
tion, and does not venture into open areas, even a
few feet from vegetation (Fisler 1965, Zetterquist
1977). Since most of its habitat is within the
range of the extreme high winter tides, high
ground from which to escape the rising water is a
necessity. The lower high tides of summer are
avoided by climbing into the higher vegetation of
the marsh or by swimming to floating objects
(Fisler 1965). Fills and habitat alteration along
the upper edge of marshes that have no cover are
unsuitable as refugia.
NESTING OR BEDDING
The subspecies R. r. halicoetes builds a nest of
dry grasses or sedges, usually located on the
ground or in a hummock of vegetation. Nests are
often used by several individuals, and are aban-
doned when fouled (Fisler 1965). Reithrodonto-
mys r. raviventris is not known to build nests, but
merely huddles in an accumulation of vegetation
or nesting material (Fisler 1965). It is also reported
to use abandoned nests of other species as tempo-
rary shelters, including those of song sparrows
Qohnston 1956).
RITUAL REQUIREMENTS
Not known. Its secretive habits render field
observation difficult.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMNTS
Fisler (1965) stressed the behavioral adapta-
tions in R. raviventris that restrict it to areas of
high cover: a placid temperament and loss of the
frenetic escape behavior typical of R. megalotis.
R. raviventris is strongly dependent on the
presence of densely vegetated salt marsh habitat
for survival.
°l
CHV^ l<==:^x
o
"o
a
n
i
0
1
1
miles
10
i
Range of the salt marsh harvest mouse.
POPULATION NUMBERS AND TRENDS
It occurs throughout the salt marshes of the
San Francisco Bay area, except for very small
marshes and those extensively modified by
human activity (Fisler 1968). The southern sub-
species (R. r. raviventris) seems the most immedi-
ately threatened, for its range includes marshes
surrounded by heavily populated areas. It has
already been eliminated from the Corte Madera
marshes in Marin County (Fisler, 1965).
Density estimates are unavailable for this spe-
cies, partly because of the rapidly changing pat-
tern of marsh occupation by harvest mouse popu-
lations. The few remaining large marshes support
moderately high populations (Schaub 1971),
especially in San Pablo and Suisun Bays. The rate
of population turnover is high; few mice reach the
age of 12 months, and most survive less than 6
months (Fisler 1971, Wondolleck et al. 1976).
The main influence on the decreasing num-
bers of these mice is habitat destruction rather
than any direct removal or direct mortality result-
ing from human activities. Seasonal changes in
population distribution occur in these mice. Dur-
ing the winter months there is a movement
toward the upper edge of the marsh to escape the
highest tides, whereas the population spreads
throughout the marsh during the summer. In
some high tides, the populations move to higher
ground.
REPRODUCTION
Some males are reproductively active during
most of the year, with the low point in the winter
months. The season for pregnancy and lactation
in females is March through November (Fisler
1965). Females may produce only one litter per
year in the field (Fisler 1965). Average litter size
is 3.7 in R. r. raviventris and 4.2 in R. r. halico-
etes (Fisler 1965).
MANAGEMENT AND CONSERVATION
This small, secretive mouse is little noticed by
humans. It poses no economic problems. Habitat
protection is the critical factor in its management.
The dense cover available in the preferred salt
marsh habitat would seem to effectively protect
the species from predation by human commensals
during most of the yearly cycle. Occasionally,
mice forced onto levees during high tides may be
captured by feral cats and birds of prey. Increas-
ing pressure for land in the San Francisco Bay
area has resulted in outright destruction of much
of the original range of this species. Its continued
survival depends directly on the persistence of the
remaining salt-marsh habitat.
AUTHORITIES
George F. Fisler
Department of Biology
California State University, Northridge
Northridge, California 91330
Howard S. Shellhammer
Department of Biological Sciences
San Jose State University
San Jose, California 95192
PREPARER'S COMMENTS
The two subspecies of this mouse appear to
have evolved different physiological and genetic
traits and they may be considered incipient spe-
cies (Fisler 1965, Shellhammer 1967, 1977).
While some populations of the northern subspe-
cies (R, r. halicoetes) appear to occupy secure
habitat, others are restricted to small areas. The
remaining populations of the southern subspecies
are all restricted to small refugia of native salt
marsh and need careful management.
Habitat occupied by the salt marsh harvest
mouse is much diminished and fragmented from
its original condtion. Remaining populations are
isolated genetically, living on natural islands in a
sea of human-altered land. Under such circum-
stances, populations can be predicted to become
extinct (McArthur and Wilson 1967). Since these
mice avoid areas without cover, a single levee or
wide space may effectively isolate two adjoining
marshes, reducing the effective genetic pool. At-
tention should be given to the maintenance of
larger refugia of continuous salt-marsh habitat,
rather than equal areas of habitat dissected by
access roads, walkways, or barren ground.
LITERATURE CITED/SELECTED
REFERENCES
Dixon, J. 1908. A new harvest mouse from the
salt marshes of San Francisco Bay, California.
Proc. Biol. Soc. Washington 21:197-198.
. 1909. A new harvest mouse from Peta-
luma, California. Univ. Calif. Publ. Zool.
5:271-273.
Fisler, G. F. 1963. Effects of salt water on food
and water consumption and weight of harvest
mice. Ecology 44:604-606.
-. 1965. Adaptations and speciation in har-
vest mice of the marshes of San Francisco
Bay. Univ. Calif. Publ. Zool. 77:1-108.
1968. Adaptations in movement patterns
._ • . _r._i» u 1 t„ D..1I c„
of two species of salt-marsh rodents. Bull. So.
Calif. Acad. Sci. 67:96-103.
. 1969. Mammalian organizational systems.
Contrib. Sci., Los Angeles County Mus. 167:
1-32.
1971. Age structure and sex ratio in
populations of Reithrodontomys . ] . Mammal.
52; 653-662.
Grinnell, J. 1933. Review of the recent mammal
fauna of California. Univ. Calif. Publ. Zool.
40:71-234.
Hadaway, H. C, and J. R. Newman. 1971. Differ-
ential responses of five species of salt marsh
mammals to innundation. J. Mammal. j2:
818-820.
Haines, H. 1964. Salt tolerance and water require-
ments in the salt-marsh harvest mouse.
Physiol. Zool. 37:266-272,
Hall, E. R., and K. R. Kelson. 1959. The mammals
of North America. 2 vols. Ronald Press Co.,
New York. 1,083 pp.
Hooper, E. T. 1944. San Francisco Bay as a factor
influencing speciation in rodents. Misc. Publ.
Mus. Zool., Univ. Michigan 59: 1-89.
Howell, A. H. 1914. Revision of the American
harvest mice (Genus Reithrodontomys). No.
Am. Fauna 36:1-97.
Leach, H. R. 1976. Salt marsh harvest mouse
[Reithrodontomys raviventris) . Pages 4-5 in
At the crossroads: a report on California's en-
dangered and rare fish and wildlife. Calif.
Dep. Fish Game Biannual Rep. 103 pp.
Johnston, R. F. 1956. Population structure in salt
marsh song sparrows. Part I. Environment and
annual cycle. Condor 58:24-44.
1957. Adaptation of salt marsh mam-
mals to high tides. J. Mammal. 38:529-531,
MacArthur, R. H., and E. O. Wilson. 1967. The
theory of island biogeography. Princeton
Univ. Press, Princeton. 203 pp.
Rice, V. C. 1974. The population ecology of the
salt marsh harvest mouse at Triangular Marsh.
M.A. Thesis, San Jose State Univ., San Jose,
Cahf. 128 pp.
Schaub, D. B. 1971. Salt marsh harvest mouse
survey, 1971. Calif. Dep. Fish Game, Spec.
Wildl. Invest. Final Rep. 11 pp.
Shellhammer, H. S. 1967. Cytotaxonomic studies
of the harvest mice of the San Francisco Bay
region. J. Mammal. 48:549-556.
Wondolleck, J. T., W. Zolan, and G. L. Stevens.
1976. A population study of the harvest mice
[Reithrodontomys raviventris Dixon) in the
Palo Alto Baylands salt marsh. Wasmann J.
Biol. 34:52-64.
Zetterquist, D. K. 1977. The salt marsh harvest
mouse [Reithrodontomys reviventris) in mar-
ginal habitats. Wasmann J. Biol. 35:68-76.
PREPARED/UPDATED BY
National Fish and Wildlife Laboratory
1300 Blue Spruce Drive
Fort Collins, Colorado 80524
Biological Services Program
FWS/OBS-80/01.29
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
PUERTO RICAN PARROT
Fish and Wildlife Service
U.S. Department of the Interior
prefacp:
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as ainendcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of F.nginccrs in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report shiiuld be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.29
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
PUERTO RICAN PARROT
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Shdell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
PUERTO RICAN PARROT
Amazona vittata (Boddaert)
KINGDOM Animalia
CLASS Aves
ORDER Psittaciformes
FAMILY Psittacidae
OTHER COMMON
NAMES Puerto Rican Amazon (Forshaw
1973); Cotorra de Puerto Rico (Leopold
1963); Cotorra Puertorriquena (Rules and
regulations concerning wildlife and hunt-
ing in Commonwealth of Puerto Rico).
DATE
Entered into SWIS To be determined
Update To be determined
LEGAL STATUS
Federal: Endangered (42 FR 36426, 14 July
1977). Listed Appendix I, Convention
on International Trade in Endangered
Species of Wild Fauna and Flora (42
FR 10478, 22 February 1977).
States: Protected against hunting, killing, or
capture permanently by Rules and
Regulations concerning the Wildlife and
Hunting in the Commonwealth of
Puerto Rico, 1972, Article 8a.
REASONS FOR CURRENT STATUS
Deforestation, widespread hunting, and taking
of parrots for pets almost certainly brought about
the original decline of this species. Now that the
population is so small, all adverse pressures are
very serious. Severe hurricanes in 1928 and 1932
decimated the population, probably by destruc-
tion of fruit (Noel Snyder pers. comm.). Inter-
actions between parrots and red-tailed hawks,
broad-winged hawks, and sharp-shinned hawks
have been seen (Cameron Kepler in Forshaw
1973). The pearly-eyed thrasher {Margarops fus-
cata), which competes with parrots for nesting
sites and also preys on eggs and chicks, has be-
come much more abundant and widespread in re-
cent years (Nathan Leopold 1968 pers. comm.;
Cameron Kepler in Forshaw 1973). Pearly-eyed
thrashers and red-tailed hawks are believed to be
the most serious predators; rats and screech owls
are not considered serious (Noel Snyder pers.
comm.).
A critical shortage of nesting holes in hollow
trees combined with fierce competition from the
exploding population of pearly-eyed thrashers
continue to threaten parrots. lUegcil hunting and
nest robberies by humans are still threats (Noel
Snyder pers. comm.).
PRIORITY INDEX
65'
DESCRIPTION
The adult parrot is about 29 cm long, with
predominantly green plumage, paler and more
yellowish on underparts; feathers are edged with
dusky black, p2U"ticularly on head and neck ; lores
and frontal band are red; there is a distinct white
eye ring; under tail coverts are yellowish-green;
feathers of abdomen are sometimes slightly tinged
with dull red; primary-coverts and primaries axe
dark blue; outer webs of outermost secondaries
are blue narrowly edged with dull green; under
wing-coverts are green; undersides of flight fea-
thers are bluish-green; tail is green narrowly tipped
with yellowish-green, bases of lateral feathers are
marked with red on inner webs and outermost
feathers are edged with blue; bill is yellowish horn;
iris is brown; legs are yellowish brown. Immatures
^u■e similar to adults (Forshaw 1973).
The extinct subspecies A. v. gracilipes of
nearby Culebralsland was very similar, but smaller
and with smaller, more slender feet (Forshaw
1973).
Measurements. A. v. vittata: (8 males) wing
182-193 mm (av. 188.5 mm); taU 90-103 (96.9);
exposed cuhnen 27-30 (28.5);tarsus 21-24 (22.1);
(5 females) wing 178-196 (av. 185.6 mm); tail 93-
104 (98.2); exposed cuhnen 27-28 (27.2); tarsus
22-24 (23.0). A. v. gracilipes: (2 males) wing 169-
173 (av. 171.0 mm); taU 93-95 (94.0); exposed
culmen 26 (26.0); tarsus 20-21 (20.5); (1 female)
wing 175 mm; tail 100; exposed culmen 23; tar-
sus 20 (Forshaw 1973).
RANGE
The present range is confined to Puerto Rico;
it formerly included nearby Culebra, Vieques and
Mona Islands. This parrot was last recorded on
Culebra Island in 1899; it disappeared from other
offshore islands of Puerto Rico earlier.
It has not been recorded from the mangrove
swamp at the mouth of the Mameyes River since
Wetmore (1927) found it there. It was known to
be in Guanjataca Forest at medium altitudes up
to 1910, in Rio Abaja Forest also at medium
elevations up to 1920's, and in Carite Forest at
high elevations up until the 1930's. It is now
virtually confined to Luguillo National Forest,
which comprises 11,200 ha of relatively high-
elevation tropical rainforest in extreme eastern
Puerto Rico (Noel Snyder pers. comm.).
RANGE MAP
The range of the parrot has included most
areas of Luguillo Forest above 400 m elevation.
STATES/COUNTIES
Commonwealth of Puerto Rico.
HABITAT
Mature wet forests with high rainfall between
about 400 and 800 m elevation are required by
this species. It is now confined in breeding areas
having the largest number of old Colorado trees,
o
o
I— I
o
(£!
W
A,
u
o
u
S
u
J3
which supply nesting cavities. It formerly fre-
quented more diversified habitat, particularly at
lower elevations (Noel Snyder pers. comm.).
Dwarf forest at higher elevations and second
growth lowland forest are not used by parrots
(Cameron Kepler in lit. 8 Feb. 1971).
Parrots occupy the Tabonuco, Sierra Palm,
and Colorado forest types of Wadsworth (1952).
FOOD AND FORAGING BEHAVIOR
Rodriguez-Vidal (1959) lists more than 50
species of fruiting plants used by parrots. Most
commonly used was sierra palm {Prestoea mon-
tana [Euterpe geobosa] ) with a long fruiting pe-
riod from November tojune (chiefly Feb. through
March). Other important food plants were bejuco
de rana vine {Marcgravia sintenisii), camasey de
paloma [Miconia sintenisii), tabonuco {Dacryodes
excelsa), cabeilo {Casearia guianensis), guara
[Cupania triquetra) and hueso bianco (Mayepea
domingensis) trees. Kepler (1970) has seen par-
rots extracting nectar from fleshy bracts below
flower clusters on marcgravia vines and noted
that, in some areas parrots feed more heavily on
Clusia krugiana than any other tree.
Rodriguez-Vidal (1959) also stressed the im-
portance of the tabonuco tree in providing fruit
during August to November because parrot food
is scarce during that period of heavy rains. How-
ever the largest flocks he counted (200) were
feeding on sierra palm, bejuco de rana and cama-
sey de paloma (with no mention of tabonuco); on
8 November 1953 and 31 October 1954, both at
Valle Hicaco on El Yunque. He never observed
parrots feeding on the ground. They fed chiefly
on pericarps of wild fruits but silso on flowers
and tender shoots. If fruit is in clusters, parrots
cut off the entire small supporting branch, held it
in one foot, and picked off ripe fruit, letting un-
ripe fruit fall to ground. They fed in groups and
if frightened while eating would fly off, some
carrying one piece or clusters of fruit in their bills.
He did not see any parrots fighting over food.
Parrots move about widely to feed especially be-
tween September and December when tabonuco
is in fruit. They are highly regular in their daily
flights of from 1 to 5 km to and from food sources
and night roosts (Kepler 1973). Kepler (1970)
points out the possibility that an adequate supply
of their essential foods is not found within the
Forest Reserve, accounting for reported flights
from the forest during the summer months. They
forage in semi-social groups. The normal foraging
range of a nesting pair is about 1.6 km, but some-
times 8 km or farther. There is no evidence of a
shortage of food. With failure of the sierra palm
crop in 1974, parrots shifted to other foods (Noel
Snyder pers. comm.).
SHELTER REQUIREMENTS
Cavities for nesting are the only shelter re-
quired.
NESTING OR BEDDING
The nest is a hollow in a tree. In Luquillo
Forest Reserve nearly all nests found have been in
cavities in large Colorado trees {Cyrilla racemiflora)
formed by rotting of wood of trunk after branches
have been lost to wind or other accidents. Parrots
prepare nests by clearing out the interior of the
cavity, but do not add lining material. Most sites
chosen have been more than 5 m from the ground.
Cavities have been random in compass orientation,
with minimum observed entrance diameters of
10 cm and minimum observed internal diameters
of about 23 cm. Cavity depths have ranged to
240 cm with the deepest cavities apparently pre-
ferred. All recent nests have been about 500 m
elevation.
Historically, parrots nested in holes in cliffs as
well as hollow trees, but recently located nests
have all been in rotted out cavities in large color-
ado trees. Most such cavities are unsuitable for
nesting because they are too wet or too small.
Suitable nest sites are scarce and limiting to breed-
ing of parrots in their traditional areas, which
they seem to be reluctant to leave for other areas
where suitable nest sites exist (Noel Snyder pers.
comm.).
RITUAL REQUIREMENTS
Kepler (1973) says there are distinct take-off
calls, flight calls, and series of contact calls, in-
cluding duetting between pair members. The lat-
ter probably serve as "station identification"
(Helen Snyder pers. papers 9 May 1975).
Calls are pair specific. Pairs are very territorial
and aggressively defend their territory. There is
evidence that pairs identify one another by calls,
and don't pay much attention to nonterritorial
pairs (Noel Snyder pers. comm.).
OTHER CRITICAL REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
Wetmore (1927) points out that in 1836,
Moritz reported great flocks of parrots in Puerto
Rico, and in 1864 Taylor found them to be
common. It seems that until the turn of the cen-
tury they were plentiful, particularly in the in-
terior, but in 1911-12, Wetmore located popula-
tions only in the karst of western Puerto Rico,
in the lowlands near the mouth of the Mameyes
River, and in the Luquillo Forest of eastern
Puerto Rico (Forshaw 1973). Counts in Luquillo
National Forest Reserve from August 1953 to
March 1956 never exceeded approximately 200
individuals (Rodriguez-Vidal 1959). By December
1966, the highest count achieved by Victor
Marquez in a several month effort was 70 indivi-
duals (pers. comm.), and by 1968 the highest
count achieved by Kepler was only 24. The wild
population dropped to a low point of only 13 or
14 in 1974, but has been showing some signs of
recovery since that time. By summer 1979, there
were 26 to 28 birds in the wild, including 4 breed-
ing pairs.
REPRODUCTION
With but one exception— a 1974 cavity in a
laurel sabino (Magnolia splendens)~dll recent
nests of the parrots have been in cavities in palo
colorados. The predominant use of palo colorados
is probably a reflection of the greater abundance of
natural cavities in this species. Nevertheless, good
nesting cavities are not abundant in general. Sys-
tematic climbing and checking of over 1200 trees
within the parrot nesting areas from 1973-1976
reve8ded a dearth of good cavities, and several
pairs of recent years have failed to lay eggs be-
cause of apparent failures to locate good sites, a
problem that has been alleviated in the last 4
years by provision of artificial nest sites (Snyder
arid Wiley pers. comm.).
Incubation is approximately 26 days; clutch
size 2 to 4 (average 3), and nestling period 8 to 11
weeks (average 9 weeks). Only femjiles incubate
and males provide all food to females during the
incubation period. Both adults provide food for
young. Sexual maturity is reached at 3 to 4 years
of age.
Before 1973, when intensive nest manage-
ment efforts were begun, nesting success for all
nests found at the egg stage was between 1 1% and
26%. Since 1973, success has increased to 71%,
primarily due to efforts to alleviate nest predation
by pearly-eyed thrashers and to maintain the
quality of nest sites. Sources of nesting failure in
addition to nest-site disintegration and predation
by thrashers have been parasitism of nestlings by
bot flies and predation on adults and nestlings by
red-tailed hawks {Buteo jamaicensis), but neither
of these factors appears to have been a major
cause of the historical decline of the species. Nest-
robbing by man accounted for a large fraction of
the nestlings that survived other pressures up until
the late 1960's, but has not been a major problem
since that time.
Numbers of young fledged in the wild in re-
cent years have run: 1973,3; 1974,3; 1975,6;
1976,8; 1977,3 ;1978,9;and numbers of egg-laying
pairs have run: 1973,3; 1974,2; 1975,5; 1976,4;
1977,3; 1978,4.
MANAGEMENT AND CONSERVATION
A program of study conducted by the Fishery
and Wildhfe Section of the Puerto Rico Dept. of
Agriculture and Commerce financed by the
Pitman-Robertson Federal Aid to Wildlife Pro-
gram from 18 August 1953 to 30 June 1956, with
Jose A. Rodriguez-Vidal as chief investigator, was
followed by a project developed in 1968 by the
U.S. Fish and WildUfe Service, U.S. Forest Ser-
vice, World Wildlife Fund, and the Commonwealth
of Puerto Rico. The latter project is still continu-
ing with primary funding from the U.S. Forest
Service and the U.S. Fish and Wildlife Service.
Studies from 1968 to 1971 were under the direc-
tion of Cameron Kepler; from 1972 to 1976 were
under the direction of Noel Snyder; and from
1977 to the present are under James Wiley.
Management efforts began in 1956 with rat-
poisoning with warfarin in the parrot nesting
areas. Rat control has been continued in most
years of study since that date. However, recent
evidence strongly suggests that rats are only a
secondary threat to nests (Snyder pers. comm.),
and in recent years management emphasis has
shifted from rat control to efforts to reduce the
impacts of thrashers and to provide and enhance
the quality of nest sites.
In the years just prior to 1973, thrashers were
apparently responsible for the majority of parrot
nest failures, although this species was not present
in Luquillo Forest in any abundance before 1950
and thus was not involved in the early decline of
the parrots there. Starting in 1973, intensive
guarding of parrot nests has prevented any further
cases of nest failure from this cause. In 1973,
1974, and 1975, thrashers were eliminated by
shooting whenever they threatened parrot nests,
and in addition some nests were protected by
artificial incubation of eggs and later replacement
of young into nests. Parrots were maintained in
these nests by dummy eggs made from plaster. At
least 2, very likely 4, and possibly as many as 6
parrot nests were saved from thrasher predation
during these years by these efforts.
Studies of nest-site preferences of thrashers
conducted in 1974, 1975, and 1976 demonstrated
a reluctance of this species to enter deep structures
with bottoms not visible from entrances. Parrots,
in contrast, appear to prefer such structures, and
in 1976 efforts to convert parrot nests into such
structures were begun. All parrot nests were
deepened and provided with baffles - changes
which the parrots accepted without obvious dif-
ficulties. Thrashers have been provided with
alternative nest sites close to each parrot nest and
once established in these alternative sites, have
greatly reduced the frequency with which thrash-
ers have attempted to enter parrot nests by their
intraspecific territorial behavior. Since 1977 there
have been no significant thrasher threats to any
parrot nests, and it has not been necessary to
eliminate any thrashers.
The provision of artificially created nest sites
has also met with considerable success, and all
currently active parrot pairs are using such sites.
As a hedge against loss of the wild population
and to provide a source of birds for future release
into the wild, a number of parrots have been taken
into captivity in recent years, primarily as eggs.
The current (1979) captive population stands at
15 individuals. As yet, no successful reproduction
has taken place in the captive flock, although
several females have laid infertile eggs and one
pair laid fertile eggs that did not hatch in 1978.
By chance, the captives taken consist largely of
females so their number of potential egg-laying
pairs has remained low. Success in breeding the
closely related Hispaniolan Parrot {Amazona
ventralis) at Patuxent and at other facilities pro-
vides optimism that the captive program may
soon begin to produce young.
Other management efforts of recent years
have been directed at reducing the impact of bot
fly parasitism of young parrots. Experiments
are currently underway with methods of prevent-
ing this parasitism. Nestlings of the past few
years have been closely monitored for parasitism
and treated when such parasitism has occurred.
AUTHORITIES
James Wiley
Box 21
Palmer, Puerto Rico 00721
Noel Snyder
Patuxent Wildlife Research Center
Laurel, MD 20810
Cameron Kepler
RR Maalaea Road
Kula, Hawaii 96790
Frank H. Wadsworth
Institute of Tropical Forestry
University of Puerto Rico
Box AQ
Rio Piedras, Puerto Rico 00928
PREPARER'S COMMENTS
It appears that we now have the technology
for ensuring satisfactory reproduction of Puerto
Rican Parrots in the wild by appropriate effort
in nest site enhancement, brood care, and pearly-
eyed thrasher management. Future parrot man-
agement should concentrate on providing and
maintaining suitable nest sites combined with
continued routine censusing of the species.
LITERATURE CITED/SELECTED
REFERENCES
Danforth, S. J. 1935. Supplementary account of
the birds of the Virgin Islands, including
Culebra and adjacent islands, pertaining to
Puerto Rico, with notes on their food habits.
J. Agric. Univ. Puerto Rico 19(4):430-472.
Forshaw, J. M. 1973. Parrots of the World. New
York, Doubleday & Company, p. 522.
Kepler, C. B. 1970. The Puerto Rican Parrot in
Chapter E-14: Preliminary comparison of bird
species diversity and density in Luquillo and
Guanica Forests, H. J. Odum ed. A Tropical
Rain Forest, Oak Ridge, Tenn. U.S. Atomic
Energy Commission Div. of Tech. Info.
Leopold, N. J. 1963. Checklist of birds of Puerto
Rico and the Virgin Islands. Bull. 168, Uni-
versity of Puerto Rico Agric. Exp. Sta. 119 pp.
Rodriguez-Vidal, J. A. 1959. Puerto Rican Parrot
{Amazona vitata vitata) study. Monog. Dept.
Agric. and Commerce, Puerto Rico, No. 1:1-.
15.
Wadsworth, F. H. 1952. Forest Management in
the Luquillo Mountains. Caribbean Forests,
Ann. Rep. Vol. 14 ( 1&2) p. 40.
Wetmore, A. 1916. The birds of Vieques Island,
Puerto Rico. Auk 33:403-419.
Wetmore, A. 1927. The birds of Puerto Rico and
the Virgin Island-Psittaciformes to Passeri-
formes. Scient. Surv. Puerto Rico and Virgin
Islands, New York Acad. Sci. 9(4):409-598.
Biological Services Program
FWS/OBS-80/01.30
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
KEMP'S (ATLANTIC) RIDLEY SEA TURTLE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoasl of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the I'.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and W'ildlifc Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal P^cosystems Team
U.S. Fish and Wildlife Ser\ ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704r)8
u
FWS/OBS-80/01.30
March 1980
SELECTKD VER 1 EBRAIK ENDANGERED SPECIES
OF IHE SEACOAS I OF 1 HE UNITED STATES-
KEMP'S (ATLANTIC) RIDLEY SEA TURTLE
A Cooperative Effort
by the
National Fish and WildHfe Laboratory,
the Office of Endangered Species
and the
National Coastal FJcosystenis Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
KEMP'S (ATLANTIC) RIDLEY SEA TURTLE
Lepidochelys kempii Carman
KINGDOM Animalia
CLASS Reptilia
ORDER Testudinata
FAMILY Cheloniidae
OTHER COMMON
NAMES Mexican ridley,
Atlantic ridley, Tortuga lora.
DATE
Entered into SWIS to be determined.
Updates. .22 September 1976, 25 February 1977.
LEGAL STATUS
Federal: Endangered (35 FR 18310, 2 De-
cember 1970)
States: Endangered: Florida, Georgia, Mary-
land, New Jersey, South Carolina,
Texas.
CREDIT: P. C. H. PRITCHARD
Protected: Alabama, North Carolina.
REASONS FOR CURRENT STATUS
Overuse by man is the primary factor respon-
sible for the present small numbers of sea turtles.
Diurnal nesting on a single beach in Mexico makes
this turtle particularly susceptible to predation by
man and wild animals. Commercial harvesting of
eggs and skin has played a significant role in the
decline (lUCN 1968). Harvesting has recently
(1966) been prohibited by the Mexican goven-
ment, but no upward trend in population num-
bers has been observed (Pritchard and Marquez M.
1973). Predation by wild animals is acute, espe-
cially on hatchlings. Crabs, fish, reptiles, birds,
and mammals are predators; adult predation is
limited to sharks (Rebel 1974).
Because of aggregate nesting on a single beach,
Rancho Nuevo in TamauHpas State, Mexico, any
habitat modification there could result in loss of
the entire breeding population.
Sea turtles are caught incidental to commer-
cial fishing activities. Some turtles drown in
trawls, some are eaten by fishermen, sold in local
markets, or mutilated as a result of entanglement
in the trawls (U.S. Department of Commerce
1976). According to Pritchard (1976), ridleys are
caught north and south of the Rio Grande off the
States of Florida, Louisiana, and Texas, as well as
in Tabasco, Veracruz, and off the Campeche Bank
in Mexico.
PRIORITY INDEX
Not assigned.
DESCRIPTION
The ridley is a small sea turtle with an unusu-
ally broad (compared with other sea turtles),
heart-shaped, keeled carapace that is serrated be-
hind the bridge. It has a triangular head and a
somewhat hooked beak with large crushing sur-
faces. The plastron has several small pores on each
side, leading to Rathke's glands (secretory struc-
tures).
Hatchlings are black on both sides. As the tur-
tle matures, the bridge and hingeless plastron
change to white, then yellow; and the carapace
changes to gray and then olive green. The head
and paddle-like limbs are gray.
Adults weigh between 35 and 42 kg, and have
a carapace length of 56 tp 70 cm.
Close examination of the carapace reveals five
pairs of pleural shields with the nuchal shield
touching the first costals. There are 12 to 14 mar-
ginals on each side of the carapace and pores in
the four bridge shields.
Black-and-white photographs appear in Carr
(1952, 1967), Ernst and Barbour (1972), Bustard
(1973), Rebel (1974) and Pritchard (1976).
RANGE
Adults are restricted to the Gulf of Mexico.
Immatures may be observed along the Atlantic
coast as far north as Massachusetts and are sighted
infrequently along European shores.
The former range was probably equivalent to
the present range (lUCN 1968, Witham 1976).
RANGE MAP
On the following pages distribution is shown
by shading, and U.S. nesting sites by dots.
STATES/COUNTIES
Alabama: Baldwin, Mobile.
Delaware: Sussex;
Florida: Bay, Brevard, Broward, Charlotte,
Citrus, Collier, Dade, Dixie, Duval,
Flagler, Franklin, Gulf, Hernando,
Hillsborough, Indian River, Jefferson,
Lee, Levy, Manatee, Martin, Monroe,
Nassau, Okaloosa, Palm Beach, Pasco,
Pinellas, St. Johns, St. Lucie, Santa
Rosa, Sarasota. Volusia, WakuUa, Wal-
ton.
Bryan, Camden, Chatham, Glynn, Li-
berty, Mcintosh.
Cameron, Iberia, Jefferson, Lafourche,
Plaquemines, St. Bernard, St. Mary,
Terrebonne, Vermilion.
Worchester.
Barnstable, Dukes, Essex, Middlesex,
Nantucket, Norfolk, Plymouth.
: Hancock, Harrison, Jackson.
: Atlantic, Cape May, Monmouth,
Ocean.
Nassau, Suffolk.
Brunswick, Carteret, Currituck, Dare,
Hyde, New Hanover, Onslow, Pender.
Newport, Washington.
Beaufort, Charleston, Colleton,
GeorgetowTi, Horry.
Aransas, Brazoria, Calhoun, Cameron,
Chambers, Galveston, Jefferson,
Kenedy, Kleberg, Matagorda, Nueces,
Willacy.
Accomack, Northampton.
Georgia:
Louisiana:
(parishes)
Maryland:
Massa-
chusetts:
Mississippi
New Jersey
New York:
North
Carolina:
Rhode
Island:
South
Carolina:
Texas:
Virginia:
HABITAT
The ridley inhabits shallow coastal and es-
tuarine waters; it is often associated with sub-
tropical shorelines of red mangrove [Rhizophora
mangle) (Witham 1976).
FOOD AND FORAGING BEHAVIOR
The ridley 's diet consists primarily of inverte-
brates, mostly crabs {Arenaeus, Calappa, Callinec-
tes, and Hepatus), but also shrimp, snails, sea ur-
chins, sea stars, fish, and, occasionally, marine
plants (Ernst and Barbour 1972, Pritchard and
Marquez M. 1973, R. Marquez M. personal
communication).
WUT LOHCITUDC
Distribution of Kemp's ridley sea turtle in the eastern United States.
Distribution (shading) and nesting sites (dots) of the Kemp's ridley sea turtle in the western United States.
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
With the exception of occasional females
known to nest on Padre Island, Texas (Werler
1951), the entire population nests on about 24
km of beach between Barra del Tordo and Ostio-
nal in State of Tamaulipas, Mexico. They prefer
sections of beach backed up by extensive swamps
or large bodies of open water having seasonal,
narrow ocean connections (Pritchard and Mar-
quezM. 1973).
A well-defined and elevated dune area is ne-
cessary for successful nesting. Pritchard and Mar-
quez M. (1973) suggest that this provides a land-
mark for the turtle to dig a nest that will be above
mean high tide.
RITUAL REQUIREMENTS
Not Known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Females land in large numbers only when
strong or moderate north winds blow. Pritchard
(1976) suggests that this may cover the turtle's
tracks and/or dissipate nesting smell.
POPULATION NUMBERS AND TRENDS
In 1947, a Mexican, Sr. Herreva, filmed nest-
ing. Estimates based on this film put the breeding
population at 40,000 at that time. Despite the
protection of the nesting beach since 1966, the
breeding population now is reported to be be-
tween 1,500 and 3,000 adults, with only about
500 females nesting during the 1978 season
(Anon. 1978).
The 1947 film shows what is believed to be
the entire breeding population swarming ashore at
once. Such a massive landing is called an 'arriba-
da.' R. Marquez M. (personal communication) has
observed nesting since 1966 and reports that the
number within arribadas is declining. In 1976, the
largest was made up of approximately 150 fe-
males, and there are usually between five and
seven arribadas per season.
Failure to rebuild population numbers in spite
of beach protection may result from low survivor-
ship (Pritchard 1976).
REPRODUCTION
Ridleys nest from April to June, during which
time turtles appear off Tamaulipas. After strong
winds, females swarm ashore to nest in daylight
hours. A female nests a maximum of three times a
season with an intemesting interval of 10 to 28
days. Individuals often nest annually with an ave-
rage clutch size of 110 eggs (Pritchard 1969a,
Lund 1976).
Copulation takes place offshore near the
nesting beach, and some pairs remain embraced
for hours. Black-and-white photographs of court-
ship and mating activities appear in Bustard
(1973).
MANAGEMENT AND CONSERVATION
Since 1966, the Mexican government has con-
ducted a tagging program for adults at Rancho
Nuevo and has protected the beach with military
personnel. About 330 clutches of eggs have been
relocated annually to fenced compounds (R. Mar-
quez M. personal commnication).
Pritchard (1976) argued for the perfection
and deployment of a modified trawl net to pre-
vent the incidental catch of sea turtles, and work
on this project is well underway by the U.S.
National Marine Fisheries Service.
Lund (1974) reported on a private effort to
stock Texas waters with eggs taken from the Ran-
cho Nuevo beach in Mexico. In 1978, a multi-
agency effort was initiated by the U.S. Fish and
Wildlife Service, National Park Service, National
Marine Fisheries Service, and Texas Parks and
Wildlife Department in coordination with the
Mexican government (Anon. 1978). This program
incorporates a wide variety of techniques, in-
cluding more intensive protection for the Rancho
Nuevo beach, an attempt to establish a breeding
population at Padre Island National Seashore by
transplanting 2,000 eggs annually, and head-
starting of 2,000 hatchlings from the Rancho
Nuevo beach and the hatchlings from Padre Island
at the NMFS laboratory in Galveston, Texas
(Anon. 1978,Wauer 1978).
Marquez M. (1976b) recommends formation
of seven natural reserves for Mexican coasts,
which include Playa de Rancho Nuevo, Tamauli-
pas, the Kemp's ridley's nesting beach..
AUTHORITIES
Archie Carr
Department of Zoology
University of Florida
GainesvUlcFL 32611
Henry Hildebrand
Department of Zoology
Texas A&M University
Kingsville.TX 78363
Frank Lund
P.O. Box 541
Jupiter Island, FL 33458
Rene Marquez M.
Apdo Postal 79-052
Col. Doctares
Mexico 7. D.F.
Peter Pritchard
Florida Audubon Society
P.O. Drawer 7
Maitland.FL 32751
PREPARER'S COMMENTS
None.
LITERATURE CITED/SELECTED
REFERENCES
Adams, D. E. 1966 More about the ridley opera-
tion: Padre Island egg transporting. Int. Turt.
and Tort. Soc. J. l(l):18-20, 40-43, 45.
Anon. 1978. U.S. -Mexico restoration efforts may
be only hope for Kemp's ridley. End. Sp.
Tech. Bull, U.S. Fish Wildl. Serv. 3(10): 6-8.
Brongersma, L. D. 1968. Miscellaneous note, on
turtles. I. Kon. Ned. Akad. Wetensch. Proc.
Serv. C. Biol. Med. Sci. 71:439-442.
Bustard, R. 1973. Sea turtles: natural history and
conservation. Taplinger Publ. Co., New York.
220 pp.
Carr, A. F. 1952. Handbook of turtles. Turtles of
the United States, Canada, and Baja California.
Comstock Publ. Assoc, Cornell Univ. Press,
Ithaca, N.Y. 542 pp.
. 1955. The riddle of the ridley. Anim.
Kingd. 58(5): 146-156.
. 1956. The windward road. Alfred Knopf.
Inc.,New York. 258 pp.
. 1961. The ridley mystery today. Anim.
Kingd. 64:7-12.
. 1967. So excellent a fishe. The Natural
History Press, Garden City, N. Y. 248 pp.
Chavez. H. 1969. Tagging and recapture of the
lora turtle {Lepidochelys kempii). Int. Turt.
and Tort. Soc. J. 3(4):14-19, 32-36.
Chavez, H., M. Contreras G., and T. P. E. Hernan-
dez D. 1968. On the coast of Tamaulipas,
Parts I and II. Int. Turt. Tort. Soc. J. 2(4) :20-
29, 37and2(5):19,27-34.
Ernst, C. H., and R. W. Barbour. 1972. Turtles of
the United States. Univ. of Kentucky Press,
Lexington. 347 pp.
lUCN. 1968. Red data book. Vol. 3, Amphibia
and reptilia. Compiled by R. E. Honegger.
lUCN, Morges, Switzerland.
Lund, F. 1974. A survey of marine turtle nesting
in the United States. Unpubl. 39 pp.
. 1979. Atlantic ridley. Pages 25-26 in R.
W. McDairmid ed. Rare and endangered biota
of Florida, Vol. 3, Amphibians and reptiles.
Univ. Presses of Florida, Gainesville.
Marquez M., R. 1973. Instructivo para la protec-
cion de las tortugas marina. Serie Dioulgacion.
INP/SD 2:1-34.
. 1976a. Estado actual de la pesquina de
tortugas marina en Mexico, 1974. INP/SI
1-27.
. 1976b. Reservas naturales para la con-
servacion de las tortugas marinas en Mexico.
INP/SIi 83:1-22.
Marquez M., R., A. Villanueva O., and C. Pena-
flores S. 1976. Sinopsis de dates biologicos
sobre la tortuga golfma. Lepidochelys oliva-
cea (Eschscholtz, 1829). INP Sinop. Pesca 2:
1-61.
Montoya, A. E. 1969. Programas de investigacion
y conservacion de las tortugas marinas en
Mexico. Pages 34-53 in Marine turtles. lUCN.
New Publ. Ser. Suppl. Paper. 20.
Pritchard, P. C. H. 1969a. Studies of the systema-
tics 3S\A reproductive cycles of the genus Lepi-
dochelys. Ph. D. Dissert. Univ. of Florida,
Gainesville. 226 pp.
. 1969b. Summary of world sea turtle sur-
vival situation. lUCN Bull. 2(1):90-91.
. 1969c. The survival status of ridley sea
turtles in American waters. Biol. Cons. 2(1):
13-17.
. 1976. Endangered species: Kemp's ridley
turtle. Florida Nat. 49(3): 15-19.
Pritchard, P. C. H., and R. Marquez M. 1973.
Kemp's ridley turtle or Atlantic ridley Lepi-
dochelys kempii. lUCN Monogr. 2. Marine
turtle series. Morges, Switzerland.
Rebel, t. P. 1974. Sea turtles and the turtle indus-
try of the West Indies, Florida, and the Gulf
of Mexico. Rev. ed. Univ. of Miami Press,
Coral Gables. 250 pp.
Svkreat, D. E. 1968. Capture of a tagged ridley tur-
tie. Q. J. Florida Acad. Sci. 31(l):47-48.
U.S. Department of Commerce. 1976. Proposed
listing of the green sea turtle {Chelonia my-
das), loggerhead [Caretta caretta), and pacific
ridley sea turtle [Lepidochelys olivacea) as
threatened species under the Endangered
Species Act of 1973. Natl. Oceanic Atm. Ad-
min., Natl. Marine Fish. Serv., Washington,
D.C.
Walker, W. F. 1971. Swimming in sea turtles of
the family Cheloniidae. Copeia 1971(2):229-
233.
Wauer, R. 1978. Headstart for endangered turtles.
Natl. Parks Conserv. Mag. 52(11): 16-20.
Werler, J. E. 1951. Miscellaneous notes on the
eggs and young of Texas and Mexican reptiles.
Zoologica36(3):37-48.
Witham, R. 1976. Lepidochelys kempii account.
In H. O. Hillestad, D. B. Means, and W. W.
Baker, eds. Endangered and threatened verte-
brates of the southeastern United States. Tall
Timbers Res. Stn. Misc. Publ. 4.
Zim, H. S., and H. M. Smith. 1953. Reptiles and
amphibians. A guide to familiar American spe-
cies. Simon and Schuster, New York. 147 pp.
GATEKEEPER
To be designated by the Office of Endangered
Species.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
GainesvUle, FL 32601
Biological Services Program
FWS/OBS-80/01.31
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
BACHMAN'S WARBLER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-SHdell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.31
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
BACHMAN S WARBLER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-SIidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
DWG. BY WALTER DAWN
BACHMAN'S WARBLER
Vermivora bachmanii Audubon
KINGDOM Animalia
CLASS Aves
ORDER Passeriformes
FAMILY Parulidae
OTHER COMMON NAMES None
DATE
Entered into SWIS To be determined.
Updates 5 April 1978; 10 October 1978.
LEGAL STATUS
Federal: Endangered: 32 FR 4001; 11 March
1967.
States: Endangered: Florida, Georgia, Missis-
sippi, North Carolina, South Carolina,
Texas.
REASONS FOR CURRENT STATUS
Stevenson (1938), USDI (1973), Shuler
(1977b, 1977c, personal communication) and
Arthur Wayne's (approximately 1910) unpublish-
ed statement, "I never expect to see another spec-
imen as the great swamp in which I found it
breeding has been deforested by a lumbering
company," all suggest that the clearing of the
southern swamp forests contributed to the de-
cline of the Bachman's warbler. On the other
hand, Stevenson (1972) and personal communica-
tions from P. Hamel, R. Hooper, and D. Urbston
insist that the reasons for the current status of
this species are unknown.
Other suggestions for the species' current
status have included: (1) competition with other
species on the breeding grounds (Hamel 1977a);
(2) excessive collecting along migration route in
Florida (USDI 1973); (3) climatic catastrophe
such as storms (R. Hooper personal communica-
tions); and (4) "genetic or evolutionary probl-
blems" (Hamel 1977a: R. Hooper personal com-
munications).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The Bachman's warbler is 10 to 11 cm long,
with a small, sharp-pointed bill. Their color
pattern is somewhat variable. Males usually have a
bright yellow shoulder patch, forehead, eye ring,
chin, and belly and a black throat patch with
some black on crowm. Crown is otherwise gray
and black olive. Wings and tail aie dusky with
subterminal white patches on tail. Females
usually lack black on throat (although some may
be present); the breast is shaded with gray; white
markings on tail are reduced, and underparts are
often much less yellowish.
Distinguishing field marks are the yellow un-
derparts and the black crown and throat. Color il-
lustrations appear in Howell (1932), the Decem-
ber 1977 cover of Birding, £ind the inside back
cover of South Carolina WUdlife 23(2), 1976.
RANGE
Bachman's warblers are known to have nested
in Alabama, Kentucky, Missouri, and South
Carolina (Hooper and Hamel 1977). The last nest
known in Alabama was discovered in 1937
(Stevenson 1938).
Their present range is unknown; the species
may be extinct (Hamel 1977b). Shuler (1977a)
reported sightings of individual males in South
Carolina's I'On Swamp (Francis Marion National
Forest) in 1974, 1975, and 1976, one in each
year. Shuler et al. (1978) reported sightings of a
male and a female in I'On Swamp in 1977, in-
cluding reported verification by subsequent
sightings by different individuals. Other recent
reported sightings include Berkeley County, S. C.
(1967), Louisiana (1973), Long County, Georgia
(1975), Kentucky (1977), Maryland (1977), and
Cameron Parish, Louisiana (1977). Hamel
(1977b) considers all of these sightings as uncon-
firmed and D. Urbston (personal communication)
considers them all 'questionable.'
The last sighting accepted by Hamel (1977b)
as valid was in Charleston County, S. C. in 1962.
If the Bachman's warbler still exists, most au-
thorities agree that it is most likely in the I'On
Swamp area in Charleston and Berkeley Counties,
South Carolina.
RANGE MAP
Not provided.
STATES/COUNTIES
South Carolina: Berkeley (?), Charleston (?).
HABITAT
Nested in bottomland hardwood swamps.
FOOD AND FORAGING BEHAVIOR
Five stomach samples contained caterpillars
and Hymenoptera (Meanley and Mitchell 1958).
SHELTER REQUIREMENTS
Dense understory of shrubs and vines within
hardwood swamps.
NESTING OR BEDDING
They nest in low bushes or vines, 0.7 to 1 m
off the ground (Wayne 1907), in nests con-
structed of dried weed and grass stalks and dead
leaves, lined with black threads of pendent lichens
{Ramalina sp.) (Widmann 1897).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
Formerly locally abundant (Widmann 1897;
Embody 1907). Formerly abundant during migra-
tion along the Suwannee River, Florida (Brewster
1891).
Present population, if any, unknowoi.
REPRODUCTION
Eggs are laid March through June with three
to five eggs per set (Wayne 1907).
MANAGEMENT AND CONSERVATION
Shuler (1977b, 1977c, personal communica-
tion) suggests discontinuing of harvesting of bot-
tomland hardwoods in the I 'On Swamp, Francis
Marion National Forest, South Carolina. R.
Hooper (personal communication) suggests in-
creased cutting to open up the canopy and thus
allow secondary growth of shrubs and vines.
Extensive studies by U.S. Forest Service biol-
ogists Urbston and Hooper and Clemson Universi-
ty zoologist Hamel are underway to better define
the status and requirements of the Bachman's
warbler.
AUTHORITIES
Paul B. Hamel
Department of Zoology
Clemson University
Clemson, SC 29631
Robert G. Hooper
U. S. Forest Service
Southeastern Forest Experiment Station
Forestry Sciences Laboratory
2730 Savannah Highway
Charleston, SC 29407
Jay Shuler
P.O. Box 288
McCleUanville, SC 29458
David F. Urbston
USDA Forest Service
Ouachita National Forest
Hot Springs, AR 71901
PREPARER'S COMMENTS
None.
LITERATURE CITED/SELECTED
REFERENCES
Barnes, I. R. 1954. A new look at Bachman's
warblers. Atlantic Nat. 10: 18-30.
Bent, A. C. 1953. Life histories of North Ameri-
can wood warblers. U.S. Natl. Mus. Bull. 203.
734 pp.
Brewster, W. 1891. Notes on Bachman's warbler
{Helminthophila bachmanii). Auk 8:149-157.
Chamberlain, G. B. 1958. Bachman's warbler in
South Carolina. Chat 22:73-74, 77.
Embody, G. C. 1907. Bachman's warbler breeding
in Logan County, Kentucky. Auk 24:41-42.
Evenden, F., D. Marshall, and W. Zeedgk. 1977.
Revised final report of the Bachman's warbler
Panel. 30 pp.
Hamel, P. B. 1977a. Problem analysis for research
to determine the status of Bachman's warbler,
and the evolutionary, ecological, and histori-
cal factors associated with its decline. U.S.
For. Serv. Southeast. For. Exper. Stn.,
Charleston, SC. Contract 18-409. 25 pp.
. 1977b. Progress report, research to de-
termine the status of Bachman's warbler, the
ecological, evolutionary, and historical factors
responsible for its decline. U.S. For. Serv.
Southeast, For. Exper. Stn., Charleston, SC.
Contract 18-409. 24 pp.
Hamel, P. B., and R. G. Hooper. In press. The
status of Bachman's warbler, a progress re-
port. Proc. Endangered Species Symp.,
Athens, Ga.
Hamel, P. B., R. G. Hooper, D. Urbston, and A.
McDonald. 1977. Bachmem's warbler breeding
habitat: A hypothesis. Presented before 95th
Annu. Mtg. Am. Omithol. Union, Berkeley,
Ca., 25 August 1977.
Hamel, P. B., R. G. Hooper, and L. M. Wright.
1976. Where is the Reverend Bachman's war-
bler? S.C. WUdl., March-April 1976: 9-13.
Holt, G. G. 1920. Bachman's warbler breeding in
Alabama. Auk 37:103-104.
Hooper, R. G., and P. B. Hamel. 1977. Nesting
habitat of Bachman's warbler: A review. Wil-
son Bull. 89(3):373-379.
Howell, A. H. 1910. Notes on the birds of the
sunken lands of southeastern Missouri. Auk
27:381-384.
Howell, A. H. 1932. Florida bird life. Coward-
McCann, Inc., New York. 597 pp.
Meanley, B. and R. T. Mitchell. 1958. Food hab-
its of Bachman's warbler. Atlantic Nat.
236-238.
Shuler, T. 1977a. Three recent sight records of
Bachman's warbler. Chat 41(1):11-12.
. 1977b. Bachman's phantom warbler.
Birding9(6): 245-250.
. 1977c. Bachman's warbler habitat. Chat
41(2):19-23.
Shuler, T., P. Nugent, J. Trochet, and J. Vanos.
1978. Bachman's warbler observations contin-
ue in the I'On Swamp. Chat 42(2):23-24.
Stevenson, H. M. 1938. Bachman's warbler in Ala-
bama. Wilson Bull. 50:36-41.
1972. A recent history of Bachman's
warbler. Wilson Bull. 84:344-347.
U. S. Department of the Interior. 1973. Threat-
ened Wildlife of the United States. Compiled
by Office of Endangered Species and Inter-
national Activities, Bur. Sport Fish. Wildl.,
Resour. Publ. 114. U.S. Gov. Printing Office,
Washington, D.C. 289 pp.
Wayne, A. T. 1907. The nest and eggs of Bach-
man's warbler, Helminthophila bachmanii
(Aud.), taken near Charleston, South Caro-
lina. Auk 24:43-48.
Widman, O. 1897. The summer home of Bach-
man's warbler no longer unknown. Auk 14:
305-309.
Biological Services Program
FWS/OBS-80/01.32
March 1980
Selected Vertebrate
Endangered Species
Of the Seacoast of the
United States-
THE FLORIDA
EVERGLADE KITE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data are not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.32
March 1980
SELECTKD VLR 1 EBRA 1 K ENDANGERED SPECIES
OF THE SEACOAS r OF I HE UNITED STATES-
THE FLORIDA EVERGLADE KITE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
FLORIDA EVERGLADE KITE
Rostrhamus sociabilis plumbeus Ridgway
KINGDOM Animalia
CLASS Aves
ORDER Falconiformes
FAMILY Accipitridae
OTHER COMMON
NAMES. . . . Everglade kite, snail kite, Florida
snail kite, snail hawk.
DATE
Entered into SWIS to be determined
Updates 15 October 1976, 13 April 1977
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 March
1967).
States: Endangered: Florida.
REASONS FOR CURRENT STATUS
Reduction of suitable habitat due to drainage
of marshes is the primary reason for the kite's en-
dangered status. By 1967, only 3,981 km^ of the
original 7,112 km^ of Everglades remained un-
drained (Stieglitz and Thompson 1967, U.S. De-
partment of the Interior 1973, Sykes 1979).
Droughts and water management have also
affected kite habitat by reducing populations of
apple snails (Pomacea paludosa), the kite's only
food source (U.S. Department of the Interior
1973).
The introduced water hyacinth [Eichornia
crassipes) now covers the water surface in many
areas, making it impossible for kites to locate
apple snails (Sykes 1979).
Excessive human disturbance and malicious
killings have also contributed to population de-
clines (Stieglitz and Thompson 1967, Sykes
1979).
Potential factors of unknovwi importance in-
clude parasites transmitted by snails, weather pat-
tern changes, and inbreeding due to reduced
population size (Stieghtz and Thompson 1967).
Pesticide concentrations in snails, kites, and
kite eggs are very low and probably do not pre-
sent a threat (Steiglitz and Thompson 1967).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The kite is a medium-sized hawk, 41 to 46 cm
long, with a wingspan of 114 cm. Its beak is slen-
der and strongly hooked. Adult males are slate
gray with black head and wing tips, and a white
patch at the base of the tail. Legs are orange-red.
Females and immatures are buffy, heavily streaked
vsath dark lines, with a white rump and yellow
legs.
RANGE
The species ranges wddely in the Neotropics,
with three recognized subspecies. The Florida
race is restricted to peninsular Florida and Cuba.
It formerly ranged over much of Florida,
wherever suitable habitat occurred, including the
following counties: Brevard, Broward, Collier,
Dade, Flagler, Glades, Hendry, Highlands, Indian
River, Jefferson, Lake, Monroe, Okeechobee, Or-
ange, Osceola, Palm Beach, St. Lucie, Seminole,
Sumter, Volusia, and Wakulla (Sykes 1979).
It is now restricted to the headwaters of the
St. Johns River, the southwest side of Lake Okee-
chobee, a portion of Everglades National Park, a
portion of the Loxahatchee National Wildlife Re-
fuge, and small areas in Broward, Dade, and Palm
Beach Counties (Sykes 1979).
RANGE MAP
Present distribution is adapted from Sykes
(1979).
STATES/COUNTIES
Florida Broward, Dade, Glades, Indian River,
Palm Beach, St. Lucie.
HABITAT
The species is typically found in fresh-
water marshes with a distant horizon. Favorable
areas consist of shallow open water vegetated
with sawgrass [Cladium jamaicensis) and spike-
rushes {Eleocharis sp.). Flats, often interspersed
with tree islands or small groups of shrubs and
trees including dahoon holly (Ilex cassine), coastal
plain willow [Saltx caroliniana), wax myrtle [My-
rica cerifera), and buttonbush [Cephalanthus occi-
dentalis). Other common aquatics include water-
lily [Nymphaea odorata), big floating heart [Nym-
phoides aquatica), maidencane {Pantcum hemito-
mon), bulltongue [Sagittaria lancifolia), pickerel-
weed [Pontederia lanceolata), cattail [Typha do-
mingensis), waterlettuce [Pistia stratiotes), and
water hyacinth (Stieglitz and Thompson 1979).
Water levels may fluctuate so long as the sur-
face does not completely dry (Sykes 1979). Habi-
tat photographs appear in Stieglitz and Thompson
(1967), Snyder and Snyder (1969), and Sykes
(1979).
FOOD AND FORAGING BEHAVIOR
Snail kites feed only on the freshwater apple
snail. They fly 1.5 to 9 m above the surface, de-
scending when a snail is sighted. Snails are grasped
with a single talon and are usually transferred to
the beak in flight. They also sometimes hunt from
perches. Snails are extracted and the shells dis-
carded.
Captives will survive on horse meat and will
take other species of snails. Their dependence up-
on apple snails in nature probably results from
this snail's occurrence near the water's surface,
unlike other large snails (Snyder and Snyder 1969).
SHELTER REQUIREMENTS
Not known.
NORTHWESTERN PART OF
FLORIDA
Z 3
e 9 10
la 13 1* 15 16 n 18 19 » ai 22 23 2« !^
The present distribution of the Everglades kite in Florida is depicted by shading.
NESTING OR BEDDING
Kites normally nest in loose aggregations,
with 150 m or less between nests (Howell 1932,
Bent 1937, Stieglitz and Thompson 1967). They
often nest among or near other colonial nesters
such as herons and egrets (Ardeidae) and An-
hingas (Anhtnga anhinga).
Nests are usually 1 to 3 m above the water in
low trees or shrubs, although occasionally sawgrass
or cattails are used for support (Howell 1932,
Stieghtz and Thompson 1967, Sykes 1976). In
the Loxahatchee marshes, nests are built in small
cypress trees {Taxodium sp.) 2 to 3 m above the
surface.
The nest is loose and bulky, 30 to 38 cm in
diameter, and 20 to 30 cm deep. It is built of
twigs and Spanish moss (Tillandsia usneoides)
and lined with green grasses and vines (Bent
1937).
RITUAL REQUIREMENTS
An important part of the courtship display
consists of the male carrying a small twig in his
beak while flying and making a series of swooping
dives wdth the wings folded (Stieglitz and Thomp-
son 1967).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Adequate water levels are essential, as snail
populations are drastically reduced when the
marsh dries.
POPULATION NUMBERS AND TRENDS
Snail kite numbers have dropped drastically.
Bent (1937) stated that kites were common in
1909, but that population levels had declined
sharply by 1937, due to the draining of the Ever-
glades. Estimates for 1973 were slightly over 100
individuals (Sykes 1979).
The total population of Florida Everglade
kites has probably averaged close to 100 birds for
the past 4 to 5 years, with 80% of the reproduc-
tion occurring on the southwest side of Lake
Okeechobee (T. Martin personal communication).
Seventeen young were fledged from 7 of 9
nests in 1968 (U.S. Department of the Interior
1973).
REPRODUCTION
The breeding season is variable and probably
correlated vwth snail availability, rainfall, and/or
temperature; generally, from February to June
(Howell 1932, Bent 1937). Average clutch size is
two to four. Eggs are oval or elliptical and smooth,
colored dull white with brown mottling (Bent
1937); average size is 44.2 by 36.2 mm (Bent
1937).
The male builds the nest, and both sexes incu-
bate eggs and assist in rearing young (Bent 1937).
Courtship displays consist of soaring about
150 m above the marsh, folding the wings for sud-
den short dips, stretching the legs while diving as
if to grapple, somersaulting while diving, and
stick-carrying (Stieghtz and Thompson 1967).
MANAGEMENT AND CONSERVATION
In a 2-year habitat management study by
Martin and Doebel (1973), water levels were raised
on a 142-ha site in the Loxahatchee National
Wildlife Refuge; plant density was controlled and
snails were stocked. The first two techniques were
successful, but stocking of snails had little effect
on population level. Other methods will be imple-
mented in a projected second experiment. Arti-
ficial nesting platforms have been successfully
used to support nests that would otherwise be
subject to damage from high winds or heavy
rains (Sykes and Chandler 1974).
Kite response to management efforts was
rapid and extremely favorable. Martin and Doe-
bel (1973) concluded that continued research in-
to life history of both snail and kite is critical to
management success. They believe the kite can re-
main a part of our natural fauna only through
sound management practices and maintenance of
natural habitats. The American Ornithologists'
Union (1975) states that the only solution for
saving the kite is a very expensive land-aquisition
program. Sykes (1979) makes several recommen-
dations for preservation of the Everglade kite.
These include purchasing and managing suitable
kite habitat, providing protection during the
hunting season, establishing a 0.4 km buffer zone
closed to entry around nesting areas, maintaining
adequate surface water in the kite areas, and re-
searching both kite and apple snail biology.
Critical habitat has been designated (42 FR
47841, 22 September 1977) as parts of Dade,
Broward, and Palm Beach counties and the west-
em shore of Lake Okeechobee in Glades County.
AUTHORITIES
Thomas Martin
Loxahatchee National Wildlife Refuge
Route l,Box 278
Delray Beach, FL 33444
4
Alexander Sprunt, IV
Natl. Audubon Society
115 Indian Mound Trail
Tavemier, FL 33070
PaulW. Sykes.Jr.
U.S. Fish and Wildlife Service
P.O.Box 2077
Delray Beach, FL 33444
PREPARERS COMMENTS
None.
Sykes, P. W., Jr., and R. Chandler. 1974. Use of
artificial nest structures by Everglade kites.
Wilson Bull. 86:282-284.
U.S. Department of the Interior. 1973. Threat-
ened wildlife of the United States. Compiled
by Office of Endangered Species and Inter-
national Activities, Bur. Sport Fish. Wild!.
Resour. Publ. 114. U.S. Govt. Printing Office,
Washington, D.C. 289 pp.
LITERATURE CITED/ SELECTED
REFERENCES
Amadon, D. 1975. Variation in the Everglade kite.
Auk 92:380-382.
American Ornithologists' Union. 1975. Report of
the committee on conservation. Auk 92 (4,
Suppl.):lB-16B.
Bent, A. C. 1937. Life histories of North Ameri-
can birds of prey. Part I. U.S. Natl. Mus. Bull.
267.409 pp.
Howell, A. H. 1932. Florida bird life. Coward-
McCann, Inc. New York. 527 pp.
Martin, T. W., and J. H. Doebel. 1973. Manage-
ment techniques for the Everglade Kite, pre-
liminary report. Proc. S.E. Assoc. Game Fish
Comm. Annu. Conf. 27:225-236.
Peterson, R. T. 1947. A field guide to the birds.
Houghton Mifflin, Boston. 230 pp.
Snyder, N. F. R., and H. R. Snyder. 1969. A com-
parative study of mollusk predation by limp-
kins, Everglade kites, and boat-tailed grackles.
Living Bird 8:177-223.
Sprunt, A.,Jr. 1945. The phantom of the marshes.
Audubon Mag. 47:15-72.
. 1954. Florida bird Ufe. Coward-McCann,
Inc., and Natl. Audubon Soc, New York. 527
pp.
Stieglitz, W. O., and R. L. Thompson. 1967.
Status and life history of the Everglade kite in
the United States. Bur. Sport Fish. Wildl.
Spec. Sci. Rep. Wildl. 198. 21 pp.
Syke%, P. W., Jr. 1979. Florida Everglade kite.
Pages 4-7 in H. W. Kale ed. Rare and endan-
gered biota of Florida, Vol. 2, Birds. Univ.
Presses of Florida, Gainesville.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.33
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
PUERTO RICAN WHIP POOR WILL
>'4X
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensiti\c Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.33
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
PUERTO RICAN WHIP POOR WILL
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastsd Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
PUERTO RICAN WHIP-POOR-WILL
Caprimulgus noctitherus (Wetmore)
KINGDOM Animalia
ORDER Caprimulgiformes
FAMILY Caprimulgidae
OTHER COMMON
NAMES Pajaro Bobo (Reynard 1962);
Guabairo Pequeno de Puerto Rico (Biaggi
1979 and Leopold 1963); Guabairo Chico
and Guaraiba (Bond 1936 and Wetmore
1916); Puerto Rican Nightjar (Leopold
1963).
DATE
Entered into SWIS To be determined
Updates To be determined
LEGAL STATUS
Federal: Endangered (42 FR 36428, 14 July
1977). Protected by Migratory Bird
Treaty Act of July 2, 1918 (40
Stat. 755; 16 U.S.C. 703-711) as
amended Dec. 3, 1969. Public Law
91-135.
Puerto Rico: Listed as "very endangered."
Commonwealth "Wildlife Law"
protects ail nongame bird species
including Puerto Rican Whip-poor-
WUl.
REASONS FOR CURRENT STATUS
Wetmore (1927), Wadsworth (1949), and
Wolcott (1953) assumed that the exotic mongoose
[Herpestes jarvanicus), introduced in 1877, had
stripped the whip-poor-will population from
much its former more extensive range. Some of
this former range still has habitat that appears
suitable for whip-poor-wills, but also has rainfall
and standing water sufficient to support mon-
gooses.
At present, lowland semiarid scrubby forest
habitat is being reduced by expanding human
population and developing industries. A rapidly
expanding petrochemical industry is adjacent to
the Guayanilla whip-poor-will population, only 8
km east of the Guanica Forest which harbors the
largest number of the remaining birds. The Com-
monwealth Forest system is not immune to in-
dustrial pressure and over 400 hectares were des-
troyed between 1968 and 1973 (Kepler and
Kepler 1973).
PRIORITY INDEX
33
DESCRIPTION
The Puerto Rican whip-poor-will is a small
ground-inhabiting nocturnal bird with fluffy,
mottled brown, black, and gray plumage, a very
short bill, and very wide mouth with long bristles.
There is a white band across the throat and white
spots at the ends of the taU feathers. It is similar
to the North American whip-poor-will {Caprimul-
gus vociferus) but smaller, with much shorter
wings, and plumage tending more toward brown
and less toward gray throughout. Buff breast spots
are more prominent; posterior underparts are
deep cinnamon buff rather than pale buff. White
or buff tips on three outer tail feathers are greatly
restricted. The type specimen has the following
measurements: wing 135 mm; tail 112 mm; bill
11 mm; tarsus 16.3 mm; middle toe without claw
16 mm (Wetmore 1919 and Wetmore in Reynard
1962; J. W. Aldrich personal observations 1977).
Downy young are entirely reddish buff or cinna-
mon. Toes and legs are flesh gray. The iris is dark
brown. Eyes open on the day of hatching (Kepler
and Kepler 1973).
Authorities differ as to whether noctitherus is
a distinct species or a subspecies of the North
American whip-poor-will (C. v. vociferous). Wet-
more (1919, 1922, 1927), Vincent (1966), Mayr
and Short (1920), Fish and Wildlife (1973), Kep-
ler and Kepler (1973), and Storrs Olson (pers.
comm. 1977) prefer to consider it a distinct spe-
cies, whereas Peters (1940), Bond (1945), and
Greenway (1958) consider it a subspecies of C. v.
vociferous. Vocalizations of nocfzi/ierMX, described
by Reynard (1962), are completely different in
structure from vociferous and may serve to main-
tain reproductive isolation between two closely
related species (Kepler and Kepler 1973). Storrs
Olson (pers. comm. 1977) is impressed by the
much smaller bones of noctitherus. The question
of correct classification has no completely satis-
factory answer, but the preponderance of current
thinking seems to favor classification as two dis-
tinct species.
RANGE
This species is confined to the island of Puerto
Rico. It was formerly found in the moist limestone
forest of the northwest coast. Specimens were
taken in Bayamon in 1888 (Cory 1889, Wetmore
in Reynard 1962, Peters 1940). Bones found in
cave deposits near Morovis are estimated to be
less than 2,000 years old (Wetmore 1919, 1922,
1927). A bird presumed to be this species was
sighted at Rio Piedras (Wetmore 1916, 1919,
1927). AU three of those localities were in the
moist northwestern section of the island. More
recent records are all from the dry limestone for-
est of the southwest coast, including two popula-
tions at Guanica and the hills above Guayanilla
and one in Susua Commonwealth Forest. It prob-
ably once ranged over most of the coastal plain
on both sides of the island, but now is probably
confined to an area of about 3,200 hectares, or
approximately 3% of its probable former range,
the limestone regions, and only 0.7% of the total
land surface of the island (Kepler and Kepler
1973).
Distribution of the Puerto Rican whip-poor-
will has remained stable since 1969 (American
Ornithologists' Union 1976).
MAP
Past and present distribution (after Kepler
and Kepler 1973) is shown on the following page.
STATES/COUNTIES
Commonwealth of Puerto Rico:
HABITAT
Habitat formerly included the moist limestone
forest that covered most of the northwestern
third of the island north of the Cordillera Central
described by Little and Wardsworth (1964) and
Kepler and Kepler (1973). Probably once occupied
the dry limestone forest on southwestern part of
the island as well, since they have been heard by
residents there at least since 1900 (Reynard 1962).
At present, they are known tp occur only in the
dry limestone forests of the southwestern area.
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The species frequents the drier, more open sec-
tions of the Susua Forest and has not been found
in the riparian forests or the denser scrub at eleva-
tions above 350 m (Kepler and Kepler 1973).
The Guanica forest, where whip-poor-wills occur
most commonly, has an annual rainfall of 750
mm, contrasting with an annual water loss through
evaporation and plant transpiration of 2,000
mm. The porous limestone permits no standing or
running fresh water anywhere. Elevations range
from sea level to 230 m. Semideciduous vegeta-
tion consists of hardwood trees on top of Ume-
stone hills, normally above 75 m. Below this, par-
ticularly on the southern slopes, forest changes
to mixed cactus and semideciduous growth. The
canopy is 4 or 5 m high, with some trees 6-7 m.
Important species include Pisonia alba, Swietenia
mahogoni. Acacia famesiana and Bursera sima-
ruba. There is no well-marked stratification or
epiphytic flora. Whip-poor-will population cen-
tered on higher slopes of central hills coincident
with semideciduous forest. They are largely ab-
sent from south-facing slopes below 75 m domin-
ated by cactus, open patches of grassland and
scrub, and thickets of Agave (Kepler and Kepler
1973).
Susua Commonwealth Forest is xeric scrub on
steep rounded hills of decomposed serpentine
soils. It receives 1500 mm of rain per year. The
elevation is 100 to 430 m. Two permanent streams
support a lush riparian plant life less than 100 m
in width. Scrub 4-5 m high predominates 50 or
more meters from streams and extends to hill
tops, where it becomes much denser. The Puerto
Rican whip-poor-wiU occurs here in small num-
bers but not in riparian habitat. Greatest numbers
occur in open scrub forest with horizontal visibil-
ity from 5 to 7 m. None are found in denser sec-
tions where visibility dropped to 3 m. Susua For-
est is considered suboptimal habitat for whip-
poor-wills based on only one pair in 14.1 hectares
as compared with one pair in no more than 6.9
hectares in Guanica Forest (Kepler and Kepler
1973).
FOOD AND FORAGING
The whip-poor-wUl captures flying insect prey
by flying from perches well above the
ground. It has favorite perches used regularly for
feeding in the evening and ventures from
forest canopy protection after dark to more
isolated trees for feeding perches (reported by
Cotte in Kepler and Kepler 1973).
SHELTER REQUIREMENTS
It requires the shelter of low scrubby forest
and undergrowth for daytime concealment and
nesting (Kepler and Kepler 1973).
NESTING AND BEDDING
The species nests on leaf litter of the forest
floor without additional nest material.
It roosts on the ground or on branches of
trees beneath cover of leafy canopy (Kepler and
Kepler 1973).
RITUAL REQUIREMENTS
Vocalizations are thought to proclaim terri-
tory possession and determine spacing of breeding
pairs (Kepler and Kepler 1973).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Predation by introduced mongoose and
domestic cats and dogs should be controlled ; also
overbrowsing of habitat by domestic livestock,
particularly goats, should be prevented.
POPULATION NUMBERS AND TRENDS
Subfossil bones were found in a cave near
Morovis (Wetmore 1919, 1922; Storrs Olson pers.
comm. 1977). Cory (1889) reported receipt of a
specimen later used as the type specimen. Wet-
more (1919, 1922, 1927; Wetmore in Reynard
1962) during 10 months field work in Puerto
Rico in 1911 and 1912 saw only one bird that he
thought might belong to this species. It was
thought to be extinct (Bond 1940; Greenway
1958) until rediscovered in 1961 near Guanica by
Reynard (1962) who collected one male and heard
six others nearby. Leopold (1963) estimated the
Guanica population as about 25 pairs. Bond (1962)
reported "numerous individuals heard on semi-
arid hills near the coast." J. B. McCandless esti-
mated that as many as 100 birds remained within
the Guanica Forest (Kepler and Kepler 1973).
During the period 1969-1971, total Guanica pop-
ulation was estimated at about 400 breeding pairs,
based on a density of one pair per 6.9 hectares to
one pair per 4.9 hectares (Kepler and Kepler
1973). Comparison with estimated densities of the
European whip-poor-will suggests that the Guanica
birds may be at maximum possible densities with-
in their limited range.
In the Susua Commonwealth Forest 10 km to
north of Guanica, and the hills behind Guayanilla
8 km to the east, the total population is estimated
at about 80 pairs or one pair per 14.1 hectares,
more than twice the area needed in Guanica Forest,
indicating that Susua forest is suboptimal for
whip-poor-wills (Kepler and Kepler 1973).
The total of all known populations is estimated
at 450 to 500 breeding pairs, over 80% of which
are in Guanica Commonwealth Forest.
The Puerto Rican whip-poor-will population
has remained stable since 1969 when the first ac-
curate data were obtained (American Ornitho-
logistists' Union 1976).
REPRODUCTION
Calling is at its peak in February in Guanica
Forest, suggesting that territorial encounters are
frequent and vigorous at that time, with possible
emigration of young or subordinate birds from
the central population (Kepler and Kepler 1973).
In a nest found 18 June 1969, 2 chicks were
estimated to be 14 or 15 days old. A nest with 2
eggs 1 May 1971, hatched on 4 and 5 May. A nest
with 1 egg, 1 July and 2 eggs 3 July, took 19 days
for incubation (McCandless report to Kepler and
Kepler 1973). Nesting occurs mid-April through
early July. May and June probably are peak nest-
ing months, but calling is maintained from Febru-
ary to May with low numbers calling in July in
Guanica Forest. There may be two broods per
year, although this has not been verified.
No nests are built; eggs are laid directly on
leaf litter under scrub vegetation with canopy 4
to 6 m high (never in open areas).
Young wander a short distance (20 cm) from
the nest on the third day after hatching and ir-
regularly farther thereafter. They are able to fly
on the 14th day.
Based on 3 nests, the clutch size is 2 eggs laid
on successive days. Eggs are buffy brown with
numerous brownish purple spots over the entire
surface, and are only moderately cryptic against
the substrate (Kepler and Kepler 1973) of dry
leaf litter. Protective coloration of the incubating
bird provides safety from predators. The in-
cubating bird remains on eggs even when closely
approached. When driven from nest or chicks,
adults engage in distraction display by flapping
wings, spreading tail and opening mouth while
on the ground (Kepler and Kepler 1973).
MANAGEMENT AND CONSERVATION
Although Susua and Guanica Commonwealth
Forests are nominally protected, the tremendous
industrial expansion to the east and possibly real
estate development to the west threatens them
both. These forests must be given additional
recognition and protection from human use and
domestic predatory and herbivorous animals, par-
ticularly goats, to assure the continued existence
of the Puerto Rican whip-poor-will (Fish and
Wildlife Service 1973; Kepler and Kepler 1973).
A recent favorable action has been the ruling
by the Environmental Quality Board that any
new industrial growth east of Guanica cannot be
accompanied by increased air pollution. Emissions
from existing oil refineries would have to decrease
to accomodate new growth. Also, an attempt to
use Guanica as a sanitary landfill site has been
temporarily shelved.
The most recent serious threat is a proposed
condominium complex on Punta Ballena, a pri-
vately owned tract 3.2 km south of the center of
Guanica Forest. Several thousand people would
be housed within easy walking distance of the
whip-poor-wills, necessitating road and commer-
cial development with the expected onslaught of
people, cats, rats, and mongooses and the danger
of fire (American Ornithologists' Union 1976).
The proposed sanitary landfill at Guanica
Forest and highway development along its
southern boundary remain threats (H. Rafael
pers. comm. 1977).
AUTHORITIES
Cameron B. Kepler and Angela K. Kepler
R. R. Maalaea Rd.
Kula, Hawaii 96709
Ricardo Cottee
U.S. Fish and Wildlife Service
Atlanta, Georgia
Warren King
Smithsonian Institution
Washington, B.C.
Dr. James B. McCandless
P.O. Box 1060
Mayaquez, P.R. 00709
George B. Reynard
105 Midway
Riverton, N.J. 08077
James Wiley
Box 21
Palmer, Puerto Rico 00721
Noel Snyder
Patuxent Wildlife Research Center
Laurel, Md. 20810
PREPARER'S COMMENT
Obviously, the preservation of habitat is the
key to survival of the Puerto Rican whip-poor-
wUl. Vigorous action to preserve the approxi-
mately 3,200 hectares of occupied habitat, and
particularly the 2,300 hectares of optimal habitat
in Guanica Forest, and prevention of human in-
terference there is the only way to save the spe-
cies from extinction.
LITERATURE CITED/ SELECTED
REFERENCES
American Ornithologists' Uniqp. 1976. Report of
the Committee on Conservation. Auk 93(4th
Suppl.): 1DD-19DD.
Biaggi, V. 1970. Las aves de Puerto Rico. Editorial
Universitaria. Universidad de Puerto Rico. pp.
230-231.
Bond, J. 1936. Birds of the West Indies. Acad.
Nat. Sci. Philadelphia.
Bond, J. 1940, 1945, 1962. Checklist of birds of
the West Indies. Acad. Nat. Sci. Philadelphia.
Cory,C. B. \889.Antrostomusvociferus in Puerto
Rico. Auk 6:276.
Fish and Wildlife Service. 1973. Threatened Wild-
life of the United States, 1973 Ed. U.S. Gov't.
Print. Off.
Greenway, J. C, Jr. 1958. Extinct and vanishing
birds of the world. Amer. Comm. Internat'l.
WildUfe Prot. Spec. Publ. 13, New York.
Kepler, C. B. and A. K. Kepler. 1973. The distri-
bution and ecology of the Puerto Rican whip-
poor-will, an endangered species. Living Bird,
Eleventh Annual Report Cornell Lab. Omith.
Leopold, N. F. 1963. Checklist of birds of Puerto
Rico and the Virgin Islands. Bull. 168 Univ.
Puerto Rico Agric. Exp. Sta., Rio Piedras.
Little, E. L. and F. H. Wadsworth. 1964. Com-
mon trees of Puerto Rico and the Virgin Is-
lands. U.S. Dept. of Agric. Handbook 249:
1-548.
Mayr, E. and L. Short. 1970. Species taxa of North
American birds, a contribution to compjira-
tive systematics. Publ. Nuttall Omith. Club 9.
McCandless, J. B. 1958. A field guide to the birds
of Puerto Rico. Inter-American University
Press, San German, Puerto Rico.
Peters, J. 1940. Check -list of the birds of the world.
Vol. 4:200. Harvard Univ. Press, Cambridge.
Reynard, G. B. 1962. The rediscovery of the
Puerto Rican Whip-poor-will. Living Bird, 1:
51-60.
Vincent, J. 1966. Puerto Rico Whip-poor-wiU. In:
Red data book. Vol. 2:Aves I.U.C.N. Survival
Service Comm., Morges, Switzerland.
Wadsworth, F. H. 1949. The development of the
forest land resources of the LuquiUo Moun-
tains, Puerto Rico. Ph.D. Thesis University
of Michigan.
Wetmore, A. 1916. Birds of Puerto Rico. Bull.
U.S. Dept. Agr. 326.
Wetmore, A. 1919. Description of a Whip-poor-
wdll from Puerto Rico. Proc. Biol. Soc. Wash.
32:235-237.
Wetmore, A. 1922. Bird Remains from the caves
of Puerto Rico. Amer. Mus. Nat. Hist. Bull.
46:297-333.
Wetmore, A. 1927. The birds of Puerto Rico and
the Virgin Islands. New York Acad. Sci. Vol.
9,pt.4.
Wolcott, G. N. 1953. The food of the mongoose
(Herpestes javanicus auropunctatus Hodgson)
in St. Croix and Puerto Rico. Jouml. Agric.
Univ. Puerto Rico 36:241-247.
Biological Services Program
FWS/OBS-80/01.34
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
ALEUTIAN CANADA GOOSE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and consersation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Ser\'ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Eouisiana 70458
u
FWS/OBS-80/01.34
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
ALEUTIAN CANADA GOOSE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endjingered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Shdell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
ALEUTIAN CANADA GOOSE
Branta canadensis leucopareia (Brandt)
KINGDOM Animalia
ORDER Anseriformes
FAMILY Anatidae
OTHER COMMON
NAMES Hutchins' Goose; Turner
1886; Cackling Goose; Lesser Canada
Goose; Legch; Luch or lugach or Lagix;
land goose; Shijukara gan.
DATE
Entered into SWIS To be determined
Update To be determined
LEGAL STATUS
Federal: Endangered: (42 FR 36427, 14 July
1977). Migratory Bird Treaty Act of
July 3, 1918 (40 Stat. 755; 16 U.S.C.
703-711) as amended Dec. 3, 1969.
Public Law 91-135.
States: Endangered: Alaska.
REASONS FOR CURRENT STATUS
The rapid population decline during the last
50 years and reduction of known breeding range
to one small island is due primarily to introduc-
tion of arctic or blue foxes {Alopex lagopus) on
the Aleutian Islands, although excessive hunting
along the migration route and wintering grounds
has probably contributed to the decline (Grinnell
et al. 1918, Murie 1959, Jones 1963). Blue foxes
have been introduced on almost every island in the
Aleutian chain from Yunaska westward, excluding
Attu which had an ancestral population of foxes
Qones 1963), primarily in the 1920's when fox fur
commanded high prices. Large fox populations
were produced rapidly and maintained. Buldir Is-
land, the most isolated and inaccessible of the
chain, is one of the few where foxes were not in-
troduced and the only one known to have breed-
ing Canada geese at present (Jones 1963, Springer
et al. 1978). Possibly islands of the northern Kuril
group where geese formerly bred may also be fox-
free (Snow 1897).
Recovery of bands from Buldir Island geese
(Springer et al. 1978) shows that hunting pressure
in the interior valleys of California is a consider-
able mortality factor. Avian cholera present in the
San Joaquin Valley produced some Aleutian Can-
ada goose mortality (D. W. Woolington et al.
unpubl. ms.).
PRIORITY INDEX
24
DESCRIPTION
The Aleutian Canada goose is a small goose,
slightly larger than the mallard, with grayish
brown body and black tail, neck, bill, feet and
legs; white cheeks; a ring at the base of the black
neck; and black belly, rump, and tail coverts. It
is smaller and paler than B. c. occidentalis and
about the same size and color as B. c. tavemeri,
except that adults almost always have a relatively
wide white collar which tavemeri usually lacks,
and the bUl is more tapered and pointed at the
tip when viewed from above. Legs are relatively
long. It is slightly larger than B. c. minima, with
longer legs and a more tapered or pointed bill.
The white collar is usually absent in minima (Del-
acour 1951, 1954; J. W. Aldrich ms.; Palmer
1976). Nelson (1883) commented on the con-
spicuous wide white collars of Aleutian Island
Canada geese.
Size.- Male: Wing 356 to 425 mm (av. 386.4
mm);tail llOto 137mm(av. 125.8 mm);exposed
culmen 31 to 38 mm (av. 34.4 mm); tarsus 67 to
88 mm (av. 80.5 mm); middle toe without claw
58 to 69 mm (av. 63.2 mm).
Size.- Female: wing 358 to 390 mm (av.
372.6 mm); tail 110 to 130 mm (av. 122.8 mm);
exposed culmen 30 to 35 mm (av. 32.7 mm); tar-
sus 69 to 80 mm (av. 74.2 mm); middle toe with-
out claw 53 to 66 mm (av. 59.4 mm) Q. W.
Aldrich ms.).
Shape of bill.- The more tapered bill of B. c.
leucopareia is shown by the ratio of width of bill
at base to width at tip (base of nail). Based on
averages of those measurements, bills of leu-
copareia are 7% more tapered than tavemeri and
9% more tapered than minima. The more pointed
appearance of leucopareia bills is due to greater
length of the nail in proportion to its width. Based
on ratio of nail length to width leucopareia bills
are 8% more pointed than tavemeri and 16%
more pointed than minima (J. W. Aldrich ms.).
Weight.- A female specimen from Amchitka
Island, 10 May 1959, 1927 g (Kenyon 1961); a
female specimen from Amchitka Island, 10 June
1952, 1954 g (Krog 1953). Avg. Qohnson et al.):
males - 1946 g (Buldir I.), 2110 g (California);
females - 1703 g (Buldir I.), 1863 g (California).
Juvenile plumage is somewhat paler ventraUy,
with more blended overall coloring. The white
neck ring is indistinct or absent (J. W. Aldrich
ms.. Palmer 1976, D. H. Johnson et al. unpubl.
ms.). Downy young are brownish olive above,
whitish yellow below.
RANGE
B. c. leucoparei formerly bred in the Com-
mander and northern Kuril Islands, U.S.S.R., and
the western Aleutian Islands from Attu east to
Atka and Islands of Four Mountains (Nelson
1883, Turner 1886, Stejneger 1885 and 1887,
Snow 1897, Clark 1910, Bent 1925, Jochelson
1933, Murie 1959, Delacour 1954, Johansen
1961); possibly also in Pribilof Islands (Delacour
1954), but not according to Nelson (1883). It
now breeds only on Buldir Island in Aleutian Is-
lands, as far as is known Qones 1963, Murie
1959, Springer et al. 1978). The last breeding re-
cord for other Aleutian Islands was on Agattu in
1937 (Murie 1959). The species has apparently
been extirpated on the Commander Island since
1914 Qohansen 1961).
This species formerly wintered in Japan (Aus-
tin and Kuroda 1953), and is reported also to
have wintered from British Columbia south to
California (Delacour 1951) and northwestern
Mexico (Hansen and Nelson 1964). The popula-
tion in Japan was reduced to one bird in the fall
of 1978 (Y. Yokota pers. comm.). At present, it
winters chiefly in the Sacramento and San Joaquin
Valleys, Monro Bay, on the southeast Farallon
Island of California, in the lower Colorado River
Valley of Arizona and Mexico (Wollington et al.
unpubl. ms.); and in northwestern Mexico near
Obregon, Sonora Q. W. Aldrich ms.). There are
no definite winter records north of California
(Springer et al. 1978).
In fall migration, the geese leave the Aleutian
Islands breeding area as early as late August
(Woolington and D. R. Yparraguirre unpubl. ms.)
and early September (Byrd and Springer 1976),
and move eastvi^ard along the Aleutian chain to
possible staging areas on Unimak Island. They
arrive on the northwestern California coast and
in the Sacramento Valley between October and
early November. They are not found north of
there, suggesting that they fly directly overwater
from the western Aleutians to the coast of north-
western California. From there, they move south-
ward through the interior valleys of California to
their wintering grounds (Springer et al. 1978, D.
W. Woolington et al. unpubl. ms.).
The migrants move from their San Joaquin
Valley wintering area to a staging area near Cres-
cent City in northwestern California between
middle of February and early March. They leave
that staging area for the Aleutian Islands breeding
grounds during the latter part of April, presum-
ably flying directly northwestward over the Pacific
Ocean, although several reports of unhanded
specimens presumed to be Aleutian Canada geese
have been received from the mouth of the Colum-
bia River (Byrd and Springer 1976, Springer et al.
1978).
Confusion in taxonomy of Aleutian and Com-
mander Islands populations of Branta canadensis
has led to misconceptions, including the reported
nesting of three subspecies, hutchinsii, minima,
and asiatica in those islands (Bent 1912, 1925;
Murie 1959; J. W. Aldrich 1946). ActuaUy, only
one subspecies, leucopareia, nests in the Aleutian
Islands (Delacour 1954), and B. c. asiatica of the
Commander Islands is now thought to be the same
as leucopareia (]. W. Aldrich ms.; Austin and
Kuroda 1953; Palmer 1976).
RANGE MAP
Past and present breeding areas and present
wintering area from Byrd and Springer (1976)
and Springer et al. (1978) are shown on the fol-
lowing range map.
STATES/COUNTIES
Alaska (localities): Attu I., Agattu I., Semichi
Ids., Buldir I., Kiska I.,
Amchitka I., Tanaga I.,
Kanaga I., Adak I., Atka I.,
probably taverneri on un-
derlined areas, Unimak I.
and possibly on Izembek
Bay, head of Morzhovoi
Bay, Nelson Lagoon, Port
MoUer.
California (counties) : Del Norte, Humboldt,
Mendocino, Glenn, Colusa,
Sutter, Yolo, Solano, Sac-
ramento, Contra Costa,
San Joaquin, Stanislaus,
Merced, San Luis Obispo,
Imperial, and San Fran-
cisco (southeast Farallon
Island).
Arizona: Mohave.
Sonora.
Hokkaido and Honshu I.
Mexico:
Japan:
HABITAT
Nesting habitat was reported along the banks
of small streams that enter the sea through gaps
in high bluffs and on grassy hillsides on Agattu
(Clark 1910), and in marshes and lagoons with
rank aquatic vegetation on the Semichi Islands
(Turner 1886). Nesting on rugged sea cliffs cut by
watercourses where grasses and sedges grow in
rank profusion was reported (not interior flatlands
with ponds) on Buldir (Murie 1959, Jones 1963,
G. V. Byrd and D. W. Woolington unpubl. ms.).
Nesting habitat in the Aleutian Islands is charac-
terized by polar maritime climate with high hum-
idity, fog, rain, and small diurnal annual range of
temperature Qones 1963). Molting habitat is in
the uplands. Night roosting habitat is shallow
pools on mountainsides on Attu (Turner 1886);
islands, ponds, and flooded fields in migration
and wintering areas (D. W. Woolington et al. un-
publ. ms.). Fall, winter, and spring habitats in
California are fields from which lima bean, black-
eyed bean, rice and com have been harvested;
green barley and wheat fields; and flood-irrigated
and nonirrigated land (D. W. Woolington et al.
and Springer 1977 unpubl. ms.).
FOOD AND FORAGING
The birds fatten in the fall on Vaccinium ber-
ries on Attu (Turner 1886) and on crowberries
(Empetrum migrum) at Buldir (G. V. Byrd and
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CALIFORNIA
mPCMAk
Wintering areas of the Aleutian Canada goose.
D. W. Woolington unpubl. ms.). The main food
during the breeding season on Buldir Island is
sedge (Carex sp.) and red fescue [Festuca rubra)
(Byrd and Springer 1976).
At the northwestern California spring migra-
tion staging area, geese feed on fescue [Festuca
dertonensis), velvet grass (Holcus sp.), plantain
(Plantago major) and other plants (Byrd and
Springer 1976).
SHELTER REQUIREMENTS
Flightless young and molting birds are hidden
by over 1-m-taIl rye grass {Elymus arenarius) and
umbelliferous plants [Heracleum lanatum and
Angelica lucida) on Buldir Island (Byrd and
Springer 1976).
NESTING AND BEDDING
On Agattu, nests were composed of dead
grass and a few feathers carelessly arranged (Tur-
ner 1886), plus large masses of light drab or dark
gray down mixed with whitish down, numerous
breast feathers and bits of straw (Bent 1925). On
Buldir Island, the geese place nests on steep slopes
under tall vegetation, mostly ryegrass (Byrd and
Springer 1976, G. V. Byrd and D. W. Woolington
unpubl. ms.).
RITUAL REQUIREMENTS
None Recorded.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None known other than protection from pre-
dation by introduced Arctic Foxes on breeding
grounds and excessive hunting and disease on
migration and wintering areas.
POPULATION NUMBERS AND TRENDS
Aleutian Canada geese formerly reared young
in considerable numbers on the western Aleutian
Islands and were domesticated by the natives
(Nelson 1883; Turner 1886). They were the most
abundant birds on Agattu, breeding by the thou-
sands (Clark 1910); they were also common mi-
grants throughout the Aleutian Islands area, nest-
ing on many of the islands. Populations appeared
greatly reduced by the 1936, 1937, and 1938 ob-
servations, as the species had disappeared from
most of the islands and only a few pairs remained
anywhere (Murie 1959). On Amchitka, the 7 seen
on 27 June and 2 on 5 July 1961 were the last
observed on any Aleutian Island prior to their re-
discovery on Buldir Qones 1963). The last birds
were recorded on the Commander Islands in 1914
Qohansen 1961).
In 1977 an estimated 170 breeding pairs (340
birds) and an undetermined number of non-
breeders were located on Buldir Island (G. V.
Byrd and D. W. Woolington unpubl. ms.).
A flock believed to be almost all Aleutian
Canada geese at Crescent City in northwestern
California peaked at 790 in mid-April 1975, at
about 900 in late March 1976, and at 1150 in
the spring of 1977 (Byrd and Springer 1976,
Springer et al. 1978). This indicates a total popula-
tion more than double the number of breeding
birds on Buldir. It also indicates an increase from
1976 to 1977. The fall population in 1977 was
1600 (D. W. Woolington et al. unpubl. ms.).
REPRODUCTION
Turner (1886) said clutches of eggs of Aleutian
Island geese vary from 7 to 13; Byrd and Springer
(1976) noted that Buldir Island geese averaged
about six eggs, laid in late May or early June. In-
cubation was 27-28 days, with over 80% of the
nests hatching young. Glaucous-winged gulls eat
some eggs and bald eagles take a few birds, but pre-
dation apparently is not severe (Byrd and Springer
1976). Turner (1886) noted that parent birds
complete their molt by 20 August, at which time
young are able to fly. G. V. Byrd and D. W. Wool-
ington (unpubl. ms.) made similar observations on
Buldir. Turner (1886) noted that a heavy snowfall
in the latter part of June on Agattu and Semichi
covered the ground to 3 ft (0.91 m), and incubat-
ing geese were found dead after the snow melted.
MANAGEMENT AND CONSERVATION
The Recovery Team prepared a plan with 4
objectives: 1) prepare habitat for goose reintro-
duction by removing fox; 2) produce adequate
release stock of geese in captivity; 3) acclimatize
and release birds and monitor results; 4) study
and protect the wild birds (Byrd and Springer
1976).
Control of foxes, begun by Robert D. Jones,
Jr. in the late 1940's, resulted in their complete
elimination from Amchitka. To safeguard reintro-
duced geese, 3 areas were selected for fox removal,
widely separated to reduce the danger of local
disasters such as earthquakes and tidal waves;
these are Agattu (22,000 ha) with nearby Nizki/
Alaid Islands (1300 ha), Amchitka Island (30,000
ha) and Kanaga Island (37,000 ha), all of which
formerly had goose populations. Originally, foxes
were eliminated by poison, but poisoning is now
banned by law. Fifty-four foxes were taken on
Agattu and 130 on Nizki/Alaid in 1974 and 1975.
It was estimated that one or more foxes were left
on Agattu in 1978 (D. W. Woolington and D. R.
Yparraguirre unpubl. ms.) and none on Nizki/
Alaid in 1976 (Springer et al. 1978). Fox control
on Kanaga Island in 1977 removed 142 animals,
leaving an estimated 700 plus Q. L. Martin pers.
comm.).
Captive propagation began in 1963 with cap-
ture of 18 goslings on Buldir Island for transfer to
breeding pens at Patuxent Wildlife Research Cen-
ter, Laurel, Md., in 1966. Additional goslings
were taken for that project in 1972 (21) and
1975 (20). Altogether over 451 goslings have
been produced from 1966 through 1978 (R. C.
Erickson pers. comm.).
To facilitate acclimatization and develop a
tradition for homing to Aleutian Islands, a facility
was developed on Amchitka Island in 1976 where
goslings will be held to maturity before being re-
leased, and a second propagation facility was
estabhshed on Amchitka. Twenty geese raised
from Buldir goslings at Northern Prairie Research
Center, together with 75 birds from Patuxent,
were sent to Amchitka in fall 1977 to initiate that
operation. In 1977 and 1978, 150 goslings were
raised at Amchitka.
First releast of 75 one- two- and three-year-
old geese was on fox-free Amchitka in the spring
of 1971, but all birds disappeared soon afterwards,
some because of predation by bald eagles. The se-
cond release was on Agattu in the summer of
1974, when the island was admost fox-free and
fox control was continuing. Four pairs of geese
nested and 2 pairs raised 5 young. Other released
geese remained near the release site all summer.
Nine molting geese were brought to Agattu from
Buldir to serve as guides for captive-reared birds
in the fall migration. The geese departed Agattu
on 4 September, headed east. In the winter and
spring of 1974-75, three of the released birds,
identified by white bands, were reported on the
northwest coast of California. Released birds did
not return to Agattu in 1975 nor were they found
on Buldir with the wild birds (Byrd and Springer
1976, Springer et al. 1978).
In the spring and fall 1976, a release of 30
pen-reared birds and 3 wild guide birds was again
made on Amchitka. Most of the birds disappeared
from unknown causes, but at least 5 were killed
by eagles. Eight birds were recaptured and held at
Amchitka Q. L. Martin pers. comm.). A fourth re-
lease of 117 pen-reared birds and 22 wild guide
birds was made on Agattu in 1978; it is too early
to determine the success of this effort (P. F.
Springer pers. comm.).
Monitoring of geese banded on Buldir and
Agattu has now established fairly well their migra-
tion routes, their principal wintering areas in Cali-
fornia, and their staging area for spring migration.
Further plans for monitoring include careful
counts at Castle Rock spring staging area in north-
western California and several likely spots to the
north for migrants. Some captive-reared and wild
geese are equipped with colored neck collars for
easier location.
Based on times and places of band recoveries
and sightings in 1975, the California Depairtment
of Fish and Game closed the northwest coastaJ
counties of Del Norte, Humboldt, and Mendocino,
to hunting for the entire 1976 season, and also
closed part of the Sacramento Valley in Glenn and
Colusa counties from the opening of the season in
mid-October to mid-December, and part of the
San Joaquin Valley in San Joaquin, Stanislaus,
and Merced counties from mid-December to the
close of the season in mid-January. Subsequently,
on the basis of additional recoveries, the closed
area of the Sacramento Valley was extended to
include parts of Butte and Sutter counties and the
San Joaquin Valley closure now starts during the
last week in November. Because of lack of re-
coveries or sightings of banded wild birds in Men-
docino County, this county was reopened to
Canada goose hunting in 1977. It seems probable
that the increased goose population in California
since 1975 is due to the restrictions on hunting.
AUTHORITIES
Dennis W. Woolington
Paul F. Springer
Wildlife Research Station
Humboldt State University
Areata, California 95521
G. Vernon Byrd
Aleutian Islands N.W.R.
Box 5251
Adak, Alaska 98791
Hawaiian Islands N.W.R.
Box 87
Kilauea, Kannai, Hawaii 96754
Ray C. Erickson
Patuxent Wildlife Research Center
Laurel, Maryland 20811
Robert D. Jones, Jr.
U.S. Fish and Wildlife Service
1011 E. Tudor Rd.
Anchorage, Alaska 99507
Daniel R. Yparraguirre
California Department of Fish and Game
2166 Carriro Brazos
Pleasanton, California 94566
Karl W. Kenyon
11964 Lakeside Place
N.E. Seattle, Washington 98125
PREPARER'S COMMENTS
Good progress has been made in management
of the Aleutian Canada Goose. If the recovery
plan is carried out, particularly with respect to
fox control and reintroduction of geese in Aleutian
Islands and hunting control in migration and win-
tering areas, the prospects look very good for re-
covery — John W. Aldrich.
LITERATURE CITED/SELECTED
REFERENCES
Aldrich, J. W. 1946. Speciation in the white-
checked geese. Wilson BuU. 58:94-103.
Austin, O. L., Jr. and N. Kuroda. 1953. Birds of
Japan. Bull. Mus.Comp.Zool. Harvard 109(4).
Bent, A. C. 1912. Notes on birds observed during
a brief visit to the Aleutian Islands and Bering
Sea in 1911. Smithsonian Misc. Colls. 56(32):
1-29.
Bent, A. C. 1925. Life histories of North Ameri-
can Wild Fowl. Part II. U.S. Nat. Mus. Bull.
130.
Byrd, G. V., and D. W. Woolington. Breeding
biology, habitat utilization and population
structure of Aleutian Canada geese on Buldir
Island. Unpublished manuscript. Aleutian Is-
lands National Wildlife Refuge. 110 p.
Byrd, G. V. and P. F. Springer. 1976. Recovery
program for the endangered Aleutian Canada
goose. Cal-Neva Wildlife Transactions 1976:
65-73.
Clark, A. H. 1910. The birds collected and ob-
served during the cruise of the United States
Fisheries steamer "Albatross" in the North
Pacific Ocean, and in the Bering, Okhotsk,
Japan, and Eastern Seas, from April to Decem-
ber 1906. Proc. U.S. Nat. Mus. 38:25-74.
Delacour, J. 1951. Preliminary note on the tax-
onomy of Canada Geese, Branta canadensis.
Amer. Mus. Novits. 1537:1-10.
Delacour, J. 1954. The Waterfowl of the world.
Vol. 1. London, Country Life Limited.
Grinnell, J., H. C. Bryant, and T. I. Storer. 1918.
The game birds of California. University of
California Press, Berkeley. 642 pp.
Hansen, H. A., and H. K. Nelson. 1964. Honkers
large and small. P. 109-124 in J. P. Linduska,
ed. Waterfowl Tommorrow. U.S. Fish and
Wildlife Service. 770 pp.
Jochelson, W. 1933. History, ethnology, and
anthropology of the Aleut. Carnegie Inst.
Wash. Publ. 432. 91 pp.
Johansen, H. 1961. Revised list of the birds of the
Commander Islands. Auk 78:44-56.
Johnson, D. H., D. E. Timm, and P. F. Springer.
Morphological characteristics of Canada geese
in the Pacific Flyway. Paper to be presented
at Symposium on Biology and Management of
Pacific Flyway Geese, Portland, Or., Feb. 16,
1979.24 pp.
Jones, R. D., Jr. 1963. Buldir Island, site of a
remnant breeding population of Aleutian
Canada Geese. Ann. Rep. Wildfowl Trust 14:
80-84.
Kenyon, K. W. 1961. Birds of Amchitka Island
Alaska, Auk 78:305-320.
Krog, J. 1953. Notes on the birds of Amchitka
Island, Alaska. Condor 55:299-304.
Murie, O. J. 1959. Fauna of the Aleutian Islands
and Alaska Peninsula. North Amer. Fauna.
61:364 pp.
Nelson, E. W. 1883. Birds of the Bering Sea and
the Arctic Ocean. Cruise of the Revenue-
Steamer Corvvin in Alaska and the N.W. Arc-
tic Ocean in 1881. Washington, D.C. Govern-
ment Printing Office.
Palmer, R. S. ed. 1976. Handbook of North Am-
erican Birds, Vol. 2. New Haven, Yale Univ.
Press, p. 211.
Snow, H. J. 1897. Notes in the Kuril Islands.
Royal Geographical Society, London. 91 pp.
Springer, P. F.,G. V. Byrd, and D. W. Woolington.
1978. Reestablishing Aleutian Canada geese.
P. 331-338 in S. A. Temple, ed. Endangered
birds: management techniques for threatened
species. University of Wisconsin Press, Madi-
son. 446 pp.
Stejneger, L. 1885. Results of ornithological ex-
penditions in the Commander Islands and
Kamtschatak. BuU. U.S. Nat. Mus. 29:1-382.
Stejneger, L. 1887. Revised and annotated cata-
logue of the birds inhabiting the Commander
Islands. Proc. U.S. Nat. Mus. 10:117-145.
Turner, L. M. 1886. Contributions to the natural
history of Alaska. Results of investigations
made chiefly in the Yukon, District and the
Aleutian Islands; conducted under the auspices
of the Signal Service United States Army, ex-
tending from May, 1874, to August, 1881.
No. 11. Arctic Series of Publications in con-
nection with the Signal Service, U.S. Army.
Washington, Government Printing Office.
Woolington, D. W., P. F. Springer, and D. R.
Yparraguirre. Migration and wintering distri-
bution of Aleutian Canada geese. Paper to be
presented at Symposium on Biology and Man-
agement of Pacific Flyway Geese, Portland,
Or., Feb. 16, 1979. 11pp.
Woolington, D. W., and D. R. Yparraguirre. Pro-
gress report on restoration of Aleutian Canada
geese on Agattu Island, 1978. Unpublished
manuscript. Aleutian Islands National Wildlife
Refuge. 74 pp.
Biological Services Program
FWS/OBS-80/01.35
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
WEST INDIAN MANATEE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conser\'ation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the l",ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Eouisiana 704.58
11
FWS/OBS-80/01.35
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
WEST INDIAN MANATEE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
WEST INDIAN MANTAEE
Trichechus manatus Linnaeus
KINGDOM Animalia
CLASS Mammalia
ORDER Sirenia
FAMILY Trichechidae
OTHER COMMON
NAMES sea cow; sea
siren.
DATE
Entered into SWIS To be determined.
Updates 9 October 1976;
8 March 1977; 1 June 1979
LEGAL STATUS
Federal: Endangered (32 FR 4001; 11 March
1967 and 35 FR 8491; 2 June 1970).
States: Endangered: Georgia, South Carolina,
Texas, Threatened; Florida.
REASONS FOR CURRENT STATUS
The manatee population has decHned severely
from an estimated several thousand individuals in
the 1700's and early 1800's to as few as 1,000
today (Hartman 1974).
Initial population decreases resulted from
overharvesting for meat, oil, and leather. Today,
little illegal hunting occurs, but heavy mortality
results from accidental boat and barge strikes and
entrapment in canal locks and flood control dams
(Campbell and Powell 1976; Irvine et al. 1978).
Some malicious killing by vandals occurs, but is
not considered a significant factor.
Actual physical habitat loss may be of con-
siderable importance; residential, commercial, and
recreational development is rampant in Florida.
Heavy boat traffic in some areas is a serious threat
to manatees and their habitat.
Recent data (Campbell and Irvine 1978) in-
dicate that manatees may be diverted from their
southward fall movements to industrial warm-
water effluents located north of their historic
winter range. During exceptionally cold weather,
water temperatures suitable for the animals may
not be maintained. During the prolonged and ex-
ceptionally cold winter of 1976-77, 34 manatees
died in northern and central Florida in areas
north of their historic winter range; many of
them had been associated with power plant ef-
fluents (Campbell and Irvine 1978).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The manatee is a large gray or brown fusiform
aquatic mammal that lacks a dorsal fin. It is nearly
hairless except for stiff whiskers covering the
muzzle and widely scattered, fine hairs on the
back. The forelimbs are modified into flippers,
and the paddle-shaped tail is flattened hori-
zontally. Hind limbs are absent. The sexes are dis-
tinguished by the position of the genital opening;
the distance from the anus is greatest in males.
Within its range, the manatee is the only large
aquatic mammal exhibiting these characteristics.
Manatees are usually 3 m long, with a maxi-
mum length of 4.6 m reported (Hartman 1979b).
Adult weights range between 360 and 540 kg,
with a maximum of approximately 1,000 kg
(Hartman 1979b).
Illustrations of manatees have been published
in many popular and scientific articles: Simpson
(1930), Ledbetter (1960), Walker, et al. (1964),
Hartman (1969),Stephens(1972),Okeefe (1973),
Vietmeyer (1974), Campbell and Powell (1976),
Hartman (1979a), and Reynolds (1979).
RANGE
During the wdnter months, U.S. populations
appear to be restricted to peninsular Florida,
south of Crystal River on the Gulf coast and
Titusville on the Atlantic coast. In the St. Johns
River, the animals occur from Jacksonville south
to at least Lake Monroe (Layne 1965; Hartman
1974).
During the winter, manatees in northern and
central Florida congregate around natural and in-
dustrial warm-water sources (see Hartman 1974
for a list of sites). Winter distribution has expanded
northward during the last few decades from 27°
52' to 39° 21' on the east coast and from 26° 45'
to 27° 45' on the west coast. The expansion is
probably due to the proliferation of industrial
and power-generating plant warm-water dis-
charges (Moore 1951b, 1953; Layne 1965;
Hartman 1974).
During the summer, manatees that congre-
gate at winter warm water refugia appear to
disperse, moving north along the Atlantic and
Gulf coasts, occasionally as far as the Carolinas
and extreme western Florida. Rarely, sightings
are reported in Louisiana and Mississippi and
historic records document sightings as far north
as New Jersey (Hartman 1974, Campbell and
Powell 1976). Sightings north of Georgia are
unusual and may represent vagrant individuals.
Historic information suggests that the northern
Gulf of Mexico coast may once have been more
frequented by manatees during the summer than
it is now (Lowery, 1974). It is unlikely that
manatees could survive there in the winter due to
the low water temperatures.
Occasional sightings in extreme southern
Texas may represent remanents of a population on
the northern Mexico Gulf coast (Husar 1977).
Trichechus manatus occurs around many of
the Caribbean islands: Puerto Rico, Hispaniola,
Jamaica, Cuba, and Andros (Husar 1977). Recent
sightings have been reported for Grand Bahama
Island (Odell et al. 1968).
In Central and South America, the West Indian
manatee occurs from at least central Mexico
south along the coast to the region of Alagoas
State, Brazil (Bertram and Bertram 1973, White-
head 1977). It also penetrates the Orinoco River
in Venezuela to Apure Falls (Mondolfi, 1974) and
up the Amazon River an undetermined distance
(Bertram and Bertram 1973).
RANGE MAP
The range maps (following) show the present
distribution, historic range, peripheral sightings,
and critical habitat of the West Indian manatee in
the United States.
STATES/COUNTIES
Alabama: Baldwin, Mobile.
Florida: Bay, Brevard, Broward, Charlotte, Cit-
rus, Clay, Collier, Dade, De Soto,
Dixie, Dural, Escambia, Flagler,
Franklin, Gilchrist, Glades, Gulf,
Distribution of T. manatus in the southeastern United States (shaded area: summer
distribution; dots: peripheral sightings; arrows; northern limits of historic range)
Estimated distribution of Trichechus manatus in Florida.
NORTHWESTERN PART OF
FLORIDA
S««( SC»ie *S MAIN U«P
15 16 ■ 17 19 \9
21 22 23 a* 2S
Critical habitat of Trichechus manatus in the United States.
Georgia:
Louisiana:
Missis-
sippi:
North
Carolina:
Hendry, Hernando, Highlands, HUls-
Borough. Indian River, Jefferson,
Lake, Lafayette, Lee, Levy, Manatee,
Marion, Martin, Monrow, Okaloosa,
Okeechobee, Orange, Osceola, Palm
Beach, Pasco, Pinellas, Polk, Putnam,
St. Johns, St. Lucie, Santa Rosa, Sara-
sota, Seminole, Suwannee, Taylor,
Volusia, Wakulla, Walton.
Bryan, Camden, Chatham, Glynn,
Liiaerty, Mcintosh.
(Parishes): Cameron, Iberia, Jefferson,
Lafourche, Plaquemines, St. Bernard,
St. Mary, Terrebonne, Vermilion.
Hancock, Harrison, Jackson.
Beaufort, Bertie, Brunswick, Camden,
Carteret, Chowan, Currituck, Dare,
Hyde, New Hanover, Onslow, Pamlico,
Pasquotank, Pender, Perquimans, Tyr-
rell, Washington.
Beaufort, Berkeley, Charleston, Colle-
ton, Georgetown, Horry.
Aransas, Brazoria, Calhoun, Cameron,
Chambers, Galveston, Jefferson, Ken-
edy, Kleberg, Matagorda, Nueces, Wil-
lacy.
HABITAT
Manatees inhabit both fresh and salt water of
sufficient depth (1 meter or more) throughout
their range. They may be encountered in water-
ways with access to coastal habitats, including
canals, sluggish rivers, estuaries, and saltwater
bays. They also have been seen as far as 6 km off
the coast of Florida in the Gulf of Mexico (Hart-
man 1974). Between October and AprU in Florida,
manatees concentrate near warm water (Hartman
1974, 1979), but during other months they appear
to choose areas with adequate food near sources
of fresh water (Huser 1977, Hartman 1979a).
FOOD AND FORAGING BEHAVIOR
Manatees consume almost any vegetation
available. Comprehensive lists of plant species
recorded in their diet have been published by
Hartman (1971) and Campbell and Irvine (1977).
South
Carolina:
Texas:
Emergent and floating plants, and even vegetation
along the shore are acceptable but they prefer
submergents, emergents, and floating vegetation
in that order (Hartman 1979a). A partial list of
plants known to be eaten by manatees illustrates
the variety of their diet:
Floating fern (Ceratophyllum pteridoides)
Pondweed (Diplanthera wrightii)
Water-hyacinth (Eichhomia crassipes)
Waterweed (Elodea)
Hydrilla (Hydrilla verticillata)
Water milfoil (Myriophyllum)
Widgeon grass (Ruppia maritima)
Manatee-grass (Syringodium filiforme)
Turtle grass (Thallasia testudinum)
Eelgrass (Vallisneria neotropicalis)
Saw palmetto (Serenoa repens)
Various types of algae
While generally considered completely herb-
iverous, there are instances of wild manatees
feeding on fish (Powell 1978) and captive mana-
tees will take fish if available (Jenkins 1979).
SHELTER REQUIREMENTS
During hurricanes, manatees may seek shelter
in protected bays and canals (Hartman 1971).
NESTING OR BEDDING
Not applicable.
RITUAL REQUIREMENTS
Not appHcable.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Manatees are adapted to both fresh- and salt-
water habitats. Dependance on freshwater for
drinking has been suggested by Hartman (1974,
1979a). Saltwater populations often congregate
around sources of freshwater, such as offshore
springs, river mouths, and marina water hoses.
Numerous accounts of manatee deaths fol-
lowing cold weather in Florida suggest that winter
temperatures at the margin of their range may be
a limiting factor for U.S. populations (Bangs 1895;
Cahn 1940; Hamilton 1941; Kromholz 1943;
Moore 1951b, 1956; Layne 1965). Water temper-
ature below approximately 20° C stimulates move-
ment into warm water refugia (Hartman 1974,
1979a; Powell 1978; Powell and Waldron 1978). MANAGEMENT AND CONSERVATION
POPULATION NUMBERS AND TRENDS
Hartman (1974, 1979a) estimated the U.S.
population at 750 to 850 animals with possible
extremes of 600 and 1,000. More recent surveys
(Irvine and Campbell 1978) suggest a minimum
population between 750 and 900 animals, while
Brownell et al. (1978) estimated a U.S. popula-
tion of 1,000 or more.
No accurate historical population estimates
cure available, but numbers today are believed to
be reduced compared to historical levels (Hart-
man 1971). Concern for reduced population
levels was expressed as early as 1880 by LeBaron
and 1885 by Canova. All populations throughout
the species' range are believed to be critically re-
duced (Bertram and Bertram 1973).
United States populations appear to be in-
creasing in some areas, decreasing in other areas,
and remaining stable in some regions (Hartman
1974). Husar (1977) suggested that numbers were
increasing, but Brownell et alj (1978), based on
mortality figures, thought it more likely that an
overall decline was occuring.
REPRODUCTION
Mating generally involves the pursuit of an
estrous female by a group of courting males
(Hartman 1979a). Gestation, as estimated from a
captive conception and birth, is 365 days (Dek-
ker 1977). The manatee is basically uniparous,
with the possibility that twinning may occasionally
occur (Hartman 1979a). Hartman (1979a) believed
that there was no specific breeding season, but
recovery of dead calves suggests that most of
them are bom in the spring and early summer
(Hartman 1979a, Irvine et al. 1978). Calves are
bom and nursed in the water and are believed to
remain with the female for up to 2 years. Re-
productive maturity may be reached at 7 to 10
years (Odell et al. 1978).
The most commonly observed social unit is
the cow and her calf, but larger social groups of
unknown composition are observed, especially
around limited resources such as warm water
refugia (Hartman, 1979a).
Natural longevity is unknown, although an
individual has been kept in captivity in Florida
for over 29 years (Brownell et al. 1978).
In addition to the Endangered Species Act,
manatees are protected in the United States by
the Marine Mammal Protection Act of 1972 (PL
92-522; 80 stat 1027) and by various State laws,
such as the Florida Manatee Sanctuary Act.
Manatees are also protected in every country
throughout their range, although enforcement
efforts are ineffective in most areas.
The most immediate conservation need in the
United States is for management (Wray 1976).
Especially critical is the reduction of mortality
caused by boat strikes and entrapment in water
control structures (Hartman 1974, Campbell and
Powell 1976, Brownell et al. 1978).
The State of Florida has designated Blue
Spring State Park as a manatee sanctuary and
Brevard County, Florida, has declared itself a
manatee sanctuary. De facto sanctuaries exist in
the Everglades National Park, various National
Wildlife Refuges throughout Florida, and several
Florida State Parks. Critical Habitat was designated
in 1976 (41 FR 41916, 24 September 1976) (see
range maps).
AUTHORITIES
Edward Asper
Seaworld of Florida
7007 Seaworld Drive
Orlando, FL 32809
Robert L. Brownell, Jr. (Recovery Team)
National Fish and Wildlife Laboratory
Museum of Natural History
Washington, D.C. 29560
John C. Oberheu (Recovery Team Leader)
Jacksonville Area Office
U.S. Fish and Wildlife Service
900 San Marco Blvd.
Jacksonville, FL 32207
Daniel K. Odell
School of Marine and Atmospheric Sciences
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Galen B. Rathbun
National Fish and Wildlife Laboratory
412 N.E. 16th Ave., Room 250
Gainesville, FL 32601
Patrick M. Rose
Florida Audubon Society
P.O. Drawer 7
Maitland, FL 32751
PREPARER'S COMMENTS
Liturature reviews and bibliographies on Tri-
chechus manatus are numerous.
LITERATURE CITED/SELECTED
REFERENCES
Allsopp, W. H. L. 1969. Aquatic weed control by
manatees: its prospects and problems. In L. E.
Obeng, ed. Man-made lakes. Ghana Univ.
Press, Accra.
Bangs, O. 1895. The present standing of the Flo-
rida manatee, Trichechus [manatus?^ latiros-
tris (Harlan), in the Indian River waters. Am.
Nat. 29:783-787.
Bertram, G. C. L., and G. K. R. Bertram. 1973.
The modem sirenia: their distribution and
status. Biol. J. Linn. Soc. 5:297-338.
Brownell, R. L., Jr., Kr. Ralls, and R. R. Reeves,
eds. 1978. Report of the West Indian Manatee
Workshop, Oriando, Florida, 27-29 March
1978. Nad. Fish and Wildl. Lab, U.S. Fish and
Wildl. Serv. Rept. 37 pp.
Cahn, A. R. 1940. Manatees and the Florida
freeze. J. Mammal. 21:222-223.
Campbell, H. W. 1976. The Florida manatee and
related species. Plaster Jacket 25:10 pp.
Campbell, H. W. 1977. Mammalia; Sirenia;
Trichechidae (Manatees). In: Endangered and
Threatened Plants and Animals of North Car-
olina. N.C. State Museum of Natural History.
444 pp.
Campbell, H. S., and J. A. Powell. 1976. Endan-
gered species: the Florida manatee. Florida
Nat. AprU:15-20.
Campbell, H. W., and A. B. Irvine. 1977. Feeding
ecology of the West Indian manatee. Aqua-
culture 12:249-51.
. 1979. Manatee mortality during the unu-
sually cold winter of 1976-1977. Proc. West
Indian Manatee Workshop, 27-29 March 1978,
Orlando, Florida. 12 pp.
Canova, A. 1885. Life and adventures in south
Florida. Tribune Printing Co., Tampa. 145 pp.
Gold, J. P. 1979. The Sirenia: Dugong, manatee,
Steller's sea cow, and relatives. Natl. Mus. of
Natur. Hist., Smithsonian Inst. 11 pp.
Dekker, D. 1977. Zeekoegeboorte. Artis 23(4):
111-119.
Hamilton, W. J. 1941. Notes on some mammals
of Lee County, Florida. Am. Midi. Nat. 25:
686-691.
Hartman, D. S. 1969. Florida's manatees, mer-
maids in peril. Natl. Geogr. Mag. 136(3):
342-353.
. 1971. Behavior and ecology of the Flo-
rida manatee, Trichechus manatus latirostris
(Harlan), at Crystal River, Citrus County. Ph.
D. Dissert. Cornell Univ., Ithaca, N.Y. 285 pp.
-. 1974. Distribution, status and conserva-
tion of the manatee in the United States. U.S.
Fish Wildl. Ser. Unpubl. Rep. 247 pp.
Hartman, D. S. 1979a. Ecology and Behavior of
the Manatee {Trichechus manatus) in Florida.
Am. Soc. of Mammal. Spec. Publ. 5. 153 pp.
. 1979b. West Indian manatee. Pages 27-
39 in J. N. Layne, ed. Rare and endangered
biota of Florida, Vol. 1, Mammals. Univ.
Presses of Florida, Gainesville.
Husar, S. L. 1975. An annotated bibliography of
the recent Sirenia {Trichechus, Dugong, Hy-
drodamalis). National Fish and Wildlife Lab.,
U.S. Fish and WUdlife Service Rept. 148 pp.
. 1977. The West Indian manatee {Tri-
checus manatus). U.S. Fish and Wildl. Ser.
WUdhfe Res. Rept. 7, 22 pp.
. 1978. Trichechus manatus. Mammalian
Species 93:1-5.
Irvine, A. B., and H. W. Campbell. 1978. Aerial
census of the West Indian manatee, Triche-
chus manatus, in the southeastern United
States. J. Mammal. 59(3):613-617.
Irvine, A. B., D. K. Odell, and H. W. Campbell.
1978. Manatee mortality in the southeastern
United States: 1974-1977. Proc. West Indian
Manatee Workshop, 27-29 March 1978, Or-
lando, Florida.
Jenkins, R. L. 1979. Captive husbandry of mana-
tees at Marineland of Florida. In R. L.
Brownell, K. Ralls, and R. R. Reeves, eds,
Proc. West Indian Manatee Workshop, 27-29
March 1978, Orlando, Florida.
Kromholz, L. A. 1943. Notes on manatees in Flo-
rida waters. J. Mammal. 24:272-273.
Layne, J. N. 1965. Observations on marine mam-
mals in Florida waters. Bull. Florida St. Mus.
Biol. Sci. 9:131-181.
LeBaron, J. F. 1880. The manatee or sea cow.
Forest and Stream 13:1005-1006.
Ledbetter, C. S. 1960. Amazing mammals of Blue
Springs Run. Florida WUdl. 14(2):22-24.
Lowery, G. H. 1974. Mammals of Louisiana and
its adjacent waters. Louisiana St. Univ. Press,
Baton Rouge. 565 pp.
MacDonald, K., L. J. Seeley, and E. C. Amoroso.
1978. Biological Synopsis of the manatee. In-
ternational Development Research Centre, Ot-
tawa, Canada. 1 12 pp.
Marsh, H., P. W. Channells, and J. Morrissey.
1979. A bibliography of the recent Sirenia.
James Cook Univ., Australia. 163 pp.
Mondolfi, E. 1974. Taxonomy, distribution, and
status of the manatee in Venezuela. Sci. Cien.
Nat. La SaUe 34(97) :5-23.
Moore, J. C. 1951a. The status of the manatee in
the Everglades National Park, with notes on
its natural history. J. Mammal 32(l):22-36.
. 1951b. The range of the Florida mana-
tee. Q.J. Florida Acad. Sci. 14(1):1-19.
. 1953. Distribution of marine mammals
Powell, J. A., Jr. 1978. Evidence of camivory in
manatees {Trichechus manatus). J. Mammal.
59:442.
1978a. Status of the manatee population
in Florida waters. Amer. Midi. Nat. 49(1):
117-158.
1956. Observations of manatees in aggre-
gations. Am. Mus. Novitates. 1811:1-24
Odell, D. K., D. J. Forrester, and E. D. Asper.
1978. A preliminary analysis of organ weights
and sexual maturity in the West Indian mana-
tee {Trichechus manatus). Proc. West Indian
Manatee Workshop, 27-29 March 1978,
Orlando, Florida.
OdeU, D. K., J. E. Reynolds, and G. Waugh. 1978.
New records of the West Indian manatee {Tri-
chechus manatus) from the Bahama Islands.
Biol. Conserv. 14:289-293.
O'Keefe, M. T. 1973. Blue Springs-haven for the
manatees. Florida Sportsman 5(1):10-14.
in Crystal River, Citrus County, Florida. Proc.
West Indian Manatee Workshop, 27-29 March,
1978, Orlando, Florida. 7 pp.
Powell, J. A., and J. C. Waldron. 1978b. Status of
the manatee population in Blue Spring, Volu-
sia County, Florida. Proc. West Indian Mana-
tee Workshop, 27-29 March 1978, Orlando,
Florida. 10 pp.
Reynolds, John E. III. 1979. The semi-social
manatee. Nat. Hist. 88(2):44-53.
Simpson, G. G. 1930. Sea sirens, the strange
mammals responsible for the ancient ideas of
mermaids. Nat. Hist. 30:41-47.
Stephens, W. M. 1972. Florida mermaids. Natl.
Parks. Conserv. Mag. 46(3):28-31.
United Nations Food and Agriculture Organiza-
tion Working Party on Marine Mammals.
1978. Mammals in the seas. Report of the
FAO Advisory Committee on Marine Re-
sources Research. FAO, U.N., Rome. 264 pp.
U.S. Fish and Wildhfe Service. 1979. Administra-
tion of the Marine Mammal Protection Act of
1972. April 1, 1978 to March 31, 1979. Dept.
of Interior, U.S. Fish and Wildlife Service. 86
pp.
U.S. Fish and Wildlife Service. 1979. West Indian
manatee recovery plan. Tech. draft. U.S. Fish
and WUdlife Service. 33 pp.
Vietmayer, N. 1974. The endangered but useful
manatee. Smithsonian Mag. 5(9):60-64.
Walker, E. P. et al. 1964. Mammals of the world.
Johns Hopkins Press, Baltimore. 1,500 pp.
Whitehead, P. J. P. 1977. The former southern
distribution of New World manatees {Triche-
chus spp.). Bio. J. Linn. Soc. 9:165-189.
Whitfield, W. K. and S. L. Farrington. 1975. An
armotated bibliography of Sirenia. Fla. Marine
Res. Publ. 7:1-44.
Wray, P. 1976. The North American manatee.
Endangr. Species Productions, Inc. Ayer,
Maine. 26 pp.
Wray, P. 1978. The West Indian manatee {Triche-
chus manatus) in Florida. U.S. Dept. Com-
merce, NTIS, PB-285-410, 89 pp.
Biological Services Program
FWS/OBS-80/01.36
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
ISLAND NIGHT LIZARD
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lu:
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Sen, ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.58
u
FWS/OBS-80/01.36
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
ISLAND NIGHT LIZARD
A Cooperative Effort
by the
National Fish and Wildhfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W.Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
ISLAND NIGHT LIZARD
Xantusia riversiana
KINGDOM Animalia
CLASS Reptilia
ORDER Squamata
FAMILY Xantusiidae
OTHER COMMON NAMES None
DATE
Entered into SWIS To be determined
Updates 15 October 1978
LEGAL STATUS
Federal Threatened (42 FR 40682, 11 Aug
1977).
States Not nominated
REASONS FOR CURRENT STATUS
This lizard is known only from three major
California islands: Santa Barbara, San Nicolas,
and San Clemente. Island habitats and animal
populations have been severely modified as a
result of the introduction of new species, espe-
cially feral cats, goats, pigs, and rabbits.
PRIORITY INDEX
Not assigned.
DESCRIPTION
Xantusia riversiana was once assigned to a
separate genus, Klauberina (Savage 1957). This
lizard is 5.8 to 8.0cm in snout-to-vent length,
with soft granular sccdes and folds of skin on the
neck and along the sides of the body. The back is
mottled with pale ash gray or beige and yellow
brown, darkened in varying amounts with black.
Occasional individuals have a pale gray dorso-
lateral stripe on each side, edged with brown and
black; a brown vertebral stripe may also be
present. It is pale gray below, sometimes with a
bluish cast suffused on the belly and often on the
tail with yellow. Underside of feet may be yellow-
ish (Stebbins 1966). A photograph is in the
Endangered Species Technical Bulletin (1977,
Vol. 2, No. 9).
RANGE
Island night lizards occur on Santa Barbara,
San Nicolas, and San Clemente Islands, in the
California Channel Islands group (Stebbins 1966,
Savage 1967). A small population has recently
been discovered on Sutil Island (J. M. Erode pers
comm). Stebbins (1954) indicated that X. river-
siana occurred on Santa Catalina Island, but this
record needs confirmation.
RANGE MAP
Arrows on the accompanying map indicate
islands where X. riversiana occurs.
STATES /COUNTIES
California: Los Angelos, Ventura.
HABITAT
The species inhabits grassland brushy areas,
clumps of cactus (Opuntia), cliffs, and rocky
beaches (Stebbins 1954, 1966).
FOOD AND FORAGING BEHAVIOR
X. riversiana is an omnivorous, opportunistic
feeder, eating a large variety of anthropods —
spiders, marine isopods, scorpians, centipedes,
mites, aphids, wasps, flies, moths, silverfish, etc
(Stebbins 1954). Brattstrom (1952) notes that
ants and beetles are the most important food
items for this species; 31% of the animals he
examined from San Clemente Island had also
ingested plant matter consisting of flowers, leaves,
stems, and seeds {Mesembryanthemum, etc).
These lizards are thought of as active mostly at
night, but they have been seen foraging in the
daytime (Stebbins 1954).
SHELTER REQUIREMENTS
X. riversiana seeks shelter under rocks, drift-
wood, and fallen branches (Stebbins 1966).
NESTING AND BEDDING
No information is available concerning
nesting.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
No data are available on absolute densities.
Bezy (1976) indicated that X. riversiana was
widespread on San Clemente Island, restricted but
locally common on San Nicolas Island, and
restricted by limited habitat but moderately
abundant on Santa Barbara Island. He notes seri-
ous threats to X. riversiana or to its habitat on all
islands due to populations of introduced animals
(See Management and Conservation).
REPRODUCTION
X. riversiana is viviparous. Spermiogenesis
begins in March and concludes in June; females
ovulate from late May to early June (Goldberg
and Bezy 1974). Breeding probably occurs from
May through June. Goldberg and Bezy (1974)
report a range of 3 to 9 young per breeding
female per year, with a mean of 3.76 for San
Clemente Island females. They estimate that
about half of the female population is reproduc-
tively active in a given year. Young are born in
September, following a 14-week gestation period.
Goldberg and Bezy (1974) estimate that sexual
maturity is attained in the spring of the third or
fourth year. They consider X. riversiana to be a
K-selected species, being relatively late-maturing,
long-lived, with single broods, large adult body
x:
<b'
Sonto
I
»c «e >e ■•Lfi
Distribution of the island night hzard Xantusia riversiana.
size, and large clutches, and demonstrating a low
reproductive potential.
MANAGEMENT AND CONSERVATION
The greatest threats to this lizard are preda-
tion and habitat destruction from introduction
of exotic species into these fragile island eco-
systems. Bezy (1976) notes that feral goats could
have a serious impact on this partially herbivorous
lizard and its habitat. He also notes that feral pigs
and cats undoubtedly feed on the lizard to some
extent.
There is evidence of predation by feral house-
cats on X. riversiana on San Clemente Island, but
the magnitude of this loss has not been determined.
A feral-cat removal program is being planned for
San Nicolas Island (Steele 1979). The Cahfomia
Department of Fish and Game and the U.S. Navy
are currently considering a removal program for
some feral animals on San Clemente Island. The
Navy has a program for the control of feral goats,
but it is currently inactive (Federal Register 42,
No. 155).
The alligator lizard [Gerrhonotus multicarin-
atus) has been accidentally introduced on San
Nicolas Island. Bezy (1976) feels that this lizard
may be a competitor with X. riversiana. G. multi-
carinatus and X. riversiana have exclusive distribu-
tions on the California Channel Islands (Savage
1967), which suggests that alligator lizards may
displace the smaller-sized island night lizard.
AUTHORITIES
Robert L. Bezy
Section of Herpetology
Natural History Museum
Los Angelos, California 90007
John M. Brode
California Department of Fish and Game
1701 Nimbus Road
Rancho Cordova, California 95670
PREPARER'S COMMENTS
Bezy (1972, 1976) indicated that there is
moderate morphological differentiation between
the three island populations of X. riversiana.
Variation within the species needs further study.
The impact of predation by feral animals on
San Clemente and San Nicolas Island populations
of X. riversiana, and the potential competitive
impact by Gerrhonotus multicarinatus on San
Nicolas Island populations are both in need of
immediate investigation. Control of these intro-
duced animals may be critical to the survival of
the endemic island night lizard. Remaining habi-
tat on San Nicolas and Santa Barbara Islands
should be protected from further decimation.
LITERATURE CITED/ SELECTED
REFERENCES
Bezy, R. L. 1972. Karyotypic variation and
evolution of the lizards in the family
Xantusidae. Los Angelos Nat. Hist. Mus.,
Contri. Sci., 227: 1-29.
-. 1976. Letter in response to federal pro-
posal of threatened or endangered status
for Klauberina riversiana. (42 FR 155:
40682-40683.
Brattstrom, B. H. 1952. The food habits of
the night lizard, genus Xantusia. Copeia
1952(2):168-172.
Goldberg, S. R. and R. L. Bezy. 1974. Repro-
duction in the island night lizard, Xantu-
sia riversiana. Herpetologica 30:350-360.
Savage, J. M. 1957. Studies on the lizard family
Xantusiidae. III. A new genus for Xantu-
sia riversiana Cope, 1883. Zoologica 42:
82-86.
-. 1967. Evolution of the insular herpeto-
fauna. Pages 219-227 in Philbrick, R. N.
(ed.) Proceedings of the Symposium on
the Biology of the California Islands. 363
pp.
Schwenkmeyer, R. C. 1949. Food habits of
the island night lizard, Xantusia riversiana
reticulata, from San Clemente Island. Nat.
Hist. Misc. 38:1-3.
Stebbins, R. C. 1954. Amphibians and reptiles
of western North America. McGraw-Hill,
New York. 537 pp.
•. 1966. A field guide to the western rep-
tiles and amphibians. Houghton Mifflin,
Boston. 279 pp.
Steele, J. 1979. San Nicolas Island — home of
Indian artifacts, unusual wildlife and
guided missiles. Outdoor Calif. 40(5):
29-30.
ACCOUNT PREPARED BY /UPDATED BY
National Fish and Wildlife Laboratory
1300 Blue Spruce Drive
Fort Collins, Colorado 80524
Biological Services Program
FWS/OBS-80/01.37
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE DELMARVA PENINSULA FOX SttUIRREL
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubHc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVV'IS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to;
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
SHdell, Louisiana 70458
11
FWS/OBS-80/01.37
March 1980
SLLhXI KD V1:R 1 LBRAl i: KNDANGERED SPECIES
OF THE SEACOAST OF 1 HE UNFFED STATES-
THE DELMARVA PENINSULA FOX SQUIRREL
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell. Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
DELMARVA PENINSULA FOX SQUIRREL
Sciurvs niger cinereus Linnaeus
KINGDOM Animalia
CLASS Mammalia
ORDER Rodentia
FAMILY Sciuridae
OTHER COMMON
NAMES Bryant's fox squirrel,
peninsula fox squirrel, stump-eared squirrel.
DATE
Entered into SWIS to be determined
Updates 30 September 1978
LEGAL STATUS
Federal: Endangered (32 FR 4001; 11 Mar.
1967).
States: Endangered: Maryland.
REASONS FOR CURRENT STATUS
Primary factors for current status are destruc-
tion of habitat from timber cutting, agricultural
clearing, construction, development (Flyger 1964;
lUCN 1972; Taylor and Flyger 1973; USFWS
1973; DFSRT draft); and competition with the
gray squirrel [Sciurus carolinensis) in areas of sub-
optimal habitat. Minor mortality results from
hunt kills and road kills (Delmarva Fox Squirrel
Recovery Team [DFSRT] draft).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The Delmarva Peninsula fox squirrel is a fairly
large squirrel (larger than gray squirrel) with pelage
a uniform light grizzled-grayish with steel-blue
cast dorsally and white ventrally. The feet are
white and the tail has a pronounced black
stripe on outer edge. Melanistic forms occur with
black both dorsally and ventrally. Pictures are in
Taylor (no date) and DFSRT (draft).
RANGE
The squirrel is presently restricted to local dis-
junct populations in portions of four eastern shore
counties of Maryland (Allen 1942;Masueti 1950;
Paradiso 1969; J. Taylor personal communication).
There is an introduced population on Chincoteague
National Wildlife Refuge, Virginia (lUCN 1972;
USFWS 1973).
The species' former distribution included
southeastern Pennsylvania, eastern shore of Mary-
land and Delaware (Delmarva Peninsula), the
eastern shore of Virginia into Northampton Coun-
ty, and possibly southern New Jersey (Abbot
1830, Rhoads 1903, Poole 1932, 1944, Hall and
Kelson 1959).
RANGE MAP
Diagonal hatching on the following map de-
picts present range; shading indicates former
range (Taylor and Flyger 1973, DFSRT draft).
STATES/COUNTIES
Maryland: Dorchester, Kent, Queen Annes
Virginia:
Talbot.
Accomack.
HABITAT
The squirrel prefers mature mixed timber
with a minimum of undergrowth (Taylor and
Flyger 1973; Taylor 1976). It is thought to be
closely associated with stands of loblolly pine
[Pinus taeda) (lUCN 1972). However, others argue
that this may not be correct, because its former
distribution was north of the distribution of lob-
lolly pine (Taylor and Flyger 1973, Taylor 1976).
It is often found in savanna areas, oak open-
ings and in narrow tree zones along rivers and
streams. It does occur sparingly in areas with dense
undergrowth, but is then forced to compete with
the gray squirrel.
FOOD AND FORAGING BEHAVIOR
A variety of large trees provide adequate mast
in fall. Trees utilized include oak, hickory, wal-
nut, and loblolly pine. In spring the squirrel con-
sumes buds and flowers of trees, fungi, insects,
fruits, seeds, and occasionally bird eggs and young
(Smith and Folmer 1972; DFSRT draft; Taylor no
date).
They will consume a large vjuriety of agricul-
tural products such as corn, soybeans, or other
crops in fields and orchards. They are more ter-
restrial than the gray squirrel and will venture
farther into open fields.
SHELTER REQUIREMENTS
See nesting or bedding.
NESTING OR BEDDING
Tree dens and leaf nests are built with local
materials. They appear to use den nests in the win-
ter and leaf nests in the summer (G. \. Taylor per-
sonal communication). Paradiso (1969) states
that nests are placed near the tips of branches in
old pines, from 9 to 15 m above the ground.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
In north central populations of Sciurus niger,
home ranges average about 4 ha. The seasonal
average for home range is approximately 16
ha (Allen 1943). They are not disturbed by hu-
man activity any more than gray squirrels, pro-
vided habitat is satisfactory (DFSRT draft).
This species encounters potential competition
in areas of local sympatry with the gray squirrel
in suboptimal habitat.
POPULATION NUMBERS AND TRENDS
There are no estimates of the total number of
individuals. They are considered abundant only at
Eastern Neck Island National Wildlife Refuge,
Kent County, Maryland (DFSRT draft). Radio-
collaring projects are underway to provide infor-
mation on numbers.
REPRODUCTION
Fox squirrels are polygamous and after mat-
ing, the female raises the young alone. The breed-
ing season is extended, with peaks in February
and March, and in July and August (DFSRT
draft). Gestation period is 45 days (Asdell 1964).
Litter size is 2 to 4 with an average of 3 (lUCN
1972); 1 to 2 litters are produced per year.
The amount of food available seems to govern the
size as well as the number of litters (DFSRT
draft). Young are born naked. Eyes open at 5
weeks and young are weaned at 9 to 12 weeks.
PENNSYLVANIA
EASTERNNECK'
NWR
BLACKWfiTER<^\
NWR V^
LE COMPTE WMA
CHINCOTEAGUE
NWR
VIRGINIA
KM
70
PRESENT
RANGE
HISTORICAL
RANGE
This map depicts the historical and present range of the Delmarva Peninsula fox squirrel.
Females may breed at 10 to 11 months. Young
can be disturbed and handled with minimal
adverse effects. Females will not abandon young
if disturbed but may move them to a different
location (DFSRT draft).
MANAGEMENT AND CONSERVATION
The establishment of Blackwater National
Wildlife Refuge (1933) in Maryland has helped
protect some habitat (lUCN 1972). LeCompte
Wildlife Management Area, Maryland, was desig-
nated as a refuge for this species in 1970. Mary-
land closed the hunting season on this species in
1971. The species was successfully introduced at
Chincoteague National Wildlife Refuge, Virginia,
in 1968.
Several hundred nest boxes have been placed in
selected habitats and many used in Chinco-
teague National Wildlife Refuge. Several thou-
sand nest boxes have been built, and some put out
in Maryland. Studies with gray squirrels have
indicated that provision of nest boxes nearly
doubled the population (Barkalow and Soots
1965; Burger 1969).
Management proposals by the State of Virgin-
ia include establishing a nest box program and sur-
veying for additional sites of suitable habitat for
reestablishment (Taylor and Fisher 1978). The
State of Maryland's proposals include, in addi-
tion, the use of abundance and life history studies
(G.J. Taylor personal communication).
A recovery team has been formed and a final
draft recovery plan will be formulated by late
1978. The primary objective of the recovery plan
is the restoration of the Delmarva fox squirrel to
a secure status throughout its former range
(DFSRT draft). The plan includes three primary ac-
tions: (1) preservation and management of essen-
tial denning and feeding habitat; (2) release of
Delmarva fox squirrels into suitable habitat; (3)
protection of populations through law enforce-
ment, reduction of squirrel competition and pre-
dation; and (4) promotion of public support.
Tentative plans also include aquisition of private
lands on which this species occurs. Designation of
Critical Habitat will be proposed by 30 June 1979.
The following is a list of public lands on
which the squirrel occurs and related population
estimates (F = Federal, S = State, NWR = Na-
tional Wildlife Refuge).
State
County
Location/
Ownershp
Pop.
Source
Md.
Dor-
chester
Blackwater
NWR(F)
450
W.H. Julian
pers. com.
Kent
Ejistern
Neck
NWR (F)
250
P. Feiger
pers. com.
Dor-
chester
LeCompte
Wildl. Man.
Area (S)
no
estimate
Va.
Acco-
mack
Chinco-
teague
NWR (F)
80-100
DFSRT
pers.com.
AUTHORITIES
Vagn Flyger (Recovery Team)
1222 H.J. Paterson Hall
University of Maryland
College Park, MD 20742
Bernard F. Halla (Recovery Team)
Nongame Wildlife Program,
Maryland Department of Natural Resources
Wildlife Administration
Tawes State Office Building
Annapolis, MD 21401
William H. Julian (Recovery Team)
U.S. Fish and Wildlife Service
Blackwater National Wildlife Refuge
Rt. l,Box 121
Cambridge, MD 21613
Gary J. Taylor (Recovery Team)
Nongame and Endangered Species Program
Manager
Maryland Department of Natural Resources
Wildlife Administration
Tawes State Office Building
Annapolis, MD 21401
PREPARER'S COMMENTS
Known in literature as Sciurus niger bryanti,
S. n. neglectus, and S. n. cinereus.
LITERATURE CITED/SELECTED
REFERENCES
Abbott, C. C. 1890. The geological survey of New
Jersey, 1890. Vol. 2, Part 2:500. John Mur-
phy Co., Trenton.
Allen, CM. 1942. Extinct and vanishing mammals
of the western hemisphere. Am. Comm. Int.
WUdl. Port. Spec. Publ. 11:45-46.
Allen, D. C. 1943. Michigan fox squirrel manage-
ment. Mich. Dep. Conserv. Game Div. Publ.
100. Lansing.
Asdell, S. A. 1964. Patterns of mammalian repro-
duction. Cornell Univ. Press, Ithaca. 670 pp.
Bakken, A. 1952. Interrelationships of Sciurus
caroliinensis (Gmelin) and Sciurus niger (Lin-
naeus) in mixed popula. Ph.D. Dissert. Univ.
Wisconsin. 188 pp.
Barkalow, F. S., Jr. and R. F. Soots, Jr. 1965. An
analysis of the effect of artificial nest boxes
on a gray squirrel population. Trans. N. Am.
Wildl. Natur. Resour. Conf. 30:349-360.
Burger, G. V. 1969. Response of gray squirrels to
nest boxes at Remington Farms, Maryland. J.
Wildl. Manage. 33:796-801.
DFSRT (Delmarva Fox Squirrel Recovery Team).
Draft. Delmarva fox squirrel recovery plan.
U.S. Fish Wildl. Serv. 39 pp.
Dozier, H. L., and H. E. Hall. 1944. Observations
on the Bryant's fox squirrel, Sciurus niger
bryantii, (Bailey). Md Conserv. 21:1-12.
Flyger, V. 1964. Urban sprawl endangers native
Maryland mammals. Md. Conserv. 41:6-7.
Hall, E. R., and K. R. Kelson. 1959. Mammals of
North America. 2 vols. Ronald Press, New
York. 1,083 pp.
lUCN. 1972. Red data book. Vol. I, Mammalia.
Compiled by H. A. Goodwin and C. W. Hollo-
way. lUCN, Morges, Switzerland.
Lustig, L. W., and V. Flyger. 1976. Observations
and suggested management practices for the
endangerd Delmarva fox squirrel. Proc. Annu.
Conf. Southeast. Assoc. Game Fish Comm. 29:
433-440.
Mansueti, R. 1950. Extinct and vanishing mam-
mals of Maryland and the District of Colum-
bia. Md Natur. 20:1-48.
Moore, J. C. 1946. Mammals from Welaka, Put-
nam County, Florida. J. Mammal. 27:490-59.
. 1957. The natural history of the fox
squirrel Sciurus niger shermani. Bull. Am.
Mus. Natur. Hist. 113:1-70.
Paradiso, J. L. 1969. Mammals of Maryland. N.
Am. Fauna 66:1-93.
Poole, E. L. 1932. A survey of the mammals of
Berks County, Pennsylvania. Reading Pub.
Mus. Art Gall. Bull. 13.
• 1944. The technical names of the north-
eastern fox squirrels. J. Mammal. 25:315-317.
Rhoads, S. N. 1903. The mammals of Pennsyl-
vania and New Jersey. Philadelphia.
Smith, C, D. Follmer. 1972. Food preferences of
squirrels. Ecology 53:82-91.
Taylor, G. J. 1974. Present status and habitat sur-
vey of the Delmarva fox squirrel {Sciurus
niger cinereus) with a discussion of reasons
for its decline. Proc. Annu. Conf. Southeast.
Assoc. Game Fish Comm. 27:278-189.
. 1976. Range determination and habitat
description of the Delmarva fox squirrels in
Maryland. M.S. Thesis. Univ. of Maryland,
College Park.
. No date. Maryland's Delmarva fox squir-
rel. Maryland Dep. Natur. Resour. Fact Sheet.
2 pp.
Taylor, G. J., and V. Flyger. 1973. Distribution
of the Delmarva fox squirrel (Sciurus niger
cinereus) in Maryland. Chesapeake Sci. 14:
59-60.
Taylor, W. H., and E. Fisher. 1978. Delmarva fox
squirrel investigations. Performance report
from July 1977 to June 1978. Proj. No. E-2.
Commonwealth of Virginia, Comm. Game
Fish, Richmond. 3 pp.
USFWS. 1973. Threatened wildlife of the United
States. Compiled by Office of Endangered
Species and International Activities, Bur.
Sport Fish. Wildl. Resour. Publ. 114. U. S.
Gov. Printing Office, Washington, D.C. 289 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.38
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE HOUSTON TOAD
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or ihrcalencd vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
.\,'\SA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.'>8
u
FWS/OBS-80/01.38
March 1980
SELKCl KI) Vl.Rl KBRAl i: ENDANGERED SPECIES
OF rHE SEACOAS 1 OF 1 HE UNFFED SFA FES-
THE HOUSTON TOAD
A Cooperative Effort
by tin-
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National C:()astal Ecosystems Feam,
Office of Biological Services
Project Officer
Donald \V. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
I ish anti Wildlife Service
U.S. Department of the Interior
HOUSTON TOAD
Bufo houstonensis Sanders
KINGDOM Animalia
CLASS Amphibia
ORDER Anura
FAMILY Bufonidae
OTHER COMMON NAMES Wottring's toad
DATE
Entered into SWIS to be determined
Updates 1 January 1978, 31 March 1978,
22 May 1979
LEGAL STATUS
Federal: Endangered (35 FR 16047, 13 Octo-
ber 1970)
States: Endangered: Texas.
REASONS FOR CURRENT STATUS
Populations are extremely small and their
known distribution is restricted to two or three
disjunct localities. Early field studies indicated
Houston toads preferred temporary rain pools for
breeding sites, whereas other native toads
generally used more permanent bodies of water.
Man's alteration of natural watersheds has re-
duced the availability of temporary pools, forcing
Houston toads to breed in larger ponds, lakes, and
creeks where they have hybridized with the Gulf
Coast toad {B. valliceps) and the Woodhouse's
toad (J5. woodhousei) (Brown 1971). The prob-
lem is compounded because population levels are
so low that the probability of a female Houston
toad finding a conspecific mate in a mixed chorus
of toad species is slim (Brown 1971).
The Houston toad is generally considered a
post-Pleistocene relict of the more northerly dis-
tributed American toad (B. americanus) (Blair
1958, 1965) and may be ill-adapted to southern
environments (Brown 1971). If true, this may ex-
plain the species' apparent failure to adapt to
habitat alterations.
PRIORITY INDEX
Not assigned.
DESCRIPTION
A small toad, the Houston toad is similar to
the American toad, but with heavier cranial crests,
especially behind the eyes. Males have a 49- to
66-mm snout-to-vent length; females meaure 57
to 80 mm. The dorsum is light brown to cream
with a variable number of dark brown to black
spots, each with one or more warts. The venter is
cream, usually heavily mottled. A hght mid-dorsjd
stripe is usually present. Tadpoles are virtually
indistinguishable from American toad larvae
(Altig 1970). The mating call consists of a long
(7- to 22-sec), high-pitched (1,646- to 2,300-Hz)
trill with 14 to 16 pulses per second (Brown
1973).
Photographs appear in Sanders (1953), Blair
(1959, 1972), Kennedy (1962), Brown (1971),
Thomas (1975), Thomas and Potter (1975), Eh-
renfield (1976), and Hardy (1977). Audio-
spectrograms of the mating call are given in
Brown (1971, 1973).
RANGE
The species' historical range included the fol-
lowing localities: northwest and southeast Hous-
ton, Harris County; Lake Woodrow, Burleson
County; 6 miles south of Liberty, Liberty Coun-
ty; Austin County; and 6 and 12.6 miles east of
Columbus, Colorado County (Sanders 1953).
Brown (1971) also reported populations 2 miles
west of Fresno in Fort Bend County and in the
vicinity of Bastrop and Buescher State Parks in
Bastrop County.
The species has not been found in Liberty,
Austin, Colorado, or Fort Bend counties since
originally reported. It is now known to exist only
in the vicinity of Bastrop and Buescher State
Parks, Bastrop County; Lake Woodrow, Burleson
County; and possibly southeast Houston, Harris
County.
RANGE MAP
Historic localities are designated on the follow-
ing map by triangles; present known localities are
indicated by dots. Critical Habitat as designated
in 43 FR 4022, 31 January 1978, is shown on a
separate map.
STATES/COUNTIES
Texas: Austin,' Bastrop, Burleson, Colora-
do,* Fort Bend,' Harris, Liberty.'
HABITAT
Houston toads are restricted to areas charac-
terized by sandy soils. Localities in Bastrop and
Burleson Counties are heavily wooded with lob-
lolly pine {Pinus taeda) and/or mixed deciduous
forest, interspersed with open grassy areas. The
southeast Houston locality is residential with
open, grassy fields.
Breeding habitats include roadside ditches,
temporary ponds in residential areas and pastures,
and other seasonally flooded low spots. Houston
toads also breed in Lake Woodrow and nearby
permanent ponds.
FOODS AND FORAGING BEHAVIOR
Bragg (1960) reported captives feeding on va-
rious insects and smaller toads. Thomas (unpub-
lished data) examined 17 specimens and found
unidentified red ants in one and beetle remains in
another.
SHELTER REQUIREMENTS
Apparently requires sandy loamy soils for
burrowing.
NESTING OR BEDDING
The species breeds in temporary rain pools,
flooded fields, and permanent ponds.
RITUAL REQUIREMENTS
Males call from small mounds of soil or grass
surrounded by water, or from shallow water.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
' Believed extirpated in these counties.
V
H
^
c
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u
•a
-a
o
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c
o
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3
O
X
V
J3
C
o
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Critical habitat of the Houston toad, as designated in 43 FR 4022, 31 January 1978, is depicted
by shading in this map.
POPULATION NUMBERS AND TRENDS
Populations of Houston toads in Harris County
have decreased markedly since the late 1940's, but
the rate has not been documented. John Wottring,
the naturalist responsible for recognition of the
Houston toad as a distinct species, reported collect-
ing 66 from a single chorus in 1949, and "quite a
lot" in 1953 (from his field notes). Brown (1967,
1971) found only three during the breeding sea-
sons in 1965, 1966, and 1967. Extensive surveys
in 1974, 1975, and 1976 revealed the presence of
two Houston toads in southeastern Houston (R.
Thomas unpublished data). Surveys in 1978 failed
to locate any Houston toads in Harris County
(Jameson and Handerstein unpublished).
Populations in Bastrop and Burleson Counties
have apparently remained stable during the past
10 years, estimated at a maximum of 300 indivi-
duals (Brown 1975).
Thomas and Potter (1975) estimate the total
population to be near 1,500 individuals. Mark-re-
capture studies have not been attempted.
REPRODUCTION
Breeding is initiated by heavy rains from Feb-
ruary to June and lasts only a few nights (Kenne-
dy 1962). One female laid 728 eggs (Kennedy
1962). Hybridization with Woodhouse's toads
(fertile hybrids) and Gulf Coast toads (infertile
hybrids) has been documented (Brown 1971).
MANAGEMENT AND CONSERVATION
A Recovery Team was appointed in January
1978 to develop a plan for removal of this species
from endangered status. Critical Habitat has been
designated (43 FR 4022, 31 January 1978). Areas
of land, water, and airspace in two counties were
designated: (1) Bastrop County, from the junc-
tion of line corresponding to 30°12'00"N and
Texas State Highway 95 east along a line corres-
ponding to 30°12'00"N, to where it intersects a
line corresponding to 97°17'30"W, to where it
intersects the Colorado River, west and northwest
along the north bank of the Colorado River to
the due southward extension of Texas State High-
way 95, and north along the extension and Texas
State Highway 95 to where it intersects a line
corresponding to 3O°12'00"N; (2) Burleson Coun-
ty: a circular area with a 1.6-km radius, the center
being the north entrance to Lake Woodrow from
Texas FM 2000.
About 285 ha at Buescher State Science Park
are reportedly being maintained in a condition
as favorable for Houston toads as possible (F. Pot-
ter personal communication).
The endangered Houston toad has received
considerable publicity and most local land owners
are aware of the problem.
Houston toads were reared at the Houston
Zoo in 1978 and 1979. The first project involved
collecting eggs in the wild and then releasing over
1,000 tadpoles back into the wild. In 1979, eggs
from Houston toads were spawned at the zoo and
at the time this report was prepared (May 1979),
tadpoles were being raised for subsequent release.
AUTHORITIES
Lauren E. Brown (Recovery Team)
Department of Biological Sciences
Illinois State University
Normal, IL 61761
William L. McClure (Recovery Team)
Houston Urban Project
P.O.Box 187
Houston, TX 77001
Floyd E. Potter (Recovery Team)
Texas Parks and Wildlife Department
4200 Smith School Road
Austin, TX 78744
Norman J. Scott, Jr. (Recovery Team)
US FWS
Natl. Fish & Wildl. Lab.
Albuquerque, NM
Robert A. Thomas (Recovery Team)
Louisiana Nature Center
4100 One Shell Square
New Orleans, LA 70139
Okla W. Thornton
5525 4th Street No. 99
Lubbock, TX 79416
PREPARER'S COMMENTS
There is an obvious paucity of base-line eco-
logical information about the Houston toad. Pro-
per management of the species will require such
data.
LITERATURE CITED/SELECTED
REFERENCES
Altig, R. 1970. A key to the tadpoles of the Con-
tinental United States and Canada. Herpetolo-
gica26(2):180-207.
Blair, W. F. 1956. Call difference as an isolation
mechanism in southwestern toads (genus
Bufo). Texas J. Sci. 8:87-106.
. 1958. Distributional patterns of verte-
brates in the southern United States in rela-
tion to past and present environments. Pages
433-468 in Hubbs, C. L., ed. Zoogeography.
Am. Assoc. Adv. Sci. Publ. 51.
. 1959. Genetic conpatibility and species
groups in U.S. toads (Bufo). Texas J. Sci.
427-453,
1963. Intragroup genetic compatibility
in the Bufo americanus species group of toads.
Texas J. Sci. 15:15-34.
. 1965. Amphibian speciation. Pages 543-
556 in Wright, H. E. and D. G. Frey, eds. The
Quartemary of the United States. Princeton
Univ. Press, Princeton.
. ed. 1972. Evolution in the genus Bufo.
Univ. Texas Press, Austin.
Bragg, A. N. 1960. Feeding in the Houston toad.
Southwest Natur. 5:160.
Brown, L. E. 1967. The significance of natural
hybridization in certain aspects of the specia-
tion of some North American toads (genus
Bufo). Ph. D. Dissert. Univ. Texas, Austin.
. 1971. Natural hybridization and trend
toward extinction in some relict Texas toad
populations. Southwest Natur. 16:185-199.
. 1973. Bufo houstonensis Sanders. Cat.
Am. Amphib. Rep. 133.1-1-133.2.'
. 1975. The status of the near-extinct
Houston toad (Bufo houstonensis) with
recommendations for its conservation. Herp.
Rev. 6:37-38.
Ehrenfeld, D. W. 1976. The conservation of non-
resources. Am. Sci. 64:648-656.
Guttman, S. 1. 1969. Blood protein variation in
the Bufo americanus species group of toads.
Copeia 1969:243-249.
Hardy, M. 1977. Froggie goes to court. Texas
Monthly 5:76.
Kennedy, J. P. 1962. Spawning season and experi-
mental hybridization of the Houston toad,
Bufo houstonensis. Wer^etolo^ca. 17:239-245.
Sanders, O. 1953. A new species of toad with a
discussion of the bufonid skull. Herpetologica
9:25-47.
Thomas, R. A. 1975. Investigation on the status
of the Houston toad, Bufo houstonensis
Sanders. Unpubl. Final Rep. to the U.S. Fish
Wildl. Serv. Contract 14-16-0002-3557.
Contains complete bibliography for the species through
1973.
• 1975. (Cover photo: Bufo houstonensis).
Herp. Rev. 6(2).
Thomas, R. A., and F. E. Potter, Jr. 1975. Species
status account: Bufo houstonensis Sanders.
Texas Parks Wildl. Dep. Job Perform. Rep.
70:1-11.
GATEKEEPER
To be designated by Office of Endangered
Species.
ACCOUNT PREPARED/UPDATED BY:
Louisiana Nature Center
4100 One Shell Square
New Orleans, LA 70129
and
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.39
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE AMERICAN ALLIGATOR
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data are not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps ol Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared t)r as the status of species is changed.
Suggestions or questions regarding SWIS should be directed Id:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed t(;:
Information Transfer Specialist
National Coastal Ecosystems leani
U.S. Fish and Wildlife Senice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Shdell, Louisiana 704.^)8
u
FWS/OBS-80/01.39
March 1980
SLLKCIKU VI:RI EBRAl i: KNDANGERED SPECIES
OF IHE SEACOAS 1 OF I HE UNFIED STATES-
THE AMERICAN ALLIGATOR
A Cooperative Effort
by tin-
National Fish and WildHfc Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems 1 eani,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidcll Computer Complex
1010 Cause Blvd.
Slidell. Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fisii and Wildlife Service
U.S. Department of the Intenor
CREDIT: LUTH ER C. GOLDMAN
AMERICAN ALLIGATOR
Alligator mississippiensis (Daudin)
KINGDOM Animalia
CLASS Reptilia
ORDER Crocodylia
FAMILY Alligatoridae
OTHER COMMON
NAMES alligator, gator
DATE
Entered into SWIS to be determined
Updates 21 August 1978.
LEGAL STATUS
Federal: Endangered (40 FR 44418, 26 Septem-
ber 1975), wherever found in wild ex-
cept where threatened (42 FR 2076,
10 January 1977; 44 FR 37132, 25
June 1979; 44 FR 49084, 12 October
1979) or protected by Similarity of Ap-
pearance (40 FR 44418, 26 September
1975; 44 FR 37132, 25 June 1979; 44
FR 59084, 12 October 1979).
Threatened (42 FR 2076, 10 January
1977; 44 FR 37132, 25 June 1979;
44 FR 59084, 12 October 1979) in
Florida and along the southern Coastal
Plain within the following boundaries:
from Winyah Bay near Georgetown,
S.C, west of U.S. Highway 17 to
Georgetown; thence west and south on
U.S. Alternate Highway 17 to junction
with U.S. Interstate Highway 95 near
Watersboro, S.C; thence south on U.S.
Interstate Highway 95 (including in-
complete portions) to junction with
U.S. Highway 82; thence southwest on
U.S. Highway 82 to junction with U.S.
Highway 84 at Waycross, Ga., thence
west on U.S. Highway 84 to the Ala-
bama-Georgia border; thence south
along this border to the Florida border
and following Florida border west and
south to its termination at the Gulf of
Mexico; then from the Mississippi-Lou-
isiana border at the Gulf of Mexico
north along this border to its junction
with U.S. Interstate Highway 12;
thence west on U.S. Interstate Highway
12 (including incomplete portions) to
Baton Rouge, La.; thence north and
west along corporate limits of Baton
Rouge to U.S. Highway 190; thence
west on U.S. Highway 190 to junction
with Louisiana State Highway 12 at
Ragley, La., thence west on Louisiana
State Highway 12 to the Beauregard-
Calcasieu Parish border; thence north
and west along this border to the Texas-
Louisiana State border; thence south
on this border to Texas State Highway
12; thence west on Texas State High-
way 12 to Vidor, Tex., thence west on
U.S. Highway 90 in the Houston, Tex.,
corporate limits; thence north, west,
and south along Houston corporate
limits to junction on the west with U.S.
Highway 59; thence south and west on
U.S. Highway 59 to Victoria, Tex.,
thence south on U.S. Highway 77 to
corporate limits of Corpus Christi,
Tex.; thence southeast along the south-
em Corpus Christi corporate limits to
Laguna Madre; thence south along the
west shore of Laguna Madre to the
Nueces-Kleberg county line; thence
east along the Nueces-Kleberg county
line to the Gulf of Mexico.
Threatened (Similarity of Appearance
to Endangered and Threatened popula-
tions) (40 FR 37132, 25 June 1979) in
the wild in Cameron, Vermilion, Cal-
casieu, Ibemia, St. Mary, St. Charles,
Terrebonne, Lafourche, St. Bernard,
Jefferson, St. Tammany, and Plaque-
mine Parishes in Louisiana.
States: Endangered: Delaware, Georgia, Mas-
sachusetts, Mississippi, North Caro-
lina, Texas.
Threatened: Florida.
REASONS FOR CURRENT STATUS
Although the range of the American alligator
is but slightly reduced from historic boundaries,
population levels over much of the present range
are reduced due to loss of habitat and overharvest.
The low point in population decline was reached
in the late 1950's to mid-1960's, and in 1967 the
species was listed by the United States as endan-
gered.
Man and his associated technology are the
major threats. Alligators were killed for food by
the early settlers, to a lesser extent for leather, in
many cases out of fear, and because of livestock
depredation. It was not until 1855, however, that
any attempt was made to take large numbers for
their leather. This activity was interrupted to
some extent by the Civil War, although limited
commercial use by the Confederacy for leather
and grease continued during the war (Audubon
1931). By 1870, alligator leather again became
fashionable, and alligators were killed in the
southeast from 1870 until the mid-1960's. Smith
(1893) stated that at least 2,500,000 were killed
in Florida from 1800 to 1891. Early estimates in-
dicate that the number surviving in Florida and
Louisiana in 1902 was less than 20% of what it
had been 20 years earlier (Stevenson 1904).
From the late 1940's through the mid-1960's,
the remaining wetland areas were opened up.
Improved marsh transportation such as airboats
and marsh buggies placed unreasonable pressure
on remaining alligator populations. Chabreck
(1967) estimated that between the late 1940's
and late 1950's, populations in Louisiana de-
clined by 90%.
Outright destruction of wetlands has had
serious effects upon alligators throughout their
range. Millions of hectares of wetlands that at
one time were prime alligator habitat have been
drained throughout the southeast. Areas in south
Florida that harbored impressive numbers of
alligators as late as 1954 are now dewatered and
in agricultural production (Florida Game and
Fresh Water Fish Commission, unpubl. data).
Other areas have experienced dramatic changes in
annual water-level fluctuations, which have
affected populations. Hines et al. (1968) docu-
mented severe nest losses in the Everglades due to
flooding. Man's drainage and diking activities have
increased the frequency and magnitude of water-
level fluctuations in the Everglades, resulting in
frequent nest destruction, undoubtedly an
important factor limiting reproduction in that
part of Florida.
State and Federad protection has reversed the
population decline, and alligators are now stabili-
zed or increasing in numbers in most of their
range.
The chief threat to the American alligator is
now probably the rapid urbanization underway
throughout its range and the increasing conver-
sion of habitat to recreational use and develop-
ment. The alligator is a large predatory species,
and, in close contact with human populations.
may constitute an apparent threat as both a
competitor to and predator on man, Thus, even
a few homes on a lake or river can generate con-
siderable pressure for a reduction in the local
alligator population, especially the larger indi-
viduals. Growing human intolerance could result
in serious and possibly insurmountable obstacles
to maintaining the species over much of its
present range.
PRIORITY INDEX
Not assigned.
DESCRIPTION
The alligator is a lizard-Hke reptile that
reaches lengths of more than 4 m and weights
up to 150 kg. Adults are generally dark gray or
black, but the young are brightly patterned with
black and yellow. The body is covered with rough
scales dorsally and smooth scales ventrally. The
tail is strongly compressed laterally and crested
with high, pointed scales.
The only native species with which the alliga-
tor might be confused is the American crocodile.
The alligator has a broad, rounded snout, whereas
the crocodile's snout is sharply tapered forward
of the eyes. The fourth tooth in the crocodile's
lower jaw is exposed when the mouth is closed;
in the alligator, this tooth is covered by the upper
lip. Crocodiles are generally lighter colored as
adults than alligators, being light gray or olive as
compared to the alligator's dark gray or black.
Young crocodiles are olive with black bars and
markings, rather than black and yellow. Most
standard field guides and reptile books illustrate
the differences between these species (e.g., Dit-
mars 1953, Carr and Coin 1955, Pope 1955,
Bothwell 1962, Neill 1971,Conant 1975, Perrero
1975). All these works, as well as a wide range of
texts, magazines, and general outdoor publica-
tions provide color and black-and-white photo-
graphs.
RANGE
Alligators range throughout wetland habitats
in the coastal region of southeastern United States
from central North Carolina to Texas and north
along the Mississippi River drainage into extreme
southeastern Oklahoma and southern Arkansas
(Joanen 1974).
The present range approximates the historical
range. There is evidence that the species once
ranged north of central North Carolina into the
Dismal Swamp region of southeastern Virginia
and northeastern North Carolina (Neill 1971).
The southern extent of its historic range in Texas
and possibly northeastern Mexico is unknown;
however, records exist from the Rio Grande, indi-
cating the possiblity that it once ranged into the
extensive aquatic habitats of northeastern Tamau-
lipas, Mexico.
RANGE MAP
Range is delineated by shading on the follow-
ing map (T. Joanen personal communication).
STATES/COUNTIES
Alabama: Autauga, Baldwin, Barbour, Bullock,
Butler, Choctaw, Clarke, Coffee, Co-
necuh, Covington, Crenshaw, Dale,
Dallas, Escambia, Geneva. Greene,
Hale, Henry, Houston, Lowndes,
Macon, Marengo, Mobile, Monroe,
Montgomery, Perry, Pickens, Pike,
Russell, Sumter, Tuscaloosa, Washing-
ton, Wilcox.
Arkansas: Hempstead, Lafayette, Miller.
Florida: All.
Georgia: Appling, Atkinson, Bacon, Baker,
Baldwin, Ben Hill, Berrien, Bibb,
Bleckley, Brantley, Brooks, Bryan,
Bullock, Burke, Calhoun, Camden,
Candler, Charlton, Chatham, Chatta-
hoochee, Clay, Clinch, Coffee, Col-
quitt, Cook, Crawford, Crisp, Decatur,
Dodge, Dooly, Early, Echols, Effing-
ham, Emanuel, Evans, Glynn, Grady,
Hancock, Harris, Heard, Houston, Ir-
win, Jeff Davis, Jefferson, Jenkins,
Johnson, Jones, Lanier, Laurens, Lee,
Liberty, Long, Lowndes, Macon, Ma-
rion, Mcintosh, Meriwether, Miller,
Mitchell, Montgomery, Muscogee,
Peach, Pierce, Pulaski, Putnam, Quit-
man, Randolph, Richmond, Schley,
Screven, Seminole, Stewart, Sumter,
Talbot, Tattnall, Taylor, Telfair, Ter-
rell, Thomas, Tift, Toombs, Treutlen,
Troup, Turner, Twiggs, Upson, Ware,
Washington, Wayne, Webster, Wheeler,
Wilcox, Wilkinson, Worth.
Louisiana: All.
Mississippi: Adams, Amite, Attala, Carroll, Clai-
borne, Clarke, Clay, Coahoma, Copiah,
Covington, Forrest, George, Greene,
Grenada, Hancock, Harrison, Hinds,
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Holmes, Humphreys, Issaguena, Jack-
son, Jefferson, Jones, Lafayette,
Lamar, Lauderdale, Lawrence, Leake,
Leflore, Lincoln, Lowndes, Madison,
Marion, Monroe, Montgomery, Ne-
shoba, Newton, Noxubee, Panola,
Pearl River, Perry, Pike, Quitman,
Scott, Sharkey, Simpson, Smith,
Stone, Tallahatchie, Walthall, Warren,
Washington, Wayne, Wilkinson, Yazoo.
North
Carolina: Beaufort, Bladen, Brunswick, Carteret,
Columbus, Craven, Cumberland, Dare,
Duplin, Hyde, Jones, New Hanover,
Onslow, Pamlico, Pender, Pitt, Robe-
son, Sampson, Scotland, Tyrrell,
Washington.
Oklahoma: McCurtain.
South
Carolina: Aiken, Allendale, Bamberg, Barnwell,
Beaufort, Berkeley, Calhoun, Charles-
ton, Chesterfield, Clarendon, Colleton,
Darlington, Dillon, Dorchester, Flo-
rence, Georgetown, Hampton, Horry,
Jasper, Kershaw, Lee, Lexington,
Marion, Marlboro, Orangeburg, Rich-
land, Sumter, Williamsburg.
Texas: Anderson, Angelina, Aransas, Archer,
Atascosa, Austin, Beek, Bexar, Brazo-
ria, Brazos, Burleson, Calhoun, Came-
ron, Cass, Chambers, Collin, Colorado,
Dimmit, Ellis, Fort Bend, Freestone,
Frio, Galveston, Goliad, Grimes, Har-
din, Harris, Harrison, Henderson,
Houston, Jackson, Jasper, Jefferson,
Jim Wells, Johnson, Karnes, Kimble,
Kleberg, Lamar, La Salle, Lavaca,
Leon, Liberty, Live Oak, Madison,
Marion, Matagorda, McMullen, Mont-
gomery, Morris, Nacogdoches, Navar-
ro, Newton, Nueces, Orange, Polk, Re-
fugio, Robertson, Sabine, San Augus-
tine, San Jacinto, San Patricio, Shelby,
Smith, Travis, Trinity, Tyler, Victoria,
Walker, Waller, Webb.
HABITAT
Alligators inhabit, or inhabited, practically all
fresh and brackish water habitats in their range
except as excluded by man. Large marsh-bordered
lakes, fresh and brackish marshes, and savannas
appear to provide optimal habitat and support the
densest populations (Reese 1915, Mcllhenny
1935,Fogarty 1974).
FOOD AND FORAGING BEHAVIOR
The food of alligators consists of virtually any
animal small enough to be captured and swal-
lowed (Kellogg 1929, Mcllhenny 1935). Available
data indicate some variation between size classes
(Kellogg 1929) and between habitat types (Cha-
breck 1971). Small alligators feed extensively on
invertebrates, including crayfish, aquatic and ter-
restrial insects, and mollusks (Fogarty and Albury
1967, Neill 1971, Valentine et al. 1972), while
larger individuals take vertebrate prey, including
wading birds, snakes, turtles, small mammals, and
fish. Very large alligators may attack and eat large
animals such as deer, cattle, and even man (Neill
1971, Hines and Keenlyne 1977).
SHELTER REQUIREMENTS
Adults often dig dens or 'gator holes' at the
edges of rivers or lakes and in marsh habitats. An
underground tunnel with an underwater entrance
will lead into an underground cave a short dis-
tance back from the water (Kellogg 1929;
Mcllhenny 1935; Craighead 1969; Joanen and
McNease 1971, 1972; Neill 1971).
NESTING AND BEDDING
Alligators build mound nests of heaped-up ve-
getation in clearings in marshes or along the edges
of lakes or rivers. Mounds are generally about 2 m
in diameter and 0.5 to 1 m high (Mcllhenny 1935,
Joanen 1969, Joanen and McNease 1972, Neill
1971).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
Joanen (1974) provided estimates of alligator
populations in all areas throughout the range. The
total estimated population for the species was
738,384. In addition, the estimated available
habitat was given as 116,550 km^ for an average
of six alligators per km^ of suitable habitat.
Trends in alligator populations have been
monitored by the Cooperative AHigator Survey, a
yearly night survey over standardized survey
routes throughout the range (Chabreck 1974).
These data indicate a generally increasing trend.
Joanen (1974) also presents data that support this
observation. Outside of Florida, 168 counties had
increasing populations, 157 had stable popula-
tions, and 25 had decreasing populations.
Joanen's (1974) data are summarized by States
below.
Alabama: Some 12,715 alligators were reported
from 28 counties. Only two counties report
decreasing populations, while 15 report increasing
and 11, stable populations. There is a trend for
increasing populations over much of the southern
half of Alabama, with relatively greater abun-
dance in the region east and north of Mobile Bay.
Arkansas: Most of the previous range is apparent-
ly now uninhabited; 1,900 alligators were present
in three counties in the southwestern comer of
the State. These individuals are presumed to have
been transplanted into the area from Louisiana
over the past few years.
Florida: Alligators are reported in every respond-
ing county. Over 407,500 alligators were esti-
mated for the State. No population trend esti-
mates were given. In general, peninsular Florida,
roughly south of the Suwannee Strait, supports
moderate to large alligator populations that are
either increasing or remaining stable in areas of
increasing urbanization except in intensive de-
velopment centers. The panhandle supports
generally lower populations, with local pockets
of abundance.
Georgia: 95 counties, essentially all counties with-
in the historical range of the species in the State,
report alhgator populations; 45 counties report
populations increasing and 32 report stable
populations.
Available National Wildlife Refuge, State, and
other data show that there are alligator popula-
tions in the coastal counties and the Okefenokee
Swamp area that are subjectively as large as, or
larger than, those in southeastern South Carolina
and northern Florida. As to the remainder of the
State's populations, it can only be said that they
are generally widespread and may be locally
abundant, although less so than in the south-
eastern sector.
Louisiana: A total of 200,682 alligators distribu-
ted over 63 parishes v^th 40 reporting increasing
populations, and 22 stable. The extensive coastal
marshes of southern Louisiana probably support
the largest alligator populations anywhere in the
species' range, although some other local popula-
tions, e.g., Okenfenokee Swamp in Georgia, may
have an equal density. The situation in the north-
em, non-marsh, parishes is less clear, but numbers
there are clearly low and apparently similar to
other areas of the species' range.
Mississippi: Only 4,740 alligators were found in
55 counties — 13 with increasing populations and
38 stable. Coastal counties generally support the
highest populations.
North Carolina: Reports 1,300 alligators on about
2,590 km^ of habitat. Highest populations are in
central and southern coastal counties. No alliga-
tors are reported between Albemarle Sound and
the Dismal Swamp, Virginia, an area within the
recent historical range of the species. Only 4 of
21 reporting counties show an increase in popula-
tion.
Oklahoma: Estimates 10 alligators in one south-
eastern county. Supplemental data from field
biologists and State agencies suggests that consid-
erable unoccupied habitat exists in the southeast
corner of the State.
South Carolina: Reports 48,700 alligators on
9,479 km^ of habitat. Ten of the 28 responding
units report increases, while the others have stable
populations. As in North Carolina, populations in
the southeastern coastal counties are the largest,
and are generally increasing. Reports of increases
in some inland and northern counties suggest that
the species has responded well to protection
throughout the State.
Texas: Has 26,784 alhgators in 74 counties.
Populations are reported increasing in 35
and stable in 22. Several reporting units appear to
represent introduced alligators outside of the
historic range of the species. Large, increasing
populations are recorded for counties adjacent to
the coastal Louisiana parishes, and south along
the Gulf coast to Matagorda Bay. Interior coun-
ties appear to support small scattered populations
which are generally increasing or stable.
REPRODUCTION
Alligators and other crocodilians exhibit the
most elaborate social behavior of any reptile,
most of which is related to reproduction (Mcll-
henny 1935, Carr 1967, Joanen and McNease
1971, Campbell 1972, Kushlan 1973, Carr 1976,
Pooley and Gans 1976).
Reproductive behavior begins with bellowing
by males and females, usually in March or April,
depending on local climate. The bellow has been
reported to serve as a territorial signal and as a
component of male -female interaction, but data
are not available to fully document its function
(Campbell 1973, Herzog 1974, Garrick and Lang
1977).
Copulation takes place in the water (Mcllhenny
1935, Joanen 1969). Nests are built and eggs
layed in May to June depending on local climate,
and eggs hatch in August to September (Kellog
1929, Mcllhenny 1935, Joanen 1969, Neill 1971).
Clutch size varies from 20 to 60 eggs.
At least some females are protective of the
nest and remain nearby during the incubation
period. Eggs hatch in about 9 weeks. The 23cm
hatchlings are very vocal with a high-pitched
"umph, umph" which may stimulate the female
to help open the nest. The young often stay to-
gether in a "pod" for the rest of the summer
(Mcllhenny 1935, Campbell 1973, McNease and
Joanen 1974). Tagged young have been recaptured
near the nest site as long as 3 years after hatching
(Mcllhenny 1935, Joanen 1969, Campbell 1973,
McNease and Joanen 1974,Metzen 1979).
Survival rate is unknown except in local situa-
tions, but is expected to be low in mature popula-
tions and higher in depleted populations. Variation
in water level appears to be the major limiting
factor on nesting success (Hineset al. 1968, Joanen
1969, Nichols etal. 1976).
Growth rates of the young vary widely but
may approach 0.3m per year with optimal habitat
and food availability (Mcllhenny 1935, Jo; nen
and McNease 1975). Subadults often stay togeth-
er in loose groups until they are about Im long;
those over 1.5 m long often make extensive pre-
sumed dispersal movements (Chabreck 1965,
McNease and Joanen 1974).
MANAGEMENT AND CONSERVATION
Laws protecting alligators were, with some
exceptions, ineffective until amendment of the
Lacy Act in 1969 enabled the federal government
to control interstate shipment of alligator hides.
By the late 1960's, populations began to increase
in the Louisiana coastal marshes due to better law
enforcement and excellent landowner coopera-
tion. However, populations in other States did
not begin to increase on a wide scale until inter-
state control of marketing of illegal hides was
achieved (Chabreck 1971b). With effective control
of poaching since 1969, alligator populations have
made substantial recoveries in most areas of the
species' range.
The rapid resurgence suggests that the species
is highly responsive to conservation efforts and
amenable to standard management programs.
Managed harvests are now being made in three
Louisiana parishes — Calcasieu, Cameron, and
Vermillion — which are listed as threatened (Simi-
larity of Appearance) on the Endangered Species
List of the U.S. Department of Interior. A sport
hunting program is being developed in the same
area. A control program for nuisance alligators in
Florida derives much of its overhead from com-
mercial use of the skins.
The U.S. Fish and WildHfe Service's Recovery
Plan for the species is in draft form and a com-
plex set of regulatory and management recom-
mendations for dealing with regional and local
problems and opportunities will soon be proposed.
AUTHORITIES
Frosty Anderson (Recovery Team)
National Audubon Society
Miles Wildlife Sanctuary
Sharon, CT 06069
Howard W. Campbell (Recovery Team)
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Robert Chabreck (Recovery Team)
School of Forestry and Wildlife Management
Louisiana State University
Baton Rouge, LA 70803
Tommy Hines (Recovery Team)
Florida Game and Fresh Water
Fish Commission
4005 South Main Street
Gainesville, FL 32601
Ted Joanen (Recovery Team Leader)
Rockefeller Wildlife Refuge
Route l,Box 20-B
Grand Chenier, LA 70643
Larry McNease
Rockefeller Wildlife Refuge
Route 1, Box 20-B
Grand Chenier, LA 70643
Thomas Murphy
Biologist
Coco Sabo Plantation
Greenpond, SC 29446
Ronald R. Odom (Recovery Team)
Georgia Game and Fish Division
Endangered Species Program
Route 2
Social Circle, GA 30279
A. William Palmisano (Recovery Team
Consultant)
Natl. Fish and Wildlife Lab
Alaska Field Station
4454 Business Park Blvd.
Anchorage, AL 99503
Jake Valentine (Recovery Team)
Wildlife Biologist
U.S. Fish and Wildlife Service
Box 4753, Univ. of Southwestern Louisiana
Lafayette, LA 70501
PREPARER'S COMMENTS
None.
LITERATURE CITED /SELECTED
REFERENCES
Audubon, J.J. 1931. Observations of the natural
history of the alligator. Louisiana Conserv.
Rev. 2(3):3-8.
Bothwell, D. 1962. The great outdoors book of
alligators and other crocodilia. Great Out-
doors Publ. Co. St. Petersburg. 88 pp.
Brazaitis, P. 1973. The identification of living
crocodilians. Zoologica 58:(3-4):59-101.
Campbell, H. W. 1972. Ecological or phylogenetic
interpretations of crocodilian nesting habits.
Nature 238:404-405.
. 1973. Observations on the acoustic
behavior of crocodilians. Zoologica 1973
(Spring): 1-10.
Carr, A.F. 1967. Alligators, dragons in distress.
Natl. Geogr. Mag. 131:133-148
^.1976. Excerpts from the life an alli-
gator: a reappraisal of "the alligator's life
history." Foreword to Mcllhenny, E. A. 1935.
The alligator's life history. Facsimile reprint,
1976. Soc. Study Amphib. Reptiles Misc.
Publ.
Carr, A. F. and C. J. Goins. 1955. Guide to the
reptiles, amphibians, and fresh-water fishes
of Florida. Univ. Florida Press, Gainesville
341 pp.
Chabreck, R. H. 1965. The movement of alliga-
tors in Louisiana. Proc. Annu. Conf. South-
east Assoc. Game Fish Comm. 19:102-110.
. 1967. The American alligator:past
present and future. Proc. Annu. Conf. South-
east Assoc. Game Fish Comm. 21:554-558.
. 1972. The food and feeding habits of
alligators from fresh and saline environments
in Louisiana. Proc. Annu. Conf. Southeast
Assoc. Game Fish Comm. 25:117-124.
. 1971b. The management of the Amer-
ican alligator. Proc. 1st Working Meeting of
the Crocodile Specialists. lUCN Suppl. Paper
32:137-144.
1974. Cooperative surveys of the Amer-
ican alligator in the southeastern United
States, Mimeo report.
Conant, R. 1975. A field guide to reptiles and
amphibians of eastern and central North
America. Houghton Mifflin Co., Boston. 429
pp.
Craighead, F. C. 1968. The role of the alligator in
shaping plant communities and maintaining
wildlife in the southern Everglades. Fla. Natur.
41:2-7,69-74.
Ditmars, R. L. 1953. The reptiles of North
America. Doubleday & Co., Inc., New York.
476 pp.
Fogarty, M. J. 1974. The ecology of the Ever-
glades Alligator, in P. J. Gleason. ed.. Envi-
ronments of South Florida: Past and Present.
Mem. Miami Geol. Survey 2. 482 pp.
Fogarty, M. J. and J. D. Albury. 1967. Late
summer foods of young alligators in Florida.
Proc. Annu. Conf. Southeast Assoc. Game
Fish Comm. 21:220-222.
Garrick, L. D. and J. W. Lang. 1977. Social signals
and behaviors ofadult alligators and crocodiles.
Am. Zool. 17:225-239.
GUes, L.W. and V. L. Childs. 1949. Alligator
management on the Sabine National Wildlife
Refuge.J.Wildl. Manage. 13(1): 16-28.
Herzog, H. A., Jr. 1974. The vocal communication
system and related behaviors of the American
alligator (Alligator mississippiensis) and other
crocodilians. M. S. Thesis, Univ. Tennessee,
Knoxville. 87 pp.
Hines, T. C, M. J. Fogarty, and L. C. Chappell.
1968. Alligator research in Florida: a progress
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Game Fish Comm. 22:166-180.
Hines, T. C. and K. D. Keenlyne. 1977. Two
incidents of alligator attacks on humans in
Florida. Copeia. 1977(4):735-738.
Joanen, T. 1969. Nesting ecology of alligators in
Louisiana. Proc. Annu. Conf. Southeast
Assoc. Game Fish Comm. 23:141-151.
,,1974. Population status and distribution
of alligators in the southeastern United States.
Southeastern Regional End. Sp. Workshop,
Wildl. Soc. Tallahassee, Fla. Sept. 1974.
Unpaged.
Joanen, T. and L. McNease. 1971. A telemetric
study of nesting female alligators on Rocke-
feller Pvcfuge, Louisiana. Proc. Annu. Conf.
Southeast Assoc. Game Fish Comm. 24:
175-193.
%1972. A telemetric study of adult male
alligators on Rockefeller Refuge, Louisiana.
Proc. Annu. Conf. Southeast Assoc. Game
Fish Comm. 26:252-275.
.1976. Notes on the reproductive biology
and captive propagation of the American
alligator. Proc. Annu. Proc. Southeast Game
Fish Comm. 29:407-415.
Kellogg, R. 1929. The habits and economic
importance of alligators. U.S. Dept. Agric.
Tech. Bull. 147:36 pp.
Kushlan, J. A. 1973. Observations on maternal
behavior in the American alligator. Alliga-
tor mississippiensis . Herpetologica 29(3):
256-257.
•1974. Observations on the role of the
American alligator {Alligator mississippiensis
in the southern Florida wetlands. Copeia
1974(4):993-996.
Mcllhenny, E. A. 1935. The alligator's life history.
Christopher Publ. House, Boston. 1 16 pp.
McNease, L. and T. Joanen. 1974. A study of
immature alligators on Rockefeller Refuge,
Louisiana. Proc. Annu. Conf. Southeast Game
Fish Comm. 28:482-500.
Metzen, W. D. 1979. Nesting ecology of alligators
on the Okefenokee National Wildlife Refuge.
Proc. Annu. Conf. Game Fish Comm. 31.
Neill, W. T. 1971. The last of the ruling reptiles:
alligators, crocodiles, and their kin. Columbia
Univ. Press, Columbia. 486 pp.
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B. Fenderson. 1976. Simulation of a commer-
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iana. Louisiana Agric. Exp. Stn. Bull 691. 59
pp.
O'Neil, T. 1949. The muskrat in Louisiana coastal
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Perrero, L. 1975. Alligators and crocodiles of
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281-532.
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ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.40
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
BROWN PELICAN
Eastern and California Subspecies
Fish and Wildlife Service
U.S. Department of the Interior
PREPACK
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, us amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps ol Kngineers in coordina-
tion with the Offices of Kndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions .ind deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should he direi led lo:
Office of Endangered Species
U.S. Fish and Wildlife Senicc
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be direcii'd to:
Information Transfer Specialist
National Coastal Ecosystems leam
U.S. Fish and Wildlife Ser\ ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.^)8
u
FWS/OBS-80/01.40
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
BROWN PELICAN
Eastern and California Subspecies
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
BROWN PELICAN
(Eastern subspecies— Pelc caims
occidcntalis carolinensis Gmelin)
(California s\ihspec\es—Pclcc anus
occidcntalis californicus Rigdway)
KINGDOM Animalia
CLASS Aves
ORDER Pelecaniformes
FAMILY Pelecanidae
OTHER COMMON NAMES
Eastern subspecies Eastern brown pelican
California subspecies . . . California brown pelican
DATE
Entered into SWIS to be determined
Updates
LEGAL STATUS
Federal: Endangered: (35 FR 16047, 13 October
1970:35 FR 18319, 2 December 1970).
States: Endangered: Georgia, Mississippi,
South Carolina, California.
Threatened: Florida.
REASONS FOR CURRENT STATUS
Both the eastern and California subspecies
have been greatly reduced in numbers. The main
reason for their decline is accumulation of chlorin-
ated hydrocarbon residues, which have been
shown to cause eggshell thinning and subsequent
crushing of eggs under the weight of parent birds
(Heath et al. 1969, Peakall 1970, Wiemer and
Porter 1970). The principal residues involved are
DDT compounds (DDE, DDD, and DDT), poly-
chlorinated biphenyls (PCB's),dieldrin,andendrin
(Schreiber and Risebrough 1972, BIus et al.
1974a). The correlation between eggshell thinning
and DDE concentrations is highly significant
(Schreiber and Risebrough 1972, Blus et al. 1975).
Blus et al. (1974b) demonstrated that the success
of 93 brown pelican nests was related to low levels
of DDE and dieldrin.
Other factors affecting pelican populations
are availability of food supply and human distur-
bance of nesting colonies. Parent pelicans flushed
from nests exert forces large enough to break
eggs. Normally, one of the parents remains on the
nest throughout incubation. Desertion of the nest
can resuh in losses to predation as well as tem-
perature stresses on the eggs and young (Schreiber
and Risebrough 1972, Schreiber 1979).
Mercury may also interfere with reproduction;
mercury has been found in some pelican eggs at
levels known to have adverse effects on other spe-
cies (Blus et al. 1974a).
Eastern subspecies
Along the Atlantic Coast, eggshell thinning is
more acute at the northern end of the pelican's
range (Blus et al. 1970). From 1969 to 1974, Blus
et al. (1974a) noted that 17% eggshell thinning in
North Carohna populations was associated with
subnormal reproductive success. By 1975, egg-
shell thinning had decreased to 10% (L. J. Blus
personal communication). The present Florida
population is stable, eggshell thickness has been
reduced about 9%, and about 3% of all eggs on
the west coast of Florida have been thin-shelled
and crushed. The effects of these factors on pop-
ulation levels may not be apparent, for in 1975
and 1976, eggs in Florida had little or no thinning
or crushing (R. W. Schreiber personal communica-
tion).
In Louisiana and Texas, where the pelican
populations have declined rapidly and dramat-
ically, the levels of pollution are higher (Schrei-
ber and Risebrough 1972). In Louisiana, eggs
from transplanted Florida birds showed a steady
decline in shell thickness from 1970 to 1973.
Levels of DDE were generally lower than those
associated with subnormal reproductive success
in South Carolina, but dieldrin levels were slightly
higher. Half the eggs examined contained levels of
dieldrin considered potentially detrimental to re-
productive success (Blus et al. 1975).
Forty percent of a small population trans-
planted to Louisiana from Florida died of endrin
poisoning in the spring and summer of 1975
(Winn 1975).
Freezes, hurricanes, and beach erosion may
have affected population numbers (Schreiber and
Risebrough 1972). Blus et al. (1974a) believe that
in South Carolina, storms and other naturad fac-
tors have minimal effects on pelican populations
because, while they may temporarily disrupt nest-
ing activities, the birds are persistent renesters in
that area.
Human disturbance that causes the parent
birds to desert the nests can result in serious losses
to predation by fish crows (Corvus ossifragus), as
well as in temperature stress on eggs and young
(Schreiber and Risebrough 1972, Schreiber 1979).
In Florida, a significant number of pelicans
(500 or more) are killed each year after being
caught on fishhooks or entangled in monofila-
ment fishlines. Some are also maliciously killed or
maimed by people (Schreiber 1979).
The decline of the pelicans in South Carolina
in the 1950's coincided with a decline in Atlantic
menhaden [Brevoortia spp.). There is some evi-
dence that food shortages occur, especially in
Florida, and the red tide (Gymnodimium) has
probably affected pelican food supplies on the
west coast of Florida at times (Williams et al.
1976).
California subspecies
Pelican losses in California resulting from
paratyphoid infections have probably had little
effect on the total population (Bond 1942). The
complete failure of reproductive efforts on the
California Channel Islands in 1968, reported by
Schreiber and DeLong (1969), probably resulted
from chlorinated-hydrocarbon pollution. Levels
of residues as high as 84.4 ppm were found in the
muscles of birds from Monterey Bay in December
of 1966 (Risebrough et al. 1971). Reproduction
on Anacapa Island, California, and Los Coronados
and San Martin Islands in Baja California, was al-
most entirely unsuccessful from 1969 to 1972.
Success varied on San Benito Island in Baja Cali-
fornia (400 miles south of the U.S. border) and
on remote islands in the Gulf of California. Far-
ther south, reproductive success Wcis more satis-
factory. The U.S. Fish and Wildhfe Service (1973)
attributed the poor reproductive success to egg-
shell thinning brought about by contamination
with DDE.
In 1974, eggshell thinning was much less evi-
dent than in previous years on the Pacific Coast,
and many more young were produced in some of
the colonies where egg collapse had been severe.
More adults bred in 1974, as well. However, col-
lapsed eggs and lowered productivity continued
to be evident. Production on San Martin Island
failed completely in 1974, presumably because
of human disturbance.
In the Gulf of California, where production
had been comparatively good, it was very poor in
1974. The cause appeared to be poor food condi-
tions, although the presence of DDE in the food
chain may have contributed. A decrease in the
availability of anchovies brought about by over-
fishing is thought to have affected brown pelican
reproduction (Keith 1978). Recent studies indi-
cate that minor restrictions in food supply ser-
iously impair reproduction and that the presence
of DDE intensifies the effects of food deprivation
(U.S. Fish and Wildlife Service, Denver WildHfe
Research Center ms. 1974; Keith 1978).
PRIORITY INDEX
Eastern subspecies: None assigned.
California subspecies: 13.
DESCRIPTION
Eastern subspecies
P. o. carolinensis is a large bird, with a total
length of 114 to 137 cm and a wingspan of 2 m.
It has a long bill and a large gular pouch, which is
colored greenish during the breeding season.
Adults have a black belly with gray wings and
back. The head is yellow from early autumn until
late spring, when it turns white. The neck is white
in the fall and winter, but becomes chestnut
brown during the breeding season.
Immatures are brown with white bellies for
the first 3 to 5 years (Schreiber 1979). Photo-
graphs may be found in Bent (1922), Peterson
(1947), and Palmer (1962), among others.
California subspecies
P. o. californicus is distinguished from the
eastern subspecies by its larger size and darker
coloration. The brown color of the back of the
neck is almost black, and the gular pouch is
reddish instead of greenish during the breeding
season (Baird et al. 1884, U.S. Fish and WOdlife
Service 1973, Bent 1922, Wetmore 1945). The
young have white down when first hatched.
The eggs of both subspecies are lusterless
white, becoming dirty during incubation. The
measurements of the eastern birds' eggs average
73 to 46 mm and those of the California birds,
78.5 by 50.6 mm. Pelicans typically fly by alter-
nate flapping and soaring. The California birds
can be seen to take advantage of updrafts produced
when winds axe deflected by the coastal bluffs,
soaring considerable distances without flapping
and thus reducing the energy required for flight
(Woodward 1921).
RANGE
Eastern subspecies
On the eastern seaboard, the pelican ranges
from North Carolina to Florida, around the Gulf
Coast to Texas and Mexico, and southward to
Venezuela. It also inhabits the West Indies and
many islands in the Caribbean. Casual or sporadic
sightings have been reported from Nova Scotia,
Michigan, Wyoming, Colorado, and Bermuda
(Bent 1922).
Breeding colony sites are listed in Schreiber
and Risebrough (1972), Blus et al. (1974a) and
WilHamset al. (1976).
Pelicans are usually resident near their breed-
ing grounds, although young birds tend to wander.
Schreiber (1976a) color-marked individuals and
observed that the majority of fledglings moved
south away from the colony during the first
months out of the nest. Some individuals wander
extensively.
The Florida population has been found to be
divided into east- and west-coast groups that do
not appear to mix. Birds in South Carolina and
on the east coast of Florida move south in the
fall, but remain on the east coast and in the upper
Florida Keys. Pelicans hatched at Tampa Bay also
move southward in the fall, being common at
Flamingo, upper Florida Bay, and the lower Keys.
The former range is the same as the present
one, except that the pelican was once a common
breeder on the coast of Louisiana; it has been un-
known there since 1966 except for a small group
transplanted from Florida. The Texas and northern
Mexico populations have been much reduced
(U.S. Dept. of the Interior 1973, Schreiber 1979).
Brown pelicans breed only on coastal islands and
in Florida; fewer than 50 islands are currently in
use (Schreiber 1979).
On the 45 sites occupied since 1950, 26
(58%) are in State ownership; 9 (20%) are Fed-
erally owned; 5 (11%) are in miscellaneous or un-
certain ownership. Only the Federally owned sites
are considered secure (Williams et al. 1976).
California subspecies
At present, this species breeds locally on is-
lands along the Pacific coast from Anacapa Island,
Ventura County, California to Los Coronados,
San Martin, and San Benito Islands off the coast
of Baja California; on islands in the Gulf of Cali-
fornia, and south to Tres Marias Islands off
Nayarit. Historical nesting sites are Santa Cruz
and San Miguel Islands, Santa Barbara County,
and Bird Rock, Point Lobos, Monterey County,
California. Postbreeding movement of birds
northward along Pacific Coast in late summer and
fall is common (Wetmore 1945; A.O.U. 1957;
Anderson and Anderson 1976; Fish and Wildlife
Service 1973; Jehl 1973;Wmiams 1931). Between
breeding seasons, the range extends northward
along the coast to southern British Columbia
(Burrard Inlet) and inland to Central British
Columbia (Chilcotin District) and southward
along the Pacific Coast of Mexico an unknown
distance, but at least to Colima (Bond 1942). The
main northward movements are in July.
Rarely, pelicans wander inland in California
(Stanislaus County, 19 Sept. 1913) (Wetmore
1945; Bent 1922), but usually stay very closely
restricted to the seashore. Sightings in the interior
are rare (Mailliard 1913); Grinnell and Miller
1944; Wetmore 1945). Of 71 recoveries of peli-
cans banded as young on Anacapa Island, taken
mostly along the coast from Marin County, Cali-
fornia south to Nayarit, Mexico, 4 were recovered
inland at: Potrero, San Diego Co. (40 km inland)
San Gabriel, Los Angeles Co. (34 km inland)
Santa Maria, Santa Barbara Co. (18 km inland)
amd Petrolia, Humboldt Co. (9 km inland) (Bond
1942, 1948). Of 31 band recoveries or sightings
of brown pelicans marked as nestlings in the Gulf
of California area and recorded north of the nest-
ing area, 7 (33%) were from inland areas in the
Southwest Desert; all were first-year recoveries.
Record numbers of brown pelicans were reported
during July 1972 in the Tucson-Phoenix, Arizona
vicinity and the Salton Sea in California in July
and August 1972. Anderson et al. (1972a, 1977)
suggested that such dispersal results from dom-
inant south winds, often with heavy thunder-
storms that ground the birds, rather than being
the result of searching for food.
RANGE MAPS
The distribution of the eastern brown pelican
is indicated by shading, and its breeding colonies
are represented by dots on the following maps.
The breeding rjinge of the California subspecies
is shown on the range map for this group.
STATES/COUNTIES
Eastern subspecies
Alabama: Baldwin, Mobile.
Florida: Bay, Brevard, Broward, Charlotte,
Citrus, Collier, Dade, Dixie Duval,
Escambia, Flagler, Franklin, Gulf,
Hernando, Hillsborough, Indian
River, Jefferson, Lee, Levy, Mana-
tee, Martin, Monroe, Nassau, Oka-
loosa, Palm Beach, Pasco, Pinellas,
Santa Rosa, Sarasota, Seminole, St.
Johns, St. Lucie, Taylor, Volusia,
Wakulla, Walton.
Georgia: Bryan, Camden, Chatham, Glynn,
Liberty, Mcintosh.
Louisiana: Cameron, Jefferson, LaFourclie,
Plaquemines, St. Bernard, St. Mary,
Terrebonne, Vermilion.
Mississippi: Hancock, Harrison, Jackson.
N. Carolina: Brunswick, Carteret, Currituck,
Dare, Hyde, New Hanover, Onslow,
Pamlico, Pender.
S. Carolina: Beaufort, Charleston, Colleton,
Georgetown, Horry.
Texas: Aransas, Brazoria, Calhoun, Cam-
eron, Chambers, Galveston, Jeffer-
son, Kenedy, Kleberg, Matagorda,
Nueces, San Patricio, Willacy.
California subspecies
California: Del Norte, Humboldt, Los Angeles,
Marin, Montery, Orange, San Diego,
San Luis Obispo, Santa Barbara,
Santa Cruz, Stanislaus, Ventura.
HABITAT
Eastern subspecies
Pelicans usually feed in shallow estuarine
waters, although they are sometimes seen 30 to
60 km offshore (Schreiber 1979). Groups are
often observed flying over the surf on both the
Atlantic and Gulf shores of Florida, occasionally
feeding beyond the breakers. Schreiber (1979)
states that sandspits and offshore sandbars are
used for loafing in the daytime and roosting at
night.
Brown pelicans, especially juvenile birds,
frequent fishing piers where scraps are available
(Schreiber 1979).
Habitat photographs are found in Bent (1922).
California subspecies
The primary habitat is the ocean littoral just
outside the surf line. This pelican rarely strays
either inland or far offshore. It is confined to the
semiarid western coast, bathed by the relatively
cool waters of the California current, which pro-
bably limits its range southward (Murphy 1936,
Anderson and Anderson 1976).
The preferred nesting habitat is on offshore
islands, although some individuals nest in man-
grove growing in estuarine locations along the
Sinaloa coast.
NORTHWESTEfiN PART OF
FLORIDA
15 16 17 18 19 20 II
23 2* 25
Brown pelican colony sites in Florida.
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Point Lobos
San Miguel Island
Santa C ruz Island
Santa Barbara Island
San Nicolas Island
Santa Catalina Island
Los Coronados Islands
(Border Island)
San Martin Island
San Benito Island
Tres Marias
Islands
NftpARrr
Breeding range of the California brown pelican.
8
FOOD AND FORAGING BEHAVIOR
The mode of prey capture and the type of
food are the same for both subspecies. The birds
fly low over the water, spot a potential prey fish,
and dive to capture it. The fish is then transferred
to the gular pouch. The particular species of
fishes mjiking up most of the diet of the eastern
subspecies are menhaden {Brevoortia), mullet
[Mugil), sardines [Sardinella), and pinfish {Lago-
don) (Bent 1922, Schreiber 1979).
The California subspecies feeds mostly on
anchovies and other members of the family En-
graulididea; Scombridae; and Pacific sardines
(Sardinox sagax) when available. The Pacific
sardines and another former food fish, the Pacific
mackerel [Scomber japonicus) are now essentially
gone from CaUfomia waters, but northern ancho-
vies appear to have replaced them, and most
biologists beheve that the long-term decline of
pelicans on the west coast is unrelated to changes
in food supply (Anderson and Anderson 1976).
Keith (1978), however, believes that former sar-
dine fishermen in the Gulf of California have
switched to anchovy fishing to the probable detri-
ment of pelican food supply. Bostic and Banks
(1966) found stingrays (Dasyatidae) and tongue-
fish [Cunoglossidae) in the pouch of a dead peli-
can near San Felipe in the Gulf of California. A
stingray spine embedded in the bird's throat pro-
bably caused its death.
SHELTER REQUIREMENTS
None noted.
NESTING OR BEDDING
Pelicans prefer to nest on offshore island sites
that are protected from flooding, human distur-
bance, and terrestrial predators such as raccoons
{Procyon lotor) (Blus et al. 1974a, Schreiber per-
sonal communication). Although there are dis-
tinct similarities, there are also conspicuous dif-
ferences in the types of nest sites selected by the
two subspecies.
Eastern subspecies
Eastern brown pelicans build nests on the
ground, on mud lumps, or in trees (Bent 1922).
In Florida, they nest primarily in mangrove trees
[Avicennia germinans and Rhizophora mangle)
from 1 to 10 m above the high-tide mark (Schrei-
ber 1979).
Ground nests vary greatly in size and struc-
ture from practically nothing to large, well-built
nests of sticks, reeds, straw, palmetto leaves, and
grasses. Remains of old nests are often used, and
fresh material may be stolen from newly con-
structed nests when the owner is gone. Nest dia-
meter on Pehcan Island, Florida, ranges from 46
to 61 cm, and height ranges from 10 to 13 cm.
Arboreal nests are more firmly built of similar
materials, on substantial stick platforms securely
interwoven with the branches of the supporting
tree (Bent 1922).
Pelicans are colonial nesters, and generally
select small coastal islands in salt or brackish
water and lying landward of barrier islands or
reefs where they are protected from the surf and
ocean storms (Williams et al. 1976).
California subspecies
The California brown pelican prefers to nest
on steep, rocky slopes of islands, building its nests
of sticks, grasses, and rubbish. In some areas,
nests are reused year after year, accumulating
filth and reaching heights as great as 1 m. On 12
April 1888, A. W. Anthony (m Bent 1922) found
nests in groups of 20 to 30 on San Martin Island,
located about 0.4 km from the beach and about
76 m above the sea. Most of the nests were built
on the tops of low bushes, but many were on bare
ground or blocks of lava. Sticks, twigs, kelp, sea-
grass, and even a few bird bones were used as
building material. On 16 April 1939, Bond (1942)
found several nests on Anacapa Island in island
oak [Quercus tomentella) and toyon [Photinia
arbutifolia), at heights 3.2 to 4.8 m above the
ground— an unusual nesting situation for the spe-
cies.
Williams (1927) found nests on top of mats of
poison oak at Point Lobos. Nests on mangroves
over water on the Sinaloa coast are reported in a
manuscript of the Denver Wildlife Research Cen-
ter, U.S. Fish and Wildlife Service (1974).
RITUAL BEHAVIOR
S. C. Arthur {in Bent 1922) describes court-
ship behavior of a male P. o. occidentalis as a
slow circling of the motionless female with pon-
derous elephantine tread, while he lifted his
wings slightly and tilted his head far back. It is
presumed that the courtship behavior of P. o.
califomicus is similar.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None other than noted elsewhere.
POPULATION NUMBERS AND TRENDS
Eastern subspecies
The Florida population is stable with an adult
breeding population of 6,705 to 7,690 pairs in
1968-1970 (Williams and Martin 1968, 1970) and
6,000 to 8,000 pairs for the last 6 years. The esti-
mated total population in Florida is less than
30,000 birds (Schriber 1979).
Blus et al. (1974a) state that the South Caro-
lina population declined from 5,000 to 1,000
pairs by the late 1960's.
Schreiber and Risebrough (1972) give a de-
tailed breakdown of the status of the brown peli-
can population in each State. In North Carolina,
the population on Shell Castle Island (northern-
most breeding colony on the Atlantic coast) has
fluctuated between 1 1 and 100 from 1960 through
1967. Fewer than 30 young were believed fledged
in 1970. In South Carolina, colonies are found on
the Cape Romain Wildlife Refuge and at Deveaux
Bank south of Charleston. At Cape Romain Re-
fuge, populations have fluctuated drastically from
year to year. The Deveaux Bank colony has un-
dergone a 90% reduction in the last 10 years, due
perhaps to much of the bank's washing away.
Brown pelicans have probably never nested in
Georgia. There has been no nesting in Alabama
since 1900, and in 1956-57, the local nonbreeding
population declined sharply with no recovery as
of 1972. In Louisiana, the once large population,
estimated at between 75,000 and 85,000 (Arthur
1931), had decreased to four individuals by 1960,
according to Audubon Christmas Bird Count esti-
mates. Although Florida pelicans were released in
Louisiana from 1968 to the 1970's and some
were reported nesting, no native pelicans have
bred since 1966 (Schreiber 1979). In Texas, fewer
than 10 pairs nested in 1969 and 1970. The total
population in Louisiana and Texas is now esti-
mated to be under 500 (Schreiber 1979).
Population status is essentially unrecorded in
eastern Mexico, the Caribbean, and Central Am-
erica (Schreiber and Risebrough 1972).
California subspecies
About 1,000 to 1,500 adults were breeding in
California in 1972; the California population has
been declining 14% to 18% per year. The minimum
population in Mexico and California was estimated
at 100,000 (Fish and WHdHfe Service 1973).
Anderson et al. (1972b) estimated 21,000 resi-
dents on the California coast and somewhere near
62,000 breeding in the Gulf of California.
On Santa Barbara Island, California, between
300 and 400 birds were breeding July 1912. On
Anacapa Islands, California, there were a large
number of birds which apparently were not
breeding on 5 July 1912 (Wright and Snyder
1913). There had been at least 500 nests there in
June 1910 (Willett 1912, 1933). WUlett (1910)
counted 5 nests containing young on Prince Islet
off San Miguel Island on 15 June 1910.
Wright (1909) and Stephens (1921) found
large colonies with hundreds of nests on Los
Coronados, Baja California, in early 1900's. In the
early 1940's there were still several thousand pairs
nesting there, according to Lewis Walker (in
Schreiber and DeLong 1969). In 1958, pehcans
were present in large numbers on Los Coronados
and nesting was observed by Monte Kirven (m
Schreiber and DeLong 1969). During the 1968
breeding season, on four occasions between April
and June, only a few birds were present and none
were nesting there (Schreiber and DeLong 1969).
There was no nesting in 1963 (Keith et al. 1970).
For Santa Catalina, Santa Cruz, San Miguel,
and San Nicolos, records are scanty but indicate
that in the early 1900's irregular nesting (not
every year) occurred on all of them (WUlett 1912;
Howell 1917). No active nests were found on
those islands at any time in 1968 (Schreiber and
DeLong 1969).
Historically, Santa Barbara Island has been
second only to Anacapa as the most important
pelican rookery on the Channel Islands. In 1912
several hundred birds bred there (Howell 1917).
On 19 Aug. 1967, there were 400 to 500 on Sur-
tU Rock just off Santa Barbara, where they nested
that year. In 1968, 4 adults and 33 immatures
were present on 4 April and 11 on 12 May but
there was no nesting; nor was late nesting reported
by the National Park Service. On Anacapa, breed-
ing was first reported on all three islets in 1899
(Schreiber and DeLong 1969). In 1939, Bond
(1942) reported about 2,000 pairs nesting on the
West islet, Anacapa, which would indicate at least
as many, if not more, birds present on that islet
than found by previous observers. The largest
number recorded for East Islet, in the Anacapa
group is about 500 nests, found 5 June 1910 by
Willett (1912). Pelicans nested on West Anacapa
at least up until 1968, but not on East or Middle
Anacapa (Bond 1942; Banks 1966; and Schreiber
and DeLong 1969). On 20 March 1969, R. W.
10
Risebrough (Report to Patuxent Wildlife Research
Center, 24 April 1969) counted 298 nests con-
taining fresh plant material on Anacapa Island,
but only 12 contained intact eggs.
The Monterey Bay and Peninsula area is very
important for this species, as large numbers occur
there following dieir northward postbreeding
dispersal (Alan Baldridge in lit. 1968). Birds ar-
rive there in mid-June and remain into December,
with a peak of 2,243 on 23 Aug. 1968, over
1,000 present from August to October, and 2,300
counted on 15 Sep. 1968. Williams (1927) re-
corded the first known breeding site. He found
20 to 30 nests on 25 May 1927, 8 with eggs on
16 June 1977, and 55 with eggs on 29 May 1929
(WUliams 1927, 1931). In 1966, H. L. Cogswell
{in Alan Baldridge in litt (1968) recorded at least
three nests with birds at Point Lobos, which seems
to have been the last recorded breeding there.
Between 1 and 8 May 1969, there were nest-
ing colonies in the Gulf of California at Puerto
Refugio, Isla Lorenzo Norte, Isla Salsipuedes and
Isla Poijo; numerous adults, nests, and intact eggs
were found at all these colonies. Eggshells appeared
sound, but there was evidence of abnormal repro-
duction including having many empty nests, col-
lapsed and dehydrated eggs, and many nests hav-
ing only 1 or 2 eggs instead of the usual 3 (Keith
etal. 1970).
In 1974, about 95% of Pacific coast breeding
sites were visited and found to have present about
25,000 breeding pairs and an estimated total
population of about 70,000 (U.S. Fish and Wild-
life Service, Denver Wildlife Research Center ms.
1974).
Pelican populations off the southern Cali-
fornia coast appear to have oscillated historically
in response to environmental changes, including
changes in abundance of principal food fishes.
Also the oceanic environment in diat area is a
constant state of flux. However, the overall trend
toward population decline since the mid-1950's,
with the normal oscillating pattern superimposed,
has almost certainly been due to environmental
pollution (Anderson and Anderson 1976).
REPRODUCTION
Eastern subspecies
The breeding season depends on location.
Bent (1922) states that nesting begins on the
Florida east coast in November and December,
west coast in April, Louisiana coast in February,
and South Carolina coast in May. However, L. J.
Blus (personal communication) obsei^ved nesting
in South Carolina in March and April. Anderson
and Hickey (1970) list breeding dates for the
brown pelican throughout its range.
Williams and Joanen (1974) report that peli-
cans first nest successfully when 3 years old. All
courtship activities are confined to the nest site.
The male brings nesting materials to the female
and she builds the nest. Normal clutch size is two
or three eggs (Bent 1922, Schreiber 1979). Eggs
are snowy white with blood stains when first laid,
but turn dull and dirty during incubation (L. J.
Blus personal communication). Eggs have a granu-
lar surface texture and average about 73 x 46.5
mm (Bent 1922). Both parents share in incuba-
tion and chick raising duties (Schreiber 1979).
Henny (1972) estimated that 1.2 to 1.5 fledg-
lings per nesting effort are necessary to maintain a
stable population. Schreiber and Risebrough
(1972) state that this species is long-lived. One
brown pelican banded in September 1933 was re-
captured in November 1964 (Kennard 1975).
California subspecies
Like its eastern relative, P. o. californicus is a
colonial nester. It first breeds at about 2 years of
age. The timing of nesting varies considerably
from year to year and between adjoining colonies
during the same year. Clutch size is usually 3 eggs,
less often 2. Incubation is by both sexes in turn,
and requires about 4 weeks. Young are hatched
over a span of days, and are fed by both parents
by regurgitation. Age at first flight is 9 weeks
(Palmer 1962).
On 12 April 1888, most of the nests on San
Martin Island, Baja California, contained young
ranging in age from just hatched to full-fledged
birds capable of flying (A. W. Anthony m Bent
1922). Flightless young move about in flocks or
"pods." They are pugnacious towards intruders,
but vulnerable to attack by gulls (Wright 1919,
A. B. Howell in Bent 1922). Egg dates for Los
Coronados Islands are 29 March to 22 June (33
records) and 4 April to 6 May (17 records) (Bent
1922). On Anacapa Islaiid, Cahfornia, nest con-
tents on 16 April 1939 ranged from fresh eggs to
young about half grown with primaries just be-
ginning to show (Bond 1942). A summary of re-
production on the Anacapa Islands from several
obsei^vers shows great variation in timing, with
adults carrying nesting material on 17 March
1911, while egg laying had begun 7 March 1916
on East Island and egg laying completed with one
nest hatched on West Island; by 5 June 1910 on
East Island, fresh eggs to nearly full grown were
found; in August 1898 on West Island nearly
11
grown young were still present (Bond 1942).
Adult pelicans usually stayed on nests until
approached within 5 or 6 m and returned by the
time that observers were about 30 m away, but
even so, ravens carried off at least 8 eggs when
they were left exposed (Bond 1942).
On 20 March 1969, R. W. Risebrough (Report
to Patuxent Wildlife Research Center, 24 April
1969) counted 298 nests containing fresh plant
material, indicating occupancy, on Anacapa Is-
lands, but only 12 contained intact eggs. Nine
contained a single egg and 3 contained 2 eggs.
Fifty-one nests contained a single broken egg.
Eggshells were spongy in texture and slight pres-
sure produced a change in shape. Shells appeared
to have very little or no calcium carbonate, and it
was apparent that the weight of incubating birds
would cause breakage. One of the broken eggs
had a DDE content of 68 ppm of the total con-
tents or 522 ppm of yolk lipid. In a second colony
on Anacapa on April 1969, there were 339 nests,
with only 19 containing intact eggs and 1 out of
3 nests having collapsed, dehydrated eggs (Monthly
Report, Denver Wildlife Research Center April
1969). On 26 and 27 July 1969, 635 nests were
found on Anacapa. Two nests contained large
chicks and 2 contained single incubated eggs.
Thus, out of a minimum of 1272 nests built in
1969, only 5 may have produced young (F. H.
Sibley, Report to Patuxent Wildhfe Research Cen-
ter, 28 July 1969).
There were 552 nesting attempts in two
colonies on Anacapa in 1970, with but one chick
produced. Reproductive failure was attributed to
thin eggshells that collapsed during incubation.
Aberrant behavior associated with reproduction
was also observed. A survey of all traditional nest-
ing sites in California produced no evidence of
nesting. To the best of our knowledge, then, only
one young pelican was hatched in California in
1970 (Cress 1970).
A total of 1,962 eggshells taken after 1949
for California oological collections were 26% be-
low the normal weight of egg specimens taken
before 1949, supporting the belief that shell thin-
ning began after the wide-scale use of pesticides
(Anderson and Hickey 1970).
By use of stepwise regression, Blus et al.
(1971) found that DDE was the only residue that
accounted for a significant amount of pelican egg-
shell thinning; accounted for a significant amount
of pelican eggshell thinning; PCB's appeared to be
of Uttle, if any, importance in this respect.
On Border Island, Los Coronados, Baja Cali-
fornia, only 19 intact eggs were found in 300
nests in April 1969. On Martin Island, 150 miles
south of the U.S. border, only 17 eggs were found
in 100 nests (Monthly Report of Denver Wildlife
Research Center, April 1969).
Brown pelicans showed varying reproductive
success in different parts of their range and from
1969 to 1971, with the poorest off the California
coast, improving slightly down the west coast of
Baja California, and the best in the Gulf of Cali-
fornia. About 4.4% of the population is suffering
widespread excessively low recruitment. Of the
relatively satisfactory Gulf of California popula-
tion, it is estimated that 20% suffered eggshell
thinning in excess of 1 l%o. Anacapa Islands, Los
Coronados, and west coast Baja Cahfornia pop-
ulations had the most eggshell thinning in 1968,
1970 and 1971. Here again, DDE seems to be the
major factor contributing to thin eggshells and re-
sults in some reduction in reproduction. Other
factors reducing reproduction related to human
interference of several types. Various data suggest
that the major source of DDE for brown pehcans
is off Southern California (Anderson et al. 1975).
An indication of improved production on the
California Islands was noted in 1972, when 112
nests were counted on Scorpion Rock off Santa
Cruz Island, and 31 young had been produced
there by 13 July. In the same year, 150 nests
from which 26 young had been produced were
found on Anacapa. Early nest failures were evi-
dent from the many cracked eggshells (from
Monthly Narrative Report of Activities of Denver
Wildlife Research Center, August 1972 by David
W. Anderson and L. Rodney DeWeese).
In 1974, eggshell thinning was much less evi-
dent on the Pacific coast; many more young were
produced and more adults bred in some colonies.
About 1,400 adult pelicans were present in July
on Anacapa and Santa Cruz Islands, where they
build about 400 nests and produced about 300
young. However, collapsed eggs were found and
30% of nests on the islands were empty and de-
serted. Also, 15% of the adults were incubating
addled eggs. About 0.75 young per nest were pro-
duced, which is still too little; about 1.3 to 1.5
young per adult pair are required to maintain the
population. On the Coronados Islands, Baja Cali-
fornia, the colony was much larger in 1974 than in
any recent year but productivity averaged only
0.70 young per nest and collapsed eggs and nest
desertions were common (up to 30% in some
areas). On Isla San Martin, pelicans built 112
nests, all of which failed, probably due to human
disturbance. At Isla San Benito, production was
12
excellent in 1974 with at least 1.5 young per nest,
no nest desertion and no excessive losses of young.
In the Gulf of Cahfomia only about 60% of
adults came to colonies to breed in 1974. Adults
that did come failed to establish nests and left.
In some areas, nest desertion approached 100%.
Ultimate production was about 0.80 young per
nest, and less than one half of adults established
nests. Unusually poor food conditions were
thought responsible. In recent years, severe and
widespread failures in Gulf of California produc-
tion has occurred during periods of food scarcity.
However, it is possible that the DDE known to be
present has aggravated the situation. Experiments
have shown that minor restrictions of food supply
seriously impair reproduction and that presence
of DDE in birds increases the effect of food de-
privation on reproductive condition(U.S. Fish and
Wildlife Service, Denver Wildlife Research Center
1974;Keith 1978).
An asynchronous nesting pattern on Anacapa
and Santa Cruz Islands in 1975 made estimates of
production difficult. Production was: Anacapa -
212 nests, 182 young produced; Santa Cruz - 80
nests, 74 young produced, or 0.88 young per nest
for both islands. This compares with 0.73 in 1974,
0.14 in 1973, 0.22 in 1972 and 0.007 in period
1969-71 for the same colonies. Neither DDE con-
centrations nor eggshell thickness differed signif-
icantly between 1974 and 1975, although the
concentration of the chief food fish (anchovies)
dropped. Pelican productivity is still 10% to 50%
below the level necessary to maintain long-term
population stability (Anderson et al. 1975; An-
derson and Anderson 1976; Anderson et al. 1977).
MANAGEMENT AND CONSERVATION
State, Federal, and private cooperative research
has been directed toward analysis of brown peli-
can eggshell conditions and the resulting repro-
ductive failure. Waste discharge from pesticide
manufacturing plants is being corrected. Popula-
tion surveys have been conducted by the Fish and
Wildlife Service. Continued research on the effects
of environmental pollution and on life history and
reproductive biology are recommended, but visits
to nests should be kept to a minimum; strict reg-
ulation of the use of persistent chemical pesticides
and discharge of wastes from plants that manu-
facture those products should be encouraged and
all forms of pollution that degrade the pelican's
environment should be eliminated. Sanctuary
status should be recorded for all areas with
nesting colonies (U.S. Fish and Wildlife Service
1973).
Many eastern colonies are protected on Fed-
eral and State refuges and in National Audubon
Society sanctuaries.
A recovery team for the eastern brown peli-
can has been appointed by the U.S. Fish and
Wildlife Service to draft a recovery plan for re-
establishing this species throughout its former
range (Williams et al. 1976). The recovery outline
includes restoring the pelican in vacant breeding
habitat by identifying historic distribution, iden-
tifying or creating suitable breeding habitat and
stocking selected sites.
In order to restock historic or depleted
colonies, stocking methods must be developed
and sources of birds located. The success of new
colonies should be monitored for survival, repro-
duction, and limiting factors. Natural and restored
colonies should be maintained at self-sustaining
levels. Plans for monitoring populations and
pertinent environmental factors vary from area
to area and are outlined in Williams et al. (1976).
AUTHORITIES
Eastern subspecies
Lawrence J. Blus (Recovery Team)
U.S. Fish and Wildhfe Service
Patuxent Wildlife Research Center
Laurel, MD 20810
Kirke A. King (Recovery Team)
U.S. Fish and Wildlife Service
P.O. Box 2506
Victoria, TX 77901
Larry L. McNease (Recovery Team)
Louisiana Wildlife and Fisheries Commission
Route l,Box 25
Grand Chenier, LA 70643
Burkett S. Neely (Recovery Team)
U.S. Fish and Wildlife Service
Division of Wildlife Refuges
Washington, D.C. 20240
Stephen A. Nesbitt (Recovery Team)
Florida Game and Freshwater Fish
Commission
4005 South Main Street
Gainesville, FL 32601
Ralph W. Schreiber (Recovery Team)
Natural History Museum
Los Angeles, CA 90007
13
Lovett E. Williams (Recovery Team Leader)
Florida Game and Freshwater Fish
Commission
4005 South Main Street
GainesvUle, FL 32601
Cjilifomia subspecies
Daniel W. Anderson
U.S. Fish £ind Wildlife Service
P.O. BoxC
Davis, California 91616
Alan Baldridge
Hopkins Marine Station
Pacific Grove, California 93950
Monte N. Kirven
Natural History Museum
P.O. Box 1390
San Diego, Cahfornia 12112
Robert DeLong
Department of Biology
University of California, Santa Cruz
Santa Cruz, California 95064
Fred C. Sibley
Peabody Museum
Yale University
New Haven, Connecticut 06520
Ralph W. Schreiber
Los Angeles County Museum
Exposition Park
Los Angeles, California 90007
Robert W. Risebrough
University of California
Berkeley, California 94720
Franklin Gress
University of California
Berkeley, California 94720
PREPARER'S COMMENTS
Eastern subspecies
None.
California subspecies
The California brown pelican is probably the
most obvious and clearcut example of the effect
of pesticide chemicals in the environment on the
reproduction of birds. It is also probably the most
thoroughly studied and documented example of
those effects. The brown pelican, like other spe-
cies that are members of oceanic current ecosys-
tems, normally fluctuate widely in abundance
with the primary prey species (Anderson and
Anderson 1976; Anderson et al. 1975). This is
natural and the population dynamics of affected
species is adapted to compensate for the periodic
failures of food. However, the additional lowered
reproduction caused by pesticide chemicals in the
food, augmented by reduction in food from over-
fishing (Keith 1978), could easily tip the balance
of survival toward extinction. Although the
amount of DDE, the most important of chemicals
affecting California brown pelican reproduction,
has been reduced somewhat and reproduction has
improved, the number of fledged young per nest-
ing pair is still too low to maintain the pelican
population. If chemical level is not further
improved, and if overfishing of anchovies does go
too far, it would appear that this subspecies of
brown pelican is doomed to extinction.— John W.
Aldrich.
LITERATURE CITED/SELECTED
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Condor 33:66-69.
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McNease, B. S. Neely, S. A. Nesbitt, and R.W.
Schreiber. 1976. Recovery plan for the eastern
brown pelican. Preliminary Draft.
Williams, L. E., Jr., and T. Joanen. 1974. Age of
first nesting of the brown pelican. Wilson
Bull. 86:279-280.
Williams, L. E., Jr., and L. Martin. 1968. Nesting
status of the brown pelican in Florida in 1968.
Quart. J. Florida Acad. Sci. 31:130-140.
. 1970. Nesting populations of brown peli-
cans in Florida. Proc. Annu. Conf. S.E. Assoc.
Game and Fish Comm. 24:154-169.
Winn, B. 1975. Pesticides decimate transplanted
pelicans. Audubon 75:127-129.
Woodward, C. H. 1921. The California Brown
PeHcan as a navigator. Condor 23:137-138.
Wright, H. 1909. An orithological trip to Los
Coronados Islands, Mexico. Condor 11:96-
100.
Wright, H. and G. K. Snyder. 1913. Birds observed
in the summer of 1912 among the Santa Bar-
bara Islands. Condor 15:86-92.
16
Biological Services Program
FWS/OBS-80/01.41
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE JAGUAR
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the I'.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Ser\ ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Eouisiana 704.58
FWS/OBS-80/01.41
March 1980
SELECl KD VERl KBRA 1 1: KNDANGERED SPECIES
OF THE SEACOAS I OF I HE UNITED STATES-
THE JAGUAR
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National (-oastal Ecosystems Team,
Office of Biological Services
Project Officer
Doi^ald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Ser\ices
Fish and W'ihllife Service
U.S. Department of the Interior
JAGUAR
Panthera onca Linnaeus
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivora
FAMILY Felidae
OTHER COMMON
NAMES spotted king cat,
American tiger, "el tigre"
DATE
Entered into SWIS to be determined
Updates May 1978.
LEGAL STATUS
Federal: Endangered (non-U. S. wild popula-
tions) (35 FR 18320, 2 December
1970); (44 FR 43705, 25 July 1979).
Threatened (U.S. captive population)
(42 FR 28956, 1 June 1977).
States: Endangered Arizona, New Mexico (ex-
tirpated), Texas.
REASONS FOR CURRENT STATUS
Deliberate persecution, excessive and illegal
hunting, overexploitation by fur industry, and
predator control activities have extirpated jaguars
from much of their original range and seriously
reduced numbers in most of the rest (lUCN 1972,
Culbertson and Schmidly 1974, Davis 1974).
Timber and brush clearing have degraded and des-
troyed habitat to the point where reestablishment
of populations in the northern part of the range is
doubtful (Davis 1974, Brovmlee 1978). Mining
and oil exploration and development have made
formerly remote Central and South American
areas more accessible to human activity and sub-
sequent illegal killings of jaguars (lUCN 1972).
PRIORITY INDEX
None designated.
DESCRIPTION
The jaguar is the largest (1.5 to 2.2 m) and
most robust of the American cats. The tail is
short and somewhat bristly. Females average
smaller than males. The ground color varies from
pale yellow to rusty red dorsally, paler on the
sides, and white on the underparts and inner
surfaces of the legs. Markings are irregular
blotches and rosettes, the latter centered with
black spots. Young are more heavily spotted and
their coats are woolier. Both black and albino in-
dividuals occur occasionally.
Photographs appear in Davis (1974), Guggis-
berg (1975), and Walker (1975).
RANGE
The jaguar was formerly distributed through-
out the tropical lowlands of Mexico, Central
America, and South America to about 40° S lati-
tude (Calahane 1947, Hall and Kelson 1959, Gug-
gisberg 1975). The former U.S. distribution in-
cluded southern California (Merriam 1919, Strong
1926, Seton 1937), New Mexico (Bailey 1931,
Seton 1937, Hill 1942, Halloran 1946, Findley et
al. 1975), southern Arizona (Musgrave 1921,
Schufeldt 1921, Poole and Schantz 1942, Hock
1955, Cockrum 1960), and possibly Louisiana
(Nowak 1973, Lowery 1974), and Colorado
(Seton 1920).
Today, jaguars are essentially absent from
most of Mexico, Argentina, and settled provinces
in the remaining Central and South American
countries (lUCN 1972). There is no evidence for
its present occurrence in Louisiana (Lowery 1974)
or New Mexico (M. C. Conway personal commu-
nication); however, there may possibly be indivi-
duals in the border areas of the latter (Findley et
al. 1975). It is essentially absent from other areas
north of the Mexican border except as occasional
stray individuals in the border counties of Texas
and Arizona (Cockrum 1960, Davis 1974, Findley
et al. 1975, Guggisberg 1975, Lowman 1975).
RANGE MAP
Crosshatched areas on the following map in-
dicate possible range in border areas of U.S.
(Cockrum 1960, Davis 1974, Findley et al. 1975,
Guggisberg 1975, Brownlee 1978). Dots refer to
sightings and/or kills during the past century.
STATES/COUNTIES
Arizona: Cochise, Pima, Santa Cruz.
New
Mexico: Dona Ana, Grant, Hidalgo, Luna.
Texas: Brewster, Cameron, Dimmit, Hidalgo,
Kinney, Maverick, Starr, Terrell, Val
Verde, Webb, Zapata.
HABITAT
The jaguar appears to require areas with cover
(Ewer 1973, Lowman 1975). It inhabits tropical
and subtropical forests ranging from mangrove
swamps to rain forests (Alston 1882, lUCN
1972). At the southern extreme of the range,
open savannas and deserts are used (lUCN 1972).
Chaparral and timbered areas are preferred at the
northern extreme of the range (Davis 1974). It
appears to have a preference for areas near water
(Davis 1974, Guggisberg 1975), but has been
reported from deserts (Guggisberg 1975).
FOOD AND FORAGING BEHAVIOR
Principal foods are peccaries {Tayassu sp.) and
capybaras {Hydro choerus sp.) (Guggisberg 1975).
It will also take tapirs (Tapirus sp.), agoutis
{Agouti sp., Dasyprocta sp.), otters {Lutra), deer,
small crocodilians, turtles and their eggs, large
ground-nesting birds, and occasionally livestock
(Denis 1964, Ewer 1973, Davis 1974, Guggisberg
1975, Lowman 1975).
It stalks prey until close enough to pounce
(Guggisberg 1975), then drags the kill to the
nearest thicket to be eaten. Remains are not usu-
ally covered (Hoffmeister 1971).
SHELTER REQUIREMENTS
The jaguar requires dens in rocky caves or
dense thickets (Davis 1974).
NESTING OR BEDDING
The jaguar raises its young in dens (see Shel-
ter) (Davis 1974, Guggisberg 1975).
RITUAL REQUIREMENTS
The jaguar is solitary and somewhat territorial
in its habits, except during the breeding season.
Little is known about its territorial behavior be-
yond the fact that it will mark trees (Guggisberg
1975).
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POPULATION NUMBERS AND TRENDS
Once common throughout its range, the jaguar
is now uncommon to rare. Fair numbers remain
in eastern Campeche, Selva Lacandone, eastern
Chiapas, and eastern Oaxaca in Mexico and west-
em El Peten in Guatemala (lUCN 1972). It is
scattered and considered a pest in Argentina
(lUCN 1972). North of Mexico, it occurs ex-
tremely rarely, in Texas, Arizona, and New
Mexico (Hock 1955, Cockrum 1960, Culbertson
and Schmidly 1974, Davis 1974). It is believed
unlikely that the jaguar could reestablish in
Texas, due to habitat loss (Brownlee 1978). No
current population estimates are available. There
are no viable breeding populations in the U.S. (W.
C. Brownlee personal communication); the jaguar
in the U.S. exists only as an occasional stray.
REPRODUCTION
The breeding season is year-round in tropical
areas of the range (Cahalane 1946, Ewer 1975,
Guggisberg 1975). In the more northern areas,
breeding takes place in December and January
wdth births in April and May (Davis 1974). Fe-
males are polyestrous (Ewer 1968) and males
fight for first breeding rights (Guggisberg 1975).
Gestation lasts 93 to 113 days (Asdell 1964,
Denis 1964, Davis 1974). Litter size ranges from
one to four with an average of two (Cahalane
1947, Denis 1964, Ewer 1973). The growth rate
is not well known, but 8-week-old cubs will weigh
from 2 to 3 kg (Hunt 1967). Young stay with the
mother for about 2 years, at which time sexual
maturity is reached (Denis 1964, Guggisberg
1975), Both parents help rear young, but the bulk
of the burden is on the female (Guggisberg 1975).
The family unit is maintained until the young are
at least a year old (Davis 1974). Record longevity
for a captive is 20 years (Guggisberg 1975).
MANAGEMENT AND CONSERVATION
No recovery team has been appointed for the
jaguar. The species is protected in the U.S. and
most of the Central and South American countries
within its range, but laws are not adequately
enforced (lUCN 1972). Colombia still allows
hunting of all felid species. Protection in some
Central and South American countries is provided
in National Parks (lUCN 1972). Importation of
skins to the U.S. is prohibited except by permit.
However, illicit commerce to European and Asian
markets is common (lUCN 1972).
The lUCN is surveying the jaguar's status in
Latin America; in a cooperative venture, the
U.S. Fish and Wildlife Service and the Direccion
General de la Fauna Silvestre of Mexico with sup-
port from the National Wildlife Federation, the
National Audubon Society, and the Texas and
New Mexico Departments of Game and Fish, are
surveying the population status in Mexico (USFWS
1978). A feline status survey by the Texas Parks
and Wildlife Department has found no evidence
of recent occurrence of jaguars in that State
(Brownlee 1978).
Brush clearing in national wildlife refuges in
the Lower Rio Grande Valley of Texas has been
stopped to preserve rapidly disappearing brush
habitat.
AUTHORITIES
W. C. Brovsmlee
Texas Parks and Wildlife Department
4200 Smith School Road
Austin, TX 78744
E. Leddell Cockrum
Department of Zoology
University of Arizona
Tucson, AZ 85719
W. B.Davis
Department of Wildlife and Fisheries Science
Texas A&M University
College Station, TX 77843
Roy McBride
Department of Biology
Sul Ross University
Box 725
Alpine, TX 79830
PREPARER'S COMMENTS
Literature is scarce on the habits and ecology
of wild jaguar populations. More research is
needed in this area. Status surveys are needed to
assess population levels, in addition to protection
of habitat.
LITERATURE CITED/SELECTED
REFERENCES
Alston, E. R. 1882. Biologia Centrali-Americana.
Mammalia. 219 pp.
Anthony, H. E. 1928. Field book of North Ame-
rican mammals. Putnam and Sons, New
York. 674 pp.
Asdell, S. A. 1964. Patterns of mammalian repro-
duction, 2nd ed. Cornell Univ. Press, Ithaca.
670 pp.
Bailey, V. 1931. The mammals of New Mexico.
N.Am. Fauna 53:283-285.
. 1971. Mammals of the southwestern
United States with special reference to New
Mexico. Dover, New York. 412 pp.
Brownlee, W. C. 1978. Feline status surveys. Job
performance report. Project W-103-R-7. Tex.
Parks Wildl. Dep., Austin. 4 pp.
Cahalane, V. A. 1947. Mammals of North Ameri-
ca. MacMillan Co., New York. 682 pp.
Cockrum, E. L. 1960. The recent mammals of
Arizona; their taxonomy and distribution.
Univ. Ariz. Press, Tucson. 276 pp.
Culbertson, K., and D. J. Schmidly. 1974. Sum-
mary statements on the state of the rare, en-
dangered, and peripheral mammals in Texas.
Tex. Organization for Endangered Species,
Temple. 4 pp.
Davis, W. B. 1974. The mammals of Texas. Tex.
Parks Wildl. Dep. Bull. 41: 1-294.
Denis, A. 1964. Cats of the world. Houghton Mif-
flin Co., Boston. 144 pp.
Ewer, E. F. 1968. Ethology of mammals. Logos
Press Ltd., London. 418 pp.
• 1973. The carnivores. Cornell Univ. Press.
Ithaca. 494 pp.
Findley, J. S., A. H. Harris, D. E. Wilson, and C.
Jones. 1975. Mammals of New Mexico. Univ.
New Mex. Press, Albuquerque. 360 pp.
Guggisberg, C. A. W. 1975. Wildcats of the world.
Taplinger PubL Co., New York. 382 pp.
Hall, E. R., and K.R. Kelson. 1959. The mammals
of North America. 2 vols. Ronald Press, New
York. 1,083 pp.
Halloran, A. F. 1946. Mammals of the Chiricahua
Mountains, Cochise County, Arizona. J.
Mammal. 27:154-161.
Hill,J. E. 1942. Notes on mammals of northeastern
New Mexico. J. MammaL 23:75-82.
Hock, R. J. 1855. Southwestern exotic felids.
Am. Midi. Natur. 53:324-328.
Hoffmeister, D. F. 1971. Mammals of Grand Can-
yon. Univ. Illinois Press, Chicago. 183 pp.
Hunt, H. 1967. Growth rate of a new-born, hand-
reared jaguar [Panthera onca) at Topeka Zoo.
Int. Zoo. Yearb. 7:147-148.
lUCN. 1972. Red data book. Vol. I. Mammalia.
Compiled by H. A. Goodwin and C. W. Hollo-
way. lUCN, Morges, Switzerland.
Layne, J. N. 1974. The land mammals of Florida.
In Gleason, P. J., ed. Environments of south
Florida, past and present. Mem. 2. Miami
Geol. Soc. 452 pp.
Leopold, A. S. 1959. Wildlife of Mexico: the
game birds and mammals. Univ. California
Press, Berkeley. 568 pp.
Lowery, G. E. 1974. The mammals of Louisiana
and its adjacent waters. Louisiana State Univ.
Press, Baton Rouge. 565 pp.
Lowman, G. E. 1975. A survey of endangered,
threatened, rare, status undetermined, peri-
pheral, and unique mammals of the south-
eastern National Forests and Grasslands.
USDA, For. Serv., Atlanta, Ga. 121 pp.
McCarly, H. 1959. The mammals of eastern Texas.
Tex.J.Sci. 2:385-426.
Merriam, C. H. 1919. Is the jaguar entitled to a
place in the California fauna? J. Mammal. 1:
38-40.
Musgrave, M. E. 1921. Predatory animals in Ari-
zona. Am. Game Project. Assoc. Bull. 10:
11-12.
Nelson, E. W., and E. A. Goldman. 1929. Revi-
sion of the jaguar. J. Mammal. 14:221-240.
Nowak, R. M. 1973. A possible occurrence of the
jaguar in Louisiana. Southwest Natur. 17:
430-432.
Poole, A. J., and V. S. Schantz. 1942. Catalog of
the type specimens of mammals in the United
States National Museum, including Biological
Survey collection. Bull. U.S. Natl. Mus. 178:
58.
Schufeldt, R. W. 1921. The mountain lion, ocelot,
lynx, and their kin. Am. Forest. 27:629-636.
Seton, E. T. 1920. The jaguar in Colorado. J.
Mammal. 1:241.
. 1937. Lives of game animals. Literary
Guild, New York.
Stehlik,J. 1971. Breeding jaguars; PanM^ra onca;
at Ostrava Zoo. Int. Zoo. Yearb. 11:116-118.
Strong, W. D. 1926. Indian records of California
carnivores. J. Mammal. 7:59-60.
Taylor, W. P. 1947. Recent record of the jaguar in
Texas. J. Mammal. 28:66.
USFWS. 1973. Threatened wildlife of the United
States. Compiled by Office of Endangered
Species and International Activities. Bur.
Sport Fish. Wildl. Resour. Publ. 114. U.S.
Gov. Printing Office, Washington, D.C.
289 pp.
USFWS. 1978. International activities. U.S. -Mexi-
can Cooperation. Endangered Species Tech.
Bull. lll(3):8-9.
Walker, E. P. 1975. Mammals of the world. 2 vols.
Johns Hopkins Press, Baltimore. 1,500 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.42
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE GRAY BAT
p'.
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and c()nscr\'ation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amciulcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should he directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Eouisiana 70458
FWS/OBS-80/01.42
March 1980
SELECi i:d vkri ebrati: endangered species
OF IHE SEACOAS T OE IHE UNITED STATES-
THE GRAY BAT
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National (Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. VVoodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Ser\iccs
Fish and Wildlife Service
U.S. Department of the Interior
GRAY BAT
My Otis grisescens (Howell)
KINGDOM Animalia
CLASS Mammalia
ORDER Chiroptera
FAMILY Vespertilionidae
OTHER COMMON
NAMES Gray myotis, cave bat,
Howell's bat, Tennessee brown bat
DATE
Entered into SWIS to be determined
Updates 8 August 1978
LEGAL STATUS
Federal: Endangered (41 FR 17740, 28 April
1976).
States: Endangered: Florida, Georgia, Indiana,
Illinois, Missouri, Tennessee, North
Carolina.
REASONS FOR CURRENT STATUS
The primary cause for the species current re-
ductions from former population levels is human
disturbance of caves in which the bats roost, rear
young, or hibernate (Manvilie 1962; Barbour and
Davis 1969; Tuttlel977, 1979). Gray bats are in-
tolerant of human disturbance and Tuttle (1979)
has shown a direct correlation between frequency
of human disturbances and population reductions.
Fully 95% of the entire species winters in just
nine caves, and over 60% winters in a single cave
in northern Alabama, making the species ex-
tremely vulnerable to significant population
reductions. Gray bats, with few exceptions, also
spend the summers in caves (not the same ones
used for hibernation) and are equally vulnerable
in these (Tuttle 1979). Deliberate vandalism as
well as frequent human invasion of caves where
gray bats live has eliminated many colonies and
drastically reduced most others (Tuttle 1979).
Environmental disturbances such as deforesta-
tion and chemical and pesticide contamination
may also adversely affect gray bat populations. A
very low reproductive rate makes recovery of de-
pleted colonies questionable.
PRIORITY INDEX
Not assigned.
DESCRIPTION
Adults are medium sized. Forearms are 40 to
47 mm long. Summer weights are 8 to 10 g, but
go as high as 16 g just prior to migration. Fur is
uniformly gray immediately following molting in
late June or July, and bleaches to bright russet by
the following May or June, especially in repro-
ductive females. Russett colored animals are most
conspicuous in crowded or southern roosts. The
most useful field mark is the bat's unicolored dor-
sad fur; all other southeastern bats have conspicu-
ously bi- or tricolored dorsal fur (Barbour and
Davis 1969). Illustrations appear in Barbour and
Davis (1969) and Odum et al. (1977).
RANGE
The gray bat's range extends from eastern Ok-
lahoma and Kansas eastward to southwestern Vir-
ginia and western North Carolina; southern Illi-
nois and Indiana southward to northern Florida
(Hall and Kelson 1959, Hall and Wilson 1966,
Tuttle 1966, Barbour and Davis 1969, Humphrey
and Tuttle 1979). Bats wanter in caves, primarily
in North Carolina, Alabama, Missouri, and Ten-
nessee.
RANGE MAP
Cross-hatching on the following map shows
the species' summer range. Dots indicate major
winter caves (Hall and Kelson 1959, Barbour and
Davis 1969, Tuttle and Robertson 1969, Tuttle
1976b, Elder and Gunier 1978).
STATES/COUNTIES
Alabama: Blount, Bullock, Calhoun, Chambers,
Cherokee, Clay, Cleburne, Coffee,
Colbert, Coosa, Cullman, Dale, De
Kalb, Elmore, Etowah, Geneva,
Henry, Houston, Jackson, Jefferson,
Lauderdale, Lawrence, Limestone,
Macon, Madison, Montgomery, Mor-
gan, Pike, Randolph, Russell, St. Clair,
Shelby, Talladega, Tallapoosa.
Arkansas: Baxter, Benton, Boone, Carroll, Clay,
Cleburne, Conway, Craighead, Craw-
ford, Crittenden, Cross, Faulkner,
Franklin, Fulton, Greene, Indepen-
dence, Izard, Jackson, Johnson, Law-
rence, Lee, Logan, Madison, Marion,
Mississippi, Monroe, Newton, Poinsett,
Pope, Prairie, Randolph, St. Francis,
Searcy, Sharp, Stone, Van Buren,
Washington, White, Woodruff.
Florida: Baker, Bay, Calhoun, Columbia, Dixie,
Franklin, Gadsden, Gulf, Hamilton,
Holmes, Jackson, Jefferson, Lafayette,
Leon, Liberty, Madison, Nassau, Su-
wannee, Taylor, Wakulla, Walton,
Washington.
Georgia: Bartow, Carroll, Catoosa, Chattahoo-
chee, Chattooga, Cherokee, Clay,
Clayton, Cobb, Coweta, Dade, Daw-
son, Decatur, De Kalb, Douglas, Early,
Fannin, Fayette, Floyd, Forsyth, Ful-
ton, Gilmer, Gordon, Haralson, Harris,
Heard, Lumpkin, Meriwether, Miller,
Murray, Muscogee, Paulding, Pickens,
Polk, Quitman, Randolph, Seminole,
Stewart, Towns, Troup, Union,
Walker, White, Whitfield.
Illinois: Adams, Alexander, Bond, Brown, Cal-
houn, Cass, Clay, Clinton, Edwards,
Fayette, Franklin, Gallatin, Greene,
Hamilton, Hardin, Jackson, Jefferson,
Jersey, Johnson, Macoupin, Madison,
Marion, Massac, Monroe, Montgome-
ry, Morgan, Perry, Pike, Pope, Pulas-
ki, Randolph, St. Clair, Saline, Sanga-
mon, Scott, Union, Wabash, Washing-
ton, Wayne, White, Williamson.
Indiana: Crawford, Dubois, Floyd, Gibson,
Harrison, Knox, Orange, Perry, Pike,
Posey, Spencer, Vanderburgh, Warrick,
Washington.
Kansas: Barber, Bourbon, Chautauqua, Chero-
kee, Cowley, Crawford, Elk, Harper,
Labette, Montgomery, Neosho, Sum-
ner, Wilson,
Kentucky: Adair, Allen, Anderson, Ballard, Bar-
ren, Rath, Bell, Bourbon, Boyle,
Breathitt, Breckenridge, Bullitt, Butler,
Caldwell, Calloway, Carhsle, Carter,
Casey, Christian, Clark, Clay, Clinton,
Crittenden, Cumberland, Daviess, Ed-
monson, Elliott, Estill, Fayette, Flem-
ing, Franklin, Fulton, Garrard, Grant,
Graves, Grayson, Green, Hancock,
Hardin, Harlan, Harrison, Hart, Hen-
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derson, Henry, Hickman, Hopkins,
Jackson, Jefferson, Jessamine, Knox,
Larue, Laurel, Lee, Leslie, Letcher,
Lincoln, Livingston, Logan, Lyon,
McCracken, McCreary, McLean, Madi-
son, Magoffin, Marion, Marshall,
Meade, Menifee, Mercer, Metcalfe,
Monroe, Montgomery, Morgan, Muh-
lenberg, Nelson, Nicholas, Ohio, Old-
ham, Owen, Owsley, Perry, Powell,
Pulaski, Rockcastle, Rowan, Russell,
Scott, Shelby, Simpson, Spencer, Tay-
lor, Todd, Trigg, Union, Warren,
Washington, Wayne, Webster, Whitley,
Wolfe, Woodford.
Mississippi: Tishomingo.
Missouri: Audrain, Barry, Barton, Benton, Bol-
linger, Boone, Butler, Callaway, Cam-
den, Carter, Cedar, Christian, Cole,
Cooper, Crawford, Dade, Dallas, Dent,
Douglas, Franklin, Gasconade, Greene,
Hickory, Howard, Howell, Iron, Jas-
per, Jefferson, Laclede, Lawrence,
Lincoln, McDonald, Madison, Maries,
Marion, Miller, Moniteau, Monroe,
Montgomery, Morgan, Newton, Ore-
gon, Osage, Ozark, Perry, Pettis,
Phelps, Pike, Polk, Pulaski, Ralls, Ran-
dolph, Reynolds, Ripley, St. Charles,
St. Claire, St. Francois, Ste. Genevieve,
St. Louis, St. Louis City, Saline, Shan-
non, Stone, Taney, Texas, Warren,
Washington, Wayne, Webster, Wright.
North
Carolina: Avery, Buncombe, Cherokee, Clay,
Graham, Haywood, Jackson, Macon,
Madison, McDowell, Mitchell, Swain,
Yancey.
Oklahoma: Adair, Alfalfa, Cherokee, Craig, Creek,
Delaware, Garfield, Grant, Haskel,
Hughes, Kay, Kingfisher, LeFlore,
Lincoln, Logan, Mcintosh, Major,
Mayes, Muskogie, Noble, Nowata, Ok-
fuskee, Oklahoma, Okmulgee, Osage,
Ottawa, Pawnee, Payne, Pittsburg,
Pottawatomie, Seminole, Sequoyah,
Rogers, Wagoner, Woods.
Tennessee: Anderson, Bedford, Benton, Bledsoe,
Blount, Bradley, Campbell, Cannon,
Carroll, Cheatham, Chester, Claiborne,
Clay, Cocke, Coffee, Crockett, Cum-
berland, Davidson, Decatur, DeKalb,
Dickson, Dyer, Fayette, Fentress,
Franklin, Gibson, Giles, Grainger,
Greene, Grundy, Hamblen, Hamilton,
Hancock, Hardeman, Hardin, Hawkins,
Haywood, Henderson, Henry, Hick-
man, Houston, Humphreys, Jackson,
Jefferson, Knox, Lake, Lauderdale,
Lawrence, Lewis, Lincoln, Loudon,
McMinn, McNairy, Macon, Madison,
Marion, Marshall, Maury," Meigs, Mon-
roe, Montgomery, Moore, Morgan,
Obion, Overton, Perry, Pickett, Polk,
Putnam, Rhea, Roane, Robertson, Ru-
therford, Scott, Sequatchie, Sevier,
Shelby, Smith, Stewart, Sumner,
Tipton, Trousdale, Union, Van Buren,
Warren, Wayne, Weakley, White,
Williamson, Wilson.
Virginia: Lee, Scott, Wise.
HABITAT
In summer, this migratory species inhabits
areas in which open water and the banks of
streams, lakes, or reservoirs are within manageable
distances of roosting sites and suitable caves in
which to rear young (LaVal et al. 1976, 1977;
Tutde 1976a). In winter, it inhabits caves having
suitable temperatures for its hibernation.
FOOD AND FORAGING BEHAVIOR
Major food items consist of aquatic and non-
aquatic soft-bodied insects (R. LaVal personal
communication), especially Mayflies (Ephemerop-
tera) (Tuttle 1979).
Gray bats fly directy from cave to feeding site
with few stops. They feed by continuous pursuits,
remaining in the air most of the time. Most forag-
ing is done over lakes and rivers where aquatic in-
sects are abundant (Tuttle 1976a, 1979). LaVal et
al. (1976, 1977) found foraging taking place
along the vegetated edges of bodies of water.
SHELTER REQUIREMENTS
In winter, the species requires deep, cold
caves (preferrably 6° to 9° C) for hibernation.
These caves average 10° below the mean annual
surface temperature, and function as cold-air
traps, having multiple entrances and good air
flow (Tuttle and Stevenson 1979). The caves are
already cold in September when the bats arrive
(M. Tuttle personal communication).
NESTING AND BEDDING
In summer, females need warm caves (14° to
27° C) for rearing young. Important characteristics
of bat caves include small chambers (Dvvyer
1963), high places in domed ceilings (Davis et al.
1962), or domes or small pockets within these
locations (Dwyer 1963, Dwyer and Hamilton-
Smith 1965, Dwyer and Harris 1972), and depth
of etching and porosity of the rock surface (Tut-
tle 1975b). Males, nonreproductive females,
postlactating females, and juveniles, also use
caves in summer, but not the same ones used as
maternity sites (Tuttle 1976b).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
The gray bat is perhaps the U.S. mammal
most narrowly restricted to cave habitats (Hall
and Wilson 1966; Barbour and Davis 1969; Tuttle
1976a, 1979). For all practical purposes, it lives
in caves the year round (Tuttle 1979). It is essen-
tial that the bats choose roosts, generally caves,
having temperatures appropriate to the desired
metabolic processes: warm for digestion and
growth in the summer, and cool for torpor in the
fall and winter (Twente 1955). However, in the
summer, gray bats will use man-made structures
that simulate cave conditions (e.g., storm sewers
in Kansas and Illinois [Hays and Bingham 1964,
Elder and Gunicr 1978], and a bam in Missouri
[Gunicrand Elder 1971]).
A minimal colony size is needed to maintain
an adequate roost environment; otherwise, the
colony may fail (Tuttle 1975b, 1979). Because
travel is such a high-energy activity, the bats need
summer caves close to feeding areas— usually with-
in 1 km and no more than 4 km from the nearest
feeding areas. The distances from maternity sites
to feeding areas are important in influencing post-
flight growth and survival (Tuttle 1976a).
POPULATION TRENDS
The estimated total population of the gray
bat is 1 million (Tuttle 1975a). Five major caves
have not been used in the past 20 years (Tuttle
1975a). Twenty-two summer localities in Tennes-
see and Alabama were censused in 1970 and again
in 1976. A 64% reduction in numbers was recorded
for this period (Tuttle 1979). The decline is attri-
buted mainly to a marked increase in human dis-
turbance over that period (Tuttle 1979). In 1978
R. LaVal (personal communication) censused 27
Missouri caves that Myers (1964) had censused in
1964. Adult females and young had declined ap-
proximately 80%, and 16 caves occupied in 1964
had been abandoned. Estimates for the entire
State are not available, but the trend is downward
(R. LaVal personal communication), and is likely
to continue downward because of the bats' intole-
rance to disturbance, their concentration in a few
caves, and the possible effects of pesticide poison-
ing (R. LaVal personal communication).
REPRODUCTION
Copulation occurs in the fall; females store
sperm over the winter (Barbour and Davis 1969).
Young are bom in late May to early June and fly
by late June or mid-July (Tuttle 1976a). Sexual
maturity is reached at 2 years (Tuttle 1976a), al-
though LaVal et al. (1976) suggest that yearlings
breed. The females bear one young, and generally,
only females and young occupy the nursery cave
(Tuttle 1976b). Longevity is high— to 17 years—
but survival is only about 50% to maturity, so
that it takes a female about 5 years to produce
two surviving offspring (M. Tuttle personal com-
munication).
MANAGEMENT AND CONSERVATION
Until recently, there has been no manage-
ment or conservation effort. Most caves are in pri-
vate ownership (Tuttle 1975a). Several caves have
recently come under protection by Federal and
private owners (Harvey 1975, Tuttle 1979), but
improper gating of entrances, which results in de-
creased air flow, has caused the loss of several im-
portant colonies (Tuttle 1977, Tuttle and Steven-
son 1978). Predation at cave entrances has in-
creased with many types of gate. Bats slow down,
circle, or climb through the gates, increasing their
vulnerability to predators (Tuttle 1977). The U.S.
Army Corps of Engineers Meramec Park Lake and
Union Lake Projects in Missouri, which could
ehminate 50% of the gray bats in that area and
alter 60% of the foraging habitat, have been tem-
porarily restrained (LaVal et al. 1976). A Recovery
Team is being formed (M. Tuttle personal com-
munication).
Tuttle (1979) makes the following recom-
mendations for conservation of this species:
(1) purchase severaJ major caves, which are es-
sential to the survival of the gray bat, and pro-
tect them by proper gating; (2) educate spelunkers
and persons interested in visiting bats in winter
caves; (3) disclose the locations of unprotected
bat caves only after providing information on the
procedures necessary to avoid disturbing the bats.
AUTHORITIES
R. K. LaVal
Missouri Department of Conservation
1110 College Avenue
Columbia, MO 65201
M. D. Tuttle
Milwaukee Public Museum
800 West Wells Street
Milwaukee, WI 53233
Dr. Richard LaVal (Recovery Team Leader)
2816 Mexico Gravel Road
Columbia, Missouri 65201
Dr. Merlin Tuttle (Recovery Team)
Vertebrate Division
Milwaukee Public Museum
Milwaukee, Wisconsin 53233
Dr. Tom Kunz (Recover^' Team)
Department of Biology
Boston University
2 Cummington
Boston, Massachusetts 02215
Dr. Don Wilson (Recovery Team)
National Fish and Wildlife Laboratory
National Museum of Natural History
Washington, D.C. 20560
Mr. John Brady (Recovery Team)
St. Louis District
Corps of Engineers
210 North 12th Street
St. Louis, Missouri 63101
PREPARER'S COMMENTS
None.
LITERATURE CITED/SELECTED
REFERENCES
Barbour, R. W., and W. H. Davis. 1969. Bats of
America. Univ. Press, Kentucky, Lexington.
286 pp.
Davis, R. B., C. F. Herreid, II, and H. L. Short.
1962. Mexican free-tailed bats in Texas. Ecol.
Mongr. 32:311-346.
Dwyer, P. D. 1963. The breeding biology of
Miniopterus schreibersi blepotis (Temminck)
(Chirpotera) in north-eastern New South
Wales. Australian J. Zool. 11:219-240.
Dwyer, P. D., and E. Hamilton-Smith. 1965.
Breeding caves and maternity colonies of the
bent-winged bat in south-eastern Australia.
HeHctite 4:3-21.
Dwyer, P. D., and J. A. Harris. 1972. Behavioral
acclimitization to temperature by pregnant
{Miniopterus) (Chiroptera). Physiol. Zool. 45:
14-21.
Elder, W. H., and W.J. Gunier. 1978. Sex ratios
and seasonal movements of gray bats {Myotis
grisescens) in Southwestern Missouri and ad-
jacent States. Am. Midi. Natur. 99(2):463-
472.
Gunier, W. J., and W. H. Elder. 1971. Experimen-
tal homing of gray bats to a maternity colony
in a Missouri barn. Am. Midi. Natur. 86:502-
506.
Hall, E. R., and K. R. Kelson. 1959. The mam-
mals of North America. Vol. 1. Ronald Press,
New York. 546 pp.
Hall, J. S., and N. Wilson. 1966. Seasonal pop-
ulations and movements of the gray bat in the
Kentucky area. Am. Midi. Natur. 75:317-324.
Harvey, M.J. 1975. Endangered Chiroptera of the
southeastern United States. Proc. Annu. Conf.
Southeast. Assoc. Game and Fish Comm. 29:
429-433.
Hays, H. A., and D. C. Bingham. 1964. A colony
of gray bats in southeastern Kansas. J. Mam-
mal. 45:150.
Humphrey, S. R., and M. D. Tuttle. 1979. Gray
bat. Pages 1-3 in J. N. Layne, ed. Rare and
endangered biota of Florida, Vol. 1 , Mammals.
Univ. Presses of Florida, Gainesville.
LaVal, R. K., R. L. Clawson, W. Caire, L. R. Win-
gate, and M. L. LaVal. 1976, An evaluation of
the status of Myotine bats in the proposed
Meramec and Union Lake project areas, Mis-
souri. School of Forestry, Fish, and Wildl.
Univ. Missouri, Columbia. 136 pp.
LaVal, R. K., R. L. Clawson, M. L. LaVal, and W.
Claire. 1977. Foraging behavior and nocturnal
activity patterns of Missouri bats, with em-
phasis on the endangered species Myotis gris-
escens and Myotis sodalis. J. Mammal. 58:
592-599.
U.S. Fish and Wildlife Service. 1978. Osceola
National Forest phosphate extraction and
processing: impacts on Federally listed
threatened or endangered and other species
of concern. U.S. Dep. Inter., Fish Wildl. Serv.,
Office Biological Serv., Washington, D.C. 414
pp.
Odum, R. R., J. L. McCollum, M. A. Neville, and
D. R. Ettman. 1977. Georgia's protected wild-
life. Geogria Dep. Natur. Rcsour., Game Fish
Div., End. Wildl. Progr. 84 pp.
Tuttle, M. D. 1975a. Proposal for inclusion of the
gray bat (Myotis grisescens) on the U.S. Rare
and Endangered Species List. U.S. Dep. Inter.,
Fish and Wildl. Serv., Office Endangered
Species.
. 1975b. Population ecology of the gray
bat (Myotis grisescens): Factors influencing
early growth and development. Occas. Pap.
Mus. Natur. Hist., Univ. Kansas, 36:1-24.
1976a. Population ecology of the gray
bat {Myotis grisescens): Factors influencing
growth and survival of newly volant young.
Ecology 57:587-595.
. 1976b. Population ecology of the gray
bat {Myotis grisescens): Philopatry, timing
and patterns of movement, weight loss during
migiation, and seasonal adaptive strategies.
Occas. Pap. Mus. Natur. Hist., Univ. Kansas
54:1-38.
1977. Gating as a means of protecting
cave dwelling bats. Pages 77-82 in Natl. Cave
Manage. Symp. Proc, 1976. Speleobooks, Al-
buquerque. 106 pp.
. 1978. Myotis grisescens. In Endangered
species of the Great Laices Region. U.S. Fish
Wildl. Serv. Region 3. 3 pp.
-. 1979. Status, causes of decline, and
management of endangered gray bats. J.
Wildl. Manage. 43(1): 1-17.
Tuttle, M. D., and P. B. Robertston. 1969. The
gray bat, Myotis grisescens, east of the Appa-
lachians. J. Mammal. 50:370.
Tuttle, M. D., and D. E. Stevenson. 1978. An
analysis of movement as a mortality factor in
the gray bat, based on public recoveries of
banded, bats. Am. Midi. Natur. 97:235-240.
. 1978. Variation in the cave environment
and its biological implications. Natl. Cave
Manage. Symp. Proc. 1977. In press.
Twente, J. W., Jr. 1955. Some aspects of habitat
selection and other behavior of cavern-dwell-
ing bats. Ecology 36:706-732.
GATEKEEPER
To be designated by the Office of Endangered
Species.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.43
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE OKALOOSA DARTER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource manaj^ers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the I-',ndangcred Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed lo:
Office of F^ndangcrcd Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lu:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
FWS/OBS-80/01.43
March 1980
SELECTED VER 1 EBRA 1 E ENDANGERED SPECIES
OF THE SEACOAS I OF IHE UNITED STATES-
THE OKALOOSA DARTER
A Cooperative Effort
by the
National Fish and Wildhfc Laboratory,
the Office of Endangered Species
and the
National ('oastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidcll, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
OKALOOSA DARTER
Etheostoma okloosae Fowler
KINGDOM Animalia
CLASS Osteichthyes
ORDER Perciformes
FAMILY Percidae
OTHER COMMON NAMES
DATE
Entered into SWIS to be determined
Updates 4 October 1976, 24 January 1977
21 May 1979
LEGAL STATUS
Federal: Endangered (38 FR 14678, 4 June
1973)
States: Endangered: Florida.
REASONS FOR CURRENT STATUS
The Okaloosa darter's limited range makes it a
vulnerable species. In 1964 a small number of
brown darters {Etheostoma edwini), a very close-
ly related allopatric species with similar habitat
and ecological requirements, was found at three
stations in Rocky and Swift Creeks, perhaps due
to human introduction (Yerger 1979). By 1975,
the number of Okaloosa darters markedly de-
creased at three stations along Swift Creek, and
the number of brown darters increased (Mettee
1976). Brown darters are now known from 15
sites within the range of the Okaloosa darter and
appear to be replacing the latter species at several
sites in Rocky and Swift Creeks (Mettee and
Crittenden 1977).
Other influencing factors affecting the Oka-
loosa darter's status are temporary disruption of
habitat due to road, bridge, and powerline con-
struction, increased sedimentation, and possible
construction of dams in the future (Yerger 1979).
Mettee (1976) mentioned that there has been
some interest in damming several of the streams
occupied by this darter.
There are presently 15 impoundments on the
drainage system occupied by the Okaloosa darter.
Two on Tom's Creek were made by beavers and
the rest were made by man. Most of these have
produced little effect on darter populations above
and below impoundments, but darters are unable
to use the reservoir portions. Two eutrophic im-
poundments on the portion of Mill Creek flowing
through the golf course at Eglin Air Force Base
have adversely affected populations below the
impoundments, presumably due to higher nutri-
ent content, higher temperature, and increased
turbidity (Crittenden 1974).
Some types of habitat alteration such as road
and powerline crossings may not permanently af-
fect darter populations. Road construction ap-
pears to be harmful for a time (due to excessive
siltation), but once established, the bordering
stream may support an abundant darter popula-
tion due to increased sunlight, stimulating aquatic
plant growth (Crittenden 1974).
In one instance (Swift Creek at State Highway
285) no specimens of Okaloosa darter were taken
following road construction, while the brown
darter was abundant; it is impossible to say
whether habitat destruction or competition with
the brown darter was responsible for the absence
of Okaloosa darters at this site (Mettee et al.
1976). Right of way clearing for powerlines may
be detrimental if exposure of long sections of
streams to sunlight raises the water temperature
(Crittenden 1974).
Presently, a new highway bypassing the cities
of Niceville and Valparaiso is being constructed; it
will bridge Tom's Creek and Turkey Creek. High-
way 85 bridging Juniper (Ten Mile) Creek is being
converted to four lanes. Okaloosa darter popula-
tions are being monitored at these sites by Crit-
tenden and Mettee (Mettee et al. 1976).
PRIORITY INDEX:
Not assigned.
DESCRIPTION
The Okaloosa darter is a slender, small (up to
44 mm) perch-like fish with two dorsal fins,
rounded caudal fins and a lateral line arched
slightly upward with 32 to 37 (usually 34 to 35)
lateral-line scales. There are longitudinal rows of
dark dots along the sides of the body and a series
of dark blotches immediately below the lateral
line. The body is reddish-brown to yellow-brown
becoming lighter on lower sides; the first dorsal
fin has an orange-red stripe near the margin. It
is distinguished from its nearest relative, the
brown darter, by an absence of conspicuous red
spots and a nearly complete lateral line. Photo-
graphs appear in CoUette and Yerger (1962) and
Mettee etal. (1976).
RANGE
Okaloosa darters are endemic to a series of
five small creeks in Okaloosa and Walton Counties
in west Florida that empty into Rocky and Boggy
Bayous near the western end of Choctawhatchee
Bay: Rocky Creek, Swift Creek, Turkey Creek,
Tom's Creek, Mill's Creek and their various tribu-
taries (Collette and Yerger 1962, Crittenden
1974).
There are 300 km of streams in which the
Okaloosa darter is found with a watershed area of
45,730 ha. Approximately 4,860 ha are in private
ownership (including cities of Niceville and Val-
paraiso); the remainder are located within Eglin
Air Force Base (Crittenden 1974).
The species' former distribution is the same as
the present, but it apparently occurred at more
sites on the five creeks (U.S. Department of the
Interior 1973).
RANGE MAP
Distribution illustrated by shading. Detailed
map in Draft Recovery Plan (ODRT 1977).
STATES/COUNTIES
Florida: Okaloosa, Walton.
HABITAT
The Okaloosa darter inhabits small to medi-
um-sized (1.5 to 12.2 m wide, 0.15 to 1.2 m deep)
clear streams with moderate to swift currents.
Substrate is a clean sand with mud or detritus in
areas of reduced current. Waters are neutral to
slightly acidic. Vegetation may be absent or in
scattered patches or clumps. Bulrushes (Scirpus),
bog-moss (Mayaca), golden club (Oronttum
aquaticum), spatterdock (Nuphar luteum), green
algae {Nitella), and pondweed (Potamogeton) are
typical plants. Red algae (Batrachospermum)
frequently forms thick concentrations (Collette
and Yerger 1962).
o
o
O
V
C
u
Okaloosa darters usually are found in water
from 0.15 to 0.61 m deep (Collette and Yerger
1962) associated with clumps of bulrushes and
bur-reed {Sparganium americanum) where cover
and protection are sought (Yerger 1979). They do
not occur in impoundments (ODRT 1977).
Habitat photographs may be found in Crit-
tenden (1974) and Mettee et al. (1976).
FOOD AND FORAGING BEHAVIOR
Not known.
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
Green algae clumps are used as egg-attach-
ment sites during spawning (Collette and Yerger
1962). For water depth and flow requirements
see Reproduction.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Nothing is known of territoriality or home
range size in this species. Relationships with the
brown darter are discussed under Reasons For
Current Status.
Cool, running water is one requirement, and
eutrophication has an undesirable effect (Crit-
tenden 1974).
It is possible that severe rainfall with associ-
ated flooding may affect the distribution in
streams at least temporarily (R. W. Britte personal
communication ) .
POPULATION NUMBERS AND TRENDS
Estimates of population size range from 1,500
to 10,000 individuals with the true figure proba-
bly falling somewhere in between (Yerger 1976).
Mettee (1976) estimates between 1,500 and
2,000 individuals.
Population levels may have been only slightly
reduced in the past 15 years except in areas (parts
of Rocky and Swift Creeks) where the brown
darter has been introduced. Here the numbers of
Okaloosa darters have been reduced, probably
due to competition with sibling species (Yerger
1979).
REPRODUCTION
Spawning was observed in Tom's Creek on 25
March 1961 in sv«ft water 0.30 to 0.46m deep
over a sandy bottom at the edge of clumps of
green algae. Several eggs were deposited individu-
ally in the algae and one egg hatched on 27 March
in the laboratory (Collette and Yerger 1962).
On 25 March 1961, several mature adults
were placed in aquaria with green algae; on 1 April
1961, six eggs were discovered individually
attached to the algae (Collette and Yerger 1962).
Nothing is known of longevity or survival
rates.
MANAGEMENT AND CONSERVATION
Yerger (1979) suggests that field studies be
conducted to determine the species population
status and details of interaction between the Oka-
loosa and brown darters as well as to monitor the
spread of the latter species. He also suggests
potential methods for controlling or eliminating
introduced brown darters by physical removal. He
suggests that this be done while populations are
small and either before spawning has begun or
long after it is over. Okaloosa darters would be
returned to the water after the operation is com-
pleted. Removal should be conducted annually.
Crittenden (1974) suggests that no new
impoundments be constructed on any of the Oka-
loosa darter drainages. If determined necessary,
they should be small and at the extreme head
waters of the tributaries. Measures should be
taken to prevent excessive siltation. Suggestions
include constructing roads at oblique angles to
streams, maintaining water breaks, and seeding of
road banks as soon as possible. The use of pesti-
cides and fertilizers in this drainage area should be
exercised with great caution.
A Recovery Team has been formed and a
draft Recovery Plan submitted to the U.S. Fish
and Wildlife Service for review. The draft plan
calls for studies to determine the optimum habi-
tat for Okaloosa darters, normal population levels
and fluctuations, and potential hazards to the
species' continued existence. The plan also out-
lines actions that could be taken if the results of
research indicate a potential extirpation. These
actions include defining permissible and prohib-
ited activities, land acquisition, creating more
optimum habitat, transplanting, and reducing
populations of competitors or predators (ODRP
1977).
AUTHORITIES
Robert W. Britt (Recovery Team Leader)
Chief, Natural Resources Division
Directorate of Engineering
Elgin Air Force Base, FL 32542
Edward Crittenden (Recovery Team)'
U.S. Fish and Wildlife Service
Great Smoky National Park
Gatlinburg, TN 37738
Maurice F. Mettee, Jr. (Recovery Team)
Geological Survey of Alabama
P.O. Drawer 0
University, AL 35486
Royal D. Suttkus
Museum of Natural History
Tulane University
Belle Chasse, Louisiana
Ralph W. Yerger (Recovery Team)
Department of Biological Sciences
Florida State University
Tallahassee, FL 32306
Norman Young (Recovery Team)
Florida Game and Fresh Water Fish
Commission
P.O. Box 128
DeFuniak Springs, FL 32433
PREPARERS COMMENTS
Since all but 10% of the watershed area is
under government ownership in the Eglin Air
Force Base, control of many potentially detri-
mental factors is facilitated. Development of the
area is kept to a minimum.
Mettee, M. F. Jr. 1970. A survey of the fishes of
the Choctawhatchee Bay drainage in Alabama
and Florida. M.S. Thesis. Univ of Alabama.
93 pp.
. 1976. Etheostoma okaloosae account. In
H. O. Hillestad, D. B. Means, and W. H. Baker,
eds. Endangered and threatened vertebrates of
the southeastern United States. Tall Timbers
Res. Stn.,Misc. Pub. 4.
Mettee, M. F., Jr. and E. Crittenden. 1977. A
study on the distribution o{ Etheostoma oka-
loosa (Fowler) and Etheostoma edwini
(Hubbs and Cannon) in swift and rocky
creeks, Okaloosa and Walton Counties, Flor-
ida, during 1975-1976. Report to U.S. Fish
Wildl. Serv. 130 pp.
Mettee M. F., Jr., R. W. Yerger, and E. Crittenden.
1976. A status report on the Okaloosa darter
in northwest Florida. Proc. S.E. Fishes Counc.
1:1-3.
ODRT (Okaloosa Darter Recovery Team). 1977.
Recovery plan for the Okaloosa darter. Draft.
U.S. Fish Wildl. Serv., Office of Endangered
Species 19 pp.
U.S. Department of the Interior. 1973. Threatened
wildlife of the United States. Compiled by the
Office of Endangered Species and Interna-
tional Activities, Bur. Sports Fish, Wildl.
Resour. Publ. 1 14. U.S. Govt. Printing Office.
Washington, D.C. 289 pp.
Yerger, R. W. 1979. Okaloosa darter. Pages 2-4 in
C. R. Gilbert, ed. Rare and endangered biota
of Florida, Vol 4, Fishes. Univ. Presses of
Florida, Gainesville.
LITERATURE CITED /SELECTED
REFERENCES
Collette, B. B. and R. W. Yerger. 1962. The
American percid fishes of the subgenus
Villora. Tulane Stud. Zool. 9(4):213-230.
Crittenden, E. 1974. Status report on the Okaloosa
darter, an endangered native fish. U.S. Fish
Wildl. Serv. Div. Tech. Assist. Mimeo. Rep
20 pp.
Fowler, H. W. 1941. A collection of freshwater
fishes obtained in Florida 1939-1940, by
Francis Harper. Proc. Acad. Natur. Sci. Phila-
delphia. 92:227-244.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.44
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE EASTERN COUGAR
4^ ■ «. - >
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubhc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This scries of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to;
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems learn
U.S. Fish and Wildlife Service
.NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.44
March 1980
SELECTKl) VI.R 1 EBRA 1 1: ENDANGERED SPECIES
OF THE SEACOAS 1 OE IHE UNITED STATES-
THE EASTERN COUGAR
A Cooperative Effort
by the
National Eish and Wildlife Laboratory,
the Office of Endangered Species
and the
National (Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidcll Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performcil for
Coastal l-cosystems Project
Office of Biological Services
Eish and Wildlife Ser\ice
U.S. Department of the Interior
CREDIT: COLORADO DIVISION OF WILDLIFE
EASTERN COUGAR
Felts concolor cougar Kerr
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivore
FAMILY Felidae
OTHER COMMON
NAMES cougar, mountain lion,
panther, puma, pjiinter
DATE
Entered into SWIS to be detennined
Updates 31 August 1978.
LEGAL STATUS
Federal: Endangered; Eastern United States
(38 FR 14678; 4 June 1978).
States: Endangered: Georgia, Maryland, Mas-
sachusetts, Mississippi, Missouri,
North Carolina, South Carolina, Ten-
nessee, Vermont, Virginia.
REASONS FOR CURRENT STATUS
Regarded as extirpated in the United States in
1899 (USFWS 1973) due to large habitat losses
and disruption, excessive hunting and persecution,
and decline in deer population, its major food
(Laycock 1969; lUCN 1972; USFWS 1973;
Lowman 1975).
PRIORITY INDEX
Not assigned.
DESCRIPTION
A large (1.5-3.1 m), unspotted, long-tailed
cat. Coloration is generally a uniform fulvous or
tawny. Males are larger than females. Underparts,
inner ears, lower cheeks, chin and lips are white.
Tip of tail and base of whiskers are dark. Young
are light-brown and irregularly spotted until
about 6 months old (Dekay 1842; Young and
Goldman 1946).
Eastern cougars are similar in appearance to
the southern and western subspecies {Felis
concolor coryi and F. c. hippolestes, respectively).
Photographs of Felis concolor subspp. are in
Wright (1972).
RANGE
Former range was the entire eastern United
States and Canada as far north as Maine, New
Brunswick, southern Ontario and Quebec; south-
ward through the Appalachian Mountains to
northern Georgia and Alabama where it integra-
ted with F. c. coryi (Young and Goldman 1946;
Hall and Kelson 1959).
Recently alleged sightings (by reliable observ-
ers) and unconfirmed reports (tracks, hair, scat)
are scattered throughout the cougar's former
range.
States from which recent sightings have been
reported include Connecticut (Dowhan and Craig
1976, L. Gray personal communication), Georgia
(Odum et al. 1977, R. R. Odum personal commu-
nication), Kentucky Q. Durrel personal commu-
nication) Maine (Cram 1901, Wright 1972), Mary-
land (Larson 1963, Wright 1972), Massachusetts
(Mugford 1976), Missouri (Schwartz and Schwartz
1959), North Carolina (Linzey and Linzey 1971,
Lee 1977a, 1977b, Teulings and Cooper 1977),
New Hampshire (Dearborn 1927, Wright 1972),
New York (Reilly 1964, Manville 1951), Pennsyl-
vania (Grimm and Whitehead 1950, 1952, Dout
1969), South Carolina (Lowman 1975), Tennes-
see (Kellog 1939), Vermont (Spargo 1950, Wright
1972), Virginia (Russ 1973), West Virginia
(Handly et al. 1961), and Wisconsin (Schorger
1938).
In Canada a population is reported to exist in
New Brunswick (Wright 1953, Calahane 1964,
Wright 1972).
RANGE MAP
No range map has been prepared because of
the speculative nature of this subspecies' occur-
rence in the United States.
STATES/COUNTIES
No states or counties are given because of the
speculative nature of this subspecies' occurrence
in the United States.
HABITAT
The cats show no preference for specific habi-
tat type but appear to require large areas with
adequate food supply and dense vegetation for
refugia (Wright 1972).
FOOD AND FORAGING BEHAVIOR
The cats principal food is white-tailed deer
{Odocoileus virginianus) (Wright 1972). They will
feed on small mammals, insects, and reptiles. They
are also known to take livestock (True 1891;
Hamilton 1943).
Prey is stalked until the cat is close enough to
pounce and is grabbed by the throat or back of
the neck (Hamilton 1943). Remains and unused
portions of prey are covered (Young and Gold-
man 1946). Western subspecies kill every 3 to 4
days (Young and Goldman 1946).
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
Not known.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Not known.
POPULATION NUMBERS AND TRENDS
Populations in New Brunswick, Canada, are
estimated from 25 to 100 (Cahalane 1964; Wright
1972). Recent sightings in the United States may
be of released and/or escaped captives of western
subspecies. Wright (1971, 1972) suggests that the
eastern cougar still exists in the United States and
populations are widely scattered and at a fraction
above the limit for sustaining the population.
REPRODUCTION
Little is known about reproduction of the
eastern cougar. Available information is derived
from other subspecies. Breeding season in the
southern race {F. c. coryi) is believed to be year
round with peaks in February (Schwartz 1952).
Others believe cougars breed in spring or fall
(True 1891). Females are polyestrous and ini-
tiate the courtship and mating act (Davis 1974).
Gestation period is 90 to 98 days (Young and
Goldman 1946; Asdell 1964). Litter size varies
from 1 to 6 with 2 to 3 being average (Young and
Goldman 1946; Hall and Kelson 1959; Asdell
1964). Cougars usually breed once every 2 years
(Young and Goldman 1946; Asdell 1964). There
are also cases where two litters were produced
within a period of 12 to 15 months (Asdell 1964;
Hornocker 1970).
MANAGEMENT AND CONSERVATION
No recovery team or plan has been initiated
for the eastern cougar. There is a Recovery Team
and draft plan for the southern race, Florida
panther {F. c. coryi). Several States are conduct-
ing questionnaire type surveys and some actual
field searches. Some States have created a central
clearing house for sightings of eastern cougars.
There have been sightings and unconfirmed
reports of cougars from several publicly owned
lands in the eastern United States. These include:
Georgia, Savannah River Atomic Energy Plant;
North C.trolina, Blue Ridge Parkway (also in Vir-
ginia), Great Smoky Mountain National Park, and
Uwharrie National Forest; and Virginia, George
Washington National Forest, Jefferson National
Forest, and Shennandoah National Park (Linzey
and Linzey 1971; Russ 1973; Lowman 1975;
Teulings and Cooper 1977).
AUTHORITIES
None.
PREPARER'S COMMENTS
Most of the available literature on life history
of this species is derived from findings of studies
on western subspecies. The eastern cougar and
Florida panther should be treated together in the
Recovery Team and Plan concepts.
There is a serious question as to whether the
eastern cougar exists in the United States. More
effort should be concentrated in the direction of
securing documented evidence of this subspecies'
existence or whether released and/or escaped cap-
tive western subspecies are the animals allegedly
being seen.
LITERATURE CITED/SELECTED
REFERENCES
Asdell, S. A. 1964. Patterns of mammalian repro-
duction. Cornell Univ. Press, Ithaca. 679 pp.
Cahalane, V. H. 1964. A preliminary study of the
distribution and numbers of cougar, grizzly,
and wolf in North America. New York Zool.
Soc, New York. 12 pp.
Cram, G. 1901. Panther in Maine. For. Str. 56:
123.
Davis, W. B. 1974. The mammals of Texas. Texas
Parks Wildl. Bull. No. 41:1-294.
Dearborn, N. 1927. An old account of the moun-
tain lion in New Hampshire. J. Mammal. 8:
311-312.
Dekay, J. E. 1842. Zoology of New York, or the
New York Fauna. Albany. 146 pp.
Dout, J. K. 1969. Mountain lions in Pennsylvania.
Am. Midi. Natur. 82:281-285.
Dowhan, J. J., and R. J. Craig. 1976. Rare and
endangered species of Connecticut and their
habitat. State Geo. Natur. Hist. Surv. Conn.,
Dep. Environ. Protection Rep. 6. 137 pp.
Grimm, W. C, and R. Whitehead. 1950. Mammal
survey of southwest Pennsylvania. Pa. Game
Comm., Harrisburg. 99 pp.
, 1952. Mammal survey of northeast
Pennsylvania. Pa. Game Comm., Harrisburg.
82 pp.
Hall, E. R., and K. R. Kelson. 1959. The mam-
mals of North America. 2 vols. Ronald Press,
New York. 1,083 pp.
Hamilton, W. J. 1943. The mammals of eastern
United States; an account of recent land
mammals occurring east of the Mississippi.
Cornell Univ. Press, Ithaca. 432 pp.
Handly, C. O., Jr., R. Stafford, and E. H. Geil.
1961. A West Virginia puma. J. Mammal.
42:277-278.
Hornocker, M. G. 1969. Winter territoriality in
mountain Hons. J. Wildl. Manage. 33:457-464.
-. 1970. An analysis of mountain lion
predation upon mule deer and elk in Idaho
primitive areas. Wildl. Monogr. 21. 39 pp.
lUCN. 1972. Red data book. Vol. I, Mammalia.
Compiled by H. A. Goodwin and C. W.
Holloway. lUCN, Morges, Switzerland.
Kellog, R. 1939. Annotated list of Tennessee
mammals. Proc. U.S. Natl. Mus. 86:245-303.
Larson, J. S. 1963. Panthers in Maryland. Md.
Conserv.Vol.XLlII.
Laycock, G. 1969. America's endangered wildlife.
Norton, New York. 226 pp.
Lee, D. S. 1977a. Felis concolor True. Pages
354-355 in J. E. Cooper, S. S. Robinson, and
J. B. Funderburg, eds. Endangered and
threatened plants and animals of North
Carolina. North Carolina State Mus. Natur.
Hist., Raleigh. 444 pp.
. 1977b. Unscrambling rumor: the status
of the panther in North Carolina. Wildl. N. C.
41:6-9.
Linzey, A. V., and D. W. Linzey. 1971. Mammals
of Great Smoky Mountains National Park.
Univ. Tennessee Press, Knoxville. 114 pp.
Lowman, G. E. 1975. A survey of endangered,
threatened, rare, status undetermined, peri-
pheral, and unique mammals of the south-
western national forests and grasslands. USDA
For. Serv., Atlanta. 121 pp.
Manville, R. 1951. Reports of cougars in New
York.J. Mammal. 32:227.
Mugford, P. S. 1976. Fish and wildlife in limited
numbers in Massachusetts. Mass. Fish Wildl.
10 pp.
Odum, R. R., J. L. McCuUom, M. A. Neville, and
D. R. Ettman. 1977. Georgia's protected
wildlife. Georgia Dep. Natur. Resour. Atlanta.
Reilly, E. M., Jr. 1964. New York's spooky big
cats. Conservationist 19:227.
Russ, W. R. 1973. The rare and endangered
vertebrates of Virginia. M.S. Thesis. Virginia
Polytechnic Inst, and State Univ., Blacksburg.
338 pp.
Schorger, A. W. 1938. A Wisconsin specimen of
the cougar. J. Mammal. 19:252.
Schwartz, A. 1952. The land mammals of south-
em Florida and the upper Florida Keys. Ph.
D. Dissert. Univ. Michigan, Ann Arbor. 180
pp.
Schwartz, C. W., and E. R. Schwartz. 1959. The
wild mammals of Missouri. Univ. Missouri
Press, Missouri Conserv. Comm. 341 pp.
Sparto, J. 1950. The catamount in Vermont.
Benningtion, Vt.
Teulings, R. P., and J. E. Cooper. 1977. Cluster
areas. Pages 409-433 in J. E. Cooper, S. S.
Robinson, and J. B. Funderburg, eds. En-
dangered and threatened plants and animals
of North Carolina. North Carolina State Mus.
Natur. Hist., Raleigh. 444 pp.
True, F. W. 1891. The puma or American lion:
Felis concolor of Linnaeus. Pages 591-608 in
Ann. Rep. Natl. Mus. Year ending June 30,
1889.
USFWS. 1973. Threatened wildlife of the United
States. Compiled by Office of Endangered
Species and International Activities. Bur.
Sport Fish Wildl. Resour. Publ. 114. U.S.
Gov. Printing Office, Washington, D.C.
Wright, B. S. 1953. Further notes on the panther
in the northeast. Canadian Field Natur. 67:
12-28.
. 1961. The latest specimen of the eastern
puma. J. Mammal. 42:278-279.
1971. The recovery of the panther in
eastern North America. Intl. Symp. Ecology,
Behavior, Conserv. World Cats. Laguna Hills,
California.
1972. The eastern panther, a question of
survival. Clarke, Irvin and Co., Toronto. 180
pp.
Young, S. P., and E. A. Goldman. 1946. The
puma, mysterious American cat. Am. Wildl.
Inst. 358 pp.
GATEKEEPER
To be designated by the Office of Endangered
Species.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
GainesvUle, FL 32601
Biological Services Program
FWS/OBS-80/01.45
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE JA6UARUNDI
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubhc with information about Federally Hsted endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and c()nscr\ation of the subject
species is included (range maps and other distributional data are not necessarily equivalent
to critical habitat as defined in the F.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biohjgical Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
i\'y\SA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.45
March 1980
SELIXIKI) VKRl KBRA 1 K KNDANGERED SPECIES
OF IHE SEACOAS T OE 1 HE UNEIED STATES-
THE JA6UARUNDI
A Cooperative Effort
by the
National Fish and Wildhfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. VVoodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal F,cosystems Project
Office of Biological Services
Fish and Wilillife Service
U.S. Department of the Interior
JAGUARUNDI
Felts yagouaroundi Geoffroy
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivora
FAMILY Felidae
OTHER COMMON
NAMES eyra, ottercat, ghost cat
DATE
Entered into SWIS to be determined
Updates 31 July 1978
LEGAL STATUS
Federal: Endangered (F. y. cacomitli, F. y.
fossata, F. y . panamensis, F.y. tolteca)
(41 FR 24064; 14 June 1976).
States: Endangered: Texas (F. y. cacomitli).
REASONS FOR CURRENT STATUS
Human persecution and habitat loss due to
intense habitat alteration and destruction, partic-
ularly brush-clearing operations (Culbertson and
Schmidly 1974, Davis 1974). However, recent
data may indicate a range extension in southern
Texas (Goodwyn 1970, Brownlee 1978).
PRIORITY INDEX
Not assigned.
DESCRIPTION
A small, slender-bodied, weasel-like cat 0.8 to
1.3m long, and weighing 5.6 to 7.6kg. It is some-
what larger than a housecat, with a longer tail and
unusually short legs. Primarily there are two color
phases - gray and red - with black animals occuring
in the southern part of its range. In the gray phase,
upper parts are grizzled (salt and pepper) with
underparts slightly paler, more black in winter.
The red phase shows reddish upper parts inter-
mixed with black; head and legs are more brownish
while lips and throat are usually white. Fur in
both phases is unspotted. Young are paler with no
spots. Both phases may appear in the same litter.
Photographs are in Goodwyn (1970).
RANGE
The United States distribution is restricted to
extreme southern Texas and Arizona. Recent
reports from Refugio and Hidalgo Counties,
Texas, may indicate a possible range extension
(Goodwyn 1970, Brownlee 1978). These sight-
ings probably represent escaped and/or released
captives (R. McBride personal communication).
Two subspecies are recorded in the United States:
F. y. cacomitli occurs from southern Texas to
central Vera Cruz, Mexico, and F. y. tolteca
occurs in southern Arizona, south in Mexico
along the Pacific coast, and inland to the Mexi-
can Plateau. The Mexican Plateau is thought to
form a barrier between the two subspecies and
may explain the lack of records in New Mexico
(Hock 1955). In Central and South America it
occurs transcontinentally as a poorly known
assemblage of subspecies and closely related
species. One was reported seen at Piatt National
Park in Oklahoma (USFWS 1973).
RANGE MAP
Cross-hatched areas on the following map re-
fer to presumed U.S. distribution and dots refer
to sightings and/or kills (Brownlee 1978, Good-
wyn 1970, R. McBride personal communication).
STATES/COUNTIES
Arizona Cochise, Pima, Santa Cruz.
Texas Cameron, Hidalgo, Starr, Willacy.
HABITAT
The jaguarundi inhabits thick, dense, thorny
brushlands in the lower Rio Grande Valley (Davis
1974). Thickets need not be continuous, but may
be interspersed with clear areas, as found in Mexi-
co (Goodwryn 1970). In South America and other
southern portions of the range, the jaguarundi
occurs in high mountain forests (Alston 1882),
tropical forests, savannas (Bourliere 1964), and
forest swamps (Denis 1964). It seems to prefer
areas near water (Goodwyn 1970). The most
common plants in the lower Rio Grande Valley
where the jaguarundi is known to occur are black-
bush acacia {Acacia rigidula), chillipiquin (Caosi-
cum annum), lotebush [Condalia obstusifolia),
Texas persimmon [Diospyrus texana), coyotillo
{Karwinskia humboltiana), prairie baccharis [Bac-
charis texana), allthorn goatbush {Castela texana),
common lantana {Lantana horrida), berlandier
wolfberry (Lycium berlandieri), javelinabrush
{Microrhamuus ericoides), Texas prickly pear
{Opuntia linheimeri), retama [Parkinsonia acule-
ata), mesquite {Prosopis glandulosa), cedar elm
{Ulmus crassifolia), and lime pricklyash {Zanth-
oxylum fagara). From about 1689 to 1885, the
lower Rio Grande Valley was covered with dense
brush, with a relatively treeless plain to the north
(Inglis 1964). The spread of mesquite north into
this plain (Peacock 1968) may offer suitable habi-
tat for the jaguarundi and a potential for range
extension (Goodwyn 1970).
FOOD AND FORAGING BEHAVIOR
The jaguarundi stalks its prey and then
pounces (Cutter 1957, Goodwyn 1970). It is an
excellent tree climber and will forage in trees
(Davis 1974). Primarily active at night, it will also
forage during the day (Goodwyn 1970, Davis
1974). It consumes mostly birds, but will also
take small mammals and some fish (Cahalane
1947, Ewer 1973, HuUey 1976). Goodwyn
(1970) suggests that principal avifauna prey may
be a dense-brush subtropical cohort (Wolfe 1956)
consisting of Bronzed Cowbird (Molothrus aeneus),
Chachalaca [Ortalis vetula). Green Jay [Cyano-
corax yncas). Groove-billed Ani [Crotophaga sulcir-
ostris), Kiskadee Flycatcher {Pitangus sulphuratus),
Olive-backed Tropical Warbler {Parula pitiayuma).
Red-bellied Pigeon [Columba flavirostris). White-
fronted Dove {Leptoltila verreauxi), and White-
winged Dove (Zenaida asiatica).
SHELTER REQUIREMENTS
Dense thickets are used for refugia (Davis
1974).
NESTING OR BEDDING
Dense thickets, fallen logs, tree hollows, and
thick, grassy clumps are used for den sites
(Goodwyn 1970, Davis 1974).
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RITUAL REQUIREMENTS
The jaguarundi will mark territory by uri-
nating and scratching with hind feet (Hulley
1976). Thirteen distinct calls are recognized with
the largest repertoire occurring during the mating
season (Hulley 1976). Cutter (1957) found three
calls in a captive kitten: chirp, purr, and cough.
Goodwyn (1970) also found these three calls.
Chirp was a high-pitched, bird-like sound, sono-
graph in Goodwyn (1970). Purr sounds are similar
to the domestic cat's purr, but of a higher tone
and more erratic. The cough is a throaty sound as
of air forced from the lungs.
Jaguarundis are solitary except at mating
(Denis 1964). Some captive animals show indica-
tions of gregariousness within family groups (Hul-
ley 1976).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Home range size is not known, but a captive
female was tracked a distance of 3.2 km in one
morning (Hulley 1976).
POPULATION NUMBERS AND TRENDS
No population estimates are available for
most of the range. It is extremely rare in Arizona
(Cockrum 1960).
A questionnaire survey conducted by Texas
Parks and Wildlife Department indicated that jag-
uarundi may occur on three National Wildlife Re-
fuges in Texas: 12 on Santa Ana NWR, 8 on La-
guna Atascosa NWR, and 2 on Aransas NWR
(Brownlee 1978). R. McBride (personal communi-
cation) believes jaguarundis may occur on Laguna
Atascosa, but not on the other two refuges.
REPRODUCTION
Mating is believed to take place in November
and December (Denis 1964). Young are born in
March or August and may weigh up to 22.7 g
(Denis 1964, Caras 1967, Hulley 1976). The ges-
tation period is given as 6 months (Walker 1975).
There may be two litters per year with a range of
one to four (average two) young per litter. Both
gray and red phase may occur in the same litter
(Denis 1964). There is an indication of gregarious-
ness and tolerance of different generations by
adults in captive situations (Hulley 1976), which
may indicate a larger degree of socialism than
once thought. Longevity records indicate a life
span of 8 to 10 years (Rue 1967).
MANAGEMENT AND CONSERVATION
The jaguarundi is protected in the U.S. and
some Latin American countries. Brush is no
longer cleared in the national wildlife refuges in
the lower Rio Grande Valley (lUCN 1972) in
order to maintain brush habitat in its natural
form. Santa Ana and Laguna Atascosa National
Wildlife Refuges are thought to contain habitat
most similar to the original brush habitat of the
lower Rio Grande Valley (Goodwyn 1970).
No recovery team or plan is formalized at this
time.
AUTHORITIES
William C. Brownilee
Texas Parks and Wildlife Department
4200 Smith School Road
Austin, TX 78744
W. B. Davis
Department of Wildlife and Fisheries Science
Texas A&M University
College Station, TX 77843
Roy McBride
Department of Biology
Sul Ross University
Box 725
Alpine, TX 79830
PREPARER'S COMMENTS
It is difficult to describe current population
numbers and distribution based on the question-
naire survey methods used by the Texas Parks
and Wildlife Department. Problems are encoun-
tered with sightings of escaped and/or released
captive individuals and sightings of the same in-
dividual several times. These factors, as well as
misidentification, lead to inaccurate distribution
and population data.
LITERATURE CITED/SELECTED
REFERENCES
Alston, E. R. 1882. Biologia Centralia-Americana.
Mammalia. 219 pp.
Bourliere, F. 1964. Mammals of the world, their
life and habits. A. A. Knopf Co., New York.
223 pp.
Brownlee, W. C. 1978. Feline status survey. Per-
formance Report, Proj. W-103-R-7. Texas
Parks Wildl. Dept., Austin. 4 pp.
Cahalane, V. H. 1947. Mammals of North Ameri-
ca. MacMillan Co., New York. 682 pp.
Caras, R. A. 1967. North American mammals;
furbearing animals of the United States and
Canada. Meredith Press, New York. 578 pp.
Cockrum, E. L. 1960. The recent mammals of
Arizona: their taxonomy and distribution.
Univ. Arizona Press, Tucson. 276 pp.
Culbertson, W. L. 1957. A young jaguarundi in
captivity. J. Mammal. 38:515-516.
Davis, W. B. 1957. The mammals of Texas. Texas
Parks WUdl. Dep. Bull 41 : 1-294.
Denis, A. 1964. Cats of the world. Houghton Mif-
flin Co., Boston. 144 pp.
DeVos, A., R. H. Manville, and R. G. Van Gelder.
1956. Introduced mammals and their influ-
ence on the native biota. Zoologica 41:163-
194.
Ewer, E. F. 1973. The carnivores. Cornell Univ.
Press, Ithaca. 494 pp.
Findley, J. S., A. H. Harris, D. E. Wilson, and C.
Jones. 1975. Mammals of New Mexico. Univ.
New Mexico Press, Albuquerque. 360 pp.
Goldman, E. A. 1925. Two new ocelots from
Mexico. J. Mammal. 6:122-124.
Goodwyn, F. 1970. Behavior, hfe history, and
present status of the jaguarundi. Felts yagoua-
roundi (Lacepede), in south Texas. M. A. The-
sis. Texas A&I University, Kingsville. 63 pf .
Hall, E. R., and K. R. Kelson. 1959. Mammals of
North America. 2 vols. Ronald Press, New
York. 1,083 pp.
Hock, R. J. 1955. Southwestern exotic felids.
Am. Midi. Natur. 53:324-328.
HuUey, J. T. 1976. Maintenance and breeding of
captive jaguarundis {Felis yagouaroundi) at
Chester Zoo and Toronto. Int. Zoo. Yearb.
16:120-122.
Inglis, J. M. 1964. A history of vegetation on the
Rio Grande plain. Texas Parks Wildl. Dep.
Bull. 45:1-122.
lUCN. 1972. Red data book. Vol. I. Mammalia.
Compiled by H. A. Goodwin and C. W. Hol-
loway. lUCN, Morges, Switzerland.
Layne, J. N. 1974. The land mammals of Florida.
In Gleason, P. J., ed. Environments of south
Florida, past and present. Mem. 2 Miami
Geol. Soc. 452 pp.
Little, E. L. 1938. A record of the jaguarundi in
Arizona. J. Mammal. 19:500-501.
Neil, W. T. 1961. On the trail of the jaguarundi.
Fla. Wildl. 15:10-13.
Peacock, J. T. 1968. Historical and present status
of the southern grassland, with special refer-
ence to the distribution of mesquite. Unpubl.
MS. 29 pp.
Rue, L. L. 1967. Pictorial guide to the mammals
of North America. T. Y. Crowell Co., New
York. 299 pp.
Sanderson, I. T. 1959. The living mammals of the
world. Hanover House, New York. 303 pp.
USFWS. 1973. Threatened wildhfe of the United
States. USDI, Bureau of Sport Fisheries and
Wildlife, Washington, D.C. Resource Publ.
114.289 pp.
Walker, E. P. 1975. Mammals of the world. 2 vols.
Johns Hopkins Press, Baltimore. 1,500 pp.
Wolfe, L. R. 1956. Checklist of birds of Texas. In-
teUigence Printing Co., Lancaster.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.46
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE FLORIDA PANTHER
Fish and Wildlife Service
U.S. Department of the Interior
PREPACK
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the I",ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Kndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Sewice
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.46
March 1980
SELECl i:i) VKR 1 KBRA n: ENDANGERED SPECIES
OF IHE SEACOAS I OF IHE UNITED SIATES-
THE FLORIDA PANTHER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fisii and Wildlife Service
U.S. Department of the Interior
CREDIT: R. CHRIS BEL.DEN
FLORIDA PANTHER
Felts concolor coryi Bangs
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivora
FAMILY Felidae
OTHER COMMON NAMES .... cougar, painter,
mountain lion, puma
DATE
Entered into SWIS to be determined
Updates 7 October 1976, 8 March 1977
LEGAL STATUS
Federal: Endangered (32 FR 4001, 11 March
1967)
State: Endangered: Florida, Georgia,
Mississippi
REASONS FOR CURRENT STATUS
The primary cause for their present status is
excessive hunting and habitat disruption (Laycock
1969, Anon. 1973, Nowak and McBride 1973).
Although legally protected since 1958, illegal kills,
highway mortality, and habitat loss probably con-
tinue to depress the population below potential
carrying capacity (Layne and McCauley 1976).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The panther is a large (1.5 to 3.1m), long-
tailed cat, usually rusty on the back, fulvous on
the sides, and buff underneath. The sides of the
nose, tip of the tail, and backs of the ears are dark
brown or blackish. Young panthers, until 6 months
old, arc buff colored with black spots (Audubon
and Backman 1851, Merriam 1901, Hamilton
1943, Hall and Kelson 1959, Hughes 1965).
Photographs are in Allen (1950), Tinsley (1970),
and Lowery (1974).
RANGE
The species formerly ranged throughout Flor-
ida, Georgia, Alabama, Mississippi, Louisiana, and
Arkansas (Hall and Kelson 1959). Its present dist-
ribution is uncertain. Tinsley (1970) and Layne
(1974) state that remaining major concentrations
seem to be in Florida — in Big Cypress Swamp
and the Everglades National Park — while the
extent of the population throughout the remain-
der of its range is unknown.
Although it is questionable that the panther
survives outside of Florida (Williams 1979), recent
reports suggest that they do. R. Nowak (personal
communication) in 1975 examined what may
have been a Florida panther in Logan County,
Arkansas. Lowman (1975) lists sightings near
Valdosta, Georgia; Bankhead National Forest,
Baldwin, and Clarke Counties in Alabama; the
Pascagoula Swamp region of George and Jackson
Counties, and Amite and Claiborne Counties in
Mississippi; Catahoula, Concordia, East Baton
Rouge, Madison, Natchitoches, St. Tammany, and
Webster Parishes in Louisiana; the Ouachita River
bottomlands, Ouachita Mountains, and the White
River National Wildlife Refuge in Arkansas. Also,
the Mississippi Game and Fish Commission (per-
sonal communication) has a recent report of a
sighting in Hancock County, Mississippi.
RANGE MAP
The estimated range in eastern United States
is represented by shading, and confirmed reports
are represented on the following map by stars
(kills, live captures, plaster track casts, photo-
graphs) (Goertz and Abegg 1966, Noble 1971, Sea-
lander andGipson 1973, Lowery 1974,Belden and
Williams 1976, R. C. Belden personal communica-
tion, R. Nowak personal communication).
The Florida map shows the locations of con-
firmed reports by stars, and unconfirmed reports
by dots (Belden and Wilhams 1976).
STATES /COUNTIES
Alabama Baldwin, Clarke, Greene, Mobile,
Tuscaloosa, Winston.
Arkansas Arkansas, Ashley, Carroll, Dallas,
Drew, Franklin, Jefferson, Logan,
Newton, Nevada, Pope, Saline, Scott,
Stone, Washington, White.
Florida Alachua, Baker, Brevard, Browerd,
Citrus, Collier, Columbia, Dade, Dixie,
Duval, Glades, Hendry, Highlands,
Lake, Lafayette, Levy, Manatee,
Marion, Martin, Okeechobee, Osceola,
Palm Beach, Pinellas, Polk, Santa Rosa,
Sarasota, St. Lucie, Taylor, Union,
Wakulla, Walton.
Georgia Charlton, Clinch, Lowndes, Ware.
Louisiana Caddo, Catahoula, Concordia, East
(Parishes) Baton Rouge, Madison, Natchitoches,
St, Tammany, Webster.
Mississippi Amite, Claiborne, George, Hancock,
Jackson.
South
Carolina Aiken, Barnwell.
HABITAT
The panther has been reported in every habi-
tat type. No preference or nonpreference for a
specific habitat type has been noted. A large area
is required, with adequate food supply and dense
vegetation for cover (Bangs 1899).
FOOD AND FORAGING BEHAVIOR
Its principal food is white-tailed deer (True
1891). They also eat small mammals, insects, and
reptiles (Hughes 1965, Smith 1968, Tinsley 1970)
and livestock (True 1891, Hamilton 1943, Rodgers
and Crowder 1974).
They stalk their prey until close enough to
pounce on it, grabbing the throat or back of the
neck (Hamilton 1943, Lowery 1974). If prey can
not be entirely consumed at one time, a panther
will cover it with brush and leaves, and return as
long as it is palatable (Goin 1948, Allen 1950,
Young and Goldman 1964, Lowery 1974). The
frequency of meals is not known, although hunters
in western United States report that mountain
lions (Felis concolor ssp.) kill every 3 to 4 days
(Young and Goldman 1964).
SHELTER REQUIREMENTS
Not known
NESTING OR BEDDING
Not known.
RITUAL REQUIREMENTS
Not known.
The range of the Florida panther is depicted by shading. Stars represent sites of kills, live captures,
plaster track casts or photographs.
NORTHWESIEBN PURT OF
FLORIDA
SAME SCALE as M«IH U«C
9 10 11 12 13 It IS 16 IT IB
20 31
23 2t 25
Confirmed reports of Florida panthers in Florida are indicated by stars on this map; unconfirmed
reports of panthers are depicted by dots.
4
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Because of its scarcity and secretive nature,
almost nothing is known of the behavioral and
environmental requirements of this species. It is
believed to require large expanses of undisturbed
habitat without human interference (Layne 1970).
The panther's home range is unknowTi, but
speculations vary from 150km^ (Layne 1971) to
1,000 to 2,000km^ (Leposky 1975a).
POPULATION NUMBERS AND TRENDS
Specific information on population trends are
not available, but Sealander and Gipson (1973)
and Layne and McCauley (1976) feel that the
population has been on the increase in recent
years. Current estimates of the population in
Florida are between 30 and 300 (Wilhams (1979).
REPRODUCTION
Little is known about the reproduction of the
Florida panther. Most of the information availa-
ble is from the western subspecies. Panthers begin
to breed at 2 to 3 years of age (Hall and Kelson
1959, Young and Goldman 1964). Breeding is
believed to occur year-round with births peaking
in February (Schwartz 1952, Frye et al. no date).
Other sources indicate that panthers breed in the
spring or fall (Maynard 1883, True 1891).
Davis (1966) states that females start the
courtship and mating act. Males will fight for
first breeding privileges. Copulation is followed
by subsequent unions with other males (Lowery
1974). The gestation period is 90 to 98 days
(True 1891, Asdell 1964, Young and Goldman
1964). Litter size varies from 1 to 6 (Hall and
Kelson 1959) with 2 or 3 being typical (Asdell
1964, Young and Goldman 1964).
Panthers usually breed every 2 years (Asdell
1964, Young and Goldman 1964, Rogers and
Crowder 1974). There are a few reports of two
litters being born within a 12- or 15-month
period (Asdell 1964, Hornocker 1970).
MANAGEMENT AND CONSERVATION
The Florida Panther Recovery Team was
organized in July 1976, with the main objective
to develop a recovery plan "... that delineates
and schedules those actions required for securing
or restoring . . ." the Florida panther ". . . as a
viable, self-sustaining member of its ecosystem."
A rough draft has been developed which calls for
four major efforts (R. C. Beldon personal com-
munication):
1. Find and delineate present populations.
2. Develop a captive breeding program for possi-
ble restocking in the future.
3. Develop a public education program.
4. Develop a program to evaluate present rules
and regulations regarding panthers, and form-
ulate new ones where necessary.
The Florida Game and Fresh Water Fish Com-
mission (R. C. Belden personal communication),
has outlined an investigation to locate and delin-
eate the present population, and if found, to
determine important habitat and management
needs to assure continued survival. To locate and
delineate the present population, a central "Flor-
ida Panther Record Clearinghouse" will be set up
to receive panther records. The public will be
advised whom to contact and what to report.
These records will then be analyzed and collated
into a list of priority areas to be searched for
panther sign. Intensive field searches will be con-
ducted in areas with positive sign of a panther
population. Efforts will be made to determine
habitat utilization and population dynamics. All
information will then be analyzed to determine
panther habitat needs and to develop appropriate
management strategies.
AUTHORITIES
Ken C. Alvarez (Recovery Team)
Division of Recreation and Parks
Florida Department of Natural Resources
P.O. Box 398
Osprey, FL 33559
Robert E. Baudy (Recovery Team)
Rare Feline Breeding Compound
P.O.Box 132
Center Hill, FL 33514
Robert C. Belden (Recovery Team Leader)
Florida Game and Fresh Water Fish Commis-
sion
4005 South Main Street
Gainesville, FL 32601
James N. Layne (Recovery Team)
Archbold Biological Station
Route 2, Box 180
Lake Placid, FL 33852
Brian Noles (Recovery Team)
Appalachicola Ranger District
P.O. Box 578
Bristol, FL 32321
Peter C. H. Pritchard (Recovery Team)
Florida Audubon Society
P.O. Drawer 7
Maitland,FL 32751
Lovett Williams
Florida Game and Fresh Water Fish Commis-
sion
4005 South Main Street
Gainesville, FL 32601
PREPARER'S COMMENTS
Current status, life history, and ecology of the
Florida panther are poorly known. Most biologi-
cal data are limited to general accounts or brief
notes from early Florida explorers and naturalists.
The majority of publications since the turn of the
century rehash the limited original data and in-
formation extracted from general studies done on
other subspecies. More research to determine sta-
tus, distribution, and ecological requirements is
needed for making sound conservation and man-
agement decisions.
LITERATURE CITED/SELECTED
REFERENCES
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1974. A user-accessed computer information
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Vol. 2. U.S. Army Eng. Waterways Exp. Stn.
Mob. Environ. Systems Lab. Vicksburg, I lis-
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Allen, E. R. 1950. Notes on the Florida panther,
Felis concolor coryi Bangs. J. Mammal. 31:
279-280.
Allen, E. R., and W. T. Neill. 1954. The raccoon
preyed upon by panther and rattlesnake. Ev-
erglades Natur. Hist. 2:46.
Anon. 1971. Wildlife popular in state parks. Flor-
ida Natur. 44(3 A) :9.
. 1973. Floridapanther in imninent danger.
World Wildl. News 1(3): 3-4.
Asdell, S. A. 1964. Patterns of mammahan repro-
duction. 2nd ed. Comstock Publ. Assoc, Cor-
nell Univ. Press, Ithaca, New York. 670 pp.
Audubon, J. J. and J. Bachman. 1846, 1851,
1854. The viviparous quadrupeds of North
America. 3 vols. George R. Lockwood, New
York.
Bangs, O. 1898. The land mammals of peninsular
Florida and the coast region of Georgia. Proc.
Boston Soc. Natur. Hist. 28(7):157-235.
. 1899. The Florida puma. Proc. Biol. Soc.
Washington, D.C. 13:15-17.
Barbour, T. 1943. Naturalist at large. Little,
Brown, and Co., Boston. 314 pp.
Belden, R.C., and L.E. Williams, Jr. 1976. Surviv-
al status of the Florida panther. In Proc.
of a Florida panther conference and work-
shop. March 1976. Florida Audubon Soc.
In Press.
Blair, W. F., A. P. Blair, P. Brodkorb, F. R. Cagle,
and G. A. Moore. 1968. Vertebrates of the
United States. 2nd ed. McGraw-Hill, New
York. 616 pp.
Cahalane, V. W. 1964. A preliminary study of dis-
tribution and numbers of cougar, grizzly and
wolf in North America. New York Zool. Soc.
12 pp.
Carr, A. F. 1973. The Everglades. Time-Life
Books, New York. 184 pp.
Cory, C. B. 1896. Hunting and Fishing in Florida.
Estes and Lauriat, Boston. 304 pp.
Davis, W. B. 1966. The mammals of Texas. Rev.
ed. Texas Parks and Wildl. Dept. Bull. 41:
1-267.
Dodson, P. 1973. Journey through the old Ever-
glades, the log of Minnehaha. Trend House,
Tampa, Florida. 75 pp.
Eaton, R. L. 1970. Pilot study on ecology of
Florida panther. Unpubl. M.S. Thesis.
.1973. The status, conservation, and man-
agement of the cougar in the United States.
Pages 68-86 in R. L. Eaton, ed. The world's
cats. World Wildlife Safari, Winston, Oregon.
349 pp.
Elliott, D. G. 1905. A check list of mammals of
the North American continent, the West In-
dies, and the neighboring seas. Publ. Field
Col. Mus., Zool. Ser. 6:1-761.
Florida Game and Fresh Water Fish Commission.
1976. Cross Florida barge canal restudy re-
port; endangered, threatened, rare, special
concern, status undetermined and biologically
sensitive species. Submitted to Fish Wildl. Ser.
267 pp.
Frye, O.E., B. Piper, and L. Piper. No date. The
disappearing panther. Mimeogr. 4 pp.
Goertz, J. W., and R. Abegg. 1966. Pumas in Lou-
isiana. J. Mammal. 47:727.
Goin, O. B. 1948. Cat of a dozen names. Fauna
10(4):101-105.
Hall, E. R., and K. R. Kelson. 1959. The mam-
mals of North America. 2 vols. Ronald Press,
New York. 1,083 pp.
Hamilton, W. J. 1943. The mammals of eastern
United States: an account of recent land
mammals occurring east of the Mississippi.
Comstock Publ. Assoc, Cornell Univ. Press,
Ithaca, New York. 432 pp.
Harper, F. 1927. The mammals of the Okefeno-
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Soc. Natur. Hist. 38:191-396.
Heilprin, A. 1887. Explorations on the west coast
of Florida and in the Okeechobee wilderness
with special reference to the geology and zo-
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Inst. Sci. Trans. 134 pp.
Homocker, M. G. 1969. Winter territoriality in
mountain lions. J. Wildl. Manage. 33:457-464.
. 1970. An analysis of mountain Hon
predation upon mule deer and elk in Idaho pri-
mitive area. Wildl. Monogr. 21. 39 pp.
Howell, A. H. No date. Xerox copy of unpub-
lished manuscript on Florida mammals. Con-
tained in the files of U.S. Fish Wild. Serv.
Washington, D.C. 114 pp.
Hughes, W. 1965. Florida's wild cats. Florida
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lUCN. 1972. Red data book. Vol. 1. Mammalia.
Compiled by H. A. Goodwin and C. W. Hol-
loway. lUCN, Merges, Switzerland.
Jenkins, J. H. 1971. The status and management
of the bobcat and cougar in the southeastern
states. Pages 87-91 in S. E. Jorgenscn and
L. D. Mech, eds. Proc. of a symposium on the
native cats of North America; their status and
management. U.S. Dept. Inter., Fish Wildl.
Serv., Bur. Sport Fish. Wildl. Region 3. 139
pp.
Lahart, D. 1973. Florida's endangered dozen.
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Laycock, G. 1969. America's endangered wildlife.
Norton, New York. 226 pp.
Layne, J. N. 1970. Terrestrial wildlife of the
Oklawaha regional ecosystem, effects of
Cross-Florida Barge Canal construction on
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and 88-91 in Environmental impact of the
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Florida Def. Environ. 115 pp.
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. 1974. The land mammals of south Flori-
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Memoir 2 Miami Geol. Soc. 452 pp.
Layne, J. N. and M. N. McCauley. 1976. Biologi-
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Nowak, R. M., and R. McBride . 1973. Feasibility
of a study on the Florida panther. Rep. to
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Pearson, P. G. 1951. Mammals of Gulf Hammock,
Levy County, Florida. M.S. Thesis. Univ. of
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617.
Ray, C. E. 1958. Additions to the Pleistocene
mammalian fauna from Melbourne, Florida.
Bull. Mus. Comp. Zool. 119:421-449.
Robertson, W. B., Jr. 1959. Everglades— the park
story. Univ. of Miami Press, Coral Gables.
88 pp.
Robinette, W. L., J. S. Gashwiler, and O. W.
Morris. 1961. Notes on cougar productivity
and life history. J. Mammal. 42:204-217.
Rodgers, D. P., and J. P. Crowder. 1974. Threat-
ened wildlife of south Florida. South Florida
Environ. Proj. Ecol. Rep. DI-SFEP-74-25.
34 pp.
Rogers, A. J. 1953. A study of the exodid ticks of
northern Florida, including the biology and
life history oi Ixodes scapularis Say (Ixodidae:
Acarina). Ph. D. Dissert. Univ. Maryland, Col-
lege Park. 176 pp.
Russ, W. P., 1973. The rare and endangered verte-
brates of Virginia. M.S. Thesis. Virginia Poly-
technic Inst, and State Univ., Blacksburg.
338 pp.
St. Amant, L. S. 1959. Louisiana wildlife inven-
tory and management plan. Louisiana Wildl.
Fish Comm. 329 pp.
Schemnitz, S. D. 1974. Populations of bear, pan-
ther, alligator, and deer in the Florida Ever-
glades. Q.J. Fla. Acad. Sci. 37:157-167.
Schwartz, A. 1952. The land mammals of south-
ern Florida and the upper Florida Keys. Ph. D.
Dissert. Univ. of Michigan, Ann Arbor.
180 pp.
Sealander, J. A., Jr. 1951. Mountain lion in Ar-
kansas. J. Mammal. 32:364.
• 1956. Provisional checklist and key to
the mammals of Arkansas (with annotations).
Am. Midi. Natur. 56:257-296.
Sealander, J. A., Jr., and P. S. Gipson. 1973. Sta-
tus of the mountain lion in Arkansas. Proc.
Arkansas Acad. Sci. 27:38-41.
Seidensticker, J. C, IV., M. G. Hornocker, W. V.
Wiles, and J. P. Messick. 1973. Mountain lion
social organization in the Idaho Primitive
Area. Wildl. Monogr. 35:1-61.
Smith, G. 1968. The Florida panther. Florida
Wildl. 21(8):30-31.
. 1970. Mystery cat. Florida Wildl. 24(3):
4-6.
Tinsley, J. B. 1970. The Florida panther. Great
Outdoors Publ. Co., St. Petersburg, Florida.
60 pp.
True, F. W. 1891. The puma, or American lion:
Felis concolor of Linnaeus. Pages 591-609
in Ann. Rep. U.S. Natl. Mus. Year ending
June 30, 1889.
U.S. Department of the Interior. 1969. Environ-
mental impact of the Big Cypress Swamp Jet-
port. 155 pp.
. 1973. Threatened wildlife of the United
States. Compiled by Office of Endangered
Species and International Activities. Bur.
Sport Fish. Wildl. Res. Publ. 114. U.S. Gov.
Printing Office, Washington, D. C. 289 pp.
Webb, S. D. 1974. Chronology of Florida Pleisto-
cene mammals. Pages 5-31 in S. D. Webb, ed.
Pleistocene mammals of Florida. Univ. of Flo-
rida Press, Gainesville. 270 pp.
Wolfe, J. L. 1971. Mississippi land mammals. Mis-
sissippi Game Fish Comm. 44 pp.
Young, S. P., and E. A. Goldman. 1964. The
puma, mysterious American cat. Dover Publ.,
Inc. New York. 358 pp.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.47
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE AMERICAN CROCODILE
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoast of the United States. In-
formation about life history, distribution, requirements and c<)nscr\'ation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as ainendcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps o( Engineers in coordina-
tion with the Offices of Kndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal F-cosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.47
March 1980
stLECi i:d vi:ri ebra 1 i: endangered species
OF THE SEACOAS 1 OF IHE UNITED STATES-
THE AMERICAN CROCODILE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. VV'oodard
National Coastal Ecosystems Team
NASA-Slidcll Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Ser\iccs
Fish and Wildlife Service
U.S. Department of the Interior
M^:."^^^
« • '■A
"^^^
^■l^^-'^P-^
CREDIT: J. ROSS WILCOX
AMERICAN CROCODILE
(Crocodylus acutus Cuvier)
KINGDOM Animalia
CLASS Reptilia
ORDER Crocodylia
FAMILY Crocodylidae
OTHER COMMON
NAMES Florida crocodile
DATES
Entered into SWIS to be determined
Updates 20 September 1976; 8 March 1977
LEGAL STATUS
Federal! Endangered (Florida populations only)
(40 FR 44149, 25 September 1975).
States: Endangered: Florida.
REASONS FOR CURRENT STATUS
Loss of habitat due to urbanization of south
Florida; habitat loss is not all physical alteration,
but includes reduction in suitability of remaining
habitats through increased human recreational
and other uses (Ogden 1979).
Excessive mortality occurs through malicious
and ill-advised killing by humans and through
accidental deaths in commercial fishing nets and
on highways (Lang 1975).
Heavy predation on hatchlings in Florida Bay,
primarily by raccoons, is believed to be hampering
recruitment (Lang 1975).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The crocodile is a large, lizard-shaped reptile.
Its back is covered with regular series of keeled,
bony scales and the belly with smooth white
scales. It is dorsally greenish or gray-brown with
irregular black mottling. Young are more con-
trastingly colored than adults. Hatchlings are
about 23 cm in length; adults may grow to 4.5 m
or larger.
The characteristics that distinguished the
crocodile from the alligator are the crocodile's
generally more slender build and its snout, which
tapers noticeably forward of the eyes, while the
alligator's snout is untapered and rounded at the
end. The fourth tooth in the crocodile's lower jaw
is exposed when the mouth is closed; this tooth is
concealed in the alligator. Coloration of the alliga-
tor is dark gray or black with yellow markings.
Illustrations of the crocodile appear in
Ditmars (1953), Carr and Coin (1955), Pope
(1955), Bothwell (1963), Neill (1971), Conant
(1975), Lang (1975), and Perrero (1975), among
other technical, semitechnical. and popular works.
RANGE
In the United States, the American crocodile
is known to breed currently only in southern
parts of the Everglades National Park, chiefly Flo-
rida Bay, and outside the park on adjacent Key
Largo and Turkey Point. Another population is
reported in the Lower Florida Keys on Big Pine,
Little Pine, and Howe Keys, with breeding
rumored on Little Pine Key (Powell 1973, Ogden
1979).
Individuals are still occasionally reported north
of Key Largo into Biscayne Bay on the east coast
of Florida, and as far north as Marco Island and
Charlotte Harbor on the west coast. Breeding in
these areas has not been documented.
Historically, the crocodile ranged north at
least to Lake Worth, Palm Beach County, on the
east coast (Dimock and Dimock 1908). Breeding
is suspected but undocumented for these northern
populations. On the west coast of Florida, no his-
toric records exist for sightings outside the areas
that still occasionally report crocodiles, although
a record does exist for Mobile Bay, Alabama
(Loding 1922).
The largest segment of the known population
in Florida is found in the Everglades National
Park. The lower Keys population, if a viable
breeding unit, lives primarily in the National Key
Deer Refuge. Crocodiles occasionally use the
cooling canals of the Florida Power and Light
Corporation's Turkey Point power plant. Individ-
uals are also occasionally seen in Homestead Bay-
front Park.
RANGE MAP
Shading indicates the known range in Florida.
STATES/COUNTIES
Florida Collier ( ?) , Dade , Monroe.
HABITAT
Primarily coastal, crocodiles use mangrove
swamps, salt and brackish bays, and brackish
creeks. They also enter coastal canals and borrow
pits. Nesting occurs primarily in hardwood thick-
ets at the heads of small sand beaches and on marl
banks along narrow coastal creeks (Ogden 1979).
Present data indicate that hatchlings cannot
tolerate seawater salinities, but must have brackish
or freshwater for the early development period
(Neill 1971, T. Ellis and W. E. Evans personal
communication), but this has been questioned
(Lang 1975). Adults are able to withstand full
seawater salinity and may wander widely in
coastal areas (Neill 1971).
Critical habitat for the American crocodile in
Florida has been designated (41 FR 41915, 24
September 1976). All land and water (excluding
man-made structures or settlements not necessary
to the normal needs or survival of the species)
within the followdng boundaries are included: be-
ginning at the easternmost tip of Turkey Point,
Dade County, on the coast of Biscayne Bay;
thence southeastward along a straight line to
Christmas Point at the southernmost tip of Elliott
Key; thence southwestward along a line following
the shores of the Atlantic Ocean side of Old
Rhodes Key, Palo Alto Key, Angelfish Key, Key
Largo, Plantation Key, Windley Key, Upper Mate-
cumbe Key, Lower Matecumbe Key, and Long
Key, to the westernmost tip of Long Key; thence
northwestward along a straight line to the western-
most tip of Middle Cape; thence northward along
the shore of the Gulf of Mexico to the north of
the mouth of Little Sable Creek; thence eastward
along a straight line to the northernmost point of
Nine Mile Pond; thence northeastward along a
straight line to the point of beginning.
FOOD AND FORAGING BEHAVIOR
No detailed information on foraging behavior
is known, but the food of adults is believed to
consist primarily of fish (Fernandez 1971, Ogden
_1 2 3 * 5 6 7
A'. J '^ xy.A\
17 ^18 19
31 a 33 2* 2S
NOBIHWESItCfJ PART OF
FLORIDA
5»»E se»i.£ as ifiis "i-
Shading on the map depicts the present range of the American crocodile in Florida.
23 24 2',
ALABAMA.
NORTHWESTERN PART OF
FLORIDA
2 3
13 14 15
Shading on this map indicates critical habitats of the American crocodile in Florida (41 FR 41915,
24 Sept. 1976).
1979). Young feed chiefly on aquatic inverte-
brates (Martin de Lucenay 1942, Alvarez del Toro
1974, Ogden 1979). In general, crocodilians feed
on any prey items that can be caught and over-
powered (Cott 1961,Neill 1971).
SHELTER REQUIREMENTS
Not known.
NESTING OR BEDDING
Nesting site preferences appear to be hardwood
thickets at the edge of small sand beaches or the
banks of narrow coastal creeks (Ogden 1979).
Loose soil that can be scraped into mounds ap-
pears to be a general requirement, although the
Florida Bay population appears variable in this re-
gard (Campbell 1972, Ogden 1979). Fifty meters
between nests may be required for successful
nestin ; (Alvarez del Toro 1974).
Dens consisting of burrows dug into creek
banks are often constructed, but the relation of
these dens to nesting areas or hibernation/estiva-
tion needs is unknown (Alvarez del Toro 1974,
Ogden 1979).
RITUAL REQUIREMENTS
Crocodihans exhibit the most elaborate nest-
protection behavior of any reptile, including assis-
tance to the young during the hatching process
and posthatching protection of young (Alvarez
del Toro 1969, 1974; Neill 1971;Kushlan 1973;
Ogden and Singletary 1973; Pooley and Gans
1976). Ogden and Singletary (1973) document
this behavior for the Florida Bay population.
Requirements for this ritualistic behavior are
unstudied, but Dixon and Staton (1976) have evi-
dence that disturbance of the female while
guarding the nest will disrupt her behavior and
may lead to higher egg mortality. Freedom from
human disturbance during the period of incuba-
tion and hatching may thus be a critical factor.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Hatchling crocodiles may require access to
freshwater during their early lives (Neill 1971,
T. Ellis andW. E. Evans personal communication);
Lang (1975) differs from this view.
Home range/territoriality requirements are
unknown, but crocodiles wander over considerable
areas at times (Neill 1971). Local or seasonal vari-
ation in food supply may prompt these moves
(Ogden 1979).
POPULATION NUMBERS AND TRENDS
Present population structure in Florida may
be slanted toward larger individuals with no evi-
dence of recruitment of young into the popula-
tion. No data are available on the structure of
natural, healthy populations of American croco-
diles.
Population trends in Florida are clearly down-
ward. The estimated population in Florida is be-
tween 200 and 400 individuals with fewer than 25
known breeding females (Ogden 1979). Loss of
several breeding females on Key Largo has been
documented over the past 5 years.
REPRODUCTION
Courtship and mating are described by Lang
(1975). Females in south Florida begin construc-
tion of nests in April and eggs are laid in April or
May. Twenty to 80 eggs may be layed in a clutch
(Ogden 1979). Hatching occurs in July and
August.
An adult crocodile, presumably the female
parent, opens the nest and assists the young in
hatching (Ogden and Singeletaryl973, Alvarez del
Toro 1974). A close early protective relationship
between the parents and young is known for
several crocodile species (Kushlan 1973, Alvarez
del Toro 1974, Pooley and Gans 1976) and has
been postulated for the Florida population
(Ogden 1979), although its duration and extent
are unknown.
The survival rate of young is unknowm under
natural conditions but is very low for the Florida
population (Lang 1975).
MANAGEMENT AND CONSERVATION
Crocodilians in general, with their high repro-
ductive potential, usually respond well to effec-
tive management/ conservation programs, as the
rapid recovery of the American alligator in the
United States indicates. However, no population
of the American crocodile has as yet received
such protection and thus its potential for response
to specific programs cannot be evaluated. The
State of Florida and the Everglades National Park
have given legal protection to crocodile popula-
tions for some time, yet the population has
continued to decline. These data indicate that
enforcement or protective legislation alone is
inadequate to rebuild the Florida crocodile
population. An active program to decrease
accidental mortahty and increase recruitment
in conjunction with a public information
program to increase the public's tolerance to large
breeding-sized individuals would appear manda-
tory.
The Recovery Plan for the American croco-
dile has been approved (12 Feb 1979) and out-
lines steps to be taken to assure a self-sustaining
population throughout suitable habitat in the
United States. The plan calls for research to
determine habitat needs, habitat distribution,
crocodile distribution, habitat ownership, and
availability of crocodile habitat. The plan also
outlines action steps that may be necessary pend-
ing research findings, including controlling man-
related mortality, educating the public, increasing
recruitment by captive propagation and release,
reducing natural mortality, and protecting nesting
sites.
AUTHORITIES
Howard W. Campbell (Recovery Team Leader)
National Fish and WildHfe Laboratory
412 N.E. 16th Ave, Room 250
Gainesville, FL 32601
Tommy Hines (Recovery Team)
Research Biologist
Florida Game and Fresh Water Fish
Commission
4005 South Main Street
GainesvUle, FL 32601
Richard Klukas (Recovery Team)
Resource Management Specialist
National Park Service
Southeast Regional Office
1895 Phoenix Blvd.
Atlanta, GA 30349
James A. Kushlan (Recovery Team)
Research Biologist
Everglades National Park
P.O. Box 279
Homestead, FL 33030
Paul Moler
Florida Game and Fresh Water Fish
Commission
4005 South Main Street
Gainesville, FL 32601
John C. Ogden (Recovery Team)
Research Biologist
National Audubon Society
115 Indian Mound Trail
Tavemier, FL 33070
PREPARER'S COMMENTS
Although no subspecies of the American
crocodile have been described, unpublished data
indicate that geographic variation in several char-
acters exists and studies are required to determine
the relationships of the Florida crocodile popula-
tion to other populations. It appears, on the basis
of available data, to be distinct on an average sub-
species level of difference from at least the Jamaica
and Pacific coast of Mexico populations. Its iso-
lated and peripheral status suggests that genetic
differentiation from other populations is a strong
possibility.
LITERATURE CITED /SELECTED
REFERENCES
Alvarez del Toro, M. 1969. Breeding the spec-
tacled caiman at Tuxtla Gutierrez Zoo.
Internat. Zoo Yearb. 9:35-36.
. 1974. Los Crocodylia de Mexico (Estudio
Comparativo). Ediciones de Inst. Mex. Re-
cursos Natur. Renovables, A.C. Mexico, D.F.
70 pp.
Bothwell, D. 1962. The great outdoors book
of alligators and other crocodiUa. Great Out-
doors Publ. Co. St. Petersburg. 88 pp.
Campbell, H. W. 1972. Ecological or phylo-
genetic interpretations of crocodilian nesting
habits. Nature 238 (5364):404-405.
Carr, A. F. and C. J. Goin. 1955. Guide to the
reptiles, amphibians, and fresh-water fishes of
Florida. Univ. of Florida Press. Gainesville.
341 pp.
Conant, R. 1975. A field guide to reptiles and
amphibians of eastern and central North
America. Houghton Mifflin Co., Boston.
429 pp.
Cott, H. B. 1961. Scientific results of an inquiry
into the ecology and economic status of the
Nile crocodile {Crocodylus niloticus) in
Uganda and northern Rhodesia. Trans. Zool.
Soc. London 29(4):21 1-357.
Craighead, F. C. 1968. The role of the alligator in
shaping plant communities and maintaining
wildlife in the southern Everglades. Florida
Natur. 41(8):2-7, 69-74.
Dimock, A. W. and J. A. Dimock. 1908. Florida
enchantments. Outing Publ. Co., New York.
289 pp.
Ditmars, R. L. 1953. The reptiles of North
America. Doubleday and Co., Inc. New York.
476 pp.
Dixon, J. and M. Staton. 1976. Breeding biology
of Caiman crocodilus crocodilus from the
Venezuelan Llanos. U.S. Fish Wildl. Res. Rep.
5:1-21.
Fernandez, W. 1971. The American crocodile.
Mus. News, Miami. 1(9):209-211.
Fittkau, E. J. 1970. Role of caimans in the
nutrient regime of mouth-lakes of Amazon
effluents. Biotropica 2(2): 138-142.
. 1973. Crocodiles and the nutrient metabo-
lism of Amazon waters. Amazonia 4(1):
103-133.
Kushlan, J. A. 1973. Observations on maternal
behavior in the American alligator. Alligator
mississippiensis . Herpetologica 29(3): 256-257.
. 1974. Observations on the role of the
American alligator [Alligator mississippiensis)
in the south Florida wetlands. Copeia 1974
(4):993-996.
Lang, J. W. 1975. The Florida crocodile: will
it survive. Bull Field Mus. Natur Hist. 46(8):
5-9.
Loding, H. P. 1922. A preliminary catalog of
Alabama reptiles and amphibians. Alabama
Geol. Surv. Natur. Hist. Mus.. Pap. 5:1-59.
Martin de Lucenay, A. 1942. El crocodrilo
mexicano II, el desarroUo y la alimentacion.
Rev. Gral. de Marina. Mexico.
Ogden, J. and C. Singletary. 1973. Night of the
crocodile. Audubon 75(3):32-37.
Perrero, L. 1975. Alligators and crocodiles of the
world. Windward Publ. Co. Miami. 64 pp.
Pooley, A. C. and C. Cans. 1976. The Nile croco-
dile. Science 234(4): 114-124.
Pope, C. H. 1955. The reptile world. Alfred A.
Knopf, New York. 325 pp.
Powell, J.A. 1973. The status of Crocodylus
acutus in southern Florida. In Proc. 2nd
working meet, of crocodile specialists. lUCN
Pub. New Sen, Suppl. Pap. 41:1-115.
ACCOUNT PREPARED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Ave., Room 250
Gainesville, FL 32601
Neill, W. T. 1971. The last of the ruling reptiles.
Columbia Univ. Press, New York and London
486 pp.
Ogden, J. 1979. Florida crocodile, in R. W.
McDiarmid ed. Rare and endangered biota of
Florida, Vol 3, Amphibians and reptiles. Univ.
Presses of Florida, Gainesville. Pages 21-22.
Biological Services Program
FWS/OBS-80/01.48
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
THE KEY DEER
ts»-^ *<■•:•<; ■'---
-"--'.■.. if'.
t^e^^^.
.:d.^im'
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangcred Species Act of 1973, as dmendcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services oi the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed lo:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Sen ice
N ASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.48
March 1980
SELECTED VERI EBRA IE ENDANGERED SPECIES
OF THE SEACOAST OF 1 HE UNFFED STATES-
THE KEY DEER
A Cooperative Effort
by tin-
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National (Coastal Ecosystems 1 earn,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidcll Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office ()f Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
KEY DEER
Odocoileus virginianus clavium Barbour and Allen
KINGDOM Animalia
CLASS Mammalia
ORDER Artiodactyla
FAMILY Cervidae
OTHER COMMON NAMES toy deer
DATE
Entered into SWIS to be determined
Updates 15 October 1976, 4 March 1977
LEGAL STATUS
Federal: Endangered (32 FR 4000, 11 March
1977)
States: Endangered: Florida.
REASONS FOR CURRENT STATUS
Loss of habitat to development of Lower Keys
is the primary reason for the deer's current status.
Big Pine Key supports some two-thirds of the
total population, and 37% of this island had been
cleared for development as of 1973 (Klimstra et
al. 1974). The human population of the keys is
projected to increase by 48% by 1990, implying
even more loss of deer habitat.
Other factors include mortality of young
falling into drainage ditches and being killed by
automobiles. Overhunting with dogs and jack-
lights (U.S. Department of the Interior 1973) was
probably an important factor in the 1940's and
1950's, and reports of poaching are still rather
frequent (Klimstra et al. 1974).
PRIORITY INDEX
Not assigned.
DESCRIPTION
The smallest race of North American deer, the
adult key deer is 63 to 76 cm high at the shoulder
with an average weight of 36 kg for males and 29
kg for females. The body is stockier, the legs
shorter, and the skull wider than other races of
white-tailed deer. The coat varies from a deep red-
dish brown to a grizzled gray color. Bucks usually
develop antlers by their second year, and eight
points by the fifth year (Klimstra 1979).
RANGE
The key deer's range is restricted to the
islands of the lower Florida Keys: Annette, Big
Munson, Big Pine, Big Torch, Cudjoe, Howe,
Johnson,Knockemdown, Little Pine, Little Torch.
Mayo, Middle Torch, No Name, Porpoise, Ram-
rod, Sugarloaf, Summerland, Toptree Hammock,
Wahoo, and Water Key (Dickson 1955, Klimstra
et al. 1974). The deer are resident only on keys
with permanent freshwater: Big Pine, Big Torch,
Cudjoe, Howe, Little Pine, Litde Torch, Middle
Torch, No Name, Sugarloaf, and Summerland
Keys.
RANGE MAP
Present distribution is indicated by shading on
the following page.
STATES/COUNTIES
Florida: Monroe
HABITAT
Only islands having permanent freshwater
are used consistently by key deer. Other islands
are used temporarily during the rainy season
(Klimstra 1979). Big Pine Key (2,428 ha) and No
Name Key (404 ha) support the largest deer pop-
ulation. Both have permanent freshwater and ex-
tensive pineland habitat. Klimstra et al. (1974)
noted use of five habitat types in descending or-
der of preference: pinelands, hardwood hammock,
buttonwood-scrub mangrove, mangrove swamp,
and developed areas. Habitat selection varies with
season, time of day, and sex and age of the ani-
mal. Pinelands, hardwood hammocks, recent
clearings, roadsides, and grassy areas are used for
feeding. Hammocks and mangrove swamps are
used for cool retreats during the day (Klimstra
1979).
The pineland community on Big Pine Key is
open as a result of fire, with abundant plant spe-
cies including Dade County Pine {Pinus elliottii
var. densa), silver palm {Coccothrinax argentata),
stopper {Myrtus verrucosa), devil's claw (Pisonia
rotundata), and grasses.
Dominant species in the hammocks include
Spanish stopper {Eugenia foetida), maiden
bush {Savia bahamensis), poisonwood {Metopium
toxiferum), white indigo berry (Randia aculeata),
and darling plum (Reynosia septentrionalis).
FOOD AND FORAGING BEHAVIOR
Red mangrove {Rhizophora mangle) is an im-
portant food source, with 63% occurrence in pel-
let analysis (Dickson 1955). Klimstra et al. (1974)
list the following food plants: joeweed {Jacquinia
keyensis); white indigo berry; devil's claw; poison
ivy {Toxicodendron radicans); morinda {Morinda
royoc); catbrier {Smilax havanensis); blolly {Pi-
sonia discolor); ground cherry {Phy sails augusti-
folia); blazing star {Liatris tenuifolia);hegg,articks
{Bidens pilosa); capweed {Lippia nodiflora);
hempvine {Mikania batatifolia); false foxglove
{Agalinis spp.); borreria {Borreria); snowberry
{Chiococca pinetorum); saffron plum {Bumelia
celastrina); Christmas berry {Crossopetalum ilici-
folium); nightshade {Solanum blodgettii); vine
milkweed {Cynancfium blodgettii); white vine
{Sarcostemma clausa); mallow {Eustoma exalta-
tum); and saw palmetto {Serenoa repens). Dick-
son (1955) adds silver palm, acacia {Acacia pine-
torum), wild dilly {Manilkara bahamensis), and
brittle thatch palm {Thrinax microcarpa).
Klimstra et al. (1974) determined caloric
values of food plants used by key deer and noted
that many are equivalent in energy content to
commercial animal feeds, alfalfa, corn, wheat, and
barley.
Food plants change seasonally, probably re-
flecting availability and nutritional needs (Klim-
stra et al. 1974). Some plants are regularly
browsed, resulting in stunting and near-extirpa-
tion. Following fire in the pinelands, new growth
immediately attracted deer and extensive brows-
ing occurred for 6 to 9 months.
Virtually no plant species is immune from
deer use at one time or another (Klimstra et al.
1974).
SHELTER REQUIREMENTS
Open areas and subdivisions are favored bed-
ding sites at night, and mangrove swamps provide
cool retreats during the day (Klimstra 1979).
NESTING OR BEDDING
Not known.
3 3
A "- >. B A M A .
_ Ihoustom \
as 24 25
3 4 5 6
e 9 10 11 12 13 M 15 16 n IB 19 aO 21 23 23 2*
The present-day distribution of key deer is depicted on this map by shading.
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Adult males maintain home ranges of about
120 ha (and larger during the breeding season),
and adult females range over about 50 ha (Klim-
stra 1979).
POPULATION NUMBERS AND TRENDS
Dickson (1955) estimated the population at
25 to 80 individuals. Numbers have increased
since then to between 300 and 400 animals, and
the population is now believed stable (Klimstra
1979).
The official estimate of the National Key
Deer Wildlife Refuge is 600 animals, but this may
be re\i:5ed downward as monthly census data are
analyzed 'D. Kosin personal communication).
REPRODUCTION
Dickson (1955) stated that there was no spe-
cial breeding season, although Klimstra (1979)
observed a breeding peak in September and Octo-
ber. Territorial activity seems limited to defend-
ing a receptive doe from other bucks (Klimstra et
al. 1974). Dickson (1955) observed evidence of
fighting between adult bucks in February. Bucks
with full racks are generally the earliest breeders,
and exclude yearling males and those with lesser
racks.
The gestation period is 204 days, with peak of
fav^ming coinciding with the rainy season in April
and May (Dickson 1955, Khmstra 1979). Birth of
fawns occurs in all habitat types, but usually in
areas of open understory.
Key deer have a relatively low reproductive
rate, with an average of 1.08 fawns per adult doe
annually (Khmstra et al. 1974). Male fawns out-
number females, but the sex ratio changes until
adult females outnumber males by 2.38 to 1. Fif-
ty percent of male fawns survive 1.5 years, and
50% of female fawns survive 2.1 years (Klimstra
et al. 1974). Longevity records are 8 years for
males and 9 years for females.
Adult females form loose matriarchal groups
with one or two generations of offspring, while
bucks feed and bed together during the nonbreed-
ing season only (Klimstra 1979).
MANAGEMENT AND CONSERVATION
In 1954, the National Key Deer Wildlife Re-
fuge was established and now includes 1,764 ha,
of which 300 ha are leased. Since 1954, the deer
population has increased by more than 600%, and
is now considered to be stable.
Recommendations for management include
continuation of closed season, acquisition of addi-
tional habitat on Big Pine, No Name, and Cudjoe
Keys, controlled burning in pinelands, mainten-
ance of existing waterholes, fencing of islands
where refuge lands and subdivisions adjoin, popu-
lation and habitat monitoring, visitor manage-
ment, and additional research on all aspects of
key deer biology (Khmstra et al. 1974, Klimstra
1979).
AUTHORITIES
Donald J. Kosin, Refuge Manager
National Key Deer Wildlife Refuge
P.O. Box 510
Big Pine Key, FL 33043
W. D. Klimstra, Director
Cooperative Wildlife Research Laboratory
Southern Illinois University
Carbondale.IL 62901
PREPARER'S COMMENTS
Khmstra et al. (1974) noted that key deer
appear to be growing to a larger size in the last
few years, perhaps due to improved nutrition.
The tiixonomic status of the key deer has been
questioned and is now under investigation by
Klimstra and others.
LITERATURE CITED/SELECTED
REFERENCES
Alexander, T. R., and J. H. Dickson. 1970. Vege-
tational changes in the National Key Deer Re-
fuge. Q. J. Florida Acad. Sci. 33(2):81-89.
. 1972. Vegetational changes in the Na-
tional Key Deer Refuge -II. Q. J. Florida
Acad. Sci. 35(2):85-96.
Barbour, T., and G. M. Allen. 1922. The white-
tailed deer of eastern United States. J. Mam-
mal. 3(2):65-78.
Dickson, J. D., III. 1955. An ecological study of
the Key deer. Florida Game Fresh Water Fish
Comm., Tech. Bull. 3. 104 pp.
Dooley, A. 1975. Foods of the Florida key deer.
M. A. Thesis. So. IlHnois Univ., Carbondale.
80 pp.
Hardin, J. W. 1974. Behavior, socio-biology, and
reproductive life history of the Florida key
deer, Odocoileus virginianus clavium. Ph. D.
Dissert. So. Illinois Univ., Carbondale. 226
pp.
Harlow, R. F., and F. K. Jones, Jr. 1965. The
white-tailed deer in Florida. Florida Game
Fresh Water Fish Comm., Tech. Bull. 9. 240
pp.
Klimstra, W. D. 1968-1972. Key deer investiga-
tions annual progress reports to the U.S.
Dept. of Interior., Bur. Sport Fish. Wildl.,
Coop. Wildl. Res. Lab., So. Illinois Univ., Car-
bondale. Unpubl.
. 1979. Key deer. Pages 15-17 in]. N.
Layne, ed. Rare and endangered biota of
Florida, Vol. I, Mammals. Univ. Presses of
Florida, Gainesville.
Klimstra, W. D., J. W. Hardin, N. J. Silvy, B. W.
Jacobson, and V. A. Terpening. 1974. Key
deer investigations final report to the U.S.
Dept. of Interior, Bur. Sports Fish. Wildl., Co-
op. Wildl. Res. Lab., So. Illinois Univ., Car-
bondale. 184 pp. Unpubl.
Sokoloff, B., J. B. Reed, and R. Dutcher. 1949
(1950). Nutritive value of mangrove leaves
(Rhizophora mangle). Q.J. Florida Acad. Sci.
12(3):191-194.
U.S. Department of the Interior. 1973. Threatened
wildlife of the United States. Compiled by Of-
fice of Endangered Species and International
Activities, Bur. Sports Fish. Wildl. Resour.
Publ. 114. U.S. Govt. Printing Office, Wash-
ington, D.C. 289 pp.
GATEKEEPER
To be designated by the Office of Endangered
Species.
ACCOUNT PREPARED/UPDATED BY:
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, FL 32601
Biological Services Program
FWS/OBS-80/01.49
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
LAYSAN DUCK
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoasl of the Unitetl States. In-
formation about life history, distribution, requirements and c()nscr\'ation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Knginccrs in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed \.u:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Coinputcr Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.49
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
LAYSAN DUCK
A Cooperative Effort
by the
National Fish and Wildhfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT: DAVID B. MARSHAUL
LAYSAN DUCK
Anas laysanensis Rothschild
KINGDOM Animalia
CLASS Aves
ORDER Anseriformes
FAMILY Anatidae
OTHER COMMON
NAMES . . . .Laysan teal, Laysan Island duck
DATE
Entered into SWIS To be determined
Update To be determined
LEGAL STATUS
Federal: Endangered (42 FR 36425, 14 July
1977; 42 FR 10476, 22 Feb 1977.
States: Endangered: Hawaii
REASONS FOR CURRENT STATUS
Palmer found these ducks exceedingly tame
(Rothschild 1900). Other observers since have
commented on their tame behavior (Ely and
Clapp 1973). Weak flight allows capture by hand
after a flight of 125 yards (Wetmore in Delacour
1956). Their dependence on the lagoon increased
their vulnerability (Warner 1963). They are now
rare, after recovering from near extinction due to
denudation of vegetation by European rabbits
{Oryctolagus caniculus) introduced about 1903.
The last rabbits were eliminated in 1923, and
vegetation recovered. The duck population also
recovered, but has fluctuated considerably since.
By 1957, it was thought to be near its saturation
point; it then declined from unknown causes to
the point where it was close to extinction again
(King in press. Fish and Wildlife Service 1973.
Ely and Clapp 1973, Berger 1972, Warner 1963,
Ripley 1960). More recent counts have varied
from 69 in July 1974 to 287 in August 1976.
The last count w^as 240 in August 1977, so the
present status is relatively good (Eugene Kridler
personal communication).
Reasons for the rapid and extreme population
fluctuation in recent years are evidently unknown,
but the main one is probability inability to count
all birds present (Eugene Kridler pers. comm.).
Ely and Clapp (1973) say adults have no known
enemies on the island and destruction of eggs by
Laysan finches and young by frigatebirds is prob-
ably negligible. The limiting factor in the past and
now must have been the environment.
PRIORITY INDEX
33
DESCRIPTION
The Laysan Duck is a small, dark brown duck
similar to the Hawaiian duck, but smaller and
redder; head and neck are blackish, often
speckled irregularly wdth white almost around the
bill and on the face and chin; there is always a
white ring around the eyes. The speculum is green
and black, bordered with white posteriorly. The
bill is narrow. Females are more boldly marked
than males; the speculum is sooty brown, some-
times with a little green (Delacour 1956).
Size — Male: wingspan 192-210 mm, tail 100-
105 mm; culmen 39-40 mm; tarsus 37-39 mm;
Female: wingspan 190-196 mm; tail 90 mm,
culmen 38-39 mm; tarsus 35-38 mm (Delacour
1956).
The downy young are like those of the com-
mon mallard, but smaller and redder above and
cinnamon below; markings on upper parts are
indistinct (Delacour 1956).
Eggs are greenish white, 55 by 38 mm (Fisher
1903).
RANGE
Laysan Island (4 km ), in the northwestern or
leeward Hawaiian Islands, is the only known
range. According to von Kittlitz in Phillips
(1923), it also occured on adjoining Lisianski
Island in 1928, but this statement has never been
verified and is considered almost certainly er-
roneous (King in press, Ely and Clapp 1973,
Berger 1972). Although 525 individuals have been
banded since 1958, no natural occurrences away
from Laysan Island have been recorded (Ely and
Clapp 1973). The details of published records of
Laysan ducks on Laysan Island from 28 March
1928 to 9 September 1960 are given by Ely and
Clapp (1973).
RANGE MAP
Total Range (Ely and Clapp 1973).
STATES /COUNTIES
Hawaii Leeward Islands
HABITAT
All plant associations on Laysan Island are
used, but concentrations are found in low sedges
and vines around the central lagoon (Ely and
Clapp 1973, Warner 1963, E. Kridler in King in
press). Fisher (1903) and Bailey (1956) found
birds concentrated at a small brackish or fresh-
water pond near the southwest corner of the
lagoon, always a favorite spot until its disappear-
ance in the 1920's. They rarely occur on the
hypersaline and almost lifeless main lagoon and
almost never on the ocean (Warner 1963, Ely and
Clapp 1973). A. Wetmore in Ely and Clapp
(1973) noted that birds rested among rocks
during the day, and at night walked inland to
the lagoon margin and patches of vegetation
which probably provided food. More recently,
ducks have concentrated around slightly brackish
water available at several points following heavy
rains. They apparently survive long periods when
neither fresh nor slightly brackish water is avail-
able. The home range of each pair usually con-
tains a strip of lagoon shore (Woodside and
Kramer in Ely and Clapp 1973). Ducks occur
throughout the island, but most of them are
found in the beach morning glory near the
lagoon during the day (Ely and Clapp 1973).
They formerly used permanent freshwater
areas that are now filled with sand as a result of
the rabbits' destruction of the vegetation cover
(Warner 1963), complete except for three patches
of sesuvium that may have enabled the remnant
population of ducks to survive (Warner 1963).
Now that vegetation has recovered, the sesuvium
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is completely ignored. Present vegetation is dis-
tributed in zones away from the lagoon. The la-
goon flat at the water's edge is covered with alter-
nating strands of sedge {Sicyos sp.), heliotrope
[Heliotropium currasavicum) , and sesuvium (Se-
suvium portulacastrum) . The next zone is a knee-
deep mat of beach morning glory [Ipomoea pes-
caprae), which almost surrounds the lagoon.
Farther up the gradual slope above the water
table, morning glory is replaced by a dense belt of
bunch grass [Erogrostis variabilis), often mixed
with puncture vine {Tribulus cistoides) and boer-
haavia (Boerhaavia diffusa). Still farther up the
slope, the plant cover thins to scattered mats of
prostrate succulent boerhaavia interspersed with
bunch grass. Occasional clumps of scaevola
{Scaevola fructescens) are scattered about the
higher ridges of the island. Ducks use all of these
vegetation zones for feeding, although they are
restricted to the morning glory and bunch grass
interphase and boerhaavia zones when molting
and flightless (Warner 1963). Fresh or slightly
brackish water seeps are used for bathing when
available, but there are long periods when these
are not available. There is some evidence of salt
secretion by the nasal glands, and also that rain
water on bushes is used for bathing (Warner
1963).
FOOD AND FORAGING BEHAVIOR
The stomach of a male collected near a pond
was gorged with small flies resembling the com-
mon housefly (Fisher 1903). Warner (1963)
noted that sesuvium, so heavily utilized when it
was the major remaining vegetation on Laysan
during rabbit infestation, is now completely
ignored as food. Most feeding and other activity
begins at dusk and continues to about midnight
(Ely and Clapp 1973. Warner 1963). These ducks
are primarily insectivorous, at least during the
summer months (Ely and Clapp 1973). Warner
(1963) describes nocturnal feeding on cutworms,
which are larvae of a nocturnal moth {Agrostis
sp.), both from vegetation and the sand beneath,
and pursuit of brine flies along the lagoon edge.
He observed feeding on littoral invertebrates in
tide pools as well as on larvae and pupae of flies
and beetles in the sand around seabird carcasses.
He believed that plant materials were eaten, if at
all, only incidentally to the animal food which
was actively sought. He noted that ducks follow
favored routes for feeding activities. He deter-
mined the home range to be about 0.81 ha, and
much smaller during the summer flightless period
while molting.
SHELTER REQUIREMENTS
When approached, the birds usually walk into
vegetation, and less often, paddle into the lagoon
or take flight (Ely and Clapp 1973).
NESTING OR BEDDING
The nest is a down-lined depression under
vegetation (Ely and Clapp 1973). One nest was
placed under a thick chenopodium bush close to
a pond. The shallow bowl was a little over 12.7
cm in diameter, formed of long, dry sedge stems
(Fisher 1903).
RITUAL REQUIREMENTS
None described.
OTHER CRITICAL REQUIREMENTS
None known.
POPULATION NUMBERS AND TRENDS
Fisher (1903) estimated their numbers at
less than 100 in 1903. Dill and Bryan (1912)
could not be sure of more than 6 individuals
in 1911. Munter (1915) counted 13 on the fresh-
water pond in 1915. J. S. Palmer (m Phillips
1923) reported about 35 in 1918. A. Wetmore
(in Phillips 1923) left 14 in 1923. Brock (1951)
found 33 in 1950.
Never very common, they declined to mini-
mum between 1920 and 1930. A thorough
search in 1930 discovered only 1 female (Ely
and Clapp 1973). By 1957, the population had
recovered to an estimated 600 (Warner 1963).
Estimates fluctuated between 200 and 300 until
1969, then dechned to only 75 birds in 1970
(Fish and Wildlife Service, Ely and Clapp, Berger
1972; King in press). In 1972, the estimated
number was 175, but in 1973, they were down
to less than 40 (E. Kridler 1974 in King in press).
More recent counts are: July 1974—69, August
1975-251, May 1976-100, August 1976-287,
August 1977— 240, (Eugene Kridler pers.comm.).
The extreme variation in counts even during the
same year shows the difficulty of observing all
birds present, probably due to the secretiveness
and nocturnal habits of the species noted by
Warner (1963).
There is a captive population of over 150 in
zoos and private collections (King in press).
REPRODUCTION
H. M. Smith found 7 downy young in broods
of 3 and 4 in 1950 (Brock 1951). Fisher (1903)
found 6 eggs in a nest. The species breeds readily
in captivity (Ripley 1960).
The nesting cycle is evidently an extended
one. Pairing is first observed in early March and
continues until mid-summer. There seems to be
no published report of courtship behavior (Ely
and Clapp 1973), Warner 1963). Usual clutch
size is 5 or 6. The earliest recorded hatching date
is 19 May (Fisher 1903). Most eggs are laid in
May, but the season extends through July.
Incubation takes 27 to 28 days, The drake
guards the incubating female. The chicks remain
in the nest with the female for 2 days. In the
wild, males sometimes tend the brood (Warner
1963, Ely and Clapp 1973).
High chick mortality, which reduced brood
size from an average of 3 at hatching to about 1
or less at 1 month, strongly suggests that this
may be a limit on population growth (Warner
1963).
Warner (1963) considered destruction of
eggs by Laysan finches and young by frigate-
birds a possible but unproved limiting factor on
population; Ely and Clapp (1973) thought it to
be probably negligible.
MANAGEMENT AND CONSERVATION
The entire range (Laysan Island) is within
the Hawaiian Islands National Wildlife Refuge.
Landings on the island are restricted by permit.
The species is protected by Federal and State
law, and Listed in Appendix 1 of the 1973 Con-
vention on International Trade in Endangered
Species of Wild Fauna and Flora. A census is
conducted periodically by the U.S. Fish and
Wildlife Service. Conservation measures pro-
posed by King (in press) are:
1. Increase patrols of Laysan Island to
prevent unauthorized landings and to prevent
introduction of additional pest plants, in-
sects, and predators such as dogs, cats, and
rats.
2. Introduce captive-reared Laysan ducks
to another Pacific island with suitable habi-
tat (Fish and Wildlife Service, King in press).
Introduction attempts include a release of
24 or 25 in the Kewalo marsh on the outskirts
of Honolulu in 1894 by a Mr. Whitney, but none
have been taken since (Ely and Clapp 1973). In
March 1968, 12 were transported from Laysan
to Southeast Island, Pearl and Hermes Reef, by
Fish and Wildlife Service personnel. All disap-
peared soon after introduction (Ely and Clapp
1973, E. Kridler 1974 in King in press, Fish and
Wildlife Service 1973). Beginning with 44 cap-
tured birds, rearing in captivity has been very
succesful in the Hawaii zoo and numerous main-
land aviaries (Warner 1963, Ripley 1960). A re-
covery team has been appointed and a recovery
plan for the Laysan duck is now in progress.
AUTHORITIES
Eugene Kridler
835 Akumu Street
Kailua, Hawaii 96743
Richard Warner
Building 737 IFAS
University of Florida
Gainesville, Florida 32611
PREPARER'S COMMENTS
The great variation in population estimates
at different times, even during the same year, is
evidence of the inadequacy of present inventory
methods. Also, the almost complete lack of in-
formation on the factors that limit the population
indicates a definite need for additional research
on both of these problems in particular.
Introduction of predatory or competing ani-
mals, particularly rats, on Laysan Island as a
result of unauthorized or unknown landings re-
mains the greatest threat to the continued sur-
vival of the duck, so the sooner another popu-
lation can be established on another island, the
safer it will be from such accidents.— John W.
Aldrich.
LITERATURE CITED/SELECTED
REFERENCES
Bailey, A. M. 1942. The portuluca flats of Lay-
san. Audubon Mag. 44:150-161.
. 1956. Birds of Midway and Laysan Is-
lands. Denver Mus. Nat. His., Mus. Pictorial
12:1-130 (80-38).
Berger, A. J. 1972. Hawaiian birdlife. Univ.
Press of Hawaii, Honolulu.
Brock, V. E. 1951. Some observations of the
Laysan Duck, Anas wyvilliana laysensis. Auk
68:371-372.
Delacour, J. 1956. The waterfowl of the World,
Vol. 2., Country Life Ltd., London, p. 49.
Dill, H. R., and W. A. Bryan. 1912. Report of an
expedition to Laysan Island in 1911. U.S.
Dep. Agric. Biol. Surv. Bull. 42. 30 pp.
Ely, C. A., and R. B. Clapp. 1973. The natural
history of Laysan Island, northwestern Ha-
waiian Islands. Atoll Res. Bull. 177. pp.
Fish and Wildlife Service. 1973. Threatened wild-
life of the United States. U.S. Bur. Sport
Fish. Wildl. Resource Publ. 114. pp.
Fisher, W. K. 1903. Notes on the birds peculiar
to Laysan Island, Hawaiian group. Auk 20:
384-397.
Greenway, J. C, Jr. 1958. Extinct and vanishing
birds of the World. Spec. Pub. 13, Amer.
Commit, for Internat. Wildl. Protect.
New York. pp. 167-168.
King, W. in press. (Revised). Red data book,
Aves. Survival Service, lUCN, Morges, Switz-
erland.
Lint, K. D. 1960. Preserving the Laysan Teal.
Zoonooz 33(8):6-7.
Munter, W. H. 1915. Report of destruction of
bird life on Laysan Island. Ann. Rep. Coast
Guard for 1915:130-140.
Phillips, J. C. 1923. Natural history of the ducks.
Vol. 2. Houghton Mifflin Co., New York,
pp. 51-52.
Ripley, S. D. 1960. Laysan Teal in captivity.
Wilson Bull. 72:244-247.
Rothschild, W. 1900. The avifauna of Laysan
and neighboring islands; with a complete his-
tory to date of the birds of the Hawaiian
possessions. Pt. 3. R. H. Porter, London.
Warner, R. E. 1963. Recent history and ecology
of the Laysan Duck. Condor 65:3-23.
Wetmore, A. 1925. Bird life among lava rock
and coral sand. Nat. Geog. Mag. 48:77-108.
Biological Services Program
FWS/OBS-80/01.50
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
RED HILLS SALAMANDER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this scries of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the F.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Kngincers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems 1 eam
U.S. Fish and Wildlife Service
iNASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.58
u
FWS/OBS-80/01.50
March 1980
/
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
RED HILLS SALAMANDER
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
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CREDIT: R.H. MOUNT
RED HILLS SALAMANDER
Phaeoffiatbus hubrichti High ton
KINGDOM Animalia
CLASS Amphibia
ORDER Caudata
FAMILY Plethodontidae
OTHER COMMON
NAMES Hubricht's salamander,
Alabama red hills salamander.
DATE
Entered into SWIS to be determined
Updates 23 July 1977,31 March 1978
LEGAL STATUS
Federal: Threatened (41 FR 53032-53034,
3 December 1976).
States: None.
REASONS FOR CURRENT STATUS
Timber harvesting by clearcutting, site prepa-
ration, and conversion of mixed hardwood stands
to pine plantations have been shown to cause lo-
calized extermination of red hills salamanders
(French 1976). The amount of suitable habitat is
steadily shrinking due to these timber manage-
ment practices (Jordan and Mount 1975).
Overcollection by amateur and professional
herpetologists has reduced populations in some
areas (41 FR 53032-53034, 3 December 1976).
Natural low fertility (Brandon 1965) inhibits
recovery of decimated populations.
PRIORITY INDEX
Not assigned.
DESCRIPTION
The red hills salamander is solid dark brown
to black, with no pattern. Palms, soles, and snout
are somewhat paler. Adults are 80 to 119 mm in
snout-to-vent length (total length up to 256 mm);
the body is elongate with 20 to 22 costal grooves
and proportionately small limbs. The eyes are
protuberant.
RANGE
The salamander is known only from south
central Alabama between the Conecuh and Ala-
bama Rivers, where it is restricted to the Talla-
hatta and Hatchetigbee geological formations. Its
known distribution includes 22,213 ha (French
1976). An additional 1,485 ha contiguous with
the present range are believed to have supported
red hills salamanders prior to timber harvesting
and conversion to pine within the last decade
(French 1976).
RANGE MAP
The total known distribution is indicated by
shading.
STATES/COUNTIES
Alabama Butler, Conecuh, Covington, Cren-
shaw, Monroe.
HABITAT
The species lives in burrows on the slopes of
cool, moist ravines shaded by an overstory of
mixed hardwood trees. It does not occur in pine
forests.
FOOD AND FORAGING BEHAVIOR
The salamander feeds at or very near the
mouth of its burrow at night, especially for the
first hour or two after sunset. Prey items include
spiders and small insects that are caught with its
sticky tongue.
SHELTER REQUIREMENTS
This species requires shaded, moist ravines
with ground litter and a friable soil for burrow-
ing. Displaced individuals apparently do not make
(or find) new burrows, and are thus permanently
eliminated from the population (R. Jordan per-
sonal communication). Burrows are apparently
not started from the surface, but only by sub-
surface branching off from other burrows.
NESTING OR BEDDING
Nesting requirements are unknown. A cap-
tive female laid a clutch of infertile eggs, attached
to the underside of a piece of bark (R. Mount per-
sonal communication).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
The very specific habitat of this species may
be its only environmental requirement.
POPULATION NUMBERS AND TRENDS
The species was discovered in 1960. No popu-
lation estimates are available. Most of the suitable
habitat is owmed by timber companies who have
been harvesting, preparing sites, and converting
the mixed hardwood forests to pine plantations.
French (1976) estimates that at least 6% of its
range has been made unsuitable by these practices
within the past 10 years.
REPRODUCTION
Little is known, but the presence of only a
few large ovarian ova in adult females suggests
that direct development occurs within the egg,
without a free-living larval stage (Brandon 1965).
MANAGEMENT AND CONSERVATION
Within the range of the red hills salamander,
most timber companies are now 'marking out' the
ravines and steep slopes and harvesting only the
largest trees (French 1976). One company has
issued a policy statement indicating its intention
to protect salamander habitat (R. Mount personal
communication) .
AUTHORITIES
Thomas W. French
Department of Life Sciences
Indiana State University
Terre Haute, IN 47809
Ralph Jordan, Jr.
Tennessee Valley Authority
Regional Heritage Program
Division of Forestry
Fisheries and Wildlife Development
Norris,TN 37828
Robert H. Mount
Department of Zoology-Entomology
Auburn University
Auburn, AL 36830
Terry D. Schwaner
Museum of Natural History
University of Kansas
Lavinrence.KS 66044
PREPARER'S COMMENTS
Although much remains to be learned about
this salamander, the knowledge required to pre-
vent threats to its existence is apparently available.
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LITERATURE CITED/SELECTED
REFERENCES
Brandon, R. A. 1965. Morphological variation
and ecology of the salamander Phaeognathus
hubrichti. Copeia 1965(1):67-71.
. 1966. Phaeognathus and P. hubrichti.
Pages 26.1-26.2 in Cat. Am. Amphib. Rep-
tiles. Am. Soc. Ichtyol. Herpetol.
French, T. W. 1976. Report on the status and
future of the red hills salamander, Phaeog-
nathus hubrichti. Office of Endangered Spe-
cies, U.S. Fish Wildl. Serv., Washington,
D.C.
French, T. W., and R. H. Mount. 1978. Current
status of the red hills salamander, Phaeog-
nathus hubrichti Highton, and factors affect-
ing its distribution. J. Ala. Acad. Sci. 49:172-
179.
Highton, R. 1961. A new genus of lungless sala-
mander from the Coastal Plain of Alabama.
Copeia 1962(l):65-58.
Jordan, J. R. 1973. Sexual dimorphism in the red
hills sa\2imai\dcx , Phaeognathus hubrichti High-
ton (Amphibia: caudata: plethodontidae),
with comments on its phylogenetic assign-
ment. Assoc. S. E. Biol. Bull. 20(2)-62.
. 1975. Observations on the natural his-
tory and ecology of the red hills salamander,
Phaeognathus hubrichti Highton (caudata:
plethodontidae). M.S. Thesis, Auburn Univ.,
Auburn, Ala. 53 pp.
Jordan, J. R., and R. H. Mount. 1975. The status
of the red hills salamander, Phaeognathus
hubrichti. J. Herpetol. 9:211-215.
Mount, R. H. 1975. The reptiles and amphibians
of Alabama. Auburn Printing Co., Auburn,
Ala. 347 pp.
Mount, R. H., and T. D. Schwaner. 1970. A tech-
nique for collecting the plethodontid salaman-
der Phaeognathus hubrichti. Copeia (1):205-
206.
Schwaner, T. C. 1970. The Alabama red hills sala-
mander. Alabama Conserv. 1970:7.
Schwaner, T. D., and R. H. Mount. 1970. Notes
on the distribution, habits, and ecology of the
salamander Phaeognathus hubrichti Highton.
Copeia 1970(3):571-573.
Valentine, B. D. 1963a. The plethodontid sala-
mander Phaeognathus: external morphology
and zoogeography. Proc. Biol. Soc. Washing-
ton 76:153-158.
1963b. The plethodontid salamander
Phaeognathus: collecting techniques and ha-
bits. J. Ohio Herpetol. Soc. 4:49-54.
Biological Services Program
FWS/OBS-80/01.51
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
ARCTIC PEREGRINE FALCON
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoasl of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the l",ndangered Species .\ct of 1973, as diiniulcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of Endangered Species and Biok)gical Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell. Louisiana 704.'')8
u
FWS/OBS-80/01.51
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
ARCTIC PEREGRINE FALCON
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
PHOTO OF CLOSELY RELATED SUBSPECIES
CREDIT: KENNETH FINK
ARCTIC PEREGRINE FALCON
Falco peregrinus tundrius White (1968)
KINGDOM Animalia
CLASS Aves
ORDER Falconiformes
FAMILY Falconidae
OTHER COMMON
NAMES Duck Hawk (AOU 1931);
Beach Peregrine, Blond Peregrine (fal-
coner's names, White 1968); Tundra Pere-
grine (Ruos 1970 and numerous authors).
DATE
Entered into SWIS To be determined
Updates To be determined
LEGAL STATUS
Federal: Endangered: (42 FR 36425, 14 July
1977 p. 36425). Protected, Migratory
Bird Treaty Act of 3 July 1918 (40
Stat. 755; 16 U.S.C. 703-711) as
amended 3 Dec. 1969. Public Law 91-
135. Listed, Appendix I, Convention
on International Trade in Endangered
Species of Wild Fauna and Flora (42
FR 10476, 22 Feb. 1977).
States: Endangered: Alaska, South Carolina,
Texas - Listed: Florida.
REASONS FOR CURRENT STATUS
Up to 1960, no long-term data are available
by which to detect either an increase or decrease
in the total population of peregrines of northern
Canada. In general, up to that time, it had been
observed as a common breeding bird in an exten-
sive area (Fyfe 1969).
In 1969, J. L. Ruos (pers. comm.) estimated
the annual harvest of tundra-breeding peregrines
(taken by falconers, mostly during migration) at
between 60 and 120 individuals, over 90% of
which were first-year birds. The effect of this
former harvest on the population is unknown, but
it has been prohibited under present migratory
bird regulations in the United States, where most
of it occurred.
Earlier investigation of the Alaskan popula-
tion (Enderson et al. 1968; Cade et al. 1968) and
the Canadian population (Enderson and Berger
1968) indicated that members of this subspecies
were reproducing normally. However, Cade et al.
(1968) found that residues of organochlorines in
tissues and eggs appeared to be near the threshold
at which abnormal reproductive effects begin.
More recent studies by Cade and Fyfe (1970),
Berger et al. (1970), and Cade et al. (1971) indi-
cated that shell thinning was pronounced in pere-
grines from the northwest territories, the Ungava
region, and Alaska. The thinning approached or
exceeded 20%, the level at which reproductive
failures seem to begin. In addition, there was evi-
dence that F. p. tundrius populations of arctic
Canada and Alaska had declined suddenly. No
other reason for this decline than the effect of
pesticide poisoning on reproduction is suggested
(Cade and Fyfe 1970:235).
PRIORITY INDEX
10
DESCRIPTION
The Arctic peregrine falcon is a medium-
sized, pointed winged, swift flying bird of prey
with prominent black and white facial markings;
adults have slate gray back and white underparts
streaked and barred with black. Immatures are
brownish above and more heavily streaked below
than adults. They are similar to American pere-
grine falcons (anatum), but smaller and in all
plumages more lightly marked with paler browns
or bluish grays and less ventral rufous wash. Black
facial markings are more restricted (White 1968).
Distinction between tundris and anatum in
adult plumage is best made by facial markings
(width of black between eye and white auriculcir,
and width of malar stripe at base) and white fore-
head. The rufous basal portion of nape feathers,
which is whiter ventrally and somewhat paler dor-
sally, are also useful characters. Immature plum-
aged tundrius is distinguished from anatum by
facial characters, including a complete white
superciliary stripe, with darker stripe running
posterior to the eye usually present; also a median
pair of tail feathers conspiciously barred with
"vinaceous buff" (M. Ralph Browning ms. 1972).
Measurements (all in mm unless otherwise
indicated; values in parentheses are averages). -
Adult male (64 specimens): wing (chord) 292-330;
tail, 134-154 (140.5); tarsus 40-50 (44.3); bHl
without cere 15-20 (18.7); bUl with cere 22-25
(24.0); weight (12 breeding specimens), 550-647
(610.9) g. Adult female (62 specimens); wing
331-368 (351.8); tail 138-180 (167.8); tarsus 42-
57 (49.8); biU without cere 21-24 (22.7); bill with
cere 26-30 (27.8); weight (19 breeding specimens)
825-1094 g (952.9 g). Immature male (27 speci-
mens): wing 295-319 (311.1); tail 135-162
(151.8); tarsus 38-50 (44.0); bill without cere 17-
19 (18.3); bill with cere 22-25 (22.7); weight (4
specimens, fuUy fledged) 477-662 (570.0). Im-
mature female (30 specimens): wing 320-367
(349.6); tail, 155-189 (175.6); tarsus 44-(?)55
(49.1); bill without cere 19-24 (21.4); bill with
cere 23-28 (26.4); weight (3 specimens, fully
fledged) 844-925 g (889.0 g) (White 1968).
RANGE
F. p. tundrius breeds in tundra areas through-
out Arctic Alaska, Canada, and western Green-
land, from the northern tip of the Mackenzie Dis-
trict south and east across the Arctic tundra to
Hudson Bay, along the north coast of Ungava and
north into the Arctic islands as far as northern
Baffin Island and Melville Island; wesward as far
as Cape Prince of Wales, Alaska, and eastward to
the western coast of Greenland. The subspecies
intergrades morphologically with anatum in open
boreal forest or taiga areas of Alaska and Canada
(Fyfe 1969; White 1968).
These peregrines migrate largely along the
Atlantic coast and to some extent through the in-
terior of the continent. The Atlantic coastal flight
includes some birds from western Greenland.
They winter north (sparingly) from at least Cape
Sable, Florida and Cameron Bayou Louisiana
(probably along entire gulf coast) and Baja Cali-
fornia (occasionally); south through West Indies,
Central and South America to 40° S in Chile and
35° S in Argentina (White 1968; Rice in Hickey
ed 1969; Shor 1970a; Mueller and Berger 1961;
Hofslud 1966;Enderson 1965, 1969).
Ward and Berry (1972), analyzing records of
sightings and bandings of fall migrants on the
outer beaches of Assateague Island, Maryland,
from 1939-1971, determined that the proportion
of immatures in different years was 81% to 91%.
Of birds trapped for banding in those years, only
8% of the adults and 30% of the immatures were
males. A decline in total numbers during the
study period was indicated (Ward and Berry
1972). Shor (1970) points out that data on pro-
duction indicate that this great preponderance of
immatures is impossible unless we assume that
about 90% of adult birds are not seen in migra-
tion. It was assumed that most of the birds re-
corded were Arctic Peregrines, the majority pos-
sibly from Greenland breeding localities, as in-
dicated by Shore (1970).
Migrants appear in numbers on an island off
Kenai Peninsula, Alaska around 1 September. It
is not known where they go from there. Some in-
dividuals, almost certainly from the Arctic migrant
population, have been taken in July and August
in mid-latitudes of the United States (White
1969).
Mueller and Berger (1961) recorded 150 pere-
grines in fall migration along the west shore of
Lake Michigan from 1952 to 1957. They were
seen with concentrations of hawks of various
species along that migration route, and their
abundance at any one time was correlated with
weather conditions that caused updrafts of air.
RANGE MAP
The Breeding range (Fyfe et al. 1976) and
migration range are shown on the following map.
STATES/COUNTIES (within main migration
routes):
Barnstable, Nantucket, Dukes
Washington
Suffolk, Nassau
Monmouth, Ocean, Atlantic,
Cape May
Delaware: Sussex
Maryland:
Virginia:
North Carolina:
South Carolina:
Georgia:
Florida:
Minnesota:
Wisconsin:
Illinois:
Texas:
Massachusetts:
Rhode Island:
New York:
New Jersey:
Worcester
Accomack, Northampton
Currituck, Dane, Hyde, Carteret,
Onslow, Pender, Hanover,
Brunswick
Horry, Georgetown, Charleston,
Colleton, Beaufort
Chatham, Liberty, Mcintosh,
Glynn, Camden
Nassau, Duval, St. Johns, Flag-
ler, Volusia, Brevard, Indian
River, St. Lucie, Martin, Palm
Beach, Broward, Dade, Collier
St. Louis
Douglas, Door, Kewaunee, Man-
itowoc, Sheboygan, Ozaukee,
Milwaukee, Racine, Kenosha
Lake
Jefferson, Chambers, Galveston,
Brazoria, Matagorda, Calhoun,
Aransas, Nueces, Kleberg,
Kenedy, Willacy, Cameron
HABITAT
Peregrines prefer cliff ledges for nesting sites,
but in the absence of these, will nest on cutbanks
along rivers or coastal areas, on elevated landforms
known as dykes, and occasionally on low mounds
or even boulders. In nearly all instances, nests are
located in the immediate vicinity of a body of
water, either salt or fresh. If cliffs are not close to
water or if no food species are available, nesting
peregrines will be scarce or absent (Fyfe 1969). In
the case of joint occupancy of bluffs by peregrines
and gyrfalcons along the Colville River in Alaska,
peregrine nests were usually at the brink of a
slope or on a bluff to which a person could walk,
whereas gyrfalcon nests were on ledges on vertical
faces of cliffs, usually under an overhang, and ac-
cesible only by rope (White 1969).
In arctic Canada, these falcons nest on cliffs
and cut banks of rivers if food is available nearby.
In the central Canadian barrens near Cotwoyto
Lake, the landforms known as dykes provide
rocky formations elevated over the surrounding
countryside that are suitable for nesting. Where
land relief is slight in the central barrens (tundra)
of Canada, peregrines nest on boulders and hum-
mocks as little as 1 m above the surrounding area.
Breeding range and migration routes of the Arctic peregrine falcon
Rivers are important in the falcon's environ-
ment; they create the nesting cliffs by erosion and
provide bathing facilities, especially on gravel
bars; rivers also provide habitat for shorebirds and
v^aterfowl for the falcon's food. Peregrines also
hunt extensively over the surrounding country,
particularly taking ptarmigan and longspurs (Cade
1960).
They seldom nest on the cliffs along the Bering
Sea coast of mainland Alaska or in the mountains
above 800 m. A cliff of some sort is the most im-
portant feature of the nesting habitat (Cade 1960).
FOOD AND FORAGING
Food would not be a limiting factor in most
of northern Canada. Throughout that area, pas-
serines, shorebirds and waterfowl are abundant
in summer, with the exception of the Arctic
desert areas or relatively barren mountain tops on
Baffin Island (Fyfe 1969; Cade 1954).
Bumham and Maddox (m Fyfe et al. 1976)
reported that four species of small passerine birds
made up over 90% of the food of peregrines in
western Greenland. J. N. Rice and R. B. Berry
(ms. 1970) also found that, with the exception of
a single ptarmigan, the peregrines nesting at
Ungava Bay in 1970 fed entirely on small pas-
serines, primarily horned larks, snow buntings,
and water pipits. Cade (1960), however, reported
that Arctic peregrines take a wide variety of avian
prey (waterfowl, shorebirds, and passerines), with
no single species predominating except possibly
ptarmigan. In his opinion, peregrines probably
feed heavily on lemmings during their cyclical
abundance, but that the availability of lemming
does not affect the peregrine population.
SHELTER REQUIREMENTS
None
NESTING OR BEDDING
In Arctic Alaska and Canada, nests are scrapes
in either earth or gravel, on bare grass, rocky cliffs,
cutbanks, dykes, boulders, or hummocks. Three
nests in old rough-legged hawks nests had eggs
laid on bare sticks with no soft material. Five
nests in old rough-legged hawk nests had eggs
and other plant material (Hohn 1955, McEwen
1957;Cade (1960); Fyfe 1969).
Cliff nests are typical for the species and are
situated on a ledge or in a hole, often under a pro-
tecting overhang. Nests on boulders and hum-
mocks have little or no protection from above
(Fyfe 1969). All nesting sites known in Alaska
were on cliffs on islands or along the coast or
river bluffs (Cade 1960). Of 57 eyries along the
Colville River, height of the nests above the river
ranged from 3.2 m to 96 m (average 35 m); the
distance below the brink of the cliff was 0 to 90
(average 14) m. Fifteen of 57 eyries were located
on shale formations, 17 on sandstone or conglo-
merate, and 27 on earth or talus banks. Eleven
were in old rough-legged hawk nests. The brinks
of the cliffs are usually overgrown with dense
thickets of alder and willow scrub (Cade 1960).
All nests found by Burnham and Mattox (in Fyfe
et al. 1976) in western Greenland were located
on rocky cliffs varying in height from 25 to 120
m. All nests found in Alaska were on cliffs, mostly
along rivers (Cade 1960).
RITUAL REQUIREMENTS
The complex courtship and mating ritual is
described under Reproduction. A strong pair
bond is essential to the reproductive success of
the species (Cade 1960).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Suitable nesting sites near an adequate food
supply and freedom from organochlorine chemi-
cals in the environment are the most critical re-
quirements.
POPULATION NUMBERS AND TRENDS
White (1969) said the number of breeding
adults on the Arctic Slope of Alaska seems to be
maintaining itself and there was no evidence of a
decrease in reproductive capacity.
Population density was approximately one
pair per 52 km^ of suitable habitat in the Bathurst
Inlet area, Canada, and about one pair per 259
km^ in areas of limited nesting habitat. In esti-
mating 7,500 breeding pairs in northern Canada
(Fyfe 1969, J. H. Enderson and D. D. Berger in
Fyfe 1969), a sharp decline of nesting peregrines
was found along the Mackenzie River and at
Campbell Lake in 1966.
Enderson (1969) calculated the mortality rate
of peregrines on the basis of band recovery (pre-
sumably mostly tundrius) as 70% for immatures
and an annual rate of 25% for adults. He assumed
that the 70% immature mortality rate is indicative
of a declining population.
Rice (1969) calculated from peregrines trapped
for banding on the beaches of the Maryland-
Virginia coast that immatures constituted 83% of
the population in the 1954-1959 period and
84% in the 1960-65 period. The 1954-59 sample
included 2 adult males, 37 adult females, 50 im-
mature males and 144 immature females. The
1960-65 sample included 4 adult males, 26 adult
females, 56 immature males and 116 immature
females. Cade (1960) estimated 200 to 250 breed-
ing pairs in Arctic Alaska.
Cade (1960) suggests that gyrfalcons can suc-
cessfully prevent peregrines from occupying the
larger river cliffs and thus affect their density and
distribution.
In the Colville River area of about 36,800
km^ on the Arctic slope of Alaska, the popula-
tion of peregrines appears to have held rather
constant in the number of breeding pairs until
about 1970-71, and then started a severe drop. It
may now have leveled off at about 30% to 35% of
its former numbers, as suggested by data from 1973
and 1975. Total numbers of pairs or single adults
were as foUows: 1952-1959, 32-36; 1967, 27;
1968,32;1969,33; 1971, 25; 1973, 14; 1975, 13.
Based on early 1950 studies by Cade, the total
population of the Colville drainage may have been
between 120 and 160 pairs (White and Cade 1975;
Peakalletal. 1976).
In the Sagavanirktok River area of about
2400 km^ on the Arctic Slope of Alaska, the
population followed a similar pattern, but appears
to have started to decrease earlier and declined
slightly more. Total numbers of pairs or single
adults were as follows: pre-1950, 11; 1958, 5;
1963, 5; 1970, 3; 1972, 2; 1973, 2; 1974,5; 1975,
3 (White and Cade 1975).
Fyfe et al. (1976) note that limited data for
other parts of Arctic Alaska indicate a population
decline similar to the well-documented decline
for the Colville drainage, and that it is doubtful
that as many as 50 pairs are still producing young
in northern Alaska where Cade (1960) estimated
the breeding population at 200 to 300 pairs in the
late 1950's.
In the Canadian Arctic, most populations
have dechned to 50% or less of their historically
known size. On Ungave Bay, the Interior Barrens,
Central Arctic Coast, Banks Island, and the North
Slope of Canada where historical records exist.
only 41% of known nesting sites were occupied in
1975. Occupancy of newly found nests declined
60% from 20 in 1973 to 8 in 1975, indicating an
accelerated decline of Canadian Arctic popula-
tions since 1973.
In western Greenland, the observations of
Bumham and Mattox (in Fyfe et al. 1975) re-
vealed a substantial density of one peregrine per
200 km^
REPRODUCTION
In the Colville River area (36,800 km^) on
the Arctic Slope of Alaska, production of young
may have started to fall off as early as 1965 or
1966. The numbers of young produced in 1952-
1959 was 40 to 50; in 1967, 34; in 1968, 34; in
1969, 26; in 1971, 14;andin 1973,9 (White and
Cade 1975; Peakal et al. 1975). The DDE residue
in eggs in that area through 1971 averaged about
190 parts per million dry weight (Peakall et al.
1975) and egg shells had thinned to about 23%
(White and Cade 1975).
Burnham and Mattox (m Fyfe et al. 1976), in
an approximately 1800 km^ area of western
Greenland, found a productivity of peregrines
averaging nearly 2.5 young per pair over a 4-year
period. DDE residues of 332 parts per million
found in addled eggs and egg-shell thinning of
14% indicate incipient problems for that popula-
tion. Eleven peregrine egg shells taken from Arc-
tic Ungava in 1967 ranged in thickness between
0.25 and 0.33 mm (mean thickness 0.291 mm),
representing a 24.4% drop from the 0.385 mean
of 30 eggshells measured by Dan Anderson from
"eastern Arctic" eggs collected during the first
third of the century. In ten eggs checked, DDE
ranged from 137 to 498 ppm on a fat basis (D. D.
Berger pers. comm. 1970).
Cade (1960) suggests that when peregrines
and gyrfalcons are in direct competition, the gyr-
falcon is the dominant competitor. On the Ander-
son River, Fyfe (1969) found the gyrfalcon to be
dominant and believed that where the two species
nest during the same season, the gyrfalcon may
displace the peregrine from the better nesting
sites such as cliff locations, relegating the peregrine
to dirt cutbanks. Natural hazards to nesting are
late spring storms or excessive erosion. Because of
the short nesting season, renesting is usually not
possible (Fyfe 1969). A significant correlation of
immature peregrine migration counts along Atlan-
tic coastal beaches in October with mean daily
minimum July temperatures in eastern Arctic
Canada indicate that minimum temperature on
the breeding grounds will cause a 31.6% (1-bird)
change in numbers of migrant immature birds
identified along the Atlantic coast per party-day
(Ruos 1970).
The Arctic peregrine is, in effect, reproduc-
tively isolated from the more southern taiga pop-
ulation by its gonad cycle. The entire courtship
cycle of tundrius and thus the factors regulating
pair bonding is at variance with more southern
breeding peregrines (Cade 1960; White 1968).
Peregrines apparently become established
along the lower ColviUe river between 4 and 10
May, with peak arrivals after the middle of May.
Falcon arrival is synchronized with the arrival
of their chief prey species. The incubation pe-
riod probably averages about 29 days for each
egg. Since there are intervals of 2 days between
egg layings and incubation begins with laying of
the first egg in the Arctic, there may be as much
as a week's difference in the ages of the youngest
and oldest nestlings. On the Colville, there was
one instance of laying in the last week of May, 12
instances in the first week of June, and 6 instances
in the second week of June. The total range for
any given event in the breeding cycle is barely
more than 3 weeks. Since the total period of the
breeding cycle from laying to the beginning of
independent existence of the young is not less
than 95 days in Arctic, peregrines must begin to
lay eggs not later than the third week in June to
ensure successful rearing of young before the
food is gone. The average clutch of Arctic birds is
3 eggs (54 samples), whereas the mean for the
U.S. is 3.74 (299 samples)showing a decreasing
size of clutch northward. Average fledging success
on the Colville River, Alaska was 1 .4 young per
nest. Predators on eggs and young, in addition to
man, are a negligible factor in the Arctic. Fairly
certain predators are the timber wolf, red fox,
arctic ground squirrel, and golden eagle (Cade
1960).
Preincubation or courtship behavior divided
into 8 parts: (1) attraction of mates to each other,
(2) mutual roosting on cliffs, (3) cooperative
hunting excursions, (4) courtship flights, (5)
"familiarities" on cliff, (6) courtship feeding, (7)
copulation, (8) nest scraping. Females are dom-
inant over males and successful mating depends
on the males' ability to adjust to that situation. In-
cubation and brooding is done mostly by the fe-
male, who sits more closely than females in more
southern latitudes. Food is brought by the male,
and even during the early fledging period, the
male continues to transfer food to the female for
feeding young. When young are about 3 weeks
old, the female begins to hunt actively again
(Cade 1960). Peregrines are definitely territorial
during the breeding season and defense of terri-
tory, which is roughly 3.22 km in diameter, has a
graded intensity from less intense at the outer
limits, where defense is only over food, to most
intense near the nesting cliff, which is always de-
fended against all other predatory birds, although
rough-legged hawks are sometimes allowed to nest
on the same cliffs within 100 yards (Cade 1960).
First-year falcons not engaged in breeding
may pass their first summer within or at the peri-
phery of their breeding range, but are not obvious
because they are not associated with nesting sites.
In 1967 one first-year female was seen on the
Colville River (White 1969).
Tradition is important in the occupancy of a
particular cliff. Once vacant, it may take a long
time for another pair to reoccupy it and establish
a new tradition. New occupancy is probably the
result of population pressure as well as attractive-
ness of the site (Cade 1960).
MANAGEMENT AND CONSERVATION
The most important actions that might be
taken in behalf of the Arctic peregrine would
probably be to designate for special protection
against land-use exploitation the areas where eyries
are concentrated in Arctic Alaska, Canada, and
western Greenland; as well as major migration
concentration areas along the Atlantic and Gulf
coasts and the shores of the western Great Lakes;
and to conduct an intensive educational and law-
enforcement program to reduce the use of chlor-
inated hydrocarbon pesticides in areas along the
major migration pathways and wintering grounds
in the eastern United States, West Indies, and
Central and South American countries.
AUTHORITIES
Thomas J. Cade
Laboratory of Ornithology
Cornell University
Ithaca, New York 14850
Clayton M. White
Department of Zoology
575 Widtsoe Bldg.
Brigham Young University
Provo.Utah 84601
Richard Fyfe
Room 1110
10025 Jasper Avenue
Edmonton, Alta. Canada
T5J 1S6
James H. Enderson
Dept. of Biology
Colorado College
Colorado Springs, Colorado 80903
David B. Peakall
Laboratory of Ornithology
Cornell University
Ithaca, New York 14850
Daniel D. Berger
1328 N.Jefferson, Apt. 313
MiWaukee, Wisconsin 53202
WiUiam H. Bumham
Dept. of Zoology
Brigham Young University
Provo,Utah 84601
Joseph J. Hickey
226 Russell Laboratories
University of Wisconsin
Madison, Wisconsin 53706
James L. Ruos
7145 Deer Valley Road
Highland, Maryland 20777
Walter R. Spofford, II
Aguila-Rancho
Portal, Arizona 85632
F. Prescott Ward
302 Belfast Court
Joppa, Maryland 21085
Wilston Shor
6614 32nd Street
Washington, D.C. 20015
James N. Rice
Contention Lane
Berwy n , Pennsylvania 19312
PREPARER'S COMMENTS
It seems obvious that by far the most im-
portant factor contributing to the reproductive
difficulties and population decline of the Arctic
peregrine, as in the case of the now-extinct eastern
Unites States population of the American pere-
grine, is high levels of chlorinated hydrocarbon
chemicals in the environment. Correction of this
situation should receive the highest priority in all
efforts in behalf of this species— J. W. Aldrich.
LITERATURE CITED/SELECTED
REFERENCES
A. O. U. 1931. Ahcek-list of North American
birds. American Ornithologists' Union, Lan-
caster, PA.
Berger, D. C, D. W. Anderson, J. D. Weaver, R.
W. Risebrough and J. A. Keith. 1970. Shell-
thinning in eggs of Ungava Peregrines. Canad.
Field-Nat. 84:256-267.
Berry, R. B. 1971. Peregrine Falcon population
survey, Assateague Island, Mary land, fall 1960.
Raptor Research News 5(ll):31-43'
Bumham, W. H. 1975. Breeding biology and
ecology of the Peregrine Falcon in West
Greenland. MSc. thesis. Brigham Young Univ.
Provo, Utah.
Bumham, W. H., M. A. Jenkins, F. P. Ward, W. G.
Mattox, D. M. Clement, and J. T. Harris. 19 74.
Falcon research in Greenland. 1973. Arctic
27:71-74.
Cade, T. J. 1954. On the biology of falcons and
the ethics of falconers. Falconry News and
Notes 1:12-19.
. 1960. Ecology of the Peregrine and Gyr-
falcon populations in Alaska. Univ. Calif.
PubLZool. 63:151-290.
Cade, T. S., and R. Fyfe. 1970. The North Amer-
ican Peregrine survey, 1970. Canad. Field-Nat.
84:231-245.
Cade, T. J., C. M. White and J. R. Haugh. 1968.
Peregrines and pesticides in Alaska. Condor
70:170-178.
Cade, T. S., J. L. Lincer, C. M. White, D. G. Ros-
eneau, and L. G. Schwartz. 1971. DDE resi-
dues and eggshell changes in Alascan falcons
and hawks. Science 172(3986):955-957.
Campbell, R. W. and B. Davies. 1973. Nesting
raptor survey in western Canadian Arctic. En-
vironmental Protection Board Report; 1973,
Sect. 1.
8
Enderson, J. 1965. A breeding and migration sur-
vey of the Peregrine Falcon. Wilson BuU. 77:
327-339.
. 1969. Coastal Migration data as popula-
tion indices for the Peregrine Falcon. Chapt.
23 in J. J. Hickey ed. Peregrine Falcon popul-
ations: their biology and decline. Univ. Wis-
consin Press, Madison.
Enderson, J., and D. Berger. 1968. Chlorinated
hydrocarbon residues in Peregrines and their
prey species from northern Canada. Condor
70:149-153.
Enderson,J.J.,D.G. Rosenau,andL. G. Schwartz.
1968. Nesting performance and pesticide resi-
dues in Alaska and Yukon Peregrines in 1967.
Auk 85:683.
Fyfe, R. 1969. The Peregrine Falcon in the Can-
adian Arctic and eastern North America.
Chapt. 8. in Hickey ed. Peregrine Falcon pop-
ulations, their biology and decline. Univ. Wis-
consin Press Madison.
1977. Status of Canadian raptor popula-
tions, ICBP World Conf. Birds of Prey Proc:
34-39.
Fyfe, R., S. A. Temple, and T. J. Cade. 1976. The
1975 North American Peregrine Falcon sur-
vey. Canad. Field-Nat. 90:228-273.
Hickey, J. J. (ed). 1969. Peregrine Falcon popula-
tions: their biology and decline. Univ. Wis-
consin Press, Madison.
Hofslund, P. B. 1966. Hawk Migration over the
western tip of Lake Superior. Wilson Bull.,
78:79-87.
Hohn, E. O. 1955. Birds and Mammals observed
on a cruise in Amundsen Gulf, N.W.T. July
29-August 16. 1953. Canad. Field-Nat. 69:
41-44.
Kessel, B. and T. J. Cade. 1958. Birds of the Col-
ville River, northern Alaska. Biol. Papers
Univ. Alaska no. 2.
Mattox, W. G., R. A. Graham, W. A. Burnham,
D. M.Clement. 1972. Peregrine Falcon survey,
west Greenland, 1972. Arctic 25:308-311.
McEwen, E. G. 1957. Birds observed at Bathhurst
Inlet, Northwest Territories, Canad. Field-
Nat. 71:109-115.
Mueller, H. C. and D. D. Berger. 1961. Weather
and fall migration of hawks at Cedar Grove,
Wisconsin. Wilson Bull. 73:171-192.
Peakall, D. B., T. J. Cade. C. M. White and J. R.
Haugh. 1975. Organochlorine residues m
Alaskan Peregrines. Pesticide Monitoring
Journ. 8:255-260.
Rice, J. N. 1969. A peregrine population index on
the Maryland-Virginia coast, pp. 279-280, in
J. J. Hickey ed. Peregrine Falcon populations:
their biology and decline. Univ. Wisconsin
Press, Madison.
Ruos, J. L. 1970. Correlation of Arctic tempera-
tures in July with numbers of Tundra Pere-
grines (F. p. tundrius) seen per partyday in
October along the Mid-Atlantic coast. Special
study Report prepared for Rare and Endan-
gered Species Committee, U.S. Fish and Wild-
hfe Service, 15 Feb. 1970.
Shor, W. 1970. Peregrine Falcon population dy-
namics deduced from band recovery data.
Raptor Research News 4(2):49-59.
. 1970a. Banding recoveries of Arctic mi-
grant Peregrines of the Atlantic Coast and
Greenland population. Raptor Research News
4(4):125-131.
Spofford, W. R. 1969. Hawk Mountain counts as
population indices in northwestern America,
pp. 323-332, in J. J. Hickey ed. Peregrine Fal-
con populations: their biology and decline.
Univ. Wisconsin Press, Madison.
Walker, W., W. G. Mattox, and R. W. Risebrough.
1973. Pollutant and shell thickness determina-
tion of Peregrine eggs from west Greenland.
Arctic 26:256-158.
Ward, F. P. and R. B. Berry. 1972. Autumn migra-
tions of Peregrine Falcons on Assateague Is-
land, 1970-72. Journ. Wildl.Mngmt. 36:484-
492.
White, C. M. 1968. Diagnosis and relationships of
the North American tundra inhabiting Pere-
grine Falcons. Auk 85:179-191.
• 1969. Breeding Alaskan and Arctic mi-
grant populations of Peregrines. Chap. 2 in
Hickey, J.J. ed. Peregrine Falcon populations
their biology and decline. Univ. Wisconsin
Press, Madison.
White, C. M., and T. J. Cade. 1971. Chff-nesting
raptors and ravens along the Colville River in
Arctic Alaska. Living Bird 10:107-150.
• 1975. Long term trends of Peregrine
populations in Alaska. ICBP World Conf.
Birds of Prey Proc.
Biological Services Program
FWS/OBS-80/01.52
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
MISSISSIPPI
SANDHILL
CRANE
^•'^T-mmr •»:,'
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704r}8
11
FWS/OBS-80/01.52
March 1980
SELECTKD VKR 1 KBRA 1 K ENDANGERED SPECIES
OF THE SEACOAS 1 0\ 1 HE UNITED STATES-
MISSISSIPPI SANDHILL CRANE
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National ('oastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell. Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
(PHOTO OF C1-OSEI.Y REL.ATED SUBSPECIES)
MISSISSIPPI SANDHILL CRANE
Grus canadensis pulla Aldrich
KINGDOM Animalia
CLASS Aves
ORDER Gruiformes
FAMILY Gruidae
OTHER COMMON
NAMES. . . . Florida Crane (Mcllhenny 1938);
Florida SandhUl Crane (Valentine 1963).
CREDIT: LUTHER C. GOLDMAN
DATE
Entered into SWIS To be determined
Updates To be determined
LEGAL STATUS
Federal: Endangered: (42FR36425, 14 July
1977). Protected (Migratory Bird Treaty
Act of July 3, 1918. (40 Stat. 755; 16
U.S.C. 703-711) as amended 3 Dec.
1969. Public Law 91-135.
States: Endangered: Mississippi (protected
under the Nongame and Endangered
Species Act of 1974;
Listed: Alabama.
REASONS FOR CURRENT STATUS:
The Mississippi Sandhill Crane is nearing ex-
tinction because of its very small and restricted
population and the deterioration of its habitat,
semiopcn wet pine savanna, because of drainage,
planting of trees for timber, urban and suburban
development, and highway construction (U.S.
Fish and Wildlife Service 1973; Morine 1975;
Recovery Team for the Mississippi Sandhill Crane
1976).
Since the mid-1950's, timber companies have
acquired or leased thousands of hectares, which
have been planted to slash pine (Pinus elliottii).
Drainage ditches dug through savannas and water-
courses speed run-off. Fire, the natural agent in
maintaining even wet prairies, has been controlled
or eliminated, encouraging pine plantation sur-
vival and also permitting natural reproduction of
pine and brush in unplanted savannas. Many ac-
cess roads, trails, and fire lanes have been built,
giving people easier access to crane habitat. Tim-
ber management has made thousands of hectares
of former crane habitat unsuitable for these birds
(Valentine and Noble 1970; Morine 1975; Re-
covery Team for the Mississippi Sandhill Crane
1976).
Housing developments and small farms are ex-
panding into crane habitat from Ocean Springs,
Fontainebleau, and Gautier communities. Recent
release of St. Regis timber land holdings allowed
the sale of about 2,000 hectares of crane environ-
ment for development.
U.S. Highway 90, a four-lane east-west avenue
through crane range, now a "strip city" extends
the entire distance between Ocean Springs and
Pascagoula. Other highways through crane habitat
are being built or improved. The adverse effects
of highways on cranes are: (1) direct loss of land
to right-of-way and borrow areas; (2) disturbance
by noise, vibration, and visual factors; (3) pollu-
tion of surrounding air and soil; (4) facilitation of
public access to crane environment; and (5) stim-
ulation of residential and commercial develop-
ments along highway route (Recovery- Team for
the Mississippi Sandhill Crane 1976).
PRIORITY INDEX
50
DESCRIPTION
This species is similar in size to the Florida
sandhill crane, Grus canadensis pratensis, but
colored portions of plumage are much darker
throughout. It differs from the greater sandhill
crane, Grus canadensis tabida, in smaller size
(except tarsus) and darker color; and from the
lesser sandhill crane, Grus canadetuis canadensis,
in larger size in all dimensions and darker color.
It can be distinguished from the Canadian sandhill
crane, Grus canadensis rowani, by longer tarsi and
darker color (Aldrich 1972).
Measurements. (8 live adult birds hatched
from Mississippi-taken eggs, sex undertermined):
wind (chord), 470-493 mm; bill (from posterior
end of nostrO), 82-91; tarsus, 216-258 (Aldrich
1972).
Eggs. Light buffy background with splotches
of reddish brown and lavender, particularly around
the large end (Mcllhenny 1938;Walkinshaw 1960).
Average measurements of 16 eggs was 95.9 (89-
104) x 58.55 (56.2-59.0) (Walkinshaw 1973).
RANGE
Sandhill cranes are permanent residents in
Jackson County Mississippi near Ocean Springs and
at Fontainebleau, between the Pascagoula River
on the east and Jackson-Harrison county line on
the west, Graveline Bayou on the south and on
the north the 30°37' latitude line (Valentine 1978,
pers. comm.).
Their former distribution is poorly known,
but presumed to have been more extensive than
at present with populations in widely separated
areas of suitable habitat in southern Louisiana
(Cook 1914; Figgins 1923; Mcllhenny 1943;
Lower>' 1960), southern Mississippi (Mcllhenny
1 93 8 ; Turcotte 1 94 7 ; Walkinshaw 1949; Valentine
1963; Valentine and Noble 1970), and southern
Alabama (Howell 1928; Imhof 1962). No speci-
mens of the populations outside southern Missis-
sippi are extant, so their identification as pulla
must be assumed. In a more detailed statement by
the Recovery Team of the Mississippi Sandhill
Crane (1976), the range is said to be confined to
southern Jackson County, Mississippi, from the
Pascagoula River to about the Harrison County
Line. The northern Hmit is the east-west line (lat.
30° 35'), about 6.4 km north of Vancleave. The
southern limit is Simmons Bayou and Graveline
Bay. Eleven nesting areas were located within
those limits during 1965 through 1978. It is not
known where unpaired birds go when paired
adults are nesting.
Although this subspecies leaves its breeding
rjinge to feed and roost, it probably does not mi-
grate far. One individual was seen as far as 1 1 miles
from the nesting area. However, some individuals
of other populations of sandhill cranes may win-
ter within the range of the Mississippi birds, as
evidenced by the fact that two cranes found dead
in 1974 within the range of G. c. pulla neither
clearly fit the original description of that subspe-
cies nor matched the specimens of cranes reared
from eggs taken in Mississippi (Aldrichm Recovery
Team for the Mississippi Sandhill Crane 1976).
Also, migrant sandhill cranes have been found
wintering at Gulf Shores in Baldwin County,
Alabama, about 80 km from the range of the Mis-
sissippi subspecies (Hamilton 1971; James 1972).
RANGE MAP
Breeding range (from Valentine and Noble
1970:763; Recovery Team for the Mississippi
Sandhill Crane 1976) is shown on the following
map. Hatched areas are recent (1965-70) nesting
grounds. Numbers indicate active nests found.
STATES/COUNTIES
Mississippi: Jackson.
HABITAT
Nesting habitat was described by Walkinshaw
(1949), Valentine (1963) and Valentine and Noble
(1970) as wet areas in semiopen pine flats. It
would appear that, although savanna-like in as-
pect, breeding areas have more trees than is usually
the case wdth sandhill cranes of other subspecies
(Walkinshaw 1949, 1960, 1973). However, the
area immediately surrounding the nest is fairly
open. In the larger savannas, there are scattered
long-leaf pines [Pinus palustris), slash pine (Pinus
elliottii), baldcypress (Taxodium distichum), and
shrubs, but the view is quite open. Grassland open-
ings in the swamps, forests and pine plantations
used for nesting may be less than 0.4 hectares in
area. Of 55 nests, 24 (44%) were in open savannas;
24 (44%), in swamp edges and openings; 4 (7%),
in pine plantations; and 3 (5%), along pine forest
edges (Valentine and Noble 1970; Valentine in
Recovery Team for the Mississippi Sandhill Crane
1976).
Habitat in the winter roost in the Pascagoula
River marsh is mainly sawgrass [Cladium) with
minor components of other marsh plants. The
creek and bayou banks are lined with baldcypress
and several brush species. The roost marsh is fresh
to slightly brackish, but fairly salty water may
run up the bayous during droughts and high tides
(Recovery Team for the Mississippi Sandhill Crane
1976).
Factors in the natural environment that can
cause mortality are severe storms with flooding,
droughts, and fires. Despite some losses of nests
from fire, it is essential to the ecology of cranes
because it checks encroachment of woody vegeta-
tion on open savannas (Recovery Team for the
Mississippi Sandhill Crane 1976; Aldrich pers.
comm.).
The most serious threat to the success of crane
breeding is probably habitat destruction through
the current timber management practices, drain-
ing marshes and planting pine (Valentine and
Noble 1970).
FOOD AND FORAGING BEHAVIOR
During the summer, cranes feed on natural
foods found in swamps, savannas, and open forest
lands. Animal foods include adult and larval in-
sects, earthworms, crayfish, frogs, and small
rodents. Plant foods are roots, tubers, seeds, nuts,
fruits and leafy parts. During the fall, winter, and
early spring, most cranes feed on waste grain and
invertebrates in small com fields and pastures
several kilometers north of the breeding range.
Such farms are scarce in that area. There have
been a few complaints of crane depredations on
com fields, but most farmers welcome the cranes
(Recovery Team for the Mississippi Sandhill
Crane 1976). In winter and spring, they also feed
on freshly sprouted grass on burned open pine
flats (Mcllhenny 1938).
NESTING OR BEDDING
Nests are buUt in savannas, sparsely forested
areas, or swamp openings that hold shallow water
or may be dry. Nests placed on ground in open
areas are built of dead vegetation gathered near the
nest site (Recovery Team for Mississippi Sandhill
Crane 1976). Walkinshaw (1960) found nests of
dried sedges that were 124 by 104 cm across and
13 cm high and were situated in water 21 cm
deep. Mcllhenny (1938) described a nest of dried
grasses and weed stalks about 128 cm across and
20 to 36 cm above water 15 cm deep. The active
nest is often within a short distance of the one
used the previous year. Old or "dummy" nests
are often in close proximity to active nests (Re-
covery Team for Mississippi Sandhill Crane 1976).
Valentine found one instance of the same nest
being used 3 consecutive years; two used for 2
years; and another use 1 year, deserted for one
season and used again the next year (Valentine
1978, pers. comm.).
The main winter roost is in the Pascagoula
River marsh, in the vicinity from Bluff Creek,
Bayou Castelle, and Paige Bayou to the West
Pascagoula River (Recovery Team for the Missis-
sippi Sandhill Crane 1976). The Pascagoula River
marsh roosts are used mainly from Ausust to
March. During the breeding season, most cranes
roost in the nesting and feeding range. Cranes
may be vulnerable to hunters as they fly into and
out of the roost, generally only during brief pe-
riods at daybreak and sundown (Recovery Team
for the Mississippi Sandhill Crane 1976).
RITUAL REQUIREMENTS
All subspecies of sandhill crane engage in
spectacular dances. Sometimes a single member of
a pair will dance, sometimes both; sometimes a
group forms a circle, facing inward. The dances
consist of a great variety of postures, particularly
bowing low and leaping 2 or more meters into the
air, accompanied by vocalizations. Walkinshaw
(1949) describes these dances in detail. They
seem to be related to courtship in some cases, but
may take place at any time of the year.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None known.
POPULATION NUMBERS AND TRENDS
Leopold (1929) estimated 50 to 100 or more
cranes in Mississippi. Mcllhenny (1938) reported
1 1 nests in the Fontainebleau area and counted
34 in one flock in April 1938. Walkinshaw (1949)
estimated more than 25 pairs in 1940. Turcotte
(1947) estimated 30 in 1947. Strong (1969) es-
timated 16 pairs and a population of 50 to 60
birds. Valentine and Noble (1970) estimated about
38 to 40. Valentine (1975) judged a minimum of
10 to a maximum of 15 breeding pairs with a
population between 30 and 50 individuals. He
found 20 fairly distinct nesting territories in 11
areas during 1965 through 1978, but the most
nests found in any one year was 8 in 1969.
During the period 19-22 January 1977, a total
of 52 cranes was counted in the wintering area of
Mississippi sandhill cranes. It is not known whether
they were all of that subspecies or some were
migrants from elsewhere (Jacob Valentine, official
trip report U.S. Fish and Wildlife Service, 27 Jan.
1977).
REPRODUCTION
Paired cranes select a breeding territory for
courtship, mating, and nesting and defend it from
other cranes. Territory size may depend on the
density of cranes and physical attributes of the
habitat. In open savannas, it appears that only
one pair has possession of a territory despite the
size of the savanna. The smallest savanna occupied
by one pair was 36 ha, the largest, over 202 ha. In
Ben Williams Swamp and other areas where open-
ings suitable for nesting are shielded from each
other by large trees and shrubs, cranes will nest
close together. In 1971, three nests in a row along
southern edge of Ben Williams Pond were only
0.8 km apart. The same nesting territories are
often used year after year. Territories have been
deserted (or searchers could not find the nest) for
years and then reoccupied. Desertions may have
been caused by brush or tree encroachment on
the open area around the nest. Three territories ap-
parently were not occupied after roads were built
nearby. One nesting site was destroyed by excava-
tion of a borrow pit in construction of Interstate
10. Of the 14 or 15 known nesting territories, 6
appear to be abandoned (Recovery Team for the
Mississippi Sandhill Crane 1976).
The egg-laying period is 4 April to 20 May.
Hatching occurs between 11 April and 20 June,
with a peak during 1-20 May. Of 54 nests found
between 1965 and 1975, 9 contained 1 egg and
45 held 2, a mean of 1.83 eggs per clutch. The
incubation period is 30 to 31 days. Both parents
incubate. The hatching success of 38 eggs in the
wild was 61%, compared to 60% of 42 eggs arti-
ficially incubated for propagation. Hatching failure
of 15 wild eggs was attributed to desertion (4);
rotten eggs (2); eggs missing (3); crow depreda-
tion (3); chick dying in eggs (1); and eggs cracked
by flushing birds (2). Human disturbance can be
credited with at least 5 egg losses; crow depreda-
tions occurred during periods when cranes were
off nests after being frightened by humans. Two
eggs were broken by cranes flushed from nests by
humans. The fate of three missing eggs is un-
known; some eggs found were outside the nest
after desertion. Heavy rains are known to destroy
some eggs. Cranes attend nests constantly and so
are not normally subject to robbing by birds or
mammals. Crows have not been abundant untU
recently, when sanitary fill dumps along the road
now attract hundreds of them. Raccoons are
present but not numerous in the breeding range
(Walkinshaw 1973; Valentine in Recovery Team
for the Mississippi Sandhill Crane 1976).
Poor hatchability of eggs, weak chicks and
foot and leg deformities in chicks have been noted
in cases of captive chicks hatched from eggs in in-
cubators. Relatively low hatchability and survival
is thought by some to be due to genetic problems
resulting from inbreeding because of the obviously
small gene pool. On the other hand, J. M. Valen-
tine has observed 10 wild chicks, all free of ap-
parent defects (Valentine 1978 pers. comm.).
Another chick died while trying to break out of
the egg (Recovery Team for the Mississippi Sand-
hill Crane 1976). Chicks are able to scramble off
the nest at 8 hours of age and can swim if neces-
sary (Walkinshaw 1973).
MANAGEMENT AND CONSERVATION
Restoration of this population of sandhill
cranes began before it was recognized as a distinct
subspecies, with the rearing of young hatched
from eggs taken from wild nests in Mississippi.
Captive propagation has been at the Patuxent
Wildlife Research Center, Laurel, Maryland. The
intent was to reintroduce the propagated stock
into the wild. Acquisition and restoration of habi-
tat in the present range and adjoining areas and
rerouting proposed sections of Interstate Highway
10 were proposed as essential protection measures
(U.S. Fish and Wildhfe Service 1973).
As of 20 September 1976, a Mississippi Sand-
hill Crane Recovery Team, with Jacob M. Valen-
tine as leader, was appointed by the U.S. Fish and
Wildlife Service, and a recovery plan approved
in September 1976 was revised in 1978 (Recovery
Team for the Mississippi Sandhill Crane 1976).
Lands available for crane occupancy are
limited. The U.S. Fish and Wildlife Service has
proposed some refuge lands which are the best
available that still contain either nesting cranes or
potential nesting habitat. With the assistance of
the Nature Conservancy, the Service acquired
3,490 ha of habitat in two units and has estab-
Ushed the Mississippi Sandhill Crane National
Wildlife Refuge. A habitat management plan for
the refuge was completed by the Service on 28
March 1975 (Morine 1975; Recovery Team for
the Mississippi Sandhill Crane 1976). In June
1977, 202 ha of forest and savanna v^dthin the
breeding range of the Sandhill Crane refuge were
burned for the purpose of opening the cover and
improving nesting habitat Qacob Valentine in lit
15 June 1977).
Competition with other animals for food or
living space need not be a concern of management,
as there are few competitors in the crane's range.
Closing of the open range policy in the 1950's
prohibited cattle and hog grazing on timber com-
pany holdings, and cattle are now confined to
better grazing lands. Deer are scarce but are in-
creasing as trees and brush invade the savannas
(Valentine, pers. comm.). Management of the
crane range should not include improvement of
the habitat for deer or game birds, as this would
create a demand for hunting there as well as make
the habitat less suitable for cranes (Recovery
Team for the Mississippi Sandhill Crane 1976).
The Fish and Wildlife Service has conducted a
study and propagation program at the Patuxent
Wildlife Research Center with the objective of
rearing 10 captive breeding pairs to produce young
cranes for transplanting to suitable habitat within
the range of the Mississippi Sandhill Crane. Up to
the present, 14 captive birds have been produced
at Patuxent from Mississippi-taken eggs; several
captive pairs reared have laid eggs and from these,
2 have been raised (Ray Erickson pers. comm,
1978).
The U.S. Forest Service, in cooperation with
Fish and Wildlife Service and the Mississippi
Game and Fish Commission, will clear and main-
tain two areas in the DeSoto National Forest in an
effort to improve crane habitat. No cranes are
nesting there at present.
The U.S. National Park Service has made sur-
veys and will recommend National Landmark
status for some lands included within the proposed
Federal refuge boundaries.
The Jackson County, Mississippi, Board of
Supervisors passed a resolution supporting pur-
chase of a county school section by the Fish and
Wildlife Service to become part of the Fountaine-
bleau Unit of the refuge.
The Mississippi Game and Fish Commission
has participated in sandhill crane studies and pro-
tection for many years. It has cooperated with
the U.S. Fish and Wildlife Service in crane pro-
pagation since 1965 by approving and assisting
taking of eggs. It approved establishment of a
Federal crane refuge in 1971.
The National Wildlife Federation and Missis-
sippi Wildlife Federation filed a federal court
action against the Department of Transportation,
Federal Highway Administration, and Mississippi
Highway Department for violations of Section 7,
Endangered Species Act for construction of Inter-
state Highway I- 10 through the Mississippi Sand-
hill Crane National Wildlife Refuge. They asked
the court for elimination of an interchange on the
Gautier-Vancleave Road, the elimination of bor-
row pits in the sensitive area, and the acquisition
of lands by the highway agencies to mitigate the
loss of critical habitat by the highway.
If it is concluded that the present population
of Mississippi sandhill cranes is doomed to extinc-
tion because of impaired reproduction resulting
from inbreeding, it may be justified to try a gene-
tic infusion of another strain such as that of the
Florida or Georgia populations of the Florida
Sandhill Crane. This would be with hope of con-
tinuing to have a crane population in the remnant
habitat now occupied by the Mississippi subspecies.
This might be accomplished by placing eggs of
Florida Sandhills in the nests of Mississippi birds,
which would act as foster parents. This action
would be taken only as a last ditch option (MSCRT
1976). Critical habitat has been designated in
Jackson County, Mississippi (42FR39985, 8
August 1977).
AUTHORITIES
Lawrence H. Walkinshaw
1145 Scenic Drive
Muskegon, Michigan 49445
Robert E. Noble
Department of Forestry and Wildlife Manage-
ment
Louisiana State University
Baton Rouge, Louisiana 70803
William H. Turcotte
Mississippi Game amd Fish Commission
P.O. Box 451
Jackson, Mississippi 39205
Jacob M. Valentine, Jr.
U.S. Fish and Wildlife Service
1 1 1 East Main St.
Lafayette, Louisiana 70501
PREPARER'S COMMENTS
The Mississippi sandhill crane has had the
benefit of adequate study to show, in detail, what
is necessary to prevent its extinction. However, its
habitat and population have been reduced to such
a small remnant by drainage and planting for pine
timber production, and human population expan-
sion and economic pressures are becoming so
great in and around the crane's habitat, that only
an overwhelming public sentiment to do every-
thing possible for it can save the Mississippi sand-
hill crane.
LITERATURE CITED/SELECTED
REFERENCES
Aldrich, J. W. 1972. A new subspecies of sandhill
crane from Mississippi. Proc. Biol. Soc. Wash.
85:63-70.
Cooke, W. W. 1914. Distribution and migration of
North American rails and their allies. U.S.
Dept. Agric. Bull. 128.
Figgins, J. D. 1923. The breeding birds of the
vicinity of Black Bayou and Bird Island,
Cameron Parish, Louisiana, Auk 40:666-667.
Hamilton, R. S. 1971. Central Southern Region.
Amer. Birds 25:590.
Howell, A. H. 1928. Birds of Alabama (2nd Ed.)
U.S. Dept. of Agric. Bur. Biol. Surv. and Ala.
Dept. of Game and Fisheries.
Imhof, T. A. 1962. Alabama birds. Dept. of Con-
servation, Game and Fish Div. Univ. Alabama
Press.
James, F. G. 1972. Central Southern Region.
Amer. Birds 26:616.
Valentine, J. M.,Jr. 1963. The status of the Flor-
ida Sandhill Crane in Jackson County, Missis-
sippi. Admin. Rept. U.S. Bur. Sport Fish. &
Wildl., Atlanta, Ga.
Valentine, J. M. 1975. The Mississippi Sandhill
Crane Grus canadensis pulla). Report to Crane
Working Group, I.C.B.P. ms.
Valentine, J. M. Jr., and R. E. Noble. 1970. A
colony of Sandhill Cranes in Mississippi.
Journ. WUdl. Mgmt. 34:761-768.
Valentine, J. M., Jr., and R. E. Noble. 1976. The
Mississippi Sandhill Crane - Endangered or
Doomed. Proc. Int. Crane Workshop 1:343-
346.
Valentine, J. M., Jr. 1978. The Mississippi Sand-
hill Crane - A Status Update. Presented at the
crane workshop, December 7, 1978.
Walkinshaw, L. H. 1949. The Sandhill Cranes.
Cranbrook Institute of Science. Bull. 29.
Walkinshaw, L. H. 1960. Some Mississippi Crane
notes. Migrant 31:41-43.
Walkinshaw, L. H. 1973. Cranes of die World.
Winchester Press, pp. 136-140.
Leopold, A. 1929. Report on a game survey of
Mississippi, ms.
Lowery, G. H., Jr. 1960. Louisiana birds. Loui-
siana Wildl. and Fisheries Comm. La. State
Univ. Press.
Mcllhenny, E. A. 1938. Florida Crane a resident
of Mississippi. Auk 55:598-602.
Mcllhenny, E. A. 1943. Major changes in the bird
life of southern Louisiana during sixty years.
Auk 60:541-549.
Morine, D. E. 1975. Saving the Mississippi Sand-
hill Crane. Nature Conservancy News. Winter
1975:15-18.
Recovery Team for the Mississippi Sandhill Crane
1976. Mississippi Sandhill Crane Recovery
Plan (Draft April 1976).
Strong, L. 1969. An investigation of the status of
the Sandhill Crane in Mississippi. Miss. Game
and Fish Comm. PR Project Report W-48-R.
Turcotte, W. H. 1947. The Sandhill Crane in Mis-
sissippi. Mississippi Game and Fish. June
1947:8-9.
U.S. Fish and Wildlife Service. 1973. Threatened
Wildlife of the United States, 1973 Edition.
U.S. Dept. of Interior. Resource Publication
114, GPO.
Biological Services Program
FWS/OBS-80/01.53
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
GRAY WOLF
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data aic not necessarily equivalent
to critical habitat as defined in the F,ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. ,\rmy Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
u
FWS/OBS-80/01.53
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
GRAY WOLF
A Cooperative Effort
by the
National Fish and WildUfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT: L. DAVID MECH
GRAY WOLF
Canis lupus Linnaeus
KINGDOM Animalia
CLASS Mammalia
ORDER Carnivora
FAMILY Canidae
OTHER COMMON
NAMES Lobo, Timber Wolf
DATE
Entered into SWIS To be determined
Updates 1 Feb. 1978; U July 1978
LEGAL STATUS
Federal: Endangered in United States (48
conterminous States other than Min-
nesota) and Mexico (43 FR 9607; 9
Mar 1978). Threatened in Minnesota
(43 FR 9607; 9 Mar 1978).
States: Endangered: Colorado, Idaho (TorE),
Massachusetts (Extinct), Michigan,
Montana, New Mexico, Rhode Island
(Extinct), Texas, Wisconsin.
REASONS FOR CURRENT STATUS
Clearing of forests and proliferation of civili-
zation in the eastern United States led to extirpa-
tion there. Habitat alteration caused depletion of
prey such as deer, moose, and beaver; loss of
natural prey probably led to predation on domes-
tic animals (Mech 1977a). Bounties were imposed
in the 1600's and wolves were essentially extir-
pated from this region by the 1800's (Ruther and
Pimlott 1968).
In the Great Plains, wolves preyed on live-
stock as bison (Bison bison) became depleted. In
the 1800's, bounties on wolves were offered by
cattlemen. In 1907, the U.S. Biological Survey
concluded that wolf predation caused an annual
loss of several million dollars. The agency deter-
mined that destruction of young in dens, together
with trapping adults, significantly reduced loss of
cattle and sheep (Young and Goldman 1944). In
1915, Biological Survey was authorized to control
wolf numbers, and they systematically reduced
wolf numbers by trapping, poisoning, and other
means (Rutter and Pimlott 1968).
Bounties are still maintained in some Cana-
dian provinces, and unorganized but effective
control continues in Mexico (Mech 1974a,
McBride 1978). Minnesota (which has the last
major concentration of gray wolves in the 48
contiguous states, Wisconsin, and Michigan offered
bounties until 1965, 1950, and 1960, respectively
(Thompson 1952, Hendrickson et al. 1975, Mech
1977a). Regulations providing bounties for
wolves in Alaska were repealed for units 1-3 in
1978, and the balance of the state in 1969-1970
(R. Rausch, personal communication).
Killing by man for commercial purposes, as
well as for sport, has been the major factor in the
decline of the wolf in the contiguous United States
and Mexico (43 FR 9611; 9 Mar. 1978). Illegal
hunting continues today, largely because of wolf
predation on livestock and game and, to a lesser
extent, because of an age-old unsubstantiated fear
of attack on human beings (not one nonrabid
wolf has been known to deliberately attack a
person in North America.— Mech 1966).
PRIORITY INDEX
Not assigned.
DESCRIPTION
Canis lupus is a dog-like canid. Males are
larger than females, weighing 20 to 70 kg and
measuring 1.27 to 1.64 m in total length. Females
weigh 18 to 55 kg and measure 1.37 to 1.52 m.
Pelage is extremely variable, usually grayish
with black extending from upper side of neck
over back. The head is more or less suffused with
cinnamon. Underparts vary from white to pinkish
buff with scattered dark hairs, becoming a clearer
white in the inguinal region. Limbs range in color
from a pinkish buff to a rich cinnamon, with
some specimens having a prominent narrow black
line on the forearms. The tail is grayish above and
buff below with a black tip. Summer colors are
similar to winter but somewhat paler. Other color
phases include all white, or all black, and any
color variation between them.
It is distinguished from the coyote (Canis
latrans) by its larger size, broader snout, shorter
ears, and proportionately smaller brain case. It
resembles a German shepherd (Canis familiaris);
however, there are subtle skull differences (Mech
1974a). Gray wolves are more massive and not as
lanky as red wolves (Canis rufus) (Riley and
McBride 1972).
Black-and-white photographs are found in
Murie (1944), Young and Goldman (1944), Mech
(1966), and McBride (1978); and color pictures,
in Mech (1974a, 1977b), Peters and Mech (1975),
and March (1977).
RANGE
The present range is essentially equal to the
former range in Alaska and much of northern
Canada. In the 48 conterminous United States
and Mexico, however, populations are limited to
Isle Royale, Michigan, the northern third of
Minnesota, the northern Rocky Mountains in
Montana, and the States of Chihuahua and
Durango (and possibly San Luis Potosi, Sonora,
and Zacatecas) in Mexico. Scattered reports of
sightings occur in three other areas:
1. Southwestern Texas, southern New Mexico,
and southeastern Arizona;
2. The Rocky Mountains in Wyoming and
Idaho; and
3. (more commonly than the other two areas)
the Upper Peninsula of Michigan and north-
ern Wisconsin.
While formely common in most of North
America from the Polar ice cap south to middle
Mexico, there were probably very few or none in
California or the southeastern United States.
RANGE MAP
Present range is delineated by shading (Mech
1974a; McBride 1978). Critical habitat is taken
from the official listing (43 FR 9607, 9 Mar.
1978).
STATES /COUNTIES
Alaska: All.
Arizona: (?)
Idaho: Fremont, Clark, Lemhi.
Michigan: Chippewa, Iron, Keweenaw, Mackinac,
Marquette.
Minnesota: Aitkin, Becker, Beltrami, Carlton,
Cass, Clay, Clearwater, Cook, Crow
Wing, Dakota, Hubbard, Itasca,
Koochiching, Lake, Lake of the Woods,
Mahnomen, Marshall, Norman, Penn-
ington, Pine, Polk, Red Lake, Roseau,
St. Louis, Wadena.
Montana: Beaverhead, Flathead, Glacier, Lewis
and Clark, Madison (?), Park (?).
Pondera, Teton.
New Mexico:(?)
North Dakota:(?)
Texas: (?)
Washington:(?)
Wisconsin: (?)
Wyoming: Fremont, Park, Teton.
HABITAT
The gray wolf has no particular habitat
preference except areas of relative wilderness
where human habitations are scarce (Stabler
1944). Mech (1974b) suggests that wild land is
not actually preferred, but is the only place to
survive human persecution.
Olson (1938) observed that wolves use a
variety of habitats, depending on prey items and
season of the year. De Vos (1950) studied wolf
movements in Ontario and concluded that topog-
raphy was also important; all topographies and
habitats are utilized except deserts and high
mountain tops (Mech 1974a). McBride (1978)
and R. Rausch (personal communication), how-
ever, also include deserts and high mountain tops.
FOOD AND FORAGING BEHAVIOR
Predominant prey in Alaska depends on loca-
tion. Sitka deer {Odocoileus hemionus sitkensis),
moose {Alces alces), and caribou {Rangifer taran-
dus) are of greatest value, with numerous other
species also taken (Murie 1944; Rausch 1967;
R. Rausch, personal communication).
Gray wolves in the United States and Canada
prey on white-tailed deer [Odocoileus virginianus),
moose, snowshoe hare [Lepus americanus),
cottontail rabbit [Sylvilagus floridanus), beaver
[Castor canadensis), various rodents, carrion, and
livestock. The white-tailed deer is the single most
important food source, particularly in winter
(Young and Goldman 1944; Thompson 1952;
Weise at al. 1975).
In Wood Buffalo Park in Canada, bison are
taken (Pimlott 1967). Tundra wolves in Canada
have a winter diet almost exclusively of caribou.
Summer diet varies and includes fish, small birds,
and rodents (Kuyt 1972).
Wolves in Mexico prey almost exclusively on
livestock, particularly weaned yearling cattle,
burros, and colls. Natural prey levels are low over
wolf range (McBride 1978).
Wolf predation is largely restricted to mal-
nourished, young, old, or otherwise weakened
individuals (Murie 1944, Crisler 1956, Mech 1966,
Pimlott 1967, Mech and Frenzel 1971, Mech
1975, Wolfe 1977). Studies by McBride (1978)
and R. Rausch (personal communication), how-
ever, indicate that wolves in Mexico and Alaska
take healthy animals in preference to unhealthy
ones.
Wolves use a variety of hunting techniques.
Murie (1944) and Mech (1966) describe many
hunts, including ambushes, chases, and stalkings.
Wolves often stalk the prey until it makes a move,
and then chase at 56.3 to 64.4 km/h for up to 3
km (Mech 1970).
^h j 'r " — r— M
Present range of the gray wolf
Methods of kill depend upon the size and
condition of the prey. On occasion, one wolf will
grab the prey by the nose while others attack
from behind. They feed side by side and gorge up
to 4.5 kg per day (Mech 1975). They can go
several days without eating (Mech 1966). Remains
of the kill are often left, but wolves seldom return
(McBride 1978) unless they are in poor condition,
extremely hungry, or feeding pups (Young and
Goldman 1944).
Hunting success, studied by Mech (1966) on
Isle Royale, Michigan, indicated that less than
8% of attacks on moose by wolf packs were suc-
cesful. Kolenosky (1972) estimated that 25% and
63% of deer hunts were successful on two winters
in Ontaria, although these values might be in-
flated. Wolves in Alaska appear to take prey in
proportion to its abundance, and are highly suc-
cessful. When conditions such as deep snow or
abundant prey are present, they kill in excess of
their needs (R. Rausch, personal communication).
SHELTER REQUIREMENTS
See NESTING OR BEDDING.
NESTING OR BEDDING
Ryon (1977) describes the den-digging of a
captive wolf. The work was started in April and
May, with the female doing most of the work
(although on one occasion, the male was observed
doing most of the work). Three dens had lengths
from 138 to 183 cm, widths of 36 to 41 cm, and
heights of 31 to 38 cm. All had domed roofs with
somewhat restricted entrances .
Wolves often occupy fox dens or make dens
in hollow logs, rock caves, bases of trees, or sides
of hills (Murie 1944, Young and Goldman 1944,
Joslin 1967). Many dens are located in sandy soils
and may be in spots where there is limited visibili-
ty or on slopes where there is a clear view of the
surrounding terrain (Young and Goldman 1944,
Jordan et al. 1967, Joslin 1967).
Photographs of dens appear in Murie (1944)
Young and Goldman (1944), Mech (1966), and
McBride (1978).
RITUAL REQUIREMENTS
Not known.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
The wolf territories studied in Superior Na-
tional Forest, Minnesota, ranged in size from 125
to 310 km^ (Mech 1974b). Territories in Alaska
and Canada, however, cover a greater range of
areas (R. Rausch, personal communication).
A pack will travel its territory irregularly, but
will cover most areas every 3 weeks or oftener in
Superior National Forest. Boundaries are scent-
marked (Peters and Mech 1975). Mech (1974b)
found that lone wolves have a nomadic range over
a large, nonterritorial area and attempt to avoid
packs.
Maximum wolf densities on Isle Royale in
Lake Superior, Algonquin Provincial Park, On-
tario, and Superior National Forest have been
estimated at one wolf per 2,590 ha. Wolf density
can be much compressed in areas of very high
prey density, such as parts of Canada and the
eastern edge of the Superior National Forest
(Kuyt 1972, Mech 1974b).
POPULATION NUMBERS AND TRENDS
On Isle Royale in Lake Superior, R. Linn
(personal communication) reports that there are
40 or more wolves. Mech (1977a) estimates 1,000
to 1,200 in Minnesota. R. Ream (personal com-
munication) reports that from 10 to 20 can be
found in the extreme northern Rocky Mountains
of Montana. Wisconsin is believed to have a few
(Anon. 1977), and there are perhaps six in the
Upper Peninsula of Michigan (Hendrickson et al.
1975).
Mech (1977b) and R. Rausch (personal com-
munication) estimate that wolves number 10,000
to 15,000 in Alaska and 17,000 to 28,000 in
Canada. Fewer than 50 adult wolves probably sur-
vive in Mexico (McBride 1978).
Murie (1944) recorded a decline in wolf num-
bers which he was unable to explain in Alaska
from 1916 to 1925; he suggested that disease may
have accounted for it.
Numbers in Minnesota increased following the
prohibition of aerial hunting in 1950, curtailment
of wolf control programs in 1955, and repeal of
bounties in 1965. However, between 1968 and
1970, the Superior National Forest population
decreased 32% based on the 1967-68 numbers,
and 55% based on the 1969-70 numbers. This
appears to be a direct result of a decrease in the
deer population, which is being stressed by
wolf predation, inclement weather, and the
decline of their habitat as cut-over forests mature
(Mech 197 7d, Mech and Karns 1977).
Studies on Isle Royale, Michigan, indicated
that successful rearing of young is dependent on
ready availability of food. Two or more pups sur-
vived rather than the usual one in years when
twinning was observed in the moose population
(Jordan et al. 1967).
REPRODUCTION
Courtship lasts from a few days to months,
with a definite mate preference. Estrus lasts 5 to
7 days, with copulation occurring in February in
Minnesota and later further north and at higher
altitudes. The gestation period is 63 days; average
litter size is six.
The female stays with the young for 1 to 2
months, while the male and other family members
provide food. Weaning is at 5 weeks, and sexual
maturity at 1 to 2 years (Medjo and Mech 1976).
In each pack, there is usually a dominant pair;
the male leads attacks on prey or intruders.
Young that survive the first winter may disperse
before the next year's litter is born, but usually
stay until after it is born (Rutter and Pimlott
1968).
An unexploited population in Ontario con-
tained 35% pups, 40% yearlings, and 25% adults.
Exploited populations in Alaska contained 42%
pups, 29% yearlings, and 30% adults (Mech
1974a).
MANAGEMENT AND CONSERVATION
Cessation of wolf controls, including boun-
ties, and prohibition of aerial hunting have had
positive effects on wolf numbers in the Supe-
rior National Forest, Minnesota (Mech 1973).
Several attempts have been made to rein-
troduce gray wolves. In 1952, four zoo animals
were released on Isle Royale, but they became
a public nuisance. In 1960, four animals were
released on Coronation Island, Alaska. By 1964,
they numbered 1 1 and had learned to prey on
black-tailed deer {Odocoileus hemionus colum-
bianus). In 1972, five laboratory animals were
unsuccessfully released near Umiat, Alaska
(Weise et al. 1975).
In 1974, four live-trapped radio-tagged
Minnesota animals were released in the Upper
Peninsula of Michigan. All four were killed by
human-related causes. However, it was demon-
strated that wolves can be translocated. For a
successful reintroduction program, Weise et al.
(1975) recommended a strong public education
campaign, abatement of all coyote bounties,
and release of a larger number of animals.
U.S. Fish and Wildlife Service reclassification
of Canis lupus (43 FR 9607, 9 Mar 1978) provides
Endangered status for all gray wolves south of
Canada, excluding Minnesota (where they are con-
sidered Threatened). This ruling supersedes a pre-
vious ruling which listed four subspecies as En-
dangered and was considered unsatisfactory be-
cause the taxonomy was outdated, wolves wander
outside recognized boundaries, and unlisted sub-
species may still occur in the lower 48 States
(possibly in Washington and North Dakota). Re-
classification of the wolf as Threatened in Minne-
sota was considered necessary to provide for the
future well-being of both the wolves and persons
living in areas inhabited by wolves. The ruling
provides for establishment of five management
zones in Minnesota, of which three (Zones 1, 2,
and 3) are considered Critical Habitat (43 FR
9607, 9 Mar 1978). Zone 1 is in the extreme
northeastern part of the State and includes parts
or all of Cook, Koochiching, Lake, and St. Louis
Counties. Zone 2 borders the southern part of
Zone 1 and includes parts of Lake and St. Louis
Counties. Zone 3 is in the north-central part of
the State and includes portions of Beltrami, Itasca,
Koochiching, Lake of the Woods, Marshall, and
Roseau Counties. Zones 4 and 5 comprise the
remainder of the State. Wolf numbers in these
zones will be held below biological potential (43
FR 9607, 9 Mar 1978).
The Eastern Timber Wolf Recovery Team
(1975) outlined the three major objectives it be-
lieves are necessary to remove this subspecies
from Endangered status:
1. Determine the status and distribution
of the population
2. Insure perpetuation in its present range
3. Reestablish populations in suitable areas
within the former range where viable
populations do not now exist.
Other considerations include establishing mechan-
isms for resolving conflicts between the interests
of the wolves and the interests of people; resolving
taxonomic questions, and identifying essential
habitat and species requirements. The plan out-
lines coordination among several Federal agencies
and the States of Idaho, Montana, and Wyoming
to achieve these objectives.
In the United States, there are a number of
captive wolves. The species breeding potential is
very good. Jack Lynch's Loboland in Gardiner,
Washington, has over 100 wolves whose primary
genetic origin is in the Northern Great Plains and
Rocky Mountain areas (area of overlap of C. I. nu-
bilus and C. I. irremotus). Reintroduction of some
of these animals to the wild in a national park has
been suggested (March 1977).
McBride (1978) believes the wolf will become
extinct in Mexico in the next 20 years. The Ari-
zona-Sonora Desert Museum in Tucson, Arizona,
however, is working to establish a breeding colo-
ny of the Mexican wolf (C /. baileyi) (McBride
1978).
Wolves are persecuted in Canada for their im-
pact on caribou. Kuyt (1972) suggests that wolves
be listed as a game animal, which would render
some protection and help insure survival.
Estimates of 1978 wolf numbers on Federal
lands, based on responses to letters of inquiry, are
listed in the table below.
AUTHORITIES
Eastern Timber Wolf Recovery Team
Ralph E. Bailey, Leader
Michigan Department of Natural Resources
P.O.Box 190
Marquette, Michigan 49855
William C. Hickling
U.S. Fish and Wildlife Service
279 Federal Building
Ashville, N. Carolina 22801
Robert M. Linn
National Park Service
Biological Science Department
Michigan Technological University
Houghton, Michigan 49931
L. David Mech
U.S. Fish and Wildlife Service
North Central Forest Experiment Station
Folwell Avenue
St. Paul, Minnesota 55101
Ron Nicotera
Wisconsin Department of Natural Resources
Box 450
Madison, Wisconsin 53701
State
Location
Status
Reference
Alaska
Arctic NWR
Not surveyed
Chugach NF
40-50 estimated to be in Unit 7
R. Walker, pers. comm.
Glacier Bay NM
28-55 estimated
B. Paige, pers. comm.
Katmai NM
30-40 estimated
W. Troyer, pers. comm.
Kenai National Moose Range
94 estimated
T. Bailey, pers. comm.
Mt. McKinley NP
35-50 estimated
D. Kuehn, pers. comm.
TongassNF
No estimate, numbers increasing.
87 harvested 1975-76
W. Overdorff
Arizona
Coronada NF
Several unconfirmed reports
of possible transient wolves
P. Karp, pers. comm.
Idaho
Salmon NF
5 estimated
L. Campbell, pers. comm.
Tar ghee NF
Numerous unauthenticated
reports
O. JOhnson, pers. comm.
Michigan
Isle Royale NP
40 or more wolves known to be
present
R. Linn, pers. comm.
State
Location
Status
Reference
Minnesota
Montana
OttowaNF
Chippewa NF
Superior NF
Voyageurs NF
Beaverhead NF
Blackfeet Indian Reservation
Custer NF
Flathead NF
Gallatin NF
Glacier NP
Kootenai NF
Lewis and Clark NP
Texas
Big Bend NP
Wisconsin
Chequamegon NF
NicoletNF
Ottawa NF
Wyoming
Bridger-Teton NF
Shoshone NF
Yellowstone NP
5 wolf activity centers
40 is present estimate
300-400
Not surveyed
Some documented observation
Not surveyed
Not surveyed
Sporadic reports of sign or
sightings of single wolves
Not available
10-20 estimated, no persistent
pack activity
R. Prause, pers. comm.
J. Mathisen, pers. comm.
D. Mech (1976)
R. Kiewit, pers. comm.
R. Hensler, pers. comm.
C. Martinka, pers. comm.
Not surveyed
Scattered reports of sightings and J. Schulstad
sign— no estimate
Occasional unauthenticated reports G. Balaz, pers. comm.
of transient wolves
Not surveyed
No estimate— some transient
visitors
No estimate, but good evidence
of wolf activity there
No estimate, but sightings and
sign reported
A. Rinaldi, pers. comm.
T. Wingle, pers. comm.
R.Jackson, pers. comm.
No estimate, but unauthenticated R. Hall, pers. comm.
sightings
Unauthenticated sightings, recent Weaver (1978)
search failed to confirm any wolves
Key:NF-National Forest; NM-National Monument; NP-National Park; NWR- National Wildlife Refuge.
Robert E. Radtke
U.S. Forest Service
633 W. Wisconsin Avenue
Milwaukee, Wisconsin 53203
Leroy Rutske
Minnesota Department of Natural Resources
Centennial Building
St. Paul, Minnesota 55155
Karl Siderits
Superior National Forest
P.O. Box 338
Duluth, Minnesota 55801
Northern Rocky Mountain Wolf Recovery Team
Dennis L. Flath, Leader
Nongame Species Biologist
Montana Department of Fish and Game
Box 5, MSU Campus
Boseman, Montana 59717
Levi Mohler
602 Michael
Boise, Idaho 83704
Roger Evans
Wildlife Biologist
Lewis and Clark National Forest
Great Falls, Montana 59401
Mary M. Meagher
Research Scientist
Yellowstone National Park, Wyoming 83020
Donald Mcintosh
Bureau of Land Management
P.O.Box 30157
Billings, Montana 59101
Robert K. Turner
National Audubon Society
P.O.Box 3232
Boulder, Colorado 80303
Robert K. Ream
School of Forestry
University of Montana
Missoula, Montana 59801
Donald S. Balser
U.S. Fish and Wildlife Service
Room 3035, Federal Building
Billings, Montana 59101
Other Authorities
Durward Allen (Eastern Timber Wolf)
Department of Wildlife Ecology
Purdue University
West Lafayette, Indiana 47907
Roy McBride (Mexican
Department of Biology
Sul Ross State University
Box 725
Alpine, Texas 79830
Douglas H. Pimlott
Department of Zoology
University of Toronto
Toronto, Ontario, Canada
Robert Rausch (Alaskan Wolf)
Alaskan Department of Fish
Division of Game
Juneau, Alaska 99801
and Gray Wolves)
and Game
PREPARER'S COMMENTS
There is extensive controversy concerning
competition between wolves and hunters. Pimlott
(1975) suggests that wolves feed on the non-
breeding sector of the population and thus do not
depress prey populations, although he states that
in areas of heavy hunting, competition probably
does exist. R. Rausch (personal communication)
states that work in Alaska has shown that in
many cases wolves show a preference for adults
and thus can depress prey populations. A recent
study by Mech and Karns (1977) indicated that,
given a particular combination of factors, wolves
could severely deplete deer numbers and keep
them at a depressed level.
LITERATURE CITED/SELECTED
REFERENCES
Anon. 1977. Timber wolf declassification debated.
End. Species Tech. Bull. 2(3): 1, 2-4, 7.
Crisler, L. 1956. Observations of wolves hunting
caribou. J. Mammal. 37:337-346.
DeVos, A. 1950. Timber wolf movements of Sib-
ney Peninsula, Ontario. J. Mammal. 31(2):
167-169.
Eastern Timber Wolf Recovery Team. 1975. Re-
covery plan for the eastern timber wolf. Preli-
minary draft. 79 pp.
Hall, E. R., and K. R. Kelson. 1959. The mammals
of North America. 2 vols. Ronald Press, New
York.
Hendrickson, J., W. L. Robinson, and L. D. Mech.
1975. Status of the wolf in Michigan, 1973.
Am. Midi. Nat. 94(l):226-232.
Ingles, L. G. 1963. Status of the wolf in Califor-
nia. J. Mammal. 44:109.
Jordan, O. A., P. C. Shelton, and D. L. Allen.
1967. Numbers, turnovers, and social struc-
ture of the Isle Royale wolf populataion. Am.
Zool. 7:233-252.
Joslin, P. W. B. 1967. Movements and home sites
of timber wolves in Algonquin Park. Am. Zo-
ol. 7:279-288.
Kolenosky, G. B. 1972. Wolf predation on winter-
ing deer in east-central Ontario. J. Wildl.
Manage. 36:357-369.
Kuyt, E. 1972. Food habits and ecology of wolves
on barren-ground caribou range in the North-
west Territories. Canadian Wildl. Serv. Rep.
Ser. 21. 36 pp.
March, J. 1977. Courage and the art of wolf main-
tenance. Audubon 79(6):80-113.
McBride, R. T. 1978. Status of the gray wolf (Ca-
ms lupus baileyi) in Mexico. A progress report
to the U.S. Fish and Wildlife Service. Contract
14-16-0002-3728. 72 pp.
Mech, L. D. 1966. The wolves of Isle Royale. U.S.
Dep. Inter. Natl. Park Serv. Fauna Ser. 7,
240 pp.
1970. The wolf: the ecology and behav-
ior of an endangered species. Doubleday, New
York. 389 pp.
. 1973. Wolf numbers in the Superior Na-
tional Forest of Minnesota. North Central
For. Exp. Sta., USDA, For. Serv. Res. Pap.
NC-97. 10 pp.
. 1974a. Canis lupus. Mammalian Species
37. Am. Soc. Mammal, pp. 1-6.
1974b. A new profile for the wolf. Nat.
Hist. 83(4):26-31.
. 1975. Hunting behavior in two similar
species of social canids. Pages 363-368 in M.
W. Fox, ed. The wild canids. Van Nostrand
Reinhold Co., New York. 508 pp.
. 1977a. A recovery plan for the eastern
timber wolf. Natl. Parks Conserv. Mag.
1977(1):17-21.
. 1977b. Where can the wolf survive? Natl.
Geogr. Mag. 152(4):518-537.
. 1977c. Wolf-pack buffer zones as prey
reservoirs. Science 198:320-321.
. 1977d. Productivity, mortality, and
population trends of wolves in northeastern
Minnesota. J. Mammal. 58(4):559-574.
Mech, L. D., and L. D. Frenzel. 1971. Ecological
studies of the timber wolf in northeastern
Minnesota. U.S. Dep. Agric. For Serv. Res.
Pap. NC-52. 62 pp.
Mech, L. D., and P. D. Kams. 1977. Role of the
wolf in a deer decline in the Superior National
Forest. USDA For. Serv. Res. Pap. NC-148.
23 pp.
Medjo, D. C, and L.D. Mech. 1976. Reproductive
activity in nine- and ten-month-old wolves.
J. Mammal. 57(2):406-408.
Murie, a. 1944. The wolves of Mt. McKinley.
Fauna of the National Parks of the United
States. U.S. Dep. Inter. Natl. Park Serv.
Fauna Series 5. pp.
Northern Rocky Mountain Wolf Recovery Team.
1978. Draft plan for the recovery of the
northern Rocky Mountain wolf. 40 pp.
Olson, S. F. 1938. A study in predatory relation-
ship with particular reference to the wolf. The
Sci. Monthly 46:323-336.
Peters, R. P., and L. D. Mech. 1975. Scent-marking
in wolves. Am. Sci. 63(6):628-637.
Pimlott, D. H. 1967. Wolf predation and ungulate
populations. Am. Zool. 7(2):267-278.
. 1975. The ecology of the wolf in North
America. Pages 280-285 in M. W. Fox, ed.
The wild canids. Van Nostrand Reinhold Co.,
New York. 508 pp.
Rausch, R. A. 1967. Some aspects of the popula-
tion ecology of wolves, Alaska. Am. Zool.
7(2):253-265.
Riley, G. A., and R. T. McBride. 1972. A survey
of the red wolf (Canis rufus). U.S. Fish Wildl.
Serv. Spec. Sci. Rep. -Wildl. 162. 15 pp.
Rutter, R. J., and D. H. Pimlott. 1968. The world
of the wolf. Lippincott, Philadelphia. 202 pp.
10
Ryon, J. C. 1977. Den digging and related behav-
ior in a captive timber wolf pack. J. Mammal.
58(l):87-89.
Schenkel, R. 1947. Expression studies of wolves.
Behavior 1:81-129.
Stebler, A. M. 1944. The status of the wolf in Mi-
chigan. J. Mammal. 25:37-43.
Stenlund, M. H. 1955. A field study of the timber
wolf (Cams lupus) on the Superior National
Forest, Minnesota. Minn. Dep. Cons. Tech.
Bull. 55 pp.
Thompson, D. Q. 1952. Travel, range, and food
habits of timber wolves in Wisconsin. J. Mam-
mal. 33(4):429-442.
Weise, T. F., W. L. Robinson, R. A. Hook, and L.
D. Mech. 1975. An experimental translocation
of the eastern timber wolf. Audubon Conser.
Rep. 5 in cooperation with U.S. Dep. Inter.,
Fish Wildl. Ser. Region 3. 28 pp.
Wolfe, M. L. 1977. Mortality patterns in the Isle
Royale Moose Population . Am. Midi. Nat.
97(2):267-279.
Wolfe, M. L., and D. L. Allen. 1973. Continued
studies of the status, socialization, and rela-
tionships of Isle Royale wolves, 1967 to 1970.
J. Mammal. 54(3):61 1-635.
Young, S. P., and E. A. Goldman. 1944. The
wolves of North America. 2 vols. Am. Wildl.
Inst., Washington, D.C. 636 pp.
Zimen.E. 1975. Social dynamics of the wolf pack.
Pages 336-363 in M. W. Fox, ed. The wild ca-
nids. Van Nostrand Reinhold Co., New York.
508 pp.
PREPARED/UPDATED BY
National Fish and Wildlife Laboratory
412 N.E. 16th Avenue, Room 250
Gainesville, Florida 32601
11
Biological Services Program
FWS/OBS-80/01.54
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
THICK BILLED PARROT
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubhc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the scacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed lo:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.58
11
FWS/OBS-80/01.54
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
THICK BILLED PARROT
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT: SAN DIEGO ZOO
THICK-BILLED PARROT
Rhynchopsitta p . pachyrhyncha (Swainson)
Rhynchopsitta p. terrisi Moore 1947
KINGDOM Animalia
ORDER Psittachiformes
FAMILY Psittacidae
OTHER COMMON NAMES
R.p. pachyrhyncha Cotora serrana,
guacamayo, guacomaja
R.p. terrisi Maroon-fronted parrot,
cotora fiente purpurea,
guacamaya, papagayo
DATE
Entered into SWIS to be determined
Update to be determined
LEGAL STATUS
Federal: Endangered. Fed. Reg. Vol. 42, No.
135, 14 July 1977, p. 36427. Listed
Appendix I, Convention on Interna-
tional Trade in Endangered Species of
Wild Fauna and Flora (Federal Register,
Vol. 42, No. 35, 22 February 1977, p.
10478).
States:
REASONS FOR CURRENT STATUS
Cutting of mature mountain pine forests, de-
priving the birds of food and nesting sites, is the
primary cause of the decline of the thick-billed
parrot (Vincent 1967, Monson 1965, Ridgley
WWF Report, Crossin pers. comm., W. King
1977).
Killing of birds and taking of eggs for food,
sport, or specimens is known to occur but is cer-
tainly of minor significance because of the inac-
cessibility of areas where these birds live and the
difficulty of finding nests (Ridgley WWF Report,
Crossin pers. comm., King 1977, Lusk 1900,
Wetmore 1935, Thayer 1906, and Bergtold 1906).
Capture for zoos and for pets occurs but is
also a minor cause of decline (Lanning and Law-
son pers. comm., Crossin pers. comm.. King in
press, Wetmore 1935). Apparently, this species is
not used much in the commercial cage bird trade,
as it is not listed as imported into the U.S. in
recent years (Clapp 1975).
Forest fires and logging are considered the
greatest potential threats to R. p. terrisi at present
in southeast Coahuila (Lanning and Lawson pers.
comm.).
PRIORITY INDEX
Not assigned.
DESCRIPTION
R. p. pachyrhyncha is mostly dull olive green;
the forehead, lores, and a stripe extending along
the sides of the crown, bend of wings, and thighs
are deep red; the tail is long and pointed, outer
feathers graduated. In flight, the bright yellow
patch on under-wing converts is conspicuous.
The bin is black and notably thick. The iris is red-
dish. R. p. terrisi is larger and darker green; the
forehead and supercilliary stripes are brownish
maroon (instead of bright red); the patch on the
underwing converts is brownish gray (not bright
yellow as in R. p. pachyrhyncha) (Blake 1953).
All color characters of pachyrhyncha and terrisi
are highly variable and not definitely distinguish-
ing, except color of under primary converts,
which are bright yellow in pachyrhyncha and
olive yellow in terrisi; terrisi is also larger (Hardy
and Dickerman 1955). All examples of terrisi seen
at close range in the field have had maroon rather
than scarlet foreheads, but five adults in the Uni-
versity of Kansas collection have a few scarlet
feathers on forehead (Ely 1962).
Length, 16 - 16.7 in. (41-42 cm); wing, 8.5 -
10.5 in. (22 - 27 cm); taU, 6.3 - 7 in. (16 - 18 cm);
bill length, 1.4 - 1.5 in. (3.6 - 3.8 cm);biU height,
1.6 - 1.7 in. (4-4.3 cm). Bill relatively very large
and greatly compressed laterally (Bailey 1938).
Weight: terrisi, SW, Coahuila, 2 males, 391.5 and
467.5 gms; female, same area, 466 gms (Urban
1959).
Whether pachyrhyncha and terrisi are con-
sidered as two distinct species or two weU-marked
subspecies of a single species is a matter of opin-
ion. Prior to Dickerman and Hardy(1955) all au-
thors who reported specimens with mixed charac-
ters considered them 2 distinct species. Since then,
subspecific status has been accepted by Dicker-
man and Hardy (1955), Ely (1962), Forshaw
(1973), Urban (1959), Vincent (1967), and King
(1977). However, Hardy (1967) changed his
mind on the basis of the hypotheticail importance
of the difference in shade of red of the forehead,
which he thought might be significant in social
recognition of the birds belonging to the two
populations, serving as effective reproductive
isolating mechanisms, thus making them two dis-
tinct species by definition.
RANGE
R. p. pachyrhyncha formerly bred in high
coniferous forests in Sierra Madre Occidental of
western Mexico from northwestern Chihuahua
(Colonia Pacheco and Colonia Garcia, Thayer
1906) south to southern Durango (Canyon Rio
San Juan, Bent 1940), from 4,000 to 10,000 ft
(1,219 - 3,048 m) (Bergtold 1906). Within the area
reported from Sierra Huachinera (Marshall 1957),
Cumbre on Barranca Cobre, SW Chihuahua (Stager
1954), Mt. Mohinora, southwest of Verjel (Fried-
mann et al. 1950), mountains west of Parral (Berg-
told 1906), Guadalupe y Calvo (Bent 1940), all in
Chihuahua, and the mountains near Ciudad Dur-
ango (Bent 1940) in Durango. They formerly wan-
dered extensively, chiefly in winter, or when pine
cone crops failed, to mountains of southeastern
Arizona and southwestern New Mexico, and, cen-
turies ago, even to central and northern Arizona
(Verde Valley and San Francisco Mountain). The
chief areas visited included the Chiricahua Moun-
tains, and occasionally nearly all the higher moun-
tains east and south of the Santa Cruz and Gila
Rivers, Arizona, and also in Animas Mts., New
Mexico (Phillips et al. 1964;Wetmore 1935; Bent
1940; Smith 1907; Vorhies 1934; Bailey 1928).
The most recent U.S. record in Animas Mts. New
Mexico was in the fall of 1964 (Charles Hanson,
pers. comm. 1977). They also wandered eastward
from the breeding area to Ciudad Chihuahua
(Bent 1940), and possibly to San Antonio de las
Alazanas, Coahuila (Ely 1962). Long movements
from the breeding area southward and southeast-
ward to Volcanesde Colima, Jalisco (Schnell et al.
1974), Cerro de Tancitaro, Michoacan (Blake and
Hanson 1942), Mt. Popocatapetl, State of Mexico
(Bent 1940), and Perote and Jalapa, Vera Cruz
(Bent 1940). At present, the entire range is evi-
dently much restricted.
R. p. terrisi is assumed to have bred formerly
in high coniferous forests of the Sierra Madre
Oriental, from southeastern Coahuila north to
west of Saltillo (Irby Davis, pers. comm.). It now
breeds east of San Antonio de las Alazanas south
of Saltillo (Ely 1962), and from Sierra Zapaliname
in the Guadalupe Range (Burleigh and Lowery
1942) through southern Nuevo Leon and Cerro
Potosi (Moore 1947) ; south to Sierra de Guatemala
in the Gomez Farias region, Tamaulipas (Ridgley
WWF Report); and La Joya de Salas, 65 mi. (105
km) south southwest of Victoria, Tamaulipas
(Robins and Heed 1951). Altitudes of observation
ranged from 1,829 to 3,658 m (Moore 1947; Ely
1962; Lanning & Lawson pers. comm.). There is
no evidence of terrisi wandering extensively, as
R. p. pachyrhyncha did.
RANGE MAP
Shown on next page.
STATES/COUNTIES
Arizona: Cochise, Graham, Pima, Pinal,
Santa Cruz, Yavapai.
New Mexico: Hidalgo.
HABITAT
Mature pine-oak, pine, and fir forests are pre-
ferred habitat, in that order with increasing alti-
tude, 1,219 to 3,658 m in mountains. The species
is found in locations varying from plateau-like
tops of mountains with open pine or pine-oak
woodland and large trees and grass below (Mon-
son 1965; Marshall 1957), to pine and fir forests
below high cliffs or rimrock outcrops that are
used by the parrots for roosting and nesting. They
are assumed to require either large dead tree
stumps or high cliffs with holes for nesting.
They have been observed most frequently
in stands of mature pine at median altitudes.
Chihuahua Pine {Pinus chihuahuana) appears
to be the most common species in occupied
habitats although a number of pine and oak
species are utilized.
FOOD AND FORAGING
Pine seeds are by far the most important food.
Seeds of both large forest pines and the smaller
lower altitude pinons are eaten, along with some
acorns (Ridgley WWF Report; Lanning & Lawson,
pers. comm.; Ely 1962, Lusk 1900, Wetmore
1935; Leopold 1937). Other foods eaten occa-
sionally are terminal buds of Chihuahua and Lum-
holtz Pines, at 8,200 ft (2,499 m) in Barranca de
Cobre, Chihuahua, 13 to 21 May (Stager 1954);
fruit of a cherry (Prunus copuli) and seeds of an
unidentified legume at Cerro de Tancitaro, Michoa-
can (Blake and Hanson 1942); and juice or nectar
of agave flowers in southeastern Coahuila (Ely
1962, Lanning & Lawson pers. comm.). Captive
birds at the Arizona-Sonora Desert Museum re-
mained healthy on standard zoo parrot food, but
did not breed unless fed pinon seeds (Charles
Hanson pers. comm. 1977).
The method of feeding on pine seeds was to
snip off a cone with the bill and support it with
a foot while extracting individual seeds with the
bill (Ridgley WWF Report), or to pull bracts from
the cone and extract seeds while the cone was still
on the tree (Lanning & Lawson pers. comm.).
Seeds were even extracted from immature green
cones. Bills and breast feathers were found smeared
with pitch (Wetmore 1935, Lusk 1900). Acorns
were used in the Chiricahua Mts., Arizona, through
fall and winter after the pine seed crop was ex-
hausted (Wetmore 1935, Leopold 1937). Parrots
go to rivers or waterfalls to drink toward evening
before going to roost (Wetmore 1935, Marshall
1957). They have been seen "eating" snow and
ice on Cerro Potosi in 1977 (P. T. Moore to Lan-
ning and Lawson pers. comm.).
SHELTER REQUIREMENT
Cavities high up either in dead trees or in
cliffs are used for roosting and nesting (Thayer
1906, Bergtold 1906, Stager 1954, Lanning and
BREEDING RANGE C
R.p. terrisi
Breeding range of the thick-billed parrot.
Lawson pers. comm., Leopold 1937).
NESTING AND BEDDING
The only occupied nests reported were in cav-
ities high up in large dead pines in the Sierra Madre
Occidental of Chihuahua, at least some of which
were abandoned imperial woodpecker nests
(Thayer 1906, Bergtold 1906). According to resi-
dents in southeast Coahuila, they used cliffs and
possibly trees for nesting. There are few suitable
trees, but many cliffholes in the region (Lanning
and Lawson, pers. comm., Burleigh and Lowery
1942); they slept in higher cliffs of high barranca
at Palmito, Sinaloa, during May 1964 (R. Crossin
pers. comm. to W. King 1977). In the morning,
flocks leave the roost in the rimrocks in Sierra
Madre of Chihuahua (Leopold 1937). Eggs were
found (10 records) 10 May to 25 Aug. 1905 and
(7 recoids) 11 to 25 Aug. (Thayer 1906, Bergtold
1906, Bent 1940). They are not dependent on
holes made by the nearly extinct imperial wood-
pecker for nesting. Parrots are capable of re-
modeling a natural cavity or hole (Ridgley WWF
Report , Charles Hanson pers. comm. 1977).
RITUAL REQUIREMENTS
The courtship display, raising the wings to
expose under surfaces, may effect reproductive
isolation of pachyrhyncha and terrisi, because of
the marked difference in color of under primary
coverts. This display is accompanied by distinc-
tive vocalizations (Charles Hanson pers. comm.
1977). The habit of soaring in circles high in the
air like a hawk has been noted in both R. p.
pachyrhyncha and R. p. terrisi (Marshall 1957,
Ridgley WWF Report); in terrisi, the purpose is to
gain altitude for long flights (Lanning & Lawson
pers. comm.). When flying in flocks grouped by
pairs, two birds fly very close together, occasion-
ally with a third behind. Several feet separate pairs;
sometimes flocks fly in V-formation. They are
very noisy and their cries carry great distances,
probably facilitating communication between
wide-ranging individuals (Marshall 1957).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None known other than those given under
food, nesting and bedding, and habitat.
POPULATION NUMBERS AND TRENDS
R. p. pachyrhyncha. - Apparently is much re-
duced from Its former population; it is seen less
frequently and in smaller numbers in the heart of
its range in the Sierra Madre Occidental and no
flights have been observed in the southwestern
U.S. since the early 1900's. Pubhshed estimates
include 100 at Palmito, Sinaloa, 7 May 1964; 200,
same place, 10 May (R. Crossin pers. comm.),
flocks of 60 and 1 5 , at Volcanes de Colima, Jalisco,
6 Jan. 1973 (Schnell et al. 1974); about 300 in
the Chiricahua Mts., Ariz., 1 Sept. 1917; 100-
1500, same place, early fall, 1917; 150, same
place, middle of May through summer and early
faU 1917 (Wetmore 1935); 35, Cumbre, Barranca
deCobre, Chihuahua, 13 May 1950 (Stager 1954);
Flocks of 50, 60 and 8 in the Sierra Huachinera,
Chihuahua, 1951 and 1952; populations shift
from year to year (Marshcdl 1957). During an ex-
tensive survey of northern Mexico 11 Oct. to 31
Dec. 1971 and 2 Mar. to June 1972, with consid-
erable time spent in thick-billed parrot habitats,
only a few were seen high overhead (R. Crossin
pers. comm. to W. King).
R. p. terrisi. - There is not much evidence of a
total decline in numbers of this subspecies, since
its existence has been known only since 1947 and
recent estimates seem as high as earlier ones. How-
ever, because of its restricted range, it probably
never had a large population and a continually de-
creasing habitat would certainly result in a popu-
lation decline. Population estimates include 91 at
La Mesa de las Tables, Coahuila; 15 and 60 at
Cerro Potosi, Nuevo Leon (Ridgley WWF Report);
400 to 500, near Saltillo, Coahuila, 1977; esti-
mate on the order of 1500 in entire Saltillo, Coa-
huila area (Dirk Lanning pers. comm. to W. King);
count of 800 to 1000 in Southeast Coahuila, 4
April 1977, (Lanning & Lawson pers. comm.).
REPRODUCTION
Following are records of nests with eggs or
young of R. p. pachyrhyncha : Collected by W. W.
Brown at Colonia Pacheco and Colonia Garcia,
Chihuahua: 9 nests, all in holes in dead pines,
heights above ground 50-80 ft (15 to 24 m) av. 71
ft (22 m); depth of nest cavity 18 to 28 in (46 to
71 cm) av. 24 in (70 cm); width of cavity 6-10 in
(15 to 25 cm) av. 8 in (20 cm); diameter of en-
trance (3 nests all) 6 in (15 cm); number of eggs
or young 1 to 3 (av. 2 eggs or young per nest).
Eggs glossy white, shell very thick for size of eggs,
which averaged 1.53 x 1.19 in (3.88 x 3.02 cm)
(Thayer 1906).
A single egg hatched in captivity after
28 days of incubation (Lint 1966 in Forshaw
1973). Captive young were brooded by the fe-
male for the first 11 days after hatching; young
left nest 59 days after hatching (Dyson 1969 in
Forshaw 1973). At the Arizona-Sonora Desert
Museum, 2 pairs have bred successfully, 1 pair in
2 successive years. They began nesting in July,
but may start later. A clutch is usually 2 eggs
(Charles Hanson pers. comm. 1977).
No nests with eggs or young of R. p. terrisi
have been reported. At La Mesa de las Tablas,
Coahuilain a flight of 91 parrots, pairs were always
in evidence, but many of the pairs seemed to be
accompanied by offspring, most often 2, some-
times only 1 and once 3. The groups of 3, 4, and
5 birds were about as frequent as unaccompanied
pairs, indicating that breeding success was good
(Ridgley WWF Report).
MANAGEMENT AND CONSERVATION
Ideas that have been presented are: To set
aside and protect adequate areas of mature ever-
green mountain forest in both the Sierra Madre
Occidental and Sierra Madre Oriental (Vincent
1966). Preservation of the rapidly disappearing
mature high mountain forests is the only obvious
method to preserve the species (Ridgley WWF
Report;King 1977).
The species is Hsted in Appendix 1 of 1973
Convention on International Trade of Endangered
Species of Wild Fauna and Flora. It is protected
by law in Mexico, but the law is unenforceable in
practice because of the remoteness of the areas
where birds still occur. In 1976, 76 Thick-billed
Parrots {R. p. pachyrhyncha) were in captivity in
24 collections; of these, 10 (13%) were bred in
captivity (Olney 1976 in King 1977 ms.). Captive
rearing might be a means of aiding the survival of
the species until adequate wild populations are
assured.
PREPARER'S COMMENTS
It would appear that both subspecies of the
thick-billed parrot are endangered because of the
steady eUmination of the high mountain conifer-
ous forests of northern Mexico, on which the
birds are completely dependent because of their
specialized feeding and nesting requirements:
large trees for nesting and pine seeds for nutrition
adequate for reproduction. There would seem to
be no remedy for this except the prompt setting
aside of a number of very large tracts of such
habitat where the parrots still occur, and exclud-
ing lumbering in those tracts. It is essential to
have a number of different areas because of the
periodic failure of pine cone crops in a particular
area. Much further investigation of nesting behav-
ior is needed, particularly to determine the im-
portance of cliff nest sites as compared with holes
in dead trees.
AUTHORITIES
Robert S. Ridgley
Department of Zoology
Duke University
Durham, North Carolina 27706
John W. Hardy
Florida State Museum
University of Florida
Gainesville, Florida 32611
Robert W. Dickerman
Department of Microbiology
Cornell University Medical College
1300 York Avenue
New York, New York 10021
L. Irby Davis
2502 Keating Lane
Austin, Texas 78703
Warren B. King
871 Dolly Madison Blvd.
McLean, Virginia 22101
Charles A. Ely
Department of Zoology
Fort Hayes Kansas State College
Hayes, Kansas 67602
Joe T. Marshall
National Fish and Wildlife Laboratory
National Museum of Natural History
Washington, D.C. 20560
Charles L. Hanson
Arizona-Sonora Desert Museum
Tucson, Arizona
Richard S. Crossin
1719N. Huachuca
Tucson, Arizona 85705
Dirk Lanning and Peter Lawson
Chihuahuan Desert Research Institute
Box 1334
Alpine, Texas 79830
LITERATURE CITED/SELECTED
REFERENCES
Bailey, F. M. 1928. Birds of New Mexico. N. Mex.
Dept. of Game and Fish, and U.S. Biol. Surv.
Bent, A. C. 1940. Life histories of North Ameri-
can cuckoos, goatsuckers, hummingbirds and
their aUies. U.S. Nat. Mus. Bull. 176:(Thick-
billed Parrot 14-18).
Bergtold, W. H. 1906. Concerning the Thick-billed
Parrot. Auk 23:425-428.
Blake, E. R. 1953. Birds of Mexico. Univ. Chicago
Press. 644 pp.
Blake, E. R., and H. C. Hanson. 1942. Notes on a
collection of birds from Michoacan, Mexico.
Publ. Field Mus. Nat. Hist. Zool. Ser. 22:513-
551.
Burleigh, T. D., and G. H. Lowery. 1942. Notes
on the birds of southeastern Coahuila, Occ.
Pap.Mus. Zool. La. State U. 12:185-212.
Clapp, R. B. 1975. Birds imported into the United
States in 1972. Spec. Sci. Rep. Wildlife 193.
U.S. Fish and Wildlife Service.
Dyson, R. F. 1969. Captive hatching and develop-
ment of a Thick-billed Parrot at Arizona-
Sonora Desert Mus. Int. Zoo Yearbook No.
9:127-129 (in Forshaw 1973).
Ely, C. A. 1962. The birds of southeastern Coa-
huila, Mexico. Condor 64:34-39.
Flemming, R. L., and R. H. Baker. 1963. Notes
on the birds of Durango, Mexico. Pubis. Mich.
St. Univ. Mus. Biol. Ser. 2:275-303 (in For-
shaw 1973).
Forshaw, J. M. 1973. Parrots of the world. Garden
City: Doubleday and Co. [Rhynchopsitta p.
411).
Friedmann, H., L. Griscom, and R. J. Moore.
1950. Distributional check-list of the birds of
Mexico, Part 1. Pacific Coast Avifauna 29, p.
127.
Hardy, J. W. 1967. Rhynchopsitta terrisi is prob-
ably a valid species: a reassessment. Condor
69:527-528.
Hardy, J. W., and R. W. Dickerman. 1955. Tax-
onomic status of the Maroon-fronted Parrot.
Condor 57:305-306.
King, W. 1977 (Revised) Red data book. Survival
Service, International Union for the Conserv-
ation of Nature and Natural Resources,
Morges.
Leopold, A. 1937. The Thick-billed Parrot in
Chihuahua. Condor 39:9-10.
Lint, K. C. 1966. Thick-billed Parrots breed in
captivity. Zoonooz 39(2) :3-6 (in Forshaw
1973).
Lusk, R. D. 1900. Parrots in the United States.
Condor 2:129.
Marshall, J. T. 1957. Birds of the pine-oak wood-
land in southern Arizona and adjacent Mexico.
Pac. Coast Avifauna 32:1-125.
Monson, G. 1965. A pessimistic view— the Thick-
billed Parrot. Audubon Field Notes 19(3):
389.
Moore, R. T. 1947. New species of parrot and
race of quail from Mexico. Proc. Biol. Soc.
Wash. 60:27-28.
Olney, P. J. S. (ed.). 1976. Census of rare animals
in captivity 1975. Inter. Zoo Yearbook 16:
411-446 (in King 1977 ms.).
Phillips, A., J. Marshall, and G. Monson. 1964.
The birds of Arizona. Univ. of Arizona Press,
Tucson.
Ridgley, R. Unpubl. interim report for 1976.
Macaw status survey. World Wildlife Fund,
U.S. App.Proj.61.
Robbins, C. R., and W. B. Heed. 1951. Bird notes
from La Joya de Salas, Tamaulipas. Wilson
BuU. 63:263-270.
SchneU, G. D., J. S. Weske, and J. J. HaUack.
1974. Recent observations of Thick-billed
Parrots in Jalisco. Wilson Bull. 86:464-465.
Smith, A. P. 1907. The Thick-bUled Parrot in
Arizona. Condor 9:104.
Stager, K. E. 1954. Birds of Barranca de Cobre
region of southwestern Chihuahua, Mexico.
Condor 56:21-32.
Thayer, J. E. 1906. Eggs and nests of the Thick-
biUed Parrot. Auk 23:223-225.
Urban, E. 1959. Birds from Coahuila, Mexico. U.
of Kansas Pubis. Mus. Nat. Hist. 11:443-516.
Vincent, J. 1967. Red Data Book. Vol. 2. Survival
Services Commission lUCN Morges, Switzer-
land.
Vorhies, C. T. 1934. Arizona records of the Thick-
billed Parrot. Condor 36:180-181.
Wetmore, A. 1935. The Thick-billed Parrot in
southern Arizona. Condor 37:18-21.
Biological Services Program
FWS/OBS-80/01.55
MARCH 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
SAN CLEMENTE SAGE SPARROW
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Endangered Species Act of 1973, as dmendcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704.58
u
FWS/OBS-80/01.55
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
SAN CLEMENTE SAGE SPARROW
A Cooperative Effort
by the
National Fish and WildUfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
.-^ v-j;
>H- .:-
UV
I
fc:'
y
1 1<
PHOTO OF CLOSELY RELATED SUBSPECIES
1ERBERT CLARKE
SAN CLEMENTE SAGE SPARROW
Amphispiza belli clementeae Ridgway (1898)
KINGDOM Animalia
CLASS Aves
ORDER Passeriformes
FAMILY Fringillidae
OTHER COMMON
NAMES Bell Finch; Bell Sparrow;
San Clemente Sparrow
DATE
Entered into system To be determined
Updates To be determined
LEGAL STATUS
Federal: "Threatened." Fed. Reg. Vol. 42, No.
155, 11 August 1977, p. 40685. Pro-
tected by Migratory Bird Treaty Act.
States : Protected by California state law.
REASONS FOR CURRENT STATUS
Stewart and Clow (1974) state that since
1934, sheep and goats, formerly fenced, have
roamed the island uncontrolled, practically elim-
inating reproduction of native shrubs. Goats eat
seedlings and young plants and strip the leaves
and bark from the lower portions of trees and
bushes. No seedling or young bushes were found
at any place visited. Reduction of ground cover
has led to erosion of topsoil necessary for seedlings
to develop. Feral swine have furthered erosion by
rooting beneath what topsoil is left. The reduc-
tion in plant cover may have facilitated predation
on sage sparrows and their nests and eggs by the
island fox and feral house cats. It would also have
greatly reduced the dense bush habitat the sage
sparrows require for nesting and feeding.
PRIORITY INDEX
None assigned.
DESCRIPTION
A. b. clementeae is a small gray bird with
black streaks on sides and single black spot on
chest, dark cheeks and "moustache" streaks on
sides of throat, white line over eye, and white
corners on tail.
This is a weakly differentiated race. Some of
the alleged characteristics are not present in series.
A longer bill and lighter ju venal plum^e seem to
be the only characteristics that are reasonably
constant in separating it from the mainland race,
A. b. belli (van Rossem 1932, Grinnell and Miller
1944).
RANGE
These birds are permanent residents on and
confined to San Clemente Island about 80 km off
southwestern California (American Ornithologists'
Union 1957, Miller 1968). San Clemente is ap-
proximately 34 km long, 2.4 to 6.4 km wide, and
34 km from the nearest island, Santa Catalina, to
the north. A. b. clementae was thought to prob-
ably occur also on San Nicholas and Santa Rosa
Islands, although specimens from those islands
were never critically determined (Grinnell and
Miller 1944).
RANGE MAP
See map on following page.
STATES/COUNTIES
California: Los Angeles.
HABITAT
The San Clemente sage sparrow habitat con-
sists of xeric species of scrubby brush on mesas
(Howell 1917) and thorny brush growing in
clumps and patches interspersed with cactus
(Grinnell 1897). Frost is unknovm and conse-
quently vegetation was rank most of the year,
jdthough there was no fresh water on the island
(Breninger 1904).
Topography is dominated by a plateau with
steeply sloping east side and gently sloping west
side, now covered with introduced grasses except
where it has been completely denuded by goats.
At present, trees and shrubs grow only on the
bottoms and sides of canyons. On the east side,
ironwood (Lynothamus), island oak (Quercus
tomentella), and lemonade bush {Rhus inter-
grifolia) are the most abundant species. West side
canyons are mostly denuded by livestock, but
have some growth of toyon (Heteromeles), lemon-
ade bush, and island cherry {Prunus ilicifolia lyonii)
(Stewart and Clow 1974).
FOOD AND FORAGING BEHAVIOR
These birds forage on the ground or low in
bushes, presumably for ^mall s.--ds and insects.
Miller (1968) saw a female of ^. b. belli on the
coast of California c u. /ing four green caterpillars
obviously intended for young birds. Since fresh-
water is very scarce or absent on San Clemente
(Breninger 1904), sage spa..jws must dej. end pri-
marily on dew or fog for drinking.
SHELTER REQUIREMENTS
Low scrubby brush is presumed to be the re-
quired shelter for /i. b. clementeae, by analogy to
the California coast subsf. jcies, A b. belli, v "ch
prefers dense and continuous brush within which
it finds all of its requirements (M.ller 1968).
NESTING AND BEDDING
These birds feed and nest among the bushes
of the hillsides, along with song sparrows (Brenin-
ger 1904). Nests are situated in scrubby brush a
few inches above the ground (Howell 1917). A
nest of A. b. belli was found on the mainland in
the fork of a heavy bush at practically ground
level. It was well made of weed stalks and lined
with fine weed stems and soft weed fibers (Miller
1968).
RITUAL REQUIREMENTS
The sage sparrow presumably maintains and
defends breeding territory, proclaimed by the song
of the male, as in other related species of sparrows.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
None are known other than those noted else-
where.
POPULATION NUMBERS AND TRENDS
Grinnell (1897) wrote that these sparrows
were quite common on hillsides and the lower
mesa where there was low thorny bush growing in
clumps and patches interspersed with cactus.
Howell (1917) considered it "common" on
mesa lands back from the shore.
It is now mostly confined to the lower west
terrace. On 3 May 1974, a total adult population
of between 20 and 30 individuals was estimated
in that area. No goats were seen in the area of ob-
servation, but feral swine were seen in two dif-
ferent localities (Stewart and Clow 1974).
SAN CLEMENTE SAGE SPARROW
SAN FRANCISCO
CALIFORNIA
'0^=^*
LOS ANGELES
^
/
SAN CLEMENTE ISLAND
TOTAL RANGE OF
Amphispiza belli clementeae
REPRODUCTION
Nests with pipped eggs and young were found
in the latter part of March 1915 (Howell 1917).
If similar to the coastal race (belli), nests with
(usually) 4 eggs would be found chiefly in April
and May, and pairs would be spaced an average of
about 50 yards (46 m) apart (Miller 1968).
MANAGEMENT
The obvious procedure in management of the
Sam Clemente Sage Sparrow would be to remove
all goats and swine from the island, or at least to
fence them away from large enough sections of
the plateau to permit regrowth of the dense low
scrub necessary for the existence of this species of
sparrow.
AUTHORITIES
Robert M. Stewart and William C. Clow
Point Reyes Bird Observatory
Box 321
Bolinas, California 94924
PREPARER'S COMMENTS
1 agree with Stewart's and Clow's appraisal in
their 1974 report that the feral goats are a men-
ace, not only to the San Clemente Sage Sparrovv,
but all of the brush-inhabiting birds of the island,
and that the only way to prevent the extinction
of several species is to remove or drastically con-
fine the goats and give the vegetation a chance to
come back. The U.S. Navy that controls the is-
land should be encouraged to take all steps pos-
sible to achieve such goat control.
LITERATURE CITED/SELECTED
REFERENCES
American Ornithologists' Union. 1957. Check-list
of North American Birds, 5th Ed. Baltimore,
Md. Amer. Ornith. Union. 591 pp.
Breninger, G. F. 1904. San Clemente Island and
itsbirds. Auk 21:218-223.
Grinnell, J. 1897. Report on the birds recorded
during a visit to the islands of Santa Barbara,
San Nicholas, and San Clemente, in the spring
of 1897. Pasadena Acad. Sci. Publ. 1.
Grinnell, J. and A. H. Miller. 1944. The distribu-
tion of the birds of California. Pac. Coast Avi-
fauna 2 7 .
Howell, A. B. 1917. Birds of the islands off the
coast of southern California. Pac. Coast Avi-
fauna. 12:79.
Miller, A. H. in Bent, A. C. 1968. Life histories
of North American cardinals, grosbecks, bunt-
ings, towhees, finches, sparrows and allies,
part 2. U. S. Nat. Mus. Bull. 237:1015-1020.
Stewart, R. M. and W. C. Clow. 1974. Part I San
Clemente Island, in "The status of the Song
Sparrow and Bewick's Wren on San Clemente
Island and Santa Barbara Island, California."
by R. M. Stewart, J. Small, W. C. Clow and
R. P. Henderson. Rep. to Endangered Species
Off. U. S. Fish and Wildl. Ser. by Point Reyes
Bird Observatory.
van Rossem, A. J. 1932. On the validity of the
San Clemente Island BeU's Sparrow. Auk 49:
490-491.
Biological Services Program
FWS/OBS-80/01.56
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
CALIFORNIA CLAPPER RAIL
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubUc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as aincndcd).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SVVIS) developed by the U.S. Army Corps of Knginccrs in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of F^ndangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 704''>8
u
FWS/OBS-80/01.56
March 1980
SELECTLD VLR lEBRA 1 K ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
CALIFORNIA CLAPPER RAIL
A Cooperative Effort
by the
National Fish and WildHfc Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald VV. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT; JACK WHITE. CALIF. FISH AND GAME COMM.
CALIFORNIA CLAPPER RAIL
Rallus longirostris obsoletus Ridgway
KINGDOM Animalia
CLASS Aves
ORDER Gruiformes
FAMILY Rallidae
OTHER COMMON
NAMES California King Rail,
Red-breasted Rail, Marsh Hen (Grin-
nell and Miller 1944).
DATE
Entered into SWIS to be determined
Update to be determined
LEGAL STATUS
Federal: Endangered: (42 FR 36427, 14 July
1977). Migratory Bird Treaty Act of 3
July 1918 (40 Stat. 755; 16 U.S.C.
703-711) as amended 3 Dec. 1969.
Public Law 91-135.
State: Endangered: California.
REASONS FOR CURRENT STATUS
Originally abundant locally within its limited
range, California clapper rail numbers and the
extent of its territory were much reduced before
market hunting was outlawed. Under legal pro-
tection, its population increased, and by 1944,
much of the former territory had been regained.
Exceptions were extensive areas of San Francisco
Bay marsh that had been converted to human use
(Bryant 1915, De Groot 1927, Grinnell and Miller
1944, Wilbur and Tomlinson 1976).
This rail is vulnerable because the population
is divided into small, isolated colonies occupying
islands of suitable habitat that are separated by
long stretches of uninhabitable rocky or sandy
coast (Van Rossem 1929). There has been no in-
dication of change in status recently, except for
continuous attrition of the most productive habi-
tat in south San Francisco Bay by conversion of
marshes for other uses (Wilbur and Tomlinson
1976). The coastal marsh south of San Francisco
1
Bay at Pescadero Creek was made unsuitable for
habitat by the closing of its connection to the
sea by a sand bar; at Watsonville Slough, habitat
was destroyed by a housing development with
associated pollution. Some uninhabited marshes
in north San Francisco Bay are contaminated
with oil scum (Gould 1973).
PRIORITY INDEX
16
DESCRIPTION
R. I. obsoletus is a chicken-sized bird; grayish
brown above with light ocraceous buff or tawney
breast, flanks of grayish brown barred with white,
and a white patch under the short, upcocked tail.
Two color phases occur, one more brownish, the
other more olivaceous. The iris is reddish orange;
the bUl is orange yellow at the base with the rest
brown; the legs and feet are brownish gray. Downy
young are jet black with greenish olive above. The
legs are long (57 mm); toes 51 mm; bill 59 mm. It
is a strong runner, but a weak flyer (Ridgway and
Friedmann 1941, Ripley 1977). It differs from R.
I. levipes principally in the more grayish (less
brownish) edgings and blacker centers of back
feathers and less reddish underparts (Van Rossem
1929). Adults weigh about 330 g (Linsdale 1936).
RANGE
The California clapper rail was formerly found
in coastal marshes at Humboldt Bay (Brooks
1940), Tomales Bay, and Bolinas Lagoon in Marin
County (Gill 1972), Monterey Bay at Watsonville
and Elkhom Sloughs (Silliman 1915, Gill 1972,
Gould 1973), and probably Morro Bay (Brooks
1940). In the San Francisco Bay area, major popu-
lations centered in salt marshes bordering the
southern arm in Alameda, Santa Clara, and San
Mateo Counties and smaller populations occurred
in the northern part of the bay and San Pablo Bay
in Marin, Sonoma, Napa, Contra Costa, and ex-
treme western Solano Counties (Grinnell et dl.
1918, Bent 1926, Grinnell and Wythe 1927,
DeGroot 1927, Grinnell and Miller 1944).
At present, the largest populations continue
to be in southern San Francisco Bay, with smaller
numbers in the northern San Francisco and San
Pablo Bay marshes, at Corte Madera Marsh near
the mouth of Gallinas Creek, Petaluma Marsh,
marsh south of San Pablo Creek; marsh north of
toll plaza of the San Francisco - Oakland Bay
Bridge, and at Tubbs Island. They are apparently
gone from all coastal marshes on Monterey Bay
except Elkhom Slough (GUI 1972, Gould 1973,
Varoujean 1972, Wilbur and Tomlinson 1976).
Although apparently nonmigratory (Wilbur
and Tomlinson 1976), the rails are known to
wander away from salt marshes in autumn (Lins-
dale 1936; Orr 1939) even to Farallon Islands
(Bryant 1888).
RANGE MAP
The range map on the following page shows
both past and present distribution.
STATES/COUNTIES
California: Alameda, Santa Clara, San Mateo,
Marin, Sonoma, Napa, Contra Costa,
Solano, Humboldt, Monterey, San
Luis Obispo.
HABITAT
R. I. obsoletus inhabits saltwater marshes
traversed by tidal sloughs, usually associated with
abundant growths of pickleweed (Salicornia)
(Grinnell and Miller 1944). GiU (1972) classified
marsh habitat in southern San Francisco Bay into
primary habitat or pure stands of cord grass
{Spartina foliosa) with 0.84 to 1.08 rails per ha,
and secondary habitat of pure stands of pickle-
weed or mixtures of cordgrass, pickleweed, and
other marsh vegetation with 0.13 to 0.17 rails
per ha. In southern San Francisco Bay, Zucca
(1954) found 69 of 87 nests (79%) in pure stands
of cordgrass, 10 in bases of gumplant {Grindelia)
bushes, 6 in pickleweed and 2 in mixed cord grass
and pickleweed. Wilbur and Tomlinson (1976)
note that early writers usually described nests as
occurring in pickleweed or at the base of gum-
weed plants, but later studies showed regular use
of cordgrass as a nest site. Zucca (1954) found
that time of nesting and tidal conditions deter-
mine, in part, the type of vegetation used for nest-
ing, with early nests placed in gumweed before
cordgrass growth occurs and nests placed in pickle-
weed, which grows at slightly higher elevations
than gumweed or cordgrass, when nesting is inter-
rupted by tidal flooding.
Moffitt (1941) points out that rails are
CALIFORNIA CLAPPER RAIL
Past and Present Distribution
umboldt Bay /
To males Bayf
Farallon Islands •
San Francisco Bay
Monterey Bay
Morro Bay
Bolinas Lagoon
Golden Gate Park
SAN FRANCISCO
Watsonvllle Slough
— Elkhorn Slough
Morro Bay Slough
Pescadero Creek ^
CALIFORNIA COAST
SAN FRANCISCO BAY REGION
Past and present distribution of the California clapper rail
abundant in south San Francisco Bay, while being
less numerous in similar marshes in northern San
Francisco Bay and the south end of Tomales Bay
and apparently absent from Suisun Bay, where
extensive pickleweed marshes exist. He noted that
the latter area has lower salinity, which may pre-
vent occurrence of organisms on which rails de-
pend for food. Wilbur (pers. coram.) adds that
habitat in northern San Francisco Bay is almost
pure pickleweed while in the southern part, where
rails are more numerous, it is mixed pickleweed-
cordgrass with cordgrass predominating.
Fall wanderers from the salt marshes have
been observed feeding in a city park, on the edges
of lawns next to shrubbery into which they re-
treated (Orr 1939).
FOOD AND FORAGING BEHAVIOR
Food consists almost entirely of animal mat-
ter, such as worms and crustaceans available in
salt marshes. Several stomachs taken in Alameda
county contained only parts of crabs (Grinnell,
et al. 1918). Rails may feed on molluscs obtained
by probing in mud-bottomed sloughs (Williams
1929, Moffitt 1941, Test and Test 1942). Fall
wanderers away from salt marshes have been noted
feeding on earthworms in a city park. One was
seen to swallow 5 worms in 5 minutes (Orr 1939).
Most feeding done in marshes is at low tide when
the most food is available. Volumetric content of
stomachs of 18 rails taken near Palo Alto, Calif.
4 February 1939 averaged 85% animal matter and
14.5% vegetable matter, all of which consisted of
Spartina seeds. No gravel was found in any stom-
ach. Plaited horse-mussel {Modiolus demissus)
amounted to 66% of the animal food. Spiders of
the family Lycosidae, common inhabitants of
marshes, made up 17.5% of animal matter (Moffitt
1944). Williams (1929) found the little macoma
clam {Macoma baltica) to be a principal item of
food near Palo Alto. Grinnell, et al. (1918) found
parts of the yellow shore-crab or mud crab {Hemi-
grapts oregonensis) were almost the only food in
clapper rails in Alameda County. Emerson
(1885) found that worms and insects made up
most of the food. Moffitt (1944) comments that
the relative amounts of plant and animal matter
in the rail's diet varies with the time of year, stage
of tide, and geographic locality.
Rails often wash their food before swallowing
it. They swallow small clams whole and peck larger
ones open with their bills and eat only the con-
tents. Crabs are dismembered and pecked open
before being eaten (Cohen 1895, Grinnell et al.
1918, Tomlinson and Wilbur 1976).
SHELTER REQUIREMENTS
Dense marsh vegetation for concealment and
nesting.
NESTING OR BEDDING
Rails nest on or near ground, usually on a
slight rise near a tidal slough; many well defined
trails lead from them to the water (Bryant 1880;
Taylor 1894, Cohen 1895, DeGroot 1927). Nest
sites are in dense patches of pickleweed {Sali-
cornia) or cordgrass {Spartina), or at the base of
gumweed {Grindelia) plants (Grinnell et al. 1918,
Zucca 1954, Gill 1972, Bent 1926, Wilbur and
Tomlinson 1976).
The nest is a platform or heap of material of
cordgrass and dead plants buUt up 8 to 15 cm
above ground, sometimes deeply cupped and
securely woven to surrounding vegetation (Bryant
1880, Cohen 1895, DeGroot 1927, Emerson
1885, Taylor 1894). Rails frequently build several
nests but use only one (Adams 1900, Gill 1972,
Grinnell et al., Bent 1926, Tomlinson and Wilbur
1976).
RITUAL REQUIREMENTS
Rails exhibit intraspecific hostility indicating
territoriality (Williams 1929).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
An unobstructed tidal flow of clean saltwater,
saltmarsh vegetation, and mudbanks for suitable
food organisms are environmental requirements
of/?. /. obsolete.
POPULATION NUMBERS AND TRENDS
No estimate of the total California clapper rail
population has been made (Wilbur and Tomlinson
1976). At least 100 were noted in an area of less
than 2.5 km^ of salt marsh near Palo Alto on
Feb. 1939 (Moffitt 1941). Gill (1972), by dragging
a rope, found densities ranging from 0.34 to 0.43
rails per ha in primary habitat (pure cordgrass) to
0.13 to 0.17 per ha in secondary habitat (mixed
cordgrass, pickleweed and other marsh vegetation)
in south San Francisco Bay. By census from a
boat in flooded marsh, he found 0.57 rails per ha
in a 14 ha section of marsh. Zucca (1954), also
using the boat count method in flooded marsh,
found 0.49 rails per ha. Based on these data Gill
(1972) estimated that the number of rails in south
San Francisco Bay ranges from 2,420 to 2,880
(average 2,750). Probably 50% or more of all
California clapper rails are in that area, where,
until recently, they appeared to be maintaining
their numbers. However, loss of their already
restricted habitat continues and total numbers are
probably being reduced. Some evidence indicates
that nesting success may also be declining (R. Gill
pers. comm. to Wilbur and TomHnson 1976).
REPRODUCTION
Nesting season begins in mid-March and ex-
tends into July (DeGroot 1927). Data for 128
museum egg sets indicate even distribution of
nesting from 1 April through 10 May, with only
six later records, but concentration of collecting
activity during periods when eggs are most likely
to be found may bias data (Wilbur and Tomlinson
1976). DeGroot (1927) recorded a peak in nest-
ing activity between 10 and 25 April, almost com-
plete cessation of nesting from 15 May to 15 June,
then renewed activity during late June and early
July. Applegarth (1938) and Gill (1972) recorded
a similar pattern but peak of activity was in May
rather than April. DeGroot (1927) estimated that
perhaps 50% of the birds raise two broods per
year. Zucca (1954) and Gill (1972) suggested
that late nestings were undertaken when first
clutches were destroyed by high tides in May and
June. Wilbur and Tomlinson (1976) suggest that
late nesting by California clapper rails may in-
clude both renesting attempts and second broods.
Apparently complete clutches are reported to
contain from 5 to 14 eggs, most commonly 6 to
10 eggs (Bryant 1880, Cohen 1895, DeGroot
1927, Applegarth 1938, Zucca 1954, Gill 1972).
Both sexes incubate the eggs, which hatch in
23 to 29 days (Applegarth 1938; Zucca 1954;
Johnston 1956). Hatching requires approximately
48 hours after pipping Qohnston 1956). Incuba-
tion probably begins with laying of the last egg
and ceases when first egg hatches (Zucca 1954).
There is no specific information on nesting
success (Wilbur and Tomlinson 1976). Norway
rats {Rattus norvegicus) are known to have
destroyed rail eggs but authorities disagree as to
the magnitude of this loss (Cohen 1895, Bryant
1880, DeGroot 1927, Zucca 1954). Other nest
losses are attributed to unknown mammals
(Bryant 1915) and nest inundation by high tides
(Zucca 1954).
MANAGEMENT
The California Department of Fish and Game
considered reintroduction of clapper rails at
Monro Bay in 1972, using stock from San Fran-
cisco Bay, but decided against it because of un-
certainty of the racial identity of the original
population and also the lack of information on
whether any of that population still existed there
(Sanford Wilbur pers. comm.).
Establishment of south San Francisco Bay
National Wildlife Refuge and acquisition of Tubbs
Island and Elkhorn Slough by the Nature Conserv-
ancy are beginnings toward habitat preservation.
Proposed management includes acquisition and/or
agreements with property owners to retain suit-
able marsh habitat for this and other water-assoc-
iated birds and further improvement in control-
ling water quality in problem areas (Fish and
Wildlife Service 1973).
AUTHORITIES
Sanford R. Wilbur
Fish and Wildlife Service
1190 East Ojai Ave.
Ojai, CaHfornia 93023
Robert D. Ohmart
Randall W. Smith
Dept. of Zoology
Arizona State University
Tempe, Arizona 85281
Robert Gill
Gordon Gould
Howard Leach
California Depart, of Fish and Game
1416 Ninth Street
Sacramento, California 95814
PREPARER'S COMMENTS
Although the total population of this sub-
species of clapper rail is still fairly substantial
(about 2,800 individuals), its very strict require-
ments for relatively unpolluted tidal salt marsh
habitat, which is rapidly disappearing within its
present range, and the disappearance of the bird
itself from much of its former range are adequate
causes for considering it in danger of extinction.
LITERATURE CITED/SELECTED
REFERENCES
Adams, E. 1900. Notes on the California Clapper
RaU. Condor 2:31-32.
American Ornithologists' Union. 1957. Check -list
of North American birds. Baltimore, MD.
A.O.U.
Applegarth, J. H. 1938. The ecology of the Cali-
fornia Clapper Rail on the south arm of San
Francisco Bay. MA Thesis, Stanford Univ.
Bent, A. C. 1926. Life histories of North Ameri-
can Marsh birds. U.S. Nat. Mus. Bull. 135:267.
Brooks, A. 1940. The Clapper Rail of Monro Bay.
Condor 42:126-127.
Bryant, H. C. 1915. California Clapper Rail breed-
ing in Alameda salt marsh. Calif. Fish and
Game 1:192.
Bryant, W. E. 1880. Notes on the habits oi Rallus
obsoletus, with a description of its eggs. Bull.
Nuttall. Omith. Club 5 : 1 24-1 25 .
Bryant, W. E. 1888. Birds and eggs from the
Farallon Islands. Proc. Calif. Acad. Sci. Ser. 2,
1:42.
Cohen, D. A. 1899. California Clapper Rail in
Alameda County. Condor 1:31.
DeGroot, D. S. 1972. The California Clapper Rail,
its nesting habits, enemies, and habitat. Con-
dor 29:259-270.
Emerson, W. O. 1885. California Clapper Rail.
Omith. and Oologist 10:142-143.
Fish and Wildlife Service 1973. Threatened WUd-
Ufe of the United States. U.S. Department of
the Interior, U.S. Govt. Printing Office.
Gill, R., Jr. 1972. South San Francisco Bay breed-
ing bird survey Wildl. Mgmt. Branch Adm.
Rep. No. 72-6 (June 1972), Fed. Aid W. L.
Restor. Proj. W-54-R, Spec. W. L. Invest.
. 1979. Status and distribution of Cali-
fornia Clapper Rail (Rallus longirostris obso-
letus). Calif. Fish Game 65(l):36-49.
Gould, G. 1973. California Rail survey, 1973. Cal-
ifornia Dept. Fish and Game, Spec. Wildlife
Invest. Job 11-10, Clapper Rail Survey, 1 July
1972 -30 June 1973.
Grinnell, J. and Miller, A. H. 1944. The distribu-
tion of the birds of California. Pac. Coast Avi-
fauna 27:126.
Grinnell, J. and M. W. Wythe. 1927. Directory to
the bird-life of the San Francisco Bay region.
Pac. Coast Avifauna 18:62-63.
Grinnell, J., H. C. Bryant and T. I. Storer 1918.
The game birds of California, p. 283.
Johnston, R. H. 1956. The incubation period of
the Clapper Rail. Condor 58:166.
Linsdale, J. M. 1936. Occurrence of the California
Clapper Rail away from marshes. Condor 38:
216.
Moffitt, J. 1941. Notes on the food of the Cali-
fornia Clapper Rail. Condor 43:270-273.
Oberholser, H. C. 1937. A revision of the Clapper
Rails {Rallus longirostris Boddaert). Proc.
U.S. Nat. Mus. 84:313-354.
Ohmart, R. D. and R. W. Smith. 1973. North Am-
erican Clapper Rail (Rallus longirostris) litera-
ture survey wdth special consideration being
given to the past and current status of
yumanensis. Report in fulfillment of Bureau
of Reclamation Contract No. 14-06-300-2409.
Orr, R. T. 1939. Fall wanderings of Clapper Rails.
Condor 41:151-152.
Ridgway, R. and H. Friedmann. The birds of
North and Middle America, Part 9. U.S. Nat.
Mus. Bull. 50:73.
Ripley, S. D. 1977. Rails of the world. Boston,
David R. Godine. p. 129.
Storer, T. I. 1915. Additional records of Cahfomia
Rail and Red Phadarope in California. Condor
17:98.
Taylor, H. R. 1894. Among the Cahfomia Clap-
per Rail. Nidologist 1:153-155.
Test, F. H. and A. R. Test. 1942. Food of the
Cahfomia Clapper Rail. Condor 44:228.
van Rossem,A.J. 1929. The status of some Pacific
Coast Clapper Rails . Condor 3 1 : 2 1 3-2 1 5 .
Varoujean, D. E. 1972. A study of the California
Clapper Rail in Elkhorn Slough. Calif. Dept.
Fish and Game. Spec. Wildlife Invest., Fed.
Aid Report W-54-R.
Wilbur, S. R. and R. E. Tomlinson. 1976. The
literature of the western Clapper Rails. U.S.
Fish and Wildlife Service, Spec. Sci. Rept. -
Wildlife No. 194.
Willett, G. 1906. The Southern Clapper Rail
breeding on fresh water. Condor 8:151.
Willet, G. 1933. A revised list of the birds of
southwestern California. Pac. Coast Avifauna
21:52.
Williams, L. 1929. Notes on the feeding habits
and behavior of the California Clapper Rail.
Condor 31:52-56.
Williman, O. P. 1915. Range of the California
Clapper Rail. Condor 17:201.
Zucca, J. H. 1954. A study of the Cahfornia Clap-
per RaU. WasmannJ. Biol. 12:135-153.
Biological Services Program
FWS/OBS-80/01.57
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States
AMERICAN PEREGRINE FALCON
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
public with information about Federally listed endangered and/or threatened %'ertebratc
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the Kndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Kngineers in coordina-
tion with the Offices of F-ndangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this series to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS should be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Ser\ ice
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.57
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES
AMERICAN PEREGRINE FALCON
A Cooperative Effort
by the
National Fish and Wildlife Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
CREDIT; KENNETH FINK
AMERICAN PEREGRINE FALCON
Falco peregrinus anatum Bonaparte
KINGDOM Aninialia
CLASS Aves
ORDER Falconiformes
FAMILY Falconidae
OTHER COMMON
NAMES Duck Hawk (AOU 1931); Great-
footed Hawk (Bent 1938); Rock
Peregrine.
DATE
Entered into SWIS To be determined
Updates To be determined
LEGAL STATUS
Federal: Endangered: (42 FR 36425, 14 July
1977).
Protected: Migratory Bird Treaty Act
of 3 July 1918 (40 Stat. 755; 16
U.S.C. 703-711) as amended 3 Decem-
ber 1969. Public Law 91-135.
States: Endangered: Alaska, South Carolina,
Georgia, Florida, Mississippi, Texas,
Alabama, Nebraska, Wyoming, Cali-
fornia, New Mexico.
Protected: Colorado.
REASONS FOR CURRENT STATUS
The peregrine falcon population decline almost
certainly resulted from a complex of environ-
mental factors, differing slightly from one area to
another (Hagar 1969; Herbert and Herbert 1969;
Hickey erf 1969). The population decline coincided
with the introduction and wide use of organo-
chlorine insecticides by 1967. The initial effect
was reproductive failure that wiped out the reserve
of nonbreeding individuals.
The second stage was the steady numerical de-
cline in the 1950's of breeding adults in many re-
gions. DDE (a derivative of DDT) seems to have
been the chief chemical involved in the United
States (Hickey and Roelle 1969).
The present total peregrine population in Utah
is believed to be only 10% of what it had been in
historic times; pesticide contamination and cli-
matic changes may have been major reasons for
the decline (Porter and White 1973).
At least as early as 1948, DDE was present in
peregrine eggs in sufficient concentration to ac-
count for eggshell thinning, and is now considered
a major factor in their decline, with DDE levels of
15 parts per million in eggs that failed to hatch
(Peakall 1974, 1976). Contamination of birds and
eggs by persistent pesticides and the killing and
capture of birds on breeding grounds, in passage,
and on wintering grounds have been the main
causes of decline (King in press).
Peregrines have continued to decrease since
1970 in most of their North American range, and
nowhere have they started to recover since reduc-
tion of DDT usage (Fyfe et al. 1976; Peakall
1976).
PRIORITY INDEX
10
DESCRIPTION
Peregrines are medium-sized hawk-like birds
with long, pointed wings and long tail. Wing-beats
are rapid and shallow. Adults are slate gray above,
with wing and tail feathers and flanks barred with
black, and black moustache marks on side of face.
The throat is white. Below is white and reddish
buffy, extensively spotted and barred with black.
Legs and feed are yellow. Immature birds are
brown above, streaked below. There are larger,
darker and black markings on face, more extensive
than on the Arctic Peregrine falcon (F. p. tund-
rius); they are paler and more reddish, less grayish
below than F. p. pealei (White 1968; Fish and
Wildhfe Service 1973).
Compared with tundrius, anatum is larger and
darker, bars on dorsal feathers more contrasting
with ground color; ventral surface more extensively
marked and washed with darker pinkish or rufous;
white auricular area less extensive in proportion
to black malar stripe (White 1968).
Ground-color of the eggs varies from creamy
white to pale pink, almost always nearly or wholly
concealed by small blotches, spots or fine dots of
rich brown or reds sometimes concentrated at one
end. Measurements of 61 eggs, range 57 x 43,
56.5 X 43.5, 48.5 x 38.5 (average 52 x 41 mm)
(Bent 1938).
RANGE
This falcon formerly bred in all ecogeographic
regions of North America south of the Arctic
tundra, which is occupied by Falco peregrinus
tundrius, to northern Mexico; but excluding the
immediate vicinity of the northwest Pacific coast,
which is occupied by Falco peregrinus pealei, the
southeastern and Gulf of Mexico coastal plains,
and most of the Great Plains from Saskatchewan
and Manitoba south to Texas. It does not now
breed anywhere in the United States east of the
Rocky Mountains or in southeastern Canada
(Fyfe et al. 1976; Porter and White 1973; Cle-
ment ed 1974).
RANGE MAP
The breeding range shown (on the following
page) is taken from Fyfe et al. 1976).
STATES/COUNTIES
Colorado: El Paso, Boulder, Jefferson,
Freemont, Montrose, Larimer
Archuleta, La Plata, Monte-
zuma, Moffat, Park, Mesa,
Garfield, Conejos, Douglas,
Pueblo (Bailey and Niedrach
1965;Enderson 1965).
Wyoming: Teton, Yellowstone Park.
Montana: Gallatin, Park, Sweet Grass,
Stillwater, Cascade, Chouteau,
Fergus, Blain, Custer, Dawson,
i
*5^.
Former breeding range of the American peregrine falcon.
Georgia:
New Mexico:
Illinois:
Indiana:
Iowa:
Kansas:
Kentucky:
Louisiana:
Maine:
New Hampshire:
Massachusetts:
Connecticut:
Michigan:
Missouri:
New Jersey:
North Dakota:
Oregon:
Pennsylvania:
Arizona:
South Carolina:
Prairie (Rocky Mountain/
Southwest Peregrine Falcon
Recovery Team 1977; Ender-
son 1965).
Dade (Burleigh 1958).
Rio Arriba, Los Alamos,
Sandoval, San Miguel, Berna-
lillo, Sierra, Eddy, Catron,
Grant, San Juan (Bailey 1928;
Hubbard 1970).
Wabash (Ridgway 1889).
Knox, Gibson (Butler 1897).
Linn (Dumont 1917).
Woodson Qohnston 1960).
Laurel (Mengel 1965).
Madison (Lowery 1955).
Penobscott, Hancock, Sumer-
set, Washington, Oxford (For-
bush 1927; Palmer 1949).
Grafton, Carroll (Forbush
1927).
Berkshire, Franklin, Hamp-
shire, Hampden (Forbush
1927).
New Haven (Forbush 1927).
Leelanau, Mackinac, Alger,
Marquette (Wood 1951).
Atchinson, St. Louis, Mont-
gomery (Widmann 1907).
Bergen (Stone 1908; Bull
1974).
Dunn, Stark, Oakdale, Billings
(Stewart 1975; Rocky Moun-
tain/Southwest Recovery
Team 1977).
Harney, Lake (Gabrielson and
Jewett 1940).
Clinton, Huntington, West-
moreland, York, Allegheny
(Todd 1940; Warren 1890).
Cochise, Coconino, Gila, Gra-
ham, Greenlee (USES & FWS
1979).
Greenville, Pickens (Sprunt
and Chamberlain 1949).
Texas:
Virginia:
Washington:
Alabama:
New York :
California:
Maryland:
Nebraska:
Arkansas:
Termessee:
Utah:
South Dakota:
Kerr, Brewster, Jeff Davis,
Culberson (Oberholser 1974).
Rock Bridge, Pulaski (Murray
1952).
Asotin (Jewett et al. 1953).
De Kalb, Marshall, Lauderdale,
Limestone (Imhoff 1962).
Washington, Clinton, Franklin,
Jefferson, Lewis, Hamilton,
Essex, Warren, Montgomery,
Albany, Greene, Columbia,
Dutchess, Putnam, Ulster,
Orange, Sullivan, Rockland,
Tompkins, Schuyler, Clinton,
New York (Bull 1974).
Marin, San Mateo, Santa Cruz,
Monterey, Santa Barbara, San
Diego, San Benito, Siskiyou,
Kern, Mendocino, Napa &
Sonoma (Bond 1946; Grinnell
and Miller 1944).
Washington (Stewart and
Robbins 1958).
Dawes (Lock 1977; Rocky
Mountain/Southwest Peregrine
Falcon Recovery Team 1977).
Cleburne (Baerg 1931).
Knox, Hamilton, Obion
(HoweU and Monroe 1957;
Spofford 1942).
Boxelder, Cache, Rich, Weber,
Morgan, Davis, Salt Lake,
Uinta, Utah, Tooele, Millard,
Sanpete, Sevier, Emery, Dag-
gett, Carbon, Grand, San Juan,
Iron, Garfield, Washington,
Cane (Twomey 1942; Porter
and White 1973).
Harding, Lawrence (Rocky
Mountain/Southwest Peregrine
Falcon Recovery Team 1977).
HABITAT
This species is adapted to a wide variety of
environments, ranging from the treeline south of
the Arctic tundra southward into the northern
Mexican mountains and the southern tip of Baja
California's tropical scrub (Porter and White
1973) up to 3,048 m in the California moun-
tains (Bond 1946) and 3,658 m in the Colorado
Rockies (T. D. Ray in Porter and White 1973).
Most eyries in Utah are located near marshes
or streams which supply waterfowl or passerine
birds for food. Most eyries all on east-and-north
facing cliffs suggesting orientation away from
exposure to overheating by afternoon sun (Porter
and White 1973). Combination of marshes ad-
jacent to suitable cliffs for nesting may be con-
sidered an "ecological magnet" (Hickey 1942).
In Alaska, habitat is in river valleys in boreal
forest area up to 800 m elevation. Western Can-
ada habitat varies from boreal forest to open
deciduous forest to arid types. In the lower 48
states, it varies from closed or semi-closed decidu-
ous forest in the east to semi-alpine and arid desert
in the west, usually overlooking water areas. In
Mexico, it is mainly along main cordilleran sys-
tems (King in press).
FOOD AND FORAGING BEHAVIOR
Its diet is mostly birds. Peregrine nestlings in
Alaska prefer birds to mice (Dixon 1908). Birds
eaten range in size from mallard ducks down to
warblers and nuthatches. Pigeons, flickers, jays,
meadowlarks, and other birds of medium size
probably constitute the bulk of food in inland loc-
alities; on the seacoast and islands, it feeds almost
entirely on the smaller seabirds. Mammals form an
insignificant part of their food. Remains of hares,
rabbits, woodchucks, squirrels and field mice have
been found near peregrine nests. They prefer to
capture prey on the wing after diving from above
at great speed. The highest speed attained by diving
is estimated at between 240 and 320 km per hour
(Bent 1938). Small prey is seized in its claws and
carried away; larger birds are stuck with the feet
with such force as to kill them. Then peregrines
descend to the ground to eat them. They pluck
birds, at least partially before eating. The male
feeds the female during the nesting season by drop-
ping food for her to catch in the air (Bent 1938).
Food of Yukon River, Alaska Peregrines in
1966 included 49 species of birds and 4 species
of mammals. Waterfowl constituted 50% oi the
diet composed by weight, and shorebirds 10% to
12%; small land birds composed 20% and small
mammals 2% to 3%. Analysis of peregrine tissue
showed that the food chain was contaminated
with significant measurable quantities of chlorin-
ated hydrocarbon pesticides in 1966 (Cade et al.
1968). In Utah eyries 79% of food identified con-
sisted of: American avocet (most frequent),
mourning dove, wUlet, western meadowlark, red-
shafted flicker, Wilson's phalarope, rock dove
(common pigeon), red-winged blackbird and
Brewer's blackbird (Porter and White 1973).
Food of peregrines of Chihuahuan Desert and
adjoining mountains included 24 species of birds,
5 species of mammals (3 bats and 2 squirrels) and
1 species of lizard. Particularly frequent food
items were dowitcher (sp.), white-winged dove,
band-tailed pigeon, mourning dove, common
nighthawk, white-throated swift, common
flicker, Steller's jay, mockingbird, and thrush
(sp.) (Hunt 1976;Johnson 1976).
In certain areas bats compose a greater part of
the diet than in others (Stager 1941).
SHELTER REQUIREMENTS
Cliff ledges for nesting and for night roosting
of young after they have fledged (Hickey cc? 1969).
NESTING OR BEDDING
Nest site chosen by female, who makes a
smooth well-rounded scrape in the accumulated
soil and rubbish on the cliff ledge, 2 or 3 cm
deep on the cliff ledge, to receive the eggs Q. A.
Hager in Bent 1938). Occasionally nests in broken
tops of tall trees (Ridgway 1889; Goss 1878;
Spofford 1942).
Cliffs are by far the favorite nesting sites, but
cut banks, ledges of tall buildings, hollows in trees,
and tree nests of other large birds such as ospreys,
are also utilized occasionally (Hickey and Ander-
son in Hickey ed 1969; Porter and White 1973).
RITUAL REQUIREMENTS
Aerial courtship display centered about the
home cliff in March, consisted of the male flying
excitedly from shelf to shelf, with much vocaliza-
tion, in presence of female, bringing of food by
male to female and then intricate aerial flight by
male with vertical loops, rolls and plunges from
great heights. After some time, both male and fe-
male engage in scratching about on ledges accom-
panied by vocalization until finally eggs are laid
in one of the ledge sites Qoseph Hagar in Bent
1938).
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Freedom from human interference at nest
sites and lack of contamination of prey with
pesticide poisons (Hickey ed 1969; Herbert and
Herbert 1965; Clement ec? 1974).
POPULATION NUMBERS AND TRENDS
In the eastern U.S., before that population
started to decline, three Hudson River, New York,
peregrines attained minimal ages of 17, 18 and 20
years (Herbert and Herbert 1965). Data indicated
a low mortality rate, a strong tradition to reoc-
cupy stable nesting sites, a tenacity to remain in
face of human proximity if not molested, and a
nonbreeding segment that was generally capable
of supplying replacements (Hickey and Anderson
in Hickey ed 1969). Hickey (1942) estimated 19
pairs in about 25,900 sq km around New York
City. By 1952 R. A. Herbert and K. G. SkeUon
reported virtually 100% of pairs in Hudson Valley
had unsuccessful reproduction (Cade 1960). In
1947 that population had 9 breeding pairs ( a
peak number for the century) and productivity
averaged 1.1 to 1.2 young per nest in 1931-40,
and 0.75 in 1941-50. No young were reared in
1950. and production ceased in 1951. Breeding
pairs dropped to 7 in 1950, to 5 in 1952, to 4
in 1956 and none in 1961. Human disturbance,
including highway construction, taking of young
by falconers, shooting of adults, and unfavorable
weather contributed to extirpation of the popula-
tion (Herbert and Herbert 1969).
In Massachusetts, reproduction seemed above
normal in 1946 but well below average in 1947
when broken eggs were observed for the first time.
By 1951, occupation of eyries was spotty and by
1955-57 only an occasional single bird was left
(Hagar 1969). Complete failure of 14 pairs in
Massachusetts was attributed by Hagar {in Hickey
ed 1969, p. 32) to raccoon predation.
In Pennsylvania, despite losses to hunters, fal-
coners and predators in 1946, there were still 17
active eyries which produced 1.25 young per nest
annually in the study area. From 1947 to 1952,
the number of active nests declined to 6 and pro-
ductivity per pair declined sharply. From 1953 to
1959 only 0.4 young were reared per pair per
year. The 3 pairs remaining in 1958 and one in
1959 reared no young despite lack of known dis-
turbance (Rice 1969).
On the upper Mississippi River in 1954 and
1955, the population was estimated at 1 pair to
30 km along one side of the river. These are now
gone. The last bird was seen in March 1964. Hu-
man disturbance did not seem to account for this
disappearance. The decrease seemed to progress
from south to north (Berger and Mueller 1969).
By 1962, it was rumored that no young pere-
grines had been reared that spring in the north-
eastern states, and this situation was confirmed in
1964 when a survey team found no birds nesting
in 133 formerly used eyries in the eastern U.S.
The peregrine falcon now appears to be extinct as
a breeding bird in the eastern U.S. and south-
eastern Canada from the Mississippi Valley to the
Atlantic Coast and from Alabama north at least
to Nova Scotia, a region that previously contained
about 300 occupied nesting sites. However, cap-
tive-reared birds are being released there (Cade et
al. 1968,Fyfeet al. 1976).
In the extreme northern part of the subspe-
cies' range within the boreal forest region of
Alaska, 200 to 250 pairs may have bred prior to
1950; less than 100 may remain (as of 1976). Per-
haps 400 to 500 pairs bred in the 1940 to 1950
period (Fyfe et al. 1976). Seventeen pairs were
found along 277 km of the upper Yukon River,
Alaska, in 1966, with an average linear distance
between occupied cliffs of 17 km (range 49 to 3
km). In 1951, average distance between 19 oc-
cupied cHffs was 15 km (range 50 to 3.5 km). On
the same stretch of river, L. B. Bishop (in Osgood
1900) estimated about 1 pair every 16 km, show-
ing quite similar density in the two periods and
demonstrating the usual static nature of peregrine
populations, each pair associated with a long-used
eyrie. It also showed that the population there
had not decreased as of 1966, even though levels
of chlorinated hydrocarbon chemicals were found
to be rather high in their eggs and tissues at that
time (Cade et al. 1968). As of 1969 White and
Cade (1977) found that populations along the
upper Yukon were still holding up rather well
with about 106 parts per milhon DDE in tissues,
while populations along the parallel Tanana River
to the south of it in Alaska with 334 ppm DDT in
1969 and 302 ppm in 1973 were pracrically gone.
Egg shells of Tanana River birds had thinned
about 17% by 1969 and 19.5% by 1973. Along
the Mackenzie River in northeast Canada (south
of the tree line), 18 known eyries were occupied
by 17 pairs (94%) in 1966 but by no more than 3
pairs and a single bird in 1973 (22% occupancy).
In northern Alberta, 12 eyries were occupied by 6
pairs in 1971, but in 1973, by only 4 pairs and 2
single birds. In an overall 1973 Canadian survey
of 83 eyries rechecked, 49 occupied by 44 pairs
(Fyfe in Clement ed 1974).
In western North America, there were formerly
densities of slightly more than 1 pair per 5,180
km^ in areas where the bird was considered
"common;" in mountains and arid regions, where
it was considered "rare," density was estimated as
1 pair per 51,800 km^ (Bond 1946). In the
Rocky Mountain region, of 28 or 29 peregrine
eyries active in 1938, not more than 4 to 6 were
found occupied later. An estimated 80% to 90%
of older sites in Utah, Idaho, Oregon, western
Wyoming and western Montana have been deserted
and the population shifted north or to higher
elevations to compensate for drought conditions
in recent times (Nelson 1969). In 1964, only
slightly more than 33% of known peregrine nest
sites in the Rocky Mountain region of New
Mexico and Colorado were being used. Many sites
were not used after 1950. An estimated 25 pairs
currently nest in Colorado and Wyoming, approx-
imately 25 pairs in Arizona, New Mexico and
Texas, nearly 25 pairs in Montana and possibly 60
pairs in Alberta (Enderson 1969a). Enderson
(1977) found that the population in the Rocky
Mountains region had changed little over the last
2 years. Egg production was typical for the species
but only half of the eggs were deemed hatchable.
About 40 eyries in Utah are active at one time or
another. Based on their density, if all eyries were
active at once there would be about 5,499 km^
per eyrie. If 11 suspected but unverified eyries
were excluded, density would be 1 eyrie for every
7.6 km^.
Peregrines are much more common around
the Great Salt Lake than in other parts of Utah.
Density appears to be directly related to avail-
ability of food and suitable nesting cliffs. Average
distance between 13 eyries along 209 km of the
west face of the Wasatch Mountains is 16.1 km.
The closest spacing was about 3.2 km apart (Por-
ter and White 1973).
In the Rocky Mountains region in general,
data indicate a 50% population decHne (Enderson
in Clement ed 1974). Annual natural mortality
there ranges from 20% to 25% for aduhs and 50%
to 55% for juveniles (Rocky Mountain/Southwest
Peregrine Falcon Recovery Team 1977). Approxi-
mately 100 pairs were reproducing successfully in
California in 1946; the decline began in the early
1950's. In 1970, 10 birds were present at 4 sites.
The California population is probably reduced at
least 50% from 25 years ago. Evidence indicates
that the total number of successful pairs in Cali-
fornia in 1970 did not exceed 5 (Herman 1971;
Herman etal. 1970).
Banks (1969) found evidence of a decline in
Baja Cahfornia and the Gulf of California, with
indication of lowered productivity in the latter
area in the 1960's. Anderson (1976) found no
peregrines on the west coast of Baja, California
between 1971 and 1975. He estimated 35 to 50
active nests in 30 to 35 local areas along the Gulf
of California and 75% of them occupied over a 5-
year period. Of 6 known successful nestings,
there were 2.2 young per nest. Thus it was as-
sumed that pesticide pollution was not too acute
at that time. Porter and Jenkins (1977) also
found no nests occupied on the west coast of
Baja, California, and the population along the
Gulf of Cahfornia much the same as former years.
Farther east in the northern Mexican high
country, three eyries on the western slope of a
single 3,000 m ridge of the Sierra Madre Occiden-
tal were separated by distances of 5 and 21 km.
In the Sierra Madre Oriental 6 pairs were found in
an 18.2-km circle, with the average distances
separating 6 eyries 5.2 km. Of 19 pairs present in
the Chihuahuan Desert region of northern Mexico
in 1976, 13 (68%) produced young, indicating a
moderately healthy population (Hunt 1976). To-
day, the mainland population in Mexico may ex-
ceed 100 pairs, while in Baja, California there
may be an additional 50 pairs, although only
about 20 to 30 are currently known (R. D. Porter,
W. G. Hunt in King in press).
REPRODUCTION
In Massachusetts, peregrines occupied breed-
ing stations before the end of February, and laid
the first eggs by 25 March. The female chose and
prepared the nest hollow on a cliff ledge. At least
one day elapsed between the first, second, and
third eggs, and sometimes 2 days between the
third and fourth egg (Joseph Hagar in Bent 1938).
A clutch ordinarily included 3 or 4 eggs, oc-
casionally 5, and very rarely 6 or 7. Regional
gradients in clutch size were reported by Hickey
(1942) and Bond (1946).
Incubation requires 33 to 35 days. The two
sexes changed places on the nest frequently from
the time the first egg was laid until incubation be-
gan if the weather was cold or stormy. Incubation
for the first 2 weeks was by the female; last half
was mostly by male (Hager m Bent 1938; Herbert
and Herbert 1965; Porter and White 1973). There
was one brood per season unless the eggs were
taken; then a second or even a third clutch was
laid (Bent 1938). Sometimes the young fell from
the nesting shelf and were killed (Eaton 1910).
From 3 nests, the first young flew on the 33rd,
35th, and 33rd days (Hagar in Bent 1938).
Some peregrines, when disturbed, may desert
their eggs and roost as much as 8.8 km away. Re-
nesting may or may not occur if the first attempt
is unsuccessful (Herbert and Herbert 1965;Hickey
1942).
Northern peregrines (Yukon River, Alaska)
lay about 3 eggs; the number of chicks ranges
around 2.3 to 2.7 per eyrie (Cade 1960). Produc-
tion of "fledglings" per occupied eyrie on the up-
per Yukon River dropped from 1.80 at 15 sites in
1966 to 1.40 at 10 sites in 1967 to 0.93 at 15
sites in 1968. This decline was associated with re-
duced eggshell thickness there in the 1960's com-
pared with those prior to 1947 Qoseph Hickey in
lit 23 Dec. 1969).
In Utah, egg laying began between 12 and 29
April (1943-1952). An average of 3.8 eggs (range
3-5) was laid per year during a 5-year period, and
2.4 (range 0-4) hatched (Porter and White 1973).
Natural mortality of first year peregrines is high.
Band returns indicate that 70% of fledglings die
in their first year (Enderson 1969).
Predation on nestlings is by raccoons in Mas-
sachusetts (Hagar 1969), great horned owls in
New York (Cade ed 1975), and ring-tails in the
southwestern United States (White and Lloyd
1962).
Reproductive success in Chihuahuan Desert
eyries of west Texas, despite significant egg shell
thinning of 17.9%, was 40% in 5 eyries in 1975
and 66.7% in 6 eyries in 1976. Reproductive suc-
cess was much better in mountain cliffs than in
river canyons (Johnson 1976).
Of a northern Mexico high-country popula-
tion in the Sierra Madre Oriental of 6 pairs, 2
failed to nest successfully in 1976. The average
fledgling ratio was 2.0 per successful pair. Egg
shell thinning was 17.8% (Hitchcock 1976).
In the Chihuahuan Desert region of northern
Mexico in 1976, 13 broods ranging in size from 1
to 4 young averaged 2.15 young per nest, compar-
ing favorably with pre-DDT eyries elsewhere.
Eyries on mountain cliffs were 78%) successful
and those in river canyons, 9% successful, indicat-
ing probable higher concentrations of pesticides
in river than in mountain environments. No dif-
ference was found in productivity of eyries of the
U.S. and Mexican sections of the Chihuahuan
Desert region. The rate of eggshell thinning was
greater in the northern and eastern sections than
in the southern part of the Chihuahuan Desert
region (Hunt 1976).
A few peregrines produce young at 1 year of
age, but it is not certain that all are breeding at
age 2 (Hickey and Anderson in Hickey ed 1969).
If they do not normally breed until at least 2 years
of age, the average fledging rate, which is less than
one per pair per year, is insufficient to replace
normal adult losses (Rocky Mountain/South-
western Peregrine Falcon Recovery Team 1977).
MANAGEMENT AND CONSERVATION
This species is listed in Appendix I, Conserva-
tion for International Trade in Endangered Species
of Wild Fauna and Flora (Federal Register, Vol.
42, no. 35, 22 February 1977, p. 10476).
Critical Habitat has been designated in five
areas in Sonoma, Lake, and Napa Counties Cali-
fornia (Federal Register, Vol. 42 no. 184, 22
September 1977, p. 47842).
Four Peregrine Falcon Recovery Teams have
now been appointed by the U.S. Fish and Wildhfe
Service. The plan of action for saving the pere-
grine includes: (1) increased legal protection; (2)
preservation of natural habitat, especially nesting
sites, and evaluation of pesticide contamination;
(3) management of wild populations; (4) captive
propagation for "domestication" and for stocking
of vacated range; (5) educational and information
program to gain pubHc support; and (6) develop-
ing national and international cooperation and
coordination. Direct killing or molestation by
man is the least important threat, although it can
play a part and must be guarded against. There
are adequate laws, but law enforcement and the
public's attitude toward birds of prey should be
improved. Preservation of essential habitat, partic-
ularly irreplaceable historic falcon eyries, is the
most essential action (Cade 1974). It should be
8
possible to increase productivity by "double-
clutching," which means taking the first clutch
to induce laying of a second. Removed eggs can
be artificially incubated or fostered to other wild
parents such as prairie falcons. Incubator-hatched
and captive reared peregrines may be held in cap-
tivity during the first critical year or two, then
released to the wild as adults. Artificial propaga-
tion and conditioning to survive in the wild could
be achieved by modification of falconry techni-
ques. Liberated falcons should be paired and psy-
chologically conditioned to accept suitable nesting
ledges or artificial structures in habitat where
food is adequate (Cade 1974).
In 1970, the Laboratory of Ornithology and
Division of Biological Sciences at Cornell Univer-
sity began a propagation and restocking program
with $100,000 allocated by the University. At the
same time, Robert B. Berry began collaboration
with a breeding project in Pennsylvania. In 1973,
20 young peregrines were reared at Cornell, using
artificial insemination. Eggs were hatched in an
incubator. Multiple clutches were obtained by re-
moving eggs as laid. One pair produced 14 young
in 2 years. Two other pairs laid 3 clutches each,
totaling 24 eggs, and only 1 pair failed in fertiliza-
tion. Only birds taken from the wild as nestlings
breed easily in confinement. Reproductive cap-
ability is attained by captive birds at 3 years of
age (Cade 1973). In 1974, three mature pairs at
Cornell produced 21 young. A total of 23 young
from all pairs was reared that season. Two young
reared at Cornell were fostered to a pair of wild
Colorado peregrines that had lost their second
clutch through thin-shelled egg breakage due to
DDE. The fostered young fledged. Experiments
were begun on returning captive reared birds to
the wild by "hacking," which involved feeding
nestlings in two locations suitable for future
nesting sites until they fledged and flew away
(Cade ed 1974). From 1973 through 1975, 68
young from captive parents were raised. Eleven
pairs were laying eggs in 1975 at Cornell, and
that year a new breeding facility was started at
Fort Collins, Colorado, stocked with birds from
Cornell. At a facility in Alberta, the Canadian
Wildhfe Service raised 18 peregrines in 1975.
Others were reared by private breedings in Can-
ada and the United States (Cade ed 1975). Over
150 young were produced in captivity in all
facilities in 1977, half of which were from western
North American anatum stock (Rocky Mountain/
Southwestern Peregrine Falcon Recovery Team
1977).
The first major restocking with captive-reared
birds was in 1975, when 16 were introduced to
the wild by hacking at 5 release sites. Two placed
on cliffs where peregrines formerly nested were
killed by great homed owls, showing need for
special protection against predators at natural
nesting sites. The remaining three sites were man-
made structures: one in a former gunnery tower
at Edgewood Arsenal on Chesapeake Bay in Mary-
land; another at Moose Hill Sanctuary of Mas-
sachusetts Audubon Society; and a third in a
specially built tower on an island near extensive
salt marshes in New Jersey. Other captive-bred
and raised individuals are being released in Canada
{Cdideed 1975 ; Cade and Dayne ed^ 1976).
If reestablishment of a peregrine population
in the East is successful, it must be with stock
genetically different from the former typical ana-
tum of the Eastern Deciduous forest areas as none
of that population remains (King in press; Drury
in Clement ed 1974). The problem of artificial
rearing and restocking was reviewed at a National
Audubon Society sponsored conference in 1974
(Clement ed 1974). Choosing ecologically adapted
stock to replace the extince "rock peregrine"
population was debated. Stock from European
populations was suggested as possibly better
adapted to the eastern deciduous forest area than
other North American stocks (White and Cade in
Clement ed 1974). Stocks of various racial origins
were suggested as best to achieve greater genetic
diversity for natural selection to work with
(Hickey in Clement ed 1974). Special protection
of nesting peregrines from human disturbance at
traditional sties is considered particularly im-
portant in preservation and reestablishment of
populations (Hickey in Clement ed 1974).
In summary, essential elements of a manage-
ment program are cooperation between govern-
ment and private agencies in the United States,
Canada, and Mexico in conducting population
surveys; rearing, fostering and introducing captive
reared birds to the wild; monitoring construction;
oil, gas, and mineral exploration; and recreational
activities that might interfere with peregrines,
with restrictions when necessary; and extending
and enforcing pesticide bans (Snow 1972; King in
press; Rocky Mountain/Southwest Recovery
Team 1977).
AUTHORITIES
Thomal J. Cade
Laboratory of Ornithology
Cornell University
Ithaca, New York 14850
Joseph J. Hickey
226 Russell Laboratories
University of Wisconsin
Madison, Wisconsin 53706
Clayton M. White
Department of Zoology
575 Widtsoe Building
Brigham Young University
Provo, Utah 84601
James H. Enderson
Department of Biology
Colorado College
Colorado Springs, Colorado 80903
D. D. Berger
1328 N.Jefferson
Apartment 311
Milwaukee, Wisconsin 53202
Walter R. Spofford II
"Aguila-Rancho"
Portal, Arizona 85632
Joseph A. Hagar
Marshfield Hills
Massachusetts 02051
R. Wayne Nelson
Department of Biology
University of Calgary
Calgary Alta.
Canada T2N 1N4
Richard Fyfe
Room 1110
10025 Jasper Avenue
Edmonton, Alta.
Canada T5J 136
Steven Herman
Department of Zoology
University of California
Davis, California 95616
Morlan Nelson
732 East Way
Boise, Idaho 83702
W. Grainger Hunt
Chihuahuan Desert Research Institute
P.O. Box 1334
Alpine, Texas 79830
PREPARER'S COMMENTS
The sudden complete loss of the eastern pop-
ulations of the American peregrine and continu-
ing decline of most other populations throughout
the remainder of its extensive range, attest to its
susceptibility to reproductive failure resulting
from chlorinated hydrocarbon pesticide pollution
of its food chain. Since the subspecies is practically
nonmigratory and since most of its range is within
the United States and Canada, correction of the
chemical poisoning problem by reducing use of
the most hazardous pesticides should be relatively
easy compared with the Arctic Peregrine, which
migrates to Latin American countries where
pesticide use is generally excessive.
The American peregrine management program
is fortunate in having an unusual amount of will-
ing, expert private citizen volunteer help, particu-
larly among active or former falconers. It gives
those persons with keen interest and expertise in
handling falcons a chance to work with their
favorite species as they would otherwise be un-
able to do under present regulations governing fal-
conry. The unusually large number of authorities
and pertinent literature references, and the fact
that there are now four Recovery Teams devoted
to the American peregrine (more than for any
other endangered species) shows the relatively
great interest in it and bodes well for its recovery
if the all important problem of chemical pollution
can be solved in time.— John A. Aldrich
LITERATURE CITED/SELECTED
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14
Biological Services Program
FWS/OBS-80/01.58
March 1980
Selected Vertebrate Endangered Species
Of the Seacoast of the United States-
SANTA CRUZ LONG TOED SALAMANDER
Fish and Wildlife Service
U.S. Department of the Interior
PREFACE
The purpose of this series of species accounts is to provide resource managers and the
pubHc with information about Federally listed endangered and/or threatened vertebrate
species that occur along, or within 100 kilometers of, the seacoast of the United States. In-
formation about life history, distribution, requirements and conservation of the subject
species is included (range maps and other distributional data arc not necessarily equivalent
to critical habitat as defined in the F.ndangered Species Act of 1973, as amended).
This series of accounts is intended to complement the computerized Sensitive Wildlife
Information System (SWIS) developed by the U.S. Army Corps of Engineers in coordina-
tion with the Offices of Endangered Species and Biological Services of the Fish and Wildlife
Service. A 3-ring binder is used for this scries to facilitate additions and deletions as new
accounts are prepared or as the status of species is changed.
Suggestions or questions regarding SWIS shoidd be directed to:
Office of Endangered Species
U.S. Fish and Wildlife Service
Interior Building
Washington, D.C. 20240
Suggestions or questions regarding this report should be directed to:
Information Transfer Specialist
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
11
FWS/OBS-80/01.58
March 1980
SELECTED VERTEBRATE ENDANGERED SPECIES
OF THE SEACOAST OF THE UNITED STATES-
SANTA CRUZ LONG TOED SALAMANDER
A Cooperative Effort
by the
National Fish and Wildhfe Laboratory,
the Office of Endangered Species
and the
National Coastal Ecosystems Team,
Office of Biological Services
Project Officer
Donald W. Woodard
National Coastal Ecosystems Team
NASA-SlideU Computer Complex
1010 Cause Blvd.
Slidell, Louisiana 70458
Performed for
Coastal Ecosystems Project
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
SANTA CRUZ LONG-TOED SALAMANDER
Ambystoma macrodactylum croceum
KINGDOM Animalia
CLASS Amphibia
ORDER Urodela
FAMILY Ambystomatidae
OTHER COMMON NAMES None
DATE
Entered into SWIS To be determined
Updates 15 November 1979
LEGAL STATUS
Federal Endangered (32 FR 4001, 11 March
1967)
States Endangered: California (21 May 1971)
REASONS FOR CURRENT STATUS
This salamander was first discovered in 1954
(Russell and Anderson 1956). It is now known to
occur in only six localities near the coast in the
Monterey Bay region of central California. This
area is undergoing accelerated residential and
commercial development that once destroyed or
threatened to disrupt populations of this salaman-
der. The type locality at Valencia Lagoon was
drastically disturbed by freeway construction and
related storm drainage channels between 1955
and 1969, and acceleration of residential con-
struction nearby threatens the upland habitat of
the species. A second population, about 6.4 km
to the southeast, occurs in and near an ephemeral
pond, which was threatened in 1971 with being
developed as a trailer park. This action was not
taken. These were the only known populations up
to 1972. Recently, a population was discovered at
Seascape, between Valencia Lagoon and Ellicott,
and three sites with a few individuals were located
near Elkhorn Slough, about 18 km south of
Valencia Lagoon. The four later sites all appear to
be threatened by agricultural or subdivision devel-
opment.
PRIORITY INDEX
Not assigned.
DESCRIPTION
This small salamander is 55 to 88 mm in snout-
to-vent length. Adults are usually less than 125
mm in total length. The toes are relatively long,
the dorsum is jet black with irregular mid-dorsal
spots or blotches of metallic yellow-gold to
orange color, undersides are sooty, and the sides
and limbs are flecked with white.
Larvae are small and usually transform at a
snout-to-vent length varying from 36 to 48 mm. If
water remains in breeding ponds through the
spring or early summer, larvae transform later, at
a larger body size. Metamorphosis may take place
from early May to mid-August.
Illustrations and description appear in Russell
and Anderson (1956) and Stebbins (1966).
RANGE
Six localities known to support populations
o{ A. m. croceum are (from north to south): Va-
lencia Lagoon, Seascape Pond, Ellicott Station,
McClusky Slough, Bennett (Struve) Slough, and
Moco Cojo Slough. These sites are all near the
coast of Monterey Bay. The populations are cen-
tered in and adjacent to the breeding ponds.
RANGE MAP
Localities are shown on the accompanying
map (from Santa Cruz Long-Toed Salamander Re-
covery Team 1976 and Reed 1978).
STATES/COUNTIES
California Santa Cruz, Monterey.
HABITAT
This salamander occurs in rolling hills of
mixed grassland, coastal chaparral and oak wood-
land, and along coastal seascapes. Adults and
transformed individuals spend most of their lives
on land in underground retreats — in mammal
burrows, beneath logs and rocks, and along the
root systems of plants (Ruth and Tollestrup
1973). It frequents heavily vegetated areas, typi-
cally live oak woodland [Quercus agrifolia) having
a few madrones [Arbutus menziesii) and Douglas
firs {Pseudostsuga menzusii). The understory
may include hazelnut [Corylus rostrata), coffee-
berry [Rhamnus californica), currant [Ribes san-
guineum), and California lilac [Ceanothus thyr-
siflorus). The ground cover and low shrubs often
include blackberry [Rubus sp.), poison oak [Rhus
diver siloba), and ferns [Polystichum munitum,
Pteridium aquilinum). Willow thickets [Salix sp.)
are used as refuges. Adults also seek shelter in
coastal scrub (Northern Coastal Scrub and Coastal
Sage Scrub), which includes coyote brush [Bac-
charis pilularis), coastal sagebrushes [Artemisia
californica, A. suksdorfii), lilac [Ceanothus sp.),
lupine [Lupinus variicolor), sage [Salvia sp.), cow-
parsnip [Heracleum lanatum), and varying ele-
ments of understory shrubs and ground cover
described above for the oak woodland commun-
ity.
Aquatic plants in and adjacent to breeding
ponds include spike rush [Eleocharis sp.), knot-
weed [Polygonum sp.), and pondweed [Potamoge-
ton sp.). Cattail [Typha sp.) occurs at the Valen-
cia site; the adjacent freeway drainage channel
was planted with kikyu grass [Pennisetum clan-
destinum). Emergent and submergent vegetation
is apparently necessary for egg-laying sites, cover,
and food.
This salamander breeds principally in shallow,
ephemeral ponds with vegetation present. A vital
factor to transformed salamanders is the availa-
bility of relatively dense vegetative cover adjacent
to or within migrating distance (presumably less
than 0.5 km) from the breeding pond (SCLTSRT
1976).
FOOD AND FORAGING BEHAVIOR
Arthropods and isopods are the most impor-
tant prey for adults throughout the year, and es-
pecially in the drier months (Anderson 1968).
In wetter seasons, the adults take more beetles,
centipedes, earthworms, and spiders. Small larvae
feed mainly on planktonic invertebrates (clado-
cerans and copepods); larger larvae principally
forage on small-sized prey such as cladocerans,
copepods, ostracods, and chironomid larvae, also
occasionally taking larger aquatic insects, snails,
earthworms, and tadpoles (Anderson 1968).
SHELTER REQUIREMENTS
Adults apparently depend on densely vege-
SANTA CRUZ
Range of the Santa Cmz Long-Toed Salamander.
tated areas or subterranean retreats for cover
while on land (see HABITAT.) Breeding ponds
have emergent or aquatic plants that provide cover
for both adults and larvae. Breeding ponds be-
come choked with vegetation soon after they fill.
NESTING OR BEDDING
Eggs are deposited singly, about 2 to 3 cm
apart, on aquatic vegetation, principally spike rush
(Anderson 1968). Ruth and ToUestrup (1974)
found some small clusters of 2 or 3 eggs and some
unattached eggs floating on the surface.
RITUAL REQUIREMENTS
Courtship behavior consists of a distinct se-
quence (Anderson 1961): clasping, rubbing, and
an elaborate walking sequence, followed by
spermatophore deposition by the male and subse-
quent recovery by the female.
OTHER CRITICAL ENVIRONMENTAL
REQUIREMENTS
Adults seem to have well-developed orienta-
tion abilities that are used to locate the breeding
pond. Transformed juveniles may spend 3 or 4
years on land before returning to the 'home' pond.
They frequent ephemeral ponds that are free
of predators, especially predatory fish and the
introduced bullfrog (Rana catesbeiana), both of
which are capable of seriously reducing salaman-
der numbers. Larval salamanders are especially
vulnerable to fish predation.
Low winter rainfall occasionally results in
small ponds or early drying out of the ponds, so
that breeding is unsuccessful or the larvae die of
desiccation (Ruth and ToUestrup 1973). Some-
times several consecutive years of low rainfall
occur, and considerable reproductive effort may
be lost.
Larval transformation usually succeeds when
there is more than 760 mm of rainfall per season,
and may succeed in years with between 635 and
760, depending on the timing of the rainfall; it is
usually unsuccesful when there is less than 635
mm of rain (Ruth and ToUestrup 1973). Migra-
tion to the breeding site occurs only on rainy
nights in winter.
POPULATION NUMBERS AND TRENDS
Once estimated to number several thousand
just at Valencia Lagoon (Anderson 1960), the
salamander population has decreased because of
habitat disruption. In 1972, field census studies
estimated that about 500 adult salamanders were
living at this site (Ruth and ToUestrup 1973), and
in 1973-74 about 1,000 salamanders were estab-
lished here (ToUestrup 1974). In 1974, Reed
estimated that there were 2,720 individuals in
the breeding area. The higher number may be due
a more comprehensive study than done by early
workers.
Several thousand salamanders migrated to
Ellicott Pond in the winter of 1971 (R. Marlow,
personal communication). About 4,000 were
marked during this period, and the total popula-
tion probably numbers 5,000 to 8,000 animals.
The other breeding sites presumably have only
small populations.
The Valencia population now uses two small
artificial ponds for breeding. These ponds enable
the salamander to survive, but at a lower and
consequently more vulnerable level (SCLTSRT
1976). The Ellicott population appears to be
stable following habitat protective measures (see
MANAGEMENT AND CONSERVATION).
REPRODUCTION
Breeding occurs from November to February
and largely depends on mass migrations during
heavy rainstorms. Adults do not remain long in
the pond. Each female lays about 200 eggs, which
hatch in 30 to 45 days, and larvae transform into
juveniles after 90 to 145 days (Reed 1978).
MANAGEMENT AND CONSERVATION
Considerable sustained effort by many indi-
viduals and agencies has provided protection for
this salamander (Bury 1971, 1972; Bury and
Ruth 1972; Ruth and ToUestrup 1973; Ruth
1974; ToUestrup 1974; Cahfornia Department of
Fish and Game 1975; SCLTSRC 1976; Reed
1978).
The Valencia site was partially restored by
CALTRANS (California Division of Highways)
following disruption due to construction of a
freeway and a drainage canal. A "salamander
pond" was dug and natural vegetation replanted
in 1970. This artifical pond is being used by the
adult salamanders as a breeding site, but the
population appears to be smaller than in times
past. The reduction in numbers is thought to be
due to the decreased size of the aquatic area
available for breeding and larval development.
CALTRANS and the California Department of
Fish and Game are now making the Valencia
site into essential habitat for the salamander,
including restoration of the original vegetation
and manipulation of the water level to increase
the area of standing water.
Santa Cruz County has adopted several
residential regulations designed to minimize the
impact of housing near Valencia Lagoon. The
California Coastal Commission, in coordination
with the state Department of Fish and Game
(DFG) has established guidelines for development
near Valencia Lagoon. A DFG habitat acquisition
program is underway at Valencia and the Nature
Conservancy is helping with the effort.
Areas at Ellicott (12 ha) and Valencia (1.1
ha) were purchased in 1973 and are now Ecolog-
ical Reserves administrated by the California De-
partment of Fish and Game. At Ellicott, the U.S.
Fish and Wildlife Service purchased 47 ha of
habitat in 1975-76, principjilly terrestrial areas
used by the adults for retreats during most of the
year. The upland habitat had been severely dis-
rupted by off -road vehicle use, which removed
hillside vegetation and resulted in erosion. Vehi-
cular travel has been significantly reduced in the
terrestrial habitat by patrols, and posting no-
trespassing signs. Erosion-control techniques are
being employed to restore the vegetation. The
reserve area was fenced in 1975-76.
There are continuing efforts to protect the
salamander. In 1978, the U.S. Fish and Wildhfe
Service proposed Critical Habitat designations for
the areas used by the salamander at Ellicott and
Valencia. Needed biological studies on the sala-
mander are being conducted through the coopera-
tive efforts of the California Department of Fish
and Game the U.S. Fish and Wildlife Service.
From 1973 to 1978, four additional localities
where some salamanders live were discovered, but
the status of these populations is not well known.
Potential threats to the recently discovered
populations at Seascape and near Elkhorn Slough
are being evaluated and, if proven serious, may
warrant protective measures.
AUTHORITIES
Elden H. Vestal
California Dept. of Fish and Game
P.O. Box 47
Yountville, California 94599
John M. Erode, Acting Leader
Santa Cruz Long-Toed Salamander
Recovery Team
California Dept. of Fish and Game
1701 Nimbus Road
Rancho Cordova, California 95670
Kristine Tollestrup
Museum of Vertebrate Zoology
University of California
Berkeley, California 94720
Richard Sage
Museum of Vertebrate Zoology
University of California
Berkeley, California 94720
PREPARER'S COMMENTS
There have been significant efforts to save the
Santa Cruz long-toed salamander from extinction.
It was one of the first animals to be included on
the Endangered Species List (1967). Thousands
of hours of volunteer and professional help pre-
vented the possible extinction of this life form.
Present interagency cooperation is assisting in the
recovery of this salamander and its associated
habitat.
Further biological studies are needed to assess
population trends at the Ellicott localities, and to
determine the status of recently discovered popu-
lations near Elkhorn Slough.
LITERATURE CITED/ SELECTED
REFERENCES
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stoma macrodactylum. Ph.D. thesis, Univ of
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. 1961. The courtship behavior oi Amby-
stoma macrodactylum croceum. Copeia 1961
(2): 132-139.
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of coastal and montane populations oi Amby-
stoma macrodactylum in California. Am.
Midi. Nat. 77(2):323-355.
. 1968. A comparison of the food habits
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Amby stoma macrodactylum croceum, and
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