(DRAFT) - Taxonomy
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
TAXONOMY
NAME - WOLF, GRAY
OTHER COMMON NAMES - WOLF, GRAY;LOAFER;LOBO;LOBO MEXICANO;WOLF, GREY;WOLF, GRAY, MEXICAN;WOLF, MEXICAN;WOLF, MOGOLLON MOUNTAIN;WOLF, ROCKY MOUNTAIN, NORTHERN;WOLF, SILVER;WOLF, GRAY, TEXAS;WOLF, TIMBER;WOLF, TIMBER, EASTERN;WOLF and TUNDRA;WOLF
ELEMENT CODE -
CATEGORY - Mammals
PHYLUM AND SUBPHYLUM - CHORDATA,
CLASS AND SUBCLASS - MAMMALIA,
ORDER AND SUBORDER - CARNIVORA,
FAMILY AND SUBFAMILY - CANIDAE,
GENUS AND SUBGENUS - CANIS,
SPECIES AND SSP - LUPUS,
SCIENTIFIC NAME - CANIS LUPUS
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Gray Wolf
Canis lupus Linnaeus, 1758
KINGDOM: Animal GROUP: Mammal
PHYLUM: Chordata CLASS: Mammalia
ORDER: Carnivora FAMILY: Canidae
The gray wolf (Canis lupus Linnaeus, 1758) is the largest member
of the Canidae except for certain breeds of domestic dogs (Canis
familiaris). Adult females weigh 18 to 55 kg and measure 1.37 to
1.52 m in total length; and males 20 to 80 kg and 1.27 to 1.64 m,
depending on subspecies. Fur long and varying from pure white through
mottled gray and brown to coal black; usually grizzled gray.
Generally resembling domestic German shepherd or husky in head and
body configuration but distinguishable from them by having orbital
angle of 40 to 45 degrees as compared with 53 to 60 degrees in dogs
(01) and having large, convex, and almost spherical tympanic bullae as
compared with smaller, compressed, and slightly crumpled bullae in
dogs. Distinguishable from coyote (Canis latrans) by larger size,
broader snout, relatively shorter ears, and proportionately larger
brain case. Canis rufus of eastern Texas and Louisiana is similar to
Canis lupus, being intermediate in many characters between wolf and
Taxonomy - 1 (DRAFT) - Taxonomy
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
coyote (02). Further distinctions among these four closely related
and similar animals were detailed by Lawrence and Bossert (03) and
others (04,05). The front foot has five toes, including a short one
with a dew claw proximal from other four; the hind foot has four toes.
Limb posture is digitigrade; the chest is narrow and keel-like with
forelimbs seemingly pressed into chest, and elbows turned inward and
paws outward (01,06). The legs are moderately long. Dentition
i 3/3, c 1/1, p 4/4, m 2/3, total 42; canines are about 26 mm long,
and carnassials are well developed. The cranium is elongate and
tapering anteriorly, with long jaws. For further details on skull and
teeth see Goldman (07). The baculum is pointed and has a ventral
groove. Other descriptive details can be found in Nowak (82), Hall
and Kelson (81), and Hall (49).
Thirty-two subspecies of the gray wolf are recognized throughout
the world; including the eastern timber wolf (C. l. lycaon), the
northern Rocky Mountain wolf (C. l. irremotus), and the Mexican wolf
(C. l. baileyi) (05). Current taxonomists believe that closer
taxonomic work would indicate far fewer subspecies in North America
than the 24 of the 32 recognized worldwide (08). Photographs are
available in (04,06,15,82). Goldman (07), Nowak (82), and Hall (49)
list locations of some specimens. Subspecies of Canis lupus Linnaeus
that are presently found in the 48 contiguous United States as listed
in Hall (49) are as follows:
Canis lupus baileyi Nelson and Goldman, 1929;
Canis lupus irremotus Goldman, 1937;
Canis lupus lycaon Schreber, 1775.
Common names used for the gray wolf (also spelled "grey") include:
Eastern timber wolf (09), loafer, lobo (57), Mexican wolf (lobo
mexicano) (58), Mexican gray wolf (56), northern Rocky Mountain wolf
(80), timber wolf, silver wolf, and tundra wolf (51). The following
is a short account for each subspecies:
Eastern timber wolf (C. l. lycaon):
A small, dark-colored subspecies; skull with remarkably slender
rostrum. Hall (49) summarizes the scientific synonyms for the eastern
timber wolf as follows: Canis lycaon Schreber, 1775; Canis lupus
lycaon, Goldman, 1937; Canis lupus Blainville, 1843; and Canis
tundrarum ungavensis Comeau, 1940. Kolenosky and Standfield (85)
distinguished between a northern "Boreal" type and a southern
"Algonquin type" within the currently recognized subspecies lycaon.
Although the ranges of these two types overlap, these authors found
no conclusive evidence of their interbreeding and significant
morphological differences between the two. C. l. nubilus, the buffalo
wolf, once ranged from southern Manitoba and Saskatchewan southward to
northern Texas; intergrades are suspected in the Minnesota population
of the eastern timber wolf (83). That there might be some affinity
between the wolves of the Great Plains and those of the western Great
Lakes region has also been suggested by others (13,84).
Northern Rocky Mountain wolf (C. l. irremotus):
Type: No. 214869, U.S. Nat. Mus. 1937. Canis lupus irremotus
Taxonomy - 2 (DRAFT) - Taxonomy
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
Goldman, Jour. Mamm., 18:41, February 14, type from Red Lodge, Carbon
County, Montana. A lighter-colored subspecies of medium to rather
large size, with narrow but flattened frontal region. In southern
British Columbia, irremotus probably intergrades with columbianus
(07).
Mexican wolf (C. l. baileyi):
Type: No. 98312, U.S. Nat. Mus. Goldman (07) describes C. l.
baileyi as smallest in size of the American subspecies of C. lupus.
McBride (43) notes, however, that baileyi skulls are often as large as
or larger than those of some specimens of the eastern timber wolf
(C. l. lycaon). In addition, the average of weights McBride (43)
records for baileyi exceeds averages recorded for lycaon by Pimlott
et al. (44). Weights of intact carcasses recorded by McBride (43)
averaged 83 pounds for males and 62 pounds for females, with weights
ranging from 68 to 91 pounds for males and 58 to 68 pounds for
females.
C. l. baileyi has a large head with broad zygomata, a short,
thick muzzle and a large nose pad, its chest cavity is deep, and its
neck and forequarters are thick (43). While pelage coloration varies
somewhat from one area to another and from one individual to another,
over-all the Mexican wolf's coat resembles that of the eastern timber
wolf. Black and white guard hairs color and pattern the back and
sides; below this, the sides are buff or tawny, fading to white on the
belly (45). Fur on the legs is light or white, and there is often a
black stripe down the front of the forelegs (06). Facial fur is dark
on the upper portion of the head, but light or white on the lower
cheeks and muzzle, and backs of the ears are usually rufous (45). In
summer coat, the distinctive patterning produced by the
black-and-white guard hairs is temporarily lost, creating generally an
over all gray-beige appearance (45). Nelson and Goldman (46) proposed
the name Canis nubilus baileyi in 1929, after Allen (47) had pointed
out, in 1920, that the name Canis mexicanus could no longer be used
for the Mexican wolf because this name had originally been given to a
domesticated Mexican hairless dog. Later, when it was recognized that
the gray wolves of North America belonged to one species only,
Goldman in 1937 (48) revised the taxa and named the Mexican wolf Canis
lupus baileyi.
Difficulties in certifying captive specimens as Mexican wolves
stem from the present lack of uniformly reliable genetic markers for
distinguishing among wolves, dogs, and coyotes on the basis of
chromosome structure or sera characteristics.
Recently, Bogan and Mehlhop (50) re-assessed the systematic
relationships of the wolves of southwestern North America and proposed
that wolves formerly assigned to the subspecies mogollonensis
(Mogollon Mountain wolf) and monstrabilis (Texas gray wolf) be
referred to baileyi.
Live specimens of the Mexican wolf can be seen at the Rio Grande
Zoological Park in Albuquerque, NM, the Alamogordo Zoo in Alamogordo,
NM, the Arizona-Sonora Desert Museum at Tucson, AZ, and the Wild Canid
Survival and Research Center near St. Louis, MO. Most photos in
publications and reports are in black and white, including those used
by McBride (43), Bailey (52), and Ames (53). New Mexico Wildlife
Taxonomy - 3 (DRAFT) - Taxonomy
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
magazine has carried some color photos (January-February 1982 issue).
Some descriptive material appears in: (07,43,52,54).
Unfortunately, positive identification of a specimen as a wolf
is often difficult even by persons considered experts. In rare cases,
this may be true even with the specimen in hand. Reported sightings
are therefore difficult to validate. Difficulties in field
identification stem from the morphological resemblances among wolves,
some dogs and wolf-dog hybrids (see photos in: (53)), and, for many
persons, large coyotes.
Taxonomy - 4 (DRAFT) - Status
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
STATUS
Coded Status
E: Federal Endangered
T: Federal Threatened
E: Federal Endangered
T: Federal Threatened
Arkansas; Federal Endangered
Arkansas; State Recognized
Georgia; Federal Endangered
Georgia; State Recognized
Kentucky; Federal Endangered
Kentucky; State Recognized
Maryland; Federal Endangered
Maryland; State Recognized
Status - 1 (DRAFT) - Status
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
Coded Status
North Carolina; Federal Endangered
North Carolina; State Recognized
South Carolina; Federal Endangered
South Carolina; State Recognized
Tennessee; Federal Endangered
Tennessee; State Recognized
Virginia; Federal Endangered
Virginia; State Recognized
West Virginia; Federal Endangered
West Virginia; State Recognized
E: Federal Endangered
Status - 2 (DRAFT) - Status
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
Coded Status
T: Federal Threatened
Pest
Commercial
Game (Consumptive Recreational)
Non-consumptive recreational
Ceremonial/Cultural
Ornamental
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The gray wolf (Canis lupus) has been designated an Endangered
species in the 48 conterminous United States (and Mexico), except for
the State of Minnesota where the species has been designated as
Threatened pursuant to the Endangered Species Act of 1973 (50 CFR
17.11; P.L. 93-205, 87 Stat. 884; 16 U.S.C. 1531-1540), as amended.
Critical Habitat (50 CFR 17.95(a)) has been designated for this
species in Isle Royale National Park, Keweenaw County, Michigan; and
the Counties of Beltrami, Cook, Itasca, Koochiching, Lake, Lake of the
Woods, Roseau, and St. Louis, Minnesota, with boundaries (4th and 5th
Principal meridians) identical to those of zones 1, 2 and 3, as
delineated in special rules in 50 CFR 17.40(d). These special rules
concern 5 regulatory zones encompassing over 85,000 square miles for
the gray wolf in Minnesota and "take" (including take of depredating
wolves) for this species.
The eastern timber wolf (C. l. lycaon) was listed as an
Endangered species November 3, 1967. The northern Rocky Mountain wolf
(C. l. irremotus) was listed as Endangered April 6, 1973. The Mexican
wolf (C. l. baileyi) was added, with the status of Endangered to the
Federal List of Endangered and Threatened Wildlife and Plants April
28, 1976. On June 6, 1976 the Texas gray wolf (C. l. monstrabilis)
was listed as Endangered. On March 9, 1978, the subspecies C. l.
baileyi, C. l. monstrabilis, C. l. irremotus, and C. l. lycaon were
deleted from the Federal list and the full species Canis lupus was
added to the list as Endangered except in Minnesota were the
population was designated Threatened with Critical Habitat.
This species is also protected by the Lacey Act (P.L. 97-79,
as amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild animal
(alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation, or foreign law; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
Status - 3 (DRAFT) - Status
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
BIA -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for management/recovery on Bureau of Indian Affairs
lands.
BLM -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public land under their control (43 CFR 4140). Also
responsible for management/recovery on Bureau of Land
Management lands.
DOD -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public land under their control. Also responsible for
management/recovery on Department of Defense lands.
NPS -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for conservation (Nat. Park System Organic Act - 16
U.S.C. 1, 2-3)/management/recovery on National Park
Service lands. Taking, possessing, or disturbing of
federally listed species is prohibited on NPS lands
(36 CFR 2.1, 2.2, and 2.3).
USFS -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for management/recovery on Forest Service lands. The
Forest Service is responsible for integrating
management, protection, and conservation of federally
listed species into the Forest Planning process
(36 CFR 219.19 and 219.20).
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
DESIGNATED STATUS: Endangered. (NM) Endangered, Group No. 1.
STATE/ADMINISTRATIVE AGENCY/STATE STATUTES:
(CO) Div. of Wildl./ Rev. Stat., Sec. 33-8-103(4);
(CT) Dept. of Environ. Prot./ Gen. Stat. Annot., Sec. 26-40d and
26-40e;
Status - 4 (DRAFT) - Status
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
(ID) Dept. of Fish and Game/ ID Code, Sec. 36-103;
(MI) Dept. of Nat. Res./ Comp. Laws Annot., Sec. 299.221 to
299.230, Admin. Code 299.1021 to .1028, Feb. 1983;
(MT) Dept. of Fish, Wildl., and Parks/ MT Code Annot. 87-5-101 to
112, MT Admin. Regist. 12.5.201;
(NM) Dept. of Game and Fish/ Reg. No. 624, Eff. July 22, 1983;
(NY) Dept. of Environ. Conserv./ 6 NY Code, Rules and Regs.
182.5, Apr. 30, 1983;
(TX) Parks and Wildl. Dept./ 31 TX Admin. Code Sec. 57.133
(127.30.09.003), July 1977;
(WA) Dept. of Game/ Admin. Code 232-12-014, June 1, 1981;
(WI) Dept. of Nat. Res./ NR 27.03 WI Admin. Code, Eff. Oct.,
1981.
DESIGNATED STATUS: Recognized Endangered.
STATE/ADMINISTRATIVE AGENCY/STATE STATUTES:
(AZ) Game and Fish Dept./ AZ Rev. Stat., Sec. 17-101;
(AR) Game and Fish Comm./ AR Stat., Sec. 47-101, 47-117, 47-118,
and 47-119;
(CA) Fish and Game Comm./ CA Fish and Game Code, Sec. 2052.
(DE) Dept. of Nat. Res. and Environ. Ctrl./ 7 DE Code Annot.,
Sec. 601;
(GA) Game and Fish Div./ Game and Fish Code, Sec. 27-3-130 to
27-3-132;
(IL) Dept. of Conserv./ 17 IL Admin. Code 1010.30, Feb. 1982;
(IN) Div. of Fish and Wildl./ IN Stat. Annot., Sec. 14-2-8.5-1,
310 IN Admin. Code 3-3-6;
(IA) Div. of Fish and Game/ IA Code Annot., Sec. 109A.1, 109A.5;
(KS) Fish and Game Comm./ KS Stat. Annot. 32-502(d), (j), 504;
(KY) Dept. of Fish and Wildl. Res./ KY Rev. Stat. 150.183,
301 KY Admin. Regulation 3:061;
(ME) Dept. of Inland Fish. and Wildl./ ME Rev. Stat. Annot.,
Sec. 7001, 7751-7756;
(MD) Dept. of Nat. Res./ Nat. Res. Code Annot., Sec. 4-2A-01 to
4-2A-09, 10-2A-01 to 10-2A-09, Code of MD Regs. 08.03.01.43,
Supp. 4;
(MA) Div. of Fisheries and Wildl./ MA Gen. Laws Annot. 131, Sec.
26A;
(MO) Dept. of Conserv./ MO Rev. Stat. 252.240, 3 Wildl. Code of
MO 10-4.111, Eff. Jan. 1, 1983;
(NE) Game and Parks Comm./ Admin. Rules and Regs., Vol. 13, 6-(4)
1984, Rev. Stat. of NE, Sec. 37-430 to 438, Admin. Rules and
Regs. Vol. 13, 6 (1)(d), 6-(10), 1981;
(NV) Dept. of Wildl./ Admin. Code, Sec. 503.015 to 503.080, Dec.
21, 1981;
(NH) Fish and Game Dept./ Code of Admin. Rules Fis. 1001.01,
1001.02, Eff. Apr. 28, 1980;
(NJ) Dept. of Environ. Prot./ Div. of Fish, Game and Wildl.,
Admin. Code 7:25-11.2, Eff. Mar. 29, 1979;
(NC) Wildl. Res. Comm./ 15 NC Admin. Code 10 I .0003, Eff. July
28, 1979;
(ND) Game and Fish Dept./ ND Century Code 20.1-01-02 and
20.1-02-05;
Status - 5 (DRAFT) - Status
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
(OH) Dept. of Nat. Res., Div. of Wildl./ OH Admin. Code
1501:31-23-01, Eff. July 11, 1980;
(OK) Dept. of Wildl. Conserv./ 29 OK Stat. Annot., Sec. 2-109,
2-135, 7-501, 7-502, 7-504, and 7-602;
(OR) Dept. of Fish and Wildl./ OR Rev. Stat. 498.026;
(PA) Game Comm./ 58 PA Code Sec. 147.1, 147.21, Mar. 1983;
(RI) Dept. of Environ. Mgmt./ Gen. Laws of RI Sec. 20-37-1 to
20-37-5;
(SC) Wildl. and Marine Res. Dept./ SC Regs. 123-150;
(SD) Dept. of Game, Fish and Parks/ SD Admin. Rules of SD
41:10:02, Eff. Mar. 16, 1978;
(TN) Wildl. Res. Ag./ TN Wildl. Proc. No. 75-15, as amended, Dec.
8, 1978;
(UT) Dept. of Nat. Res./ UT Code Annot. 23-13-2 to 23-13-4;
(VT) Fish and Game Dept./ VT Regs. Annot. 13-79.1, July 1, 1975;
(VA) Comm. of Game and Inland Fish./ Code of VA Sec. 29-230
to 29-237;
(WV) Dept. of Nat. Res./ WV Code, Sec. 20-7-7(1) and Sec. 20-1-1;
(WY) Game and Fish Dept./ WY Stat. Annot., Sec. 23-1-101.
DESIGNATED STATUS: Threatened.
STATE/ADMINISTRATIVE AGENCY/STATE STATUTES:
(CT) Dept. of Environ. Prot./ CT Gen. Stat. Annot., Sec. 26-40d
and 26-40e;
(MN) Dept. of Nat. Res./ MN Code of Agency Rules, Sec. 1.5600 to
1.5601, Eff. Dec. 1, 1983.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
All subspecies and populations of the gray wolf (Canis lupus) are
listed in Appendix II of the Convention on International Trade in
Endangered Species of Wild Fauna and Flora (CITES), specifically
including the subspecies C. l. irremotus and C. l. monstrabilis.
Canis lupus is also listed as Vulnerable in the IUCN Red Data Book
(1982) and the 1986 IUCN Red List of Threatened Animals.
C. l. baileyi, Lobo mexicano, is listed as Endangered by Mexico.
C. l. lycaon, the eastern timber wolf, is listed in the Convention on
Nature Protection and Wildlife Preservation in the Western Hemisphere.
This species is not protected, but regulated, by Canadian law.
ECONOMIC STATUSES:
Wolf fur is used for parkas, mukluks, and rugs. Pelts sell for
15 to several hundred dollars depending on locale, quality, and color.
The species provided even more significant commercial value for fur
trappers in the past. The wolf was also considered a livestock
predator and bounties were offered for its capture. Active predator
control programs continued into the 1930's. Wolves that range into
the United States from Canada have been known to be "trapped out" or
hunted legally in Canada. The species provides cultural value as a
symbol of Americas untamed wilderness and as a value to the American
Indian in ceremonial gatherings. A recreational value is demonstrated
by the willingness of the public to participate in efforts to preserve
the integrity of this rare species.
Status - 6 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
INLAND AQUATIC
SOCIETY OF AMERICAN FORESTRY TYPES
SAF TYPE STAGE CLOSURE
shrub--seedling
young tree
mature tree
Old Growth
White spruce balsam fir shrub--seedling
White spruce balsam fir young tree
White spruce balsam fir mature tree
White spruce balsam fir Old Growth
Longleaf Pine shrub--seedling
Longleaf Pine young tree
Longleaf Pine mature tree
Longleaf Pine Old Growth
Loblolly Pine-Shortleaf Pine shrub--seedling
Loblolly Pine-Shortleaf Pine young tree
Loblolly Pine-Shortleaf Pine mature tree
Loblolly Pine-Shortleaf Pine Old Growth
shrub--seedling
young tree
mature tree
Old Growth
Aspen-paper birch shrub--seedling
Aspen-paper birch young tree
Aspen-paper birch mature tree
Aspen-paper birch Old Growth
Douglas-fir-western hemlock shrub--seedling
Douglas-fir-western hemlock young tree
Douglas-fir-western hemlock mature tree
Douglas-fir-western hemlock Old Growth
Interior Ponderosa Pine shrub--seedling
Interior Ponderosa Pine young tree
Interior Ponderosa Pine mature tree
Interior Ponderosa Pine Old Growth
Western white pine shrub--seedling
Western white pine young tree
Western white pine mature tree
Western white pine Old Growth
shrub--seedling
young tree
mature tree
Old Growth
Sitka spruce-western hemlock shrub--seedling
Sitka spruce-western hemlock young tree
Sitka spruce-western hemlock mature tree
Sitka spruce-western hemlock Old Growth
Western larch shrub--seedling
Western larch young tree
Western larch mature tree
Western larch Old Growth
shrub--seedling
young tree
mature tree
Old Growth
shrub--seedling
Habitat Associations - 1 SAF TYPE STAGE CLOSURE
young tree
mature tree
Old Growth
White spruce-birch shrub--seedling
White spruce-birch young tree
White spruce-birch mature tree
White spruce-birch Old Growth
LAND USE -
Cropland and Pasture
Confined Feeding Operations
Herbaceous Rangeland
Shrub and Brush Rangeland
Mixed Rangeland
Deciduous Forest Land
Evergreen Forest Land
Mixed Forest Land
Streams and Canals
Lakes
Reservoirs
Forested Wetland
Nonforested Wetland
Beaches
Sandy Areas other than Beaches
Bare Exposed Rock
Strip Mines, Quarries, and Gravel Pits
Transitional Areas
Mixed Barren Land
Tundra Land: Shrub Brush
Tundra Land: Herbaceous
Tundra Land: Bare Ground
Tundra Land: Wet
Tundra Land: Mixed
Snow or Ice: Perrenial Snowfields
Snow or Ice: Glaciers
NATIONAL WETLAND INVENTORY CODES
NWI NWICLS NWIMOD NWISPEC
Riverine, intermittent SB4
Riverine, intermittent SB3
Riverine, intermittent SB2
Riverine, intermittent SB1
Riverine, upper perennial OW0
Riverine, lower perennial OW0
Lacustrine, limnetic OW0
Palustrine SS7
Palustrine SS6
Palustrine SS4
Palustrine SS2
Palustrine SS1
Palustrine OW0
Palustrine ML2
Palustrine ML1
Palustrine FO7
Palustrine FO6
Palustrine FO5
Habitat Associations - 2
NWI NWICLS NWIMOD NWISPEC
Palustrine FO4
Palustrine FO2
Palustrine FO1
COMMENTS ON HABITAT ASSOCIATIONS -
Wolves can live in any kind of natural habitat north of 20
degrees North latitude occupied by ungulates. This includes forests
of all types, rangelands, brush land, steppes, agricultural lands,
wetlands, mountain tops, deserts, tundra, and barren ground areas.
Wolves do not have any particular habitat requirement except that they
avoid areas with heavy human use. This is reflected in their original
circumpolar distribution. When raising pups, they seek seclusion and
protection, and proximity to drinking water (04,05,26).
Wolves avoid urban or built-up land, although they have been
known to visit dump areas in search of food (59,60,61,62). While
livestock in fenced pastures, confined feeding operations, and corrals
might tempt hungry wolves, they would be inclined to avoid areas such
as these that offer marked evidence of human presence, particularly
when prey is available in more remote areas freer of humans. Wolves
will prey on livestock on open ranges, particularly if natural prey
species are scarce (58).
Current wolf range in the contiguous 48 states is limited to
northern boreal and mixed forests, with a few individuals in western
mountainous pine forests; and possibly a few individuals in
southwestern range and woodlands. However, this is not because of
habitat preference; rather it is because wolves were extirpated from
other habitats (11).
Habitats were wolves occur in the Great Lakes region are all
forested areas sparsely intermixed with pasture lands. Jack pine
(Pinus banksiana), white pine (P. strobus), red pine (P. resinosa),
black spruce (Picea mariana), white spruce (P. alba), balsam fir
(Abies balsamea), white cedar (Thuja occidentalis), tamarack (Larix
laricina), white birch (Betula papyrifera), aspen (Populus
tremuloides), and maples (Acer sp.) are a few dominant cover types.
Wolves are good swimmers and do not hesitate to wade or swim
across rivers and lakes; they sometimes follow prey into water even in
winter (40).
The western white pine and western fir-spruce are typical of
the ecosystem mainly occurring in the high mountains (which includes
tundra land types) of the northern Rocky Mountains of western Montana
and Idaho. Western white pine is sometimes found in pure stands yet
commonly occurs in mixtures with other conifers. Shrubs and forbs are
prominent in the understory, whereas grasses and sedges are minor.
Western fir-spruce is normally the highest forest zone in the mountain
ranges, but this also varies. Douglas-fir, hemlock-sitka spruce, and
the larch ecosystems along with some western hardwoods typify the
range of the northern Rocky Mountain wolf.
Gish (56) described southwestern wolf country as including areas
from the chaparral-desert scrub country, up through grasslands, and
into the spruce-fir woodlands. He noted that records are rare of
wolves denning or establishing ranges in desert scrub below 3,000
feet. Leopold (54) refers to former wolf habitat in Mexico as the
temperate uplands. McBride (43) says: "Today [Mexican] wolves
inhabit elevations about 4,500 feet above sea level where higher
rainfall has created better grazing conditions for wolf prey." Bailey
(52) found Mexican wolves mainly in the Upper Sonoran and Transition
zones and especially in the open ponderosa pine forests and the
"juniper, nut pine, and oak foothills." Within this region exists
habitat features of potential importance to the Mexican wolf such as
Habitat Associations - 3 natural water sources including springs, seeps, pools, riffles,
vernal pools, and arid riparian habitat. Captive Mexican wolves
consume about two quarts of water daily (21).
Wolf dens may be holes dug into the ground, often enlarged fox
dens, but rock caves and crevices, old beaver lodges, hollow logs,
blow downs, and ground nests may also be used (04,06,11). Internal
characteristics of dens vary considerably, ranging from a single
burrow with no whelping chamber to complex burrow systems with several
entrances and chambers. Distance from den sites to the nearest water
sources that were likely to persist throughout the denning season
averaged 194 m. This value is maximal because ephemeral pools
resulting from snow melt were not included. Vegetation, visibility,
and wind exposure at den sites varied considerably. Forested sites
had restricted visibility and low wind exposure and tundra sites had
largely unrestricted visibility and high wind exposure. Luxuriant
stands of pioneer species of grasses and forbs were present at all den
sites. Quaking aspen, a species that prefers warm, dry sites, was
represented disproportionately in relation to its general occurrence.
In the Southwest, dens in which wolf pups are born and live for
their first couple of months are usually dug in slopes where tree
roots, rocks, or firmness of soil will lessen the likelihood of a
cave-in. McBride (43) located Mexican wolf dens under rock ledges on
the slopes of canyon walls or hills, particularly where dense stands
of oak brush and mountain mahogany hid the entrances. Observations of
denning behavior of captive Mexican wolves indicated adaptability.
One captive Mexican wolf in a densely wooded enclosure produced and
raised her litter above ground in a thick pile of tree branches (74).
The significance of all these forest/rangeland associations for
the wolf lies in the fact that these are the habitats in the lower 48
states that support populations of wild ungulates, are used for
grazing domestic ungulates, and have low concentrations of human
population. The wolf is potentially/indirectly affected by
agricultural practices only to the extent that a practice may attract
or repel concentrations of deer and other wildlife preyed on by the
wolf; or affect concentrations of domestic livestock. If there is an
adequate natural prey base, careful livestock husbandry practices, and
a selective predator damage control program, wolves and humans can
coexist. Seventy percent of the annual variation in wolf depredation
on livestock was found to be explained by the severity of the winter
previous to the depredation season (11).
Habitat Associations - 4 (DRAFT) - Food Habits
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
FOOD HABITS
TROPHIC LEVEL -
CARNIVORE
LIFESTAGE FOOD FOOD PART
General Deciduous Shrubs-Flowers/Fruit/Seed
General Deciduous Trees-Flowers/Fruit/Seed
General Arthropods
General Worms
General Fish
General Amphibia
General Reptilia
General Aves
General Carrion
General Mammalia
Food Habits - 1 (DRAFT) - Environment Associations
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Inland Wetland: vegetated stream banks
G
G
G Aquatic Features: Pool areas
G Aquatic Features: Springs [flowing]
G Inland Wetlands: Bogs
G Coastal Features: Vegetated offshore islands
G Terrestrial Features: Caves, dry
G Terrestrial Features: Burrows
G Terrestrial Features: Cliffs/ledges
G Terrestrial Features: Rock outcrops
G Terrestrial Features: Downed logs
Environment Associations - 1 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
LIFE HISTORY
FOOD HABITS:
Wolves are predators primarily on large mammals including deer
(Odocoileus sp.), moose (Alces alces), elk (Cervus elaphus), caribou
(Rangifer tarandus), bison (Bison bison), musk-oxen (Ovibos
moschatus), mountain sheep (Ovis sp.), and mountain goats (Oreamnos
americanus). Almost any species of animal may be eaten including
domestic animals, such as cattle, sheep, hogs, horses, goats,
chickens, turkeys, and dogs, although generally not forming a
significant part of the diet (04,06,12). Particular prey items taken
depend on availability and wolves can be said to be opportunistic
feeders. Most analyses of wolf predation on wild species have shown
that young, old, and otherwise inferior members of prey populations
constitute most of the animals killed by wolves (04). Domestic
ungulates often became more available to the wolf than wild prey and
are easier for the wolf to catch than wary, swift wild prey unhampered
by fences. McBride (04) tells of Mexican wolves' taking cattle,
burros and horses. Bailey (52) says wolves prefer cattle to deer,
however, ease of capture was undoubtedly a more significant factor
than taste in the choice of prey. McBride (43) reports that wolves in
Mexico pass up poor quality, sickly beef or dairy calves and prefer to
take larger, healthier calves, steers and cows. However, this is
undoubtedly not indicative of normal wild prey selection by less
persecuted groups of wolves in areas with adequate wild prey
populations. Usually all of the prey is eaten except for the larger
bones and chunks of hide. Average kill rate is about 15 to 20
adult-sized deer or equivalent per wolf per year (11).
Small vertebrate remains found in wolf droppings include those of
mice, mink, porcupine, beaver, muskrats, woodchucks, squirrels and
chipmunks, rabbits and hares, various birds, fish, lizards and snakes
(04,62). Beaver is the smallest prey consistently reported. Leopold
(54) lists rabbits, many of the rodents, and occasional plant food
such as berries and fruits as the small items found in the diet of the
wolf. Other authors also list grasshoppers and earthworms for the
wolf (04) and various berries, fruits, and seeds (04,62). However,
large mammals are the wolf's preferred food (04).
Plant food, invertebrates and small vertebrates are usually
eaten either when large mammals are unavailable or as extras when the
smaller food items are abundant, easily available and palatable.
The smaller mammals, birds, fish, garbage, and berries that are also
eaten usually do not form a high proportion of the wolf's diet (04,05,
06,11).
A large carnivore's preference for large prey is a matter of
economics. The energy expended in catching many small mammals or
other small vertebrates would offset the energy gained from the
morsels caught. The wolf's capacity for cooperative hunting enables
it to locate and kill larger prey animals than a single wolf might
usually be able to catch safely and efficiently. When hungry,
however, or when presented with an appropriate opportunity, single
wolves have killed large prey unaided, such as caribou (64).
Wolves also scavenge carcasses (18). This makes them vulnerable
to poisoning when carcasses are used for poisoned bait. Wolves also
Life History - 1 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
eat carcasses of their own species (65,66), and scavenge at garbage
dumps (59,60,61,62). Dumps scavenging may provide sustenance when
carrion and live prey are unavailable.
HOME RANGE/TERRITORY:
Wolves are territorial in most areas, but migratory in some.
They travel a great deal, usually more at night, but often during the
day if the temperature is cool. Three types of movements can be
distinguished: 1) travel within the territory; 2) dispersal; and
3) migrations. Territory sizes vary from 130 square km in Minnesota
(13) to 13,000 square km in Alaska (14). Daily travel within the
territory varies from a few kilometers per day to up to 72 km per day
(15). A dispersing wolf traveled a straight-line distance of 670 km
(16). Wolf family groups or packs each occupy a territory. Lone
wolves cover much larger areas and are chased by the resident packs
when in pack territories (11,17,18).
The entire feeding territory is defended, through howling (19)
and scent-marking (20). When desperate for food, territorial packs
may abandon territories temporarily and become nomadic or migratory
(11).
Wolves are social animals, and packs usually contain 5 to 8
members, although packs of up to 36 have been reported (21). Packs
are held together by: strong affectional ties that develop, are
reinforced, and become fixed in pups during the first 5 months of
life; or by courtship behavior between two lone mature adults. Order
is maintained in the pack by a dominance hierarchy with the adult male
dominant to the adult female and pups, the female dominant to the
pups, and a linear order among the pups. In larger packs a male
order and female order develops among adults (22).
Studies of wolves' territoriality and home ranges exist for
several areas where adequate numbers of wolves have persisted (44,62,
71,72). These studies have shown: that most wolves live in social
groups or packs although lone wolves also occur frequently; that packs
do occupy rather specific territories; that individual wolves are
often habitually found within certain home ranges; that a pack's
activity of necessity focusses on a more limited area when pregnant
and nursing females become sedentary at dens, and that specific
rendezvous sites are used in summer when pups are able to travel with
the pack; that a pack's territorial boundaries are scent-marked; that
wolves may be aggressively intolerant of wolves from other packs (66),
but packs are known to accept lone wolves (72), as well as young
wolves that are dispersing from parental territory and seeking mates
(72).
Territory size may vary in detail between subspecies of wolves
depending on sizes of pack territories and densities which are
broadly related to ungulate abundance (04). It is not clear whether
Mexican wolves would exhibit the same territorial and home range
characteristics as do northern subspecies. In the 1970's,
observations of the few, scattered remaining Mexican wolves indicated
that they wandered over large areas, constantly changed routes, and
sometimes never returned to areas they once frequently used (43).
Mexican wolves apparently do not form large packs as do northern
wolves; their prey is smaller than that of northern wolf subspecies
Life History - 2 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
and cooperative hunting is not as necessary (43).
Mexican wolves were long known to follow specific runways in
traveling between northern Mexico and southwestern New Mexico and
southeastern Arizona (52,54,58). These were often likely young wolves
dispersing into what was for them new territory, but Leopold (54)
reported that wolves from Mexico's Sierra Madre "regularly" crossed
the International Boundary in these areas to raid cattle herds in
southwestern New Mexico. In 1906, Bailey (52) camped for several
weeks on the Mimbres River in the Gila National Forest in the range of
C. l. mogollonensis, a subspecies that Bogan and Mehlhop (50)
recommend be referred to C. l. baileyi. Four wolves repeatedly passed
Bailey's camp every two or three nights, and he learned to recognize
their individual tracks. This would indicate this pack's association
with a specific range or territory.
PERIODICITY:
Wolves are active during all seasons and times of day. In
general, wolves tend to be most active in the early to late evening
and travel within their territories mostly at night (05). However,
during summer they tend to move more during cooler parts of the day.
During extremely cold weather (-40 degrees C) they may sleep for days
(11).
Bailey (52) found the wolves of southwestern New Mexico to be
active at all times of the day except at mid-day, when both young and
old wolves slept. In general, the literature on wolves indicates that
wolves are adaptable and adjust their activities more to enhance
hunting success than to satisfy any inner, time-of-day preferences of
their own. Captive wolves living closely with humans also exhibit
this same adaptability, suiting their active and rest periods to those
of their keepers (45,73).
MIGRATION PATTERNS:
While most wolves are not migratory per se, they do wander great
distances daily, within their home range, most often in search of prey
(04,43,52). In addition, when reproduction increases wolf numbers
within an area, some wolves of a group, often the young adults,
disperse to new areas (58,72). In food searches and population
dispersals, wolves have long been known to follow the same routes or
runways (52,54,58). Factors accounting for establishment of such
runways may include topography, prey availability, and knowledge
passed by experienced wolves to their inexperienced companions.
In tundra areas, wolves follow migrating caribou herds and thus
annually travel more than 160 km one way, from tundra to taiga and
back (23,24). The farthest that individual migrating wolves have been
known to travel is 360 km (25). In Alaska, the fall caribou migration
begins in late October or early November, and the wolves return to
their summer denning areas by late April or early May (24).
COVER/SHELTER REQUIREMENTS:
Vegetative cover affects the wolf's survival in that it
constitutes a factor in the welfare of the prey species on which
the wolf depends and vegetative cover is useful as a means of
screening the wolf's presence from human pursuers or from the prey
Life History - 3 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
that the wolf pursues.
Healthy adult wolves need little shelter, although they will seek
shade or windbreaks to escape hot sunshine or cold winds in order to
bed down. Under such circumstances, captive wolves seek the comfort
of shelters provided in their enclosures (45), and wild wolves may
retreat to old dens, even though it is not denning season (04).
REPRODUCTIVE SITE REQUIREMENTS:
Wolf dens are used for bearing and protecting pups, and often
are abandoned after pups are about two months old. The same den may
be used each year, or different dens may be chosen. Sometimes pups
are moved from one den to another (04). Wolf dens may be holes dug
into the ground, often enlarged fox dens, but rock caves and crevices,
old beaver lodges, hollow logs, blow downs, and ground nests may also
be used (04,06,11). Following is a summary of characteristics of
ground holes used for dens in Alaska (26):
1) Internal characteristics of dens vary considerably, ranging from a
single burrow with no whelping chamber to complex burrow systems with
several entrances and chambers. Mean height of 36 entrances was
401 mm and mean width was 478 mm. Mean height of 8 tunnels was 352 mm
and mean width was 498 mm. Mean length of 17 tunnels was 2.7 m.
Three beds were found in tunnels; them averaged 415 mm high by 830 mm
wide. Mean height of 10 whelping chambers was 500 mm and mean width
was 1.35 m. Chambers were not lined.
2) Wolf den sites were located on a variety of landforms, most of
which were glacially-deposited. Dens were excavated in slopes that
tended to face southerly directions; mean slope was 16.8 degrees.
3) Distance from den sites to the nearest water sources that were
likely to persist throughout the denning season averaged 194 m. This
value is maximal because ephemeral pools resulting from snow melt were
not included.
4) With the exception of one site, soils at den sites were sandy and
well-drained, with low organic matter and moisture contents and
relatively high bulk densities.
5) Vegetation, visibility, and wind exposure at den sites varied
considerably. Forested sites had restricted visibility and low wind
exposure and tundra sites had largely unrestricted visibility and high
wind exposure. Luxuriant stands of pioneer species of grasses and
forbs were present at all den sites. Quaking aspen, a species that
prefers warm, dry sites, was represented disproportionately in
relation to its general occurrence.
In the Southwest, dens in which wolf pups are born and live for
their first couple of months are usually dug in slopes where tree
roots, rocks, or firmness of soil will lessen the likelihood of a
cave-in. McBride (43) located Mexican wolf dens under rock ledges on
the slopes of canyon walls or hills, particularly where dense stands
of oak brush and mountain mahogany hid the entrances. Observations of
denning behavior of captive Mexican wolves indicated adaptability.
One captive Mexican wolf in a densely wooded enclosure produced and
raised her litter above ground in a thick pile of tree branches (74).
In more open enclosures, captive Mexican wolves have preferred to
excavate dens (45,74). Heights, widths, and lengths of entrance
tunnels and denning or whelping chambers are probably dependent on
Life History - 4 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
soil conditions. Free water must be available within reasonable
distance of a den site (04,58). Pups obtain water from their mother's
milk and from regurgitated food brought to the den by adult wolves
(04).
REPRODUCTIVE CHARACTERISTICS:
The alpha male and female (dominant male and female) of a pack
are the primary breeding pair. In the wild, wolves usually do not
produce young before their third year (04), but both captive and wild
wolves, male and female, are known to be fertile in their second year,
about 22 months of age (04).
After courtship lasting from days to months, wolves copulate
during an estrus of 5 to 15 days (27). Although a sexually mature
female may exhibit vaginal bleeding and swollen vulva during a
pro-estrus of over a month, she usually refuses to mate with a male
until she is in estrus (04,45). The single annual receptive period
may be from January in low latitudes to April in high latitudes (04).
Mexican wolves breed once a year, copulating in February and
delivering pups in April and May, after a normal gestation period
(43,45,53). Many courtship attempts are made, but few succeed.
Courtship may take place between adult members of packs or between
lone wolves that pair before or during the mating season (11).
Copulation involves typical canid coupling in which the bulbous base
of the penis locks into the vaginal sphincter, and the tie may last
for up to 36 minutes (04). The copulatory tie in Mexican wolves lasts
an average of 13 minutes as observed by Ames (45). In dogs, and
presumably wolves, ejaculation occurs intermittently during the tie
(04). Definite mate preferences are shown, but not all courtship or
copulation by members of a pair is directed at the mate. Maturing
females may come into heat 2 weeks later than animals that have bred
before. In Alaska, mature females shed an average of 7.3 ova and
implant 6.5 embryos, and newly maturing females, 6.1 ova and 5.4
embryos (21). Gestation lasts 63 days, and an average of 6 young
(extremes, 1 to 11) are born blind and helpless. McBride (04) records
Mexican wolf litters ranging from three to nine pups, with an average
litter size of 5.63. Eight litters in the Mexican wolf captive
breeding project have ranged from two to seven pups, with an average
of 4.5 (45). Both male and female may breed until at least 11 years
of age (11). Over their lifetime, wolves may be monogamous,
polygamous, or promiscuous (11,30), but once mating has occurred, male
and female raise the pups together (11). Although the pair may
separate at times during the rest of the year, they pair up again for
breeding and raising young and stay together most of the year (04).
In the Mexican wolf captive breeding project, the only female wolf to
have been paired successively with several mates (to 1984) has
accepted and bred with two of them and with a wild male before (45).
In this captive breeding project females were first paired in their
second year and bred successfully (45). Under appropriate conditions,
captive wolves have reproduced young in their first year (75).
Senility lowers pup production in dogs (76), but few wild canids
survive to advanced ages. Pup production from an old female in the
Mexican wolf breeding project has been lower since her 10th year (45).
Life History - 5 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
PARENTAL CARE:
The female usually stays near the young for at least 2 months,
while the male and other pack members hunt more widely and often feed
both the female and the pups. Pups' eyes open at day 11 to 15, they
are weaned at 5 weeks, and most milk teeth are present by week 8 when
they are moved to a ground nest, where they romp and play over an area
(known as a "rendezvous site") of up to 0.4 hectare. The pups may
spend up to 3 weeks at one site but are then shifted up to 8 km to
another. Probably depending on the degree of pup development, this
behavior may continue through winter. Pups in good physical condition
will join adult members of a pack in their travels as early as October
when they may weigh 27 kg and be almost adult size (11). Adult teeth
replace deciduous teeth between weeks 16 and 26 (28). Epiphyseal
cartilage calcifies about month 12 (21).
Both wolf parents care for the pups (43). The male, and the
female when she is able to leave the pups, carry meat in their
stomachs and regurgitate it for the pups (04). Mexican wolves
living in groups in captivity also exhibit the regurgitation behavior
(45). The male also feeds the female in this fashion. By October the
pups are adult size and at this time some pups remain with the pack
and others leave the territory upon maturity (04).
Mexican wolves apparently do not form large packs as do northern
wolves; their prey is smaller than that of northern wolf subspecies'
and cooperative hunting is not as necessary (43). McBride (43) says
that the parent Mexican wolves aggressively defend the pups and the
den area from hunters and dogs and also notes that pups are "on their
own by October and begin to travel away from their parents by
December."
POPULATION BIOLOGY:
The population biology of wolves is highly variable depending on
prey availability and history of the population. Generally wolf
populations are limited somewhat by the species' tendency toward
territoriality, and deferred reproduction (31), and by a tendency for
sex ratios biased toward males (32). Superimposed on these traits is
the need for the wolf to capture vulnerable prey (04) to sustain its
numbers. Thus survival and mortality rates can vary from 0 to 100
percent for individual packs during individual years, and a population
can increase up to 64 percent from year to year (33). Given adequate
stock, a wolf population can recover within a few years (33).
Although wolf populations can reach densities of up to one wolf per
11 square km (33), a more usual saturation point is about one wolf
per 26 square km in areas with high prey densities (04).
Nevertheless, if prey vulnerability is low for a prolonged period,
wolf numbers will decline and cannot be maintained at high densities
(11). Limiting factors can be related to human disturbances,
harassment, and availability of ample food supply. Population
fluctuations in northern wolves has been correlated with prey
population trends (34) and winter snowfall amounts (11). Survival
rates of the species vary depending on adequate food source but on
average are 6 to 43 percent for pups from birth to first winter;
55 percent from first to second winter; and 80 percent annually for
adults (04). Mortality rates vary from 7 to 65 percent; and 25
Life History - 6 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
percent of this rate is from malnutrition and 43 percent from natural
mortality (86). A pack usually produces only one litter per year (04)
although two litters have occasionally been raised in one year by a
pack (77). While many packs actually contain only a single mature
pair, in packs with several breeding-age adults, one or both of the
breeding pair have been reported as disrupting breeding attempts of
other adults (04). Other physiological causes may also be
accountable. When adult pack members are lost, particularly
high-ranking individuals, through hunting or predator control, the
disruption of the pack's hierarchal structure may result in production
of more litters by the remaining pack members than would have been
produced by the stable pack (78).
Similarities and differences between population biology of
northern and Mexican wolf subspecies are largely conjectural.
Recorded observations of Mexican wolves are relatively few, and almost
all seem to speak of "families" (the parents or parents and young of
the year) or of temporary associations of the year's maturing young
that are dispersing into new areas (43,52). It could be deduced that
normal undisturbed populations of Mexican wolves might seldom include
packs with more than one pair of breeding-age adults and that all
adult wolves in a population could pair and produce litters in one
year's breeding season. This situation would seem to represent a
higher breeding potential than that of the northern wolf population
given the same number of adult wolves. Smaller pack size in Mexican
wolves may represent a response to the smaller size of its prey and
the lower biomass of prey in a more arid habitat. The latter factor
also reduces the total number of wolves that an area can support. No
information is available on Mexican wolf survival rates, increase and
turnover, or longevity in the wild.
SPECIES INTERRELATIONSHIPS:
The wolf is a predator on ungulates and thus is influenced by
their numbers (04,33,11). Wolf numbers decline with inadequate prey,
or if prey is not vulnerable due to good habitat and/or weather (33).
Conversely, under some conditions, wolves can reduce their prey
populations (04,33,34,35) or, if prey is too vulnerable, wolves may
extirpate local populations and thus decline in numbers themselves
after an initial increase (33,34). The precise nature of the
wolf-prey-weather interactions is unknown (11,33,34,35).
The coyote (C. latrans) competes with the wolf for many prey
species. Ravens (Corvus corax) and magpies (Pica pica) are scavengers
often associated with wolves (45,47). The presence or lack of wolf
kills could locally affect the presence and relative abundance of
these birds though survival is not dependant on this.
Feral dogs (C. familiaris) directly affect wolf survival
prospects in areas where wolf numbers are very low. In Italy, the
remaining gray wolves suffer from competition for food with a feral
dog population of 80,000 (79). Where wolf numbers are extremely
low, they will interbreed with dogs (79). Hybridization with dogs may
threaten remaining wild Mexican wolves; the last male wolf taken in
Mexico for the captive breeding project was captured in 1980 at a
ranch where he had bred successfully with a ranch dog (63).
Life History - 7 (DRAFT) - Life History
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
OTHER LIFE HISTORY DESCRIPTORS:
Hierarchal rankings within wolf packs affect the behavioral
relationships between pack members (04). The ritual displays between
dominant and subdominant wolves serve to reduce levels of conflict and
reinforce pack cohesiveness.
Wolves have three main methods of communication: 1) howling and
other vocalizations; 2) visual displays including postures and
positions of various body parts, especially the face; and
3) scent-marking. Individual wolves may have distinctive howls (38).
The howling of pups is usually high-pitched, with considerable
yapping, whereas the pitch of adults is much lower. Functions of
howling are assembling the pack, and territory advertisement (19).
Coarse barking, whining, and growling are other vocalizations. Body
positions and facial expressions show social status primarily.
Inguinal presentation and sniffing serve in individual identification.
Scent-marking involves deposition of urine and/or feces on conspicuous
objects along trails, and often includes intense scratching of the
earth. It is usually performed by high-ranking adults often at trail
intersections and is a means of advertising territory (13,20) and pair
bonding (39).
Wolves are good swimmers and do not hesitate to wade or swim
across rivers and lakes; they sometimes follow prey into water even in
winter (40).
No differences in karyotypes have been found between the wolf and
the domestic dog or the coyote (36), or the red wolf (02). Both dog
and coyote have 39 pairs of chromosomes with the autosomes and sex
chromosomes described in detail in Hsu and Benirschke (37), also,
various types of crosses are detailed in Iljin (01).
Life History - 8 (DRAFT) - Management Practices
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Prescribed/controlled burning of habitat
Beneficial Selective Thinning
Beneficial Timber Harvesting
Beneficial Restricting/regulating human disturbance of populations
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Controlling/Restricting Agricultural Practice
Beneficial Maintaining Early Stages of Succession
Beneficial Maintaining Sperm/Seed Banks
Beneficial Stocking captive-reared wild-strain animals
Beneficial Transplanting wild animals
Beneficial Supplemental Feeding
Beneficial Restricting Poaching
Beneficial Controlling/Restricting Noncommercial Harvest
Beneficial Regulating commercial harvest levels
Beneficial Controlling/Removing Feral Animals
Beneficial Controlling/Removing Domestic Animals
Adverse Subsistence Hunting/Fishing/Trapping
Existing Subsistence Hunting/Fishing/Trapping
Adverse Incidental Capturing/Killing
Existing Incidental Capturing/Killing
Adverse Harassment/Vandalism/Indiscriminate Killing
Existing Harassment/Vandalism/Indiscriminate Killing
Adverse Poaching
Existing Poaching
Adverse Commercial Exploitation
Existing Commercial Exploitation
Adverse Sport Hunting/Fishing
Existing Sport Hunting/Fishing
Adverse Off Road Vehicles
Existing Off Road Vehicles
Adverse Food Supply Reduction
Existing Food Supply Reduction
Adverse Low Gene Pool
Existing Low Gene Pool
Adverse Hybridization
Existing Hybridization
Adverse Disease
Existing Disease
Adverse Parasites
Existing Parasites
Adverse Gas/Oil Development
Existing Gas/Oil Development
Adverse Underground Mines
Existing Underground Mines
Adverse Surface Mines
Existing Surface Mines
Adverse Rural Residential/Industrial Areas
Existing Rural Residential/Industrial Areas
Adverse Recreational development
Management Practices - 1 (DRAFT) - Management Practices
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
RESULT MANAGEMENT PRACTICE
Existing Recreational development
Adverse Highway/Railroads
Existing Highway/Railroads
Adverse Transmission Lines/Towers
Existing Transmission Lines/Towers
Adverse Soil compaction by heavy equipment in mine areas
Existing Soil compaction by heavy equipment in mine areas
Adverse Strip mining
Existing Strip mining
Adverse Competition
Existing Competition
Adverse Exotic/Feral/Introducted Species
Existing Exotic/Feral/Introducted Species
Adverse Predator control
Existing Predator control
Adverse Erosion
Existing Erosion
Adverse Grazing
Existing Grazing
Adverse
Existing
Adverse Vegetation Composition Changes
Existing Vegetation Composition Changes
Adverse Forest Alteration
Existing Forest Alteration
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
While the early European settlers of North America killed wolves
for their pelts, as did the Indians before them, they also considered
it beneficial to reduce the number of wolves that competed with humans
for wild game. Wolves came to be taken in larger numbers as the
market for their furs developed (06,55). A major reason for the
species decline was primarily due to the bounties imposed in the
1600's which almost extirpated wolves from the region by the 1800's
(69). Lack of information about the wolf, and misinformation believed
by the public, further contributed to the animals extirpation (04,06).
Habitat destruction was an indirect factor in this extirpation
because as wild habitat was destroyed and livestock introduced,
opportunities for wolves to prey on livestock increased. The decline
of wolves was hastened by the steady conversion of wild habitats,
which had supported wild prey populations, to farms, ranches, towns,
cities, recreation areas (including disturbance factors related to ORV
use parks), and similar human-utilized regions. In the Southwest,
continuing urbanization of the attractive Sun Belt area places demands
on southwestern forests for recreation, big-game hunting, increased
production of timber and livestock, and continuing attempts to utilize
the soils (which are highly erodible) and water (which is in great
demand) of southwestern arid lands for growing non-native farm crops.
Management Practices - 2 (DRAFT) - Management Practices
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
These are all threats to the few wolves that may still occur is the
Southwest or may be released in this area. In fact, there is a direct
relationship between road densities, or human access, and habitat
suitability for wolves (10). Vegetative cover affects the wolf's
survival in that it constitutes a factor in the welfare of the prey
species on which the wolf depends, and vegetative cover is useful as a
means of screening the wolf's presence from human pursuers or from the
prey that the wolf pursues. Continuing conversion of wild habitats to
human uses contributes by reducing cover, space, and the wild prey
food base for wolves (43,54,10), and also hinders the wolf's ability
to disperse and repopulate areas. The combination of increased
accessibility into wolf habitat and the human attitude toward wolves
is probably the most important factor threatening the species.
Because wild habitats were altered through substitution of
domestic livestock for native wild ungulates it became important
economically to ranchers and farmers to try to eliminate the wolves
and other wild predators that adopted to livestock as prey. The wolf
was deliberately exterminated from most of the U.S. by government
programs and private individuals because of real and perceived threats
to livestock. In 1915, the Federal government began a massive,
heavily staffed predator control campaign using chiefly poison and
traps. Of the many methods of extermination used, poison was the most
effective. Nevertheless, the extermination programs were so complete
that wolves were even eradicated from National Parks and wilderness
areas without livestock. Many subspecies of wolves became extinct and
wolves were all but eliminated in the lower 48 states (06,55,04).
At the time predator control campaigns severely reduced wolf
numbers within the U.S., the major portion of the range of the wolf
(C. l. baileyi, C. l. lycaon, and to a lesser extent C. l. irremotus)
was in Mexico and Canada, respectively. For many years, wolves
continued to come north from Mexico and enter southern Arizona,
southwestern New Mexico, and, occasionally, Trans-Pecos Texas. In
the 1930's and 1940's, however, Mexican ranchers also began to apply
more effective wolf-control measures. In the 1950's, the U.S. Fish
and Wildlife Service trained Mexican ranchers and veterinarians in
the use of 1080 (strychnine). Thereafter, wolf numbers declined
rapidly in Mexico (43,54,57). Likewise, the northern populations of
wolves in the U.S. were mostly transients from Canada and consisted
of the southern-most range limit of the Canadian populations.
Extermination of the Canadian populations was not as complete as was
the campaign south of the Canadian border.
Because remaining wild Mexican wolves are so few and scattered,
it becomes increasingly likely that they may mate with domestic dogs
in rural areas. The last male wolf taken in Mexico for the Mexican
wolf captive breeding project was captured in 1980 at a ranch where he
had bred successfully with a ranch dog (63). Feral dogs
(C. familiaris) directly affect the survival prospects of wolves in
areas where wolf numbers are very low. In Italy, the remaining gray
wolves suffer from competition for food with a feral dog population of
80,000 dogs (79). Where wolf numbers are extremely low, they will
interbreed with dogs (79). The coyote (C. latrans) competes with the
wolf for many prey species. Wolves are often antagonistic toward
coyotes, and the presence of wolves has been credited as the cause of
Management Practices - 3 (DRAFT) - Management Practices
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
coyote declines in some areas (04,53). Mech (04), however, feels this
relationship is not proven and points to cases of interbreeding
between gray wolves and coyotes in the northeastern United States and
between red wolves (C. rufus) and coyotes in the south-central states.
Outbreaks of mange and rabies also contributed to declines in an
already stressed Mexican wolf population in the Southwest (56) and
could cause loss of any remaining wild wolves. Accidental killing of
wolves by vehicles and incidental taking in traps, set for other
furbearers, are also contributing factors (11).
Deliberate killing for livestock protection still threatens
remaining wolves, even though wolves are protected by law in both
Mexico and the U.S. (43). Illegal killing of wolves in the Great
Lakes region along with heartworm and parvovirus are immediate threats
to the species recovery in the lower 48 states. In some areas the
range of a wolf pack may include both the U.S. and Canada. It is
legal in Canada to hunt or trap the species, therefore possibly
slowing the expansion of the southern-most range limit of the Canadian
populations. Control measures directed against other large predators
can also jeopardize wolves that may occur in or enter areas where
poisons or traps are used (43,54). Of all the factors that affect the
existence of the remaining wild wolves, those that significantly
affect the welfare of the wolves' prey base, particularly the
ungulates, are indirect factors that will affect wolf populations.
Currently, the species range in the lower 48 states represents
approximately three percent of its historical distribution.
Future perceived threats to the wolf include primarily continued
habitat destruction (heavy equipment, surface drainage, erosion on
overgrazed lands, etc.) through development of roads, railroads,
powerlines, mining (particularly for copper in the Southwest), gas and
oil development (particularly in the Rocky Mountain area) and land
clearing (for urbanization, agriculture, grazing, recreation, etc.).
Coupled with this factor is continued fear and misinformation about
the wolf on the part of humans living in wolf range, resulting in
illegal killing of the animal (09,10).
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1978. Eastern Timber Wolf Recovery
Plan. U.S. Fish and Wildl. Serv., Twin Cites, MN. 2 Vols.
U.S. Fish and Wildlife Service. 1982. Mexican Wolf Recovery Plan.
U.S. Fish and Wildl. Serv., Albuquerque, NM. 103 pp.
U.S. Fish and Wildlife Service. 1980. Northern Rocky Mountain Wolf
Recovery Plan. U.S. Fish and Wildl. Serv., Denver, CO. 67 pp.
UNAPPROVED PLAN:
U.S. Fish and Wildlife Service. 1984. Northern Rocky Mountain Wolf
Recovery Plan. Revised Agency Review Draft. U.S. Fish and Wildl.
Serv., Denver, CO. 84 pp.
The primary objective of the Eastern Timber Wolf Recovery Plan is
to maintain and re-establish viable populations of the eastern timber
Management Practices - 4 (DRAFT) - Management Practices
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
wolf (Canis lupus lycaon) in as much of its former range as is
feasible. The following objectives are identified in the recovery
plan:
1) Insure perpetuation of the timber wolf population at levels optimum
to the varying parts of its present Minnesota range (optimum level
includes biological carrying capacity and compatibility with man).
Maintaining early stages of succession by promoting logging and
selective thinning to maintain a healthy prey base is specifically
advised. Protection from poaching, for either commercial or
noncommercial purposes, is also identified.
2) Protect and enhance existing wolf numbers and re-establish
populations if necessary and feasible at optimum levels in Michigan
(excluding Isle Royale), Wisconsin and/or northeastern United States
and/or southern Appalachians region. Transplanting wild individuals
and captive propagation are specifically discussed in the plan.
3) Continue management to perpetuate natural conditions for the
wolf on Isle Royale National Park, Michigan. Permitting natural fires
to run their course is essential to providing browse for the ungulate
prey of the wolves here.
The primary objective of the Mexican Wolf Recovery Plan is to
conserve and ensure the survival of the Mexican wolf (C. l. baileyi)
by maintaining a captive breeding program and re-establishing a
viable, self-sustaining population of at least 100 Mexican wolves in
the middle to high elevations of a 5,000-square-mile area within the
subspecies historic range. The following objectives are identified in
the recovery plan:
1) Inventory and evaluate remaining gene pool.
2) Protect the remaining gene pool from poaching, commercial, or
noncommercial harvest.
3) Re-establish and maintain viable wild populations of Mexican wolves
in at least two areas in Mexico and/or adjoining areas of the
southwestern United States. This objective includes propagating
Mexican wolves in captivity, transplanting wild individuals if any
can be found, and maintaining sperm banks and ova collections to
preserve the genetic integrity of the subspecies. This task also
specifically calls for the planned release of Mexican wolves into the
wild and the seeking of steps to reduce conflicts between wolves and
humans such as supplemental feeding.
The primary objective of the Northern Rocky Mountain Wolf
Recovery Plan is to remove the subspecies from the endangered and
threatened species list by securing and maintaining at least 30
breeding pairs dispersed over 3 recovery areas with a minimum of 10
breeding pairs maintained in each of 3 recovery areas for a minimum of
3 successive years. The secondary objective is to reclassify the
subspecies (C. l. irremotus) to threatened status over its entire
range by securing and maintaining at least 20 breeding pairs with a
minimum of 10 breeding pairs maintained in each of 2 recovery areas
for successive years. The following objectives are identified in the
recovery plan:
1) Determine the present status and distribution, and devise a
systematic approach for compiling observations.
2) Establish population goals for a threatened and fully recovered
population, and reclassify or delist when these parameters are
Management Practices - 5 (DRAFT) - Management Practices
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
reached.
3) Delineate recovery areas, identify and develop conservation
strategies, and management plans to insure perpetuation of the
subspecies. Should recovery through natural recruitment fail; this
task may include translocating wolves into certain delineated wolf
recovery zones and the coordination of multiple-use activities with
the biological requirements of the subspecies. Conflicts and impacts
will be identified and procedures such as reparations for damages,
trapping, or relocation will be developed for resolving these
situations. The controlling of domestic or feral livestock along with
other agricultural practices such as poor animal husbandry or
practices such as allowing winter kills to decay where they drop may
have to be curtailed. The controlling of human access by restricting
development or gating logging roads, for example, may also be
compatible with multiple-use activities. Habitat maintenance
promoting early stages of succession through prescribed burning,
and timber harvesting techniques like selective thinning will help the
ungulate prey base propagate, reducing the likelihood of
wolf-livestock conflicts.
4) Monitor population recovery.
5) Develop and initiate information and education programs to prevent
poaching or accidental shooting among the other benefits.
Recovery activities, mostly for the northern Rocky Mountain
population, include implementation of a computer storage and retrieval
program for wolf sightings; and research on population status and
characteristics, habitat use, food habitats, and predator/prey
relationships, dispersal, and the impacts of human activities on wolf
behavior.
Management Practices - 6 (DRAFT) - References
Species WOLF, GRAY
Species Id ESIS059001
Date 14 MAR 96
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