MAMMALIAN SPECIES 1o.22.1-42 5
Canis rufus. By John L. Paradiso and Ronald M. Nowak.
Publi
shed 29 November 1972 by The American Si
society of Mammalogists
Canis rufus Audubon and Bachman, 1851
Red Wolf
Lupus niger Bartram, 1791:199. Type locality, Alachua
wvapna (now Payne's Prairie), Alachua Court, Florida.
‘The International Commission on Zoological Nomenclature
in Opinion 447, published 29 Janvary 1967, placed Bartram
(1791) on its official Index of Rejected and Invalid Works
in Zoological Nomenclature (see Nowak, 1967).
Canis inpus vat. rufus Audubon and Bachman, 1851, 2:240
“Type locality designated by Goldman (1957345) as 15 miles
west of Auetin, Texas.
Canis rufus: Bailey, 19052174, fret use of name combination.
CONTEXT AND CONTENT. Order Carnivora, Family
Canida, Subfamily Caninae, ‘Three subspocies are currently
recognized (Goldman, 1997345) as fllowst
C. r floridanus Milles, 1912:95, Type feom Horse Landing,
‘St Johns River, about 12 miles south of Palatka, Putnam
Coxnty, Florida
1. gregoryi Goldman, 1987:48. Type from Macks Bayou, 3
‘ales east of Tenaas River, 18 miles southwest of Tallulah,
‘Madison Parish, Louisiana
4, rafus Audubon and Bachman, 1851, Vol. 2:240, ee above.
DIAGNOSIS. As noted by Goldman (Young and Gold
‘man, 1944:900), there is such s great degree of individual,
igeographic, sexial, and age variation within each species of
North American Canis, and such wide specif overlap In most
characters, that much of the following diagnosis is necessarily
ener and quale in nature. Bostic Hemiation of sex
{mens often requires comparison with arge series. Measurements
and ratios inthis diagnosis pertain to specimens in the United
States National Museum
‘C-rajus is highly vasiable in all characters; the following,
however, are the mest diagnostic. Skull narzow anc elongated
with Tong, slendor rostrum, and flat frontal region: postorbital
constrict relatively marrow and elongated; braincase rela
Lively! small; sagittal crest usually welleveloped (sce figure
1). Canine teeth long and slender, gencrally eatending below
the level of & line drawn across the anterior mental foramina
when the. jaws are losed; pronounced deuteroeane ‘preset
tn PH (capital initials indieate upper teeth); metaconule
‘welldeveloped on M1; pronounced cingulum on upper molar
eth; M2" large. in_ proportion to size of skull Crypomatic
Dreadeh averages 113 Limes greatest transverse diameter of M2
in 158 specimens), Greatest length of skull for adult specimens
from Louisiana, Arkansas, and. Missouri, collected "prior to
1990, ranges from 2173 to 2610 mim for’ TA males, and from
2095 to 28720 mun for 69 females
Compared with. C. larans, rufus is always larger, both
‘externally and cranially whem specimens of only one sex te
‘compared, (the geeatest length of skull of larans ranges from
VARA to 2158 mm for 176.males and from T7L6 to 2065 for
ANS females). “The sagittal crest in rufus invariably exhibits
more pronounced development; the postorbital constriction is
relutiely ‘marrower and more. clongited, and the braincase
felatively smaller’ and. more. heavily owified. Tho M2. is
Smaller In proportion to size of skull than in latrans.(xygomatie
breadth averages 11.9. times greatest tansverse diameter of
Mz measured. diagonally in 390. adult specimens of latrans
collected throughost the range of the species). In all other
‘cranial and dental details, rufus and latrans show a eons
resemblance to each other.
Canis rufus resembles C. lupus in size (12 male. Jupus
ange from 250.7 to 2869 mm, and 30 females from 2240 to
2775 mm in greatest length of skull) bot differs as follows: the
Skull ix more slender and less massive (although in greatest
Length arog ‘ull may ‘be longer): ostrom longer and
narrower; canine teeth longer (in lupus they do not exten
Below'a line drawn across the anterior mental foramina when
the jaws ate closed); pronounced deuterocone present ot
PA; metaconule well matked on M1; cingulum on upper
role “he ME Farge im propor feof kul than
In lupus (aygomatic breadth averages 98 times gretest trans
verse diameter of M2 measured diagonally in 168 adult spe
tens of fupus collected throughout the range of the specie).
sim ri “ually an he erent ro
familiar’ Uys combination of pelage, cratial, and. dent
Cher raf he ten nals P, ty and M2)
fre generally larger; canine teeth longer and more slender
proportionally; rogram lave. longer and more slender,
Und frootal region flatter. ‘The dometie dog fs such a variable
imal; however, that somo spocies an elesely resemble any
af the wild apetes of Novth American Canis
‘Other diferences and smilerties between rufus, latrans,
lupus, and familias, are discussed by Goldman (Young and
Goldinan, 1948) and Lawrence and Bosser (1961)
GENERAL CHARACTERS. Doglike in general form,
with size averaging intermediate between fupus and latrans,
Tikough some large specimens of rufus overlap smeller spec.
tens of Tapus in measurements and weight. Weights and
Ieasurcments given in this section are bated on specimens and
records in the United States National Museum or upon reports
ip Bureau of Spot aberies ad Wilde Hogs the
“Total leath of rufus ranges from about 1955 to 1680 mm
(atrans abort 1080 {9 1320 mm, and lupus about 1870 to 2050
nm). ‘Recently collected. specimens ffom Chambers County,
exas, measured between 1859 and 1493 mm an total length,
Adult rufus clleeted in. Arkansas prior to 1930 weighed 3s
fellows averages fllowed by extremes) :'23 males, 60, 45
to 60'Tbs (21 to AL ke); 34 females, 476, 36 to 65 Ube (16
to 29. kg). Six" apecimens weighed recently” (1970) by
Bureau of Sport Fisheries and. Wildlife field agents in
Chambers County, Texas, were between 45 and 62 Ibs, and
averaged about 3 Ibs (24 ka). 'A recently collected (1970)
inate and female from Galveston County, Texas, weighed 50
Ths ‘and’ 45" Ibs respectively, and. the largest “animal ftom
Liberty. County” was’ 51 bs. One recently trapped (1970)
Ghambers County male had’ a shoulder height of 28 ‘inches
(07m. Young and. Goldman (1944:69) sated that the
weight of fully mature Zapus fs between 60 sad 175 Ibs (27
fand 77 ky), whereas lavrane. generally ranges {rom 18 10.30
Is (8 to 14 kg) (Young and Jackson, 1951:48)
‘A detailed description of the coloration of rufus was
iver by Goldman (Young and Goldman, 1994480). The
Most common calor phase ina large series of rufus) skins
(Gpecimnscolleeted prior to 1990 im Lauisiana, Texas, Okla
homa, Arkansas and Missouri) appears more reddish and more
spurcly haired than series of letrans and Lupus in comparable
pelage. Nevertheless, individual skins of all thre, and some of
Femur as well es be found that ace vitwallyindatinguith
abte from one. another: coloration docs not appear to be
agnostic character in North-American’ Gants, Both rufus
fand lupus often cecur in a black color phase, but according
io" Yotng, Noung, and Jackon, 85:58), "Bhack coloration
is extremely rare among edyotes
‘Young (1946:36) stated with regard to rufus: “Ie i rather
areyhound like in. appearance, with Tong, somewhat spindly
een "Although there are actually’ no comparative measure.
ments as yet to confirm this, Jong legs in rufus have. been
Commented pon hy several field biologists of the Bureau of
Spon Fisheries and Wildlife who have worked with the species
recently nthe Texas Coast. Glynn A. Riley, Jr, Principal
District Field Assistant for the Division of Wildlife Services,
Bureau of Sport Fisheries and Wildlife, Liberty, Texas, reported
(Septeniier 1970), that the legs of rufus ave strikingly Tong
tnd slender, giving the animal slmest the appearance of “being
fin stita” ‘Riley also reported thet the eats of Pufus are Tar
Targer in proportion to the size of the head than are the ears
ofthe fetrane aad apts with which he has worked,
DISTRIBUTION. Probably the original range of rufus
coincided well with the Couisinian, Cafolinian, and TexanFicune 1. Skull of Canis rufus from Cook Station, Crawford
Gounty, Missouri, collected on 2 April 1924 (USNM 244180)
Drawings by Mrs. Wilma Martin.
mammal provinces of Hagmeier (1966), but the species was
fxtipated at such an carly date in the eastern United States
that ie ip impossible to be certain, Specimens confirm that
i formerly cecursed. in Florida, Alabama, Mississippi Lau
lana, Arkaneas, southern Missouri, southern Indians, eastern
Oldahoms, and casterm Texas (ac figure 2)- Goldman (Young
and Goldman, 1944:486) aesigned a specimen from Wareaw,
Hancock Counts, ines, and another supposedly from Wabach
‘Wabash County Tndana, to rajus and thus placed the linitso
distribution ofthe species somewhat to far north The specimen
from Iligis, in the collection of the American. Murcum of
Natural History, was receled from C- K. Worthen, an animal
dlaler whose hame was in THancock County, Iinos. Apparently
the specimen, was a eaplive animal, and i is impossible to he
ferain of the locality data” (E~ Raymond Tall personal
ommunication), “The faded label of ‘the specimen trom
ndiana shows that it was actually taken in the
area of southern Indiana
Th the wester part of the range of the species, Goldman
(Young and. Goldman, 1944:188) sted a specimen of rufus
from Sheffield (22 alles north), Pecos County, Texas. ‘This
specimen is in the United States’ National Museum collection,
fad is actually & coyote, C. letras, There is no evidence that
Tajus ever oocured father west ia Texas than the Edwards
Plates
. rufus has been exterminated over most of its former
range. At the present time it is known to cccur in is pute
form (hybridization with Ferns is discuseed under section on
Genetics) only inthe coastal. praiice and ‘marshes of the
Gall) Goast counties of southeastern Texas and adjacent
Louisiana (Paradivo, 19653, Nowak, 1970, 1972). Specimens
have been obtained since 1960 and deposited in the United
tates National Museum from Brazoria (east of the Drazos
Riven, Chambers, Liberty, Harris. and Jefferson counties,
‘Texas.’ For information onthe werurrence of rufus in Lou
jana and Arkansas sce Nowak, 1967, 1970 and. Pimlatt and
Sealin, 1968
FOSSIL RECORD. No fossils have been assigned to
, rufus and there have been few attempts at direct comparison
of the modern red wolf to Pleistocene specimens. One of these
few was by Gazin (1942) who, in describing Cane edward
from the early Pleistocene of Arizona, stated: (p. S01); "The
stall and jaw of C. edwardii .\.. ate about intermediate in
bash iver
ee
\\
\
Ficus: 2 Map illustrating distibution of Canis rufus. ‘The
shaded portion shows the probable distbution prior to 1600
‘The dots represent the most marginal specimens (in museum
zllections) that can definitely be assigned to pure C. rufus
‘The stippling. depicts the area in southeastern ‘Texas where
pure populations of C. rufus now occur as indicated. by
Specimens. ‘The species also probably now exists in southern
Uooisiana,
sine between those of a gray wolf, and of a corote, about
‘equalling specimens of the'red wolf, Cans rufus frm MUsourt
land Arkantos «- The teeth are comparable to those in the
ed wif and fescmble them im stracture more closely than
they do. any other species of canida.” He did not, however,
Sugeest phylogenetic affinity Ietween the, two species, and
fonsidering the wide variation that Canis has always shown,
5 cannot be abit that ewer deine represents ony
Continau with modern rufus.
‘Another reference 10-2 fol in relation to moder rufus
involves Canis ermbraster’ Gidley, {rom Pleistocene. deposits
in" Cumberland Cave, Maryland, Goldman (Young and. Gold
‘man, 1944:399), in discussing ormbrastei, stated only that
i 'ppears to have been allied tothe red woll, Canis niger. *
ibbard(1955:82) “reported «right ramos of a. sal
volt in the late Pleistocene Upper Beserra Formation, Valley
(of Tequlaquiae, Mexico. Ile noted: "The tpecimen i smaller
than Cantt lupus ‘Linnaeus and. appears closely related. to
Canis ‘niger Bartram. But the tre relationship will not be
Jknoven until the upper dentition i found.”
Nowak (197084) considered the possibility that the pro
genitor of Cx rufus was closely related to C. leans, but had
Fecome isolated inthe southeastern United States hy Plestocene
glaciation. “He reported: A late Pleistocene. (Wisconsin)
fossil from Alachua County, Florida, appears to represent an
tinal taoatonal’ bewech “a coytehe acer andthe
modern red wo.”
FORM. Atkins and Dillon (1971) compared. the gross
smonphology of the ceebella of « number of species of the
genus Canis, including. rafos, and found that the cerebella
Sdiated a division of the genus into two groupe: = rufus
Iapas group ands detranjomuflarinjackal group.” Howerer,
‘aj lye rerun and eon characteris that i
fing fom apa.
‘Although Atkins and Dillon belived thatthe cerebellum of
rufus Sndicaten that sts closest affinities are ith Tapa, they
found that the rufus cerchellum ffers from that ofall species
of Canis examined in several portant traits, and resembles
ingame laters the crea othe fxs thy tai
(alopes Tagepas, Urocyon.cinereoargenteus, Vulpes. cham,
Weeder, nt both European and American Y. vulpes). They
‘snsidered the cecbela ofthese foxes to bo more primitive in
Sractare than those int Cans, and suggested that'& retention
‘f some foxlke characters nthe raf cerebellam may indeate
{hat ajar is nearer to the common ancestral stock of Canis
than ‘ave the other species of Modern Canis they examined
ONTOGENY AND REPRODUCTION. Nothing has
been published on. reproduction and development in rufus.
Data in the files of the Bureau of Sport Fisheries and Wildlife
(pertaining to specimens. taken prior to 1930) reveal. the
fallowing. In Arkansas, females with embryos were trappedbe 38 Fay on 10 Mas oa ith ng
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ECOLOGY. No comprehensive survey of the ecology of
gelesen oe. Th falling hcchy informatio,
“The range ‘of C. rafus was mostly within the humid
division ofthe Lower ‘Austeel Life Zone) and. the. species
apparently prefered a warm, mois, and’ densely ‘vegetated
iabita The ed wolf was equally at home in the virgin pine
forests, bottomland hardwood forests, or coastal prities and
marches of the southeart The body proportions ofthe species
may he an adaptation both for lite im swamps and. marahes
for long-distance running in coastal pravies and. in the
en pine forest of the southeast The eating overt! the
re sutastern United States, probably ontributed
forests of
to the decline of rafus (Nowak, 1970)
‘Canis ‘rufus docs not appear to have been a. major
predator of big yame. There are few records of its attacking
Farge herhivores such as are numerovs for lupus. Attempts to
Uist the food of rufus. (Young and Goldman, 1944; Davi,
1660; Beesley, 1967) refer mostly to rabbits, rodents and other
small prey. Concerning. the Louisiana red wolf, St. Amant
(1950:185)_ stated. that it it not Known to” what extent It
preys on deer, and that the major woll concentrations ate not
necessarily the areas most densely populated by deer. Even
feports sich as Howell's (1921) of ed wolf predation "on
Alomestic stock generally refer to small or younger animals
hreing taken. Tt should’ not be presumed, however, that the
red wolf exclusively preys on animals emaller than itelf
Catesby (1783, 2:26) ‘ote that wolves pursued deer in the
Carolinas, and Young "(1946:39) “noted that wild razorback
hops formed a major part of the food of the red wolves found
he Tomas River’ region of Louisiana. Jackson (961)
Stated thet groups of three or more red wolves were generally
Successful im attacking. adult cattle on” the JHIK ranch in
Chambers County, Texas. In general, it appears that the prey
fof rufus is intermediate in size between that of Zatvans and
lupus,
‘According to John Steele (personal communication,
Qetsber 1969} causes of mortality for the red woll inthe
Texas Gulf Coast area include man, hookworms, distemper,
and accidents. Most pups aequite hookwormns and ste
‘weakened by them tat they ‘cannot keep up with their
parents. They die indirectly from hookworms, and adults have
shortened Tife span de to. hookworms and heartworm
Most of the adults he ‘captured during. is surveys were
anemic and had low level infections of one sort or another
‘Man is probably the greatest enemy of rufus, and deliber.
ate killing appears to be one of the major factors In the
decline of the species (Nowak, 1970). "All personnel of the
Bureau of Sport Fisheries and Wildlife (Steele, McBride,
Riley) who have recently worked with C. rafus in the Gulf
Goast area of Texas, have commented (in personal communi
cations) on the ease with which red wolves can be trapped or
poisoned. ‘The species resembles latrans in that it is able to
survive in areas of relatively dense ‘human | populations—
rafus runways have been found in Galveston County, Texas,
within sight of housing developments (Riley, personal commu
nication, September 1970)--but it does not appear to possete
the cunning and caution widely attsbuted to letrans. ‘The
apparent ease of trapping rufus, combined with ecological
3
appear to be the major factors in
““(MeCarley, 1962; ‘Paradiso, 1968:
changes i
the decline of the sp
Nowak, 1970)
BEHAVIOR. Young (1946:36) noted the long legs and
slender build of rufus and felt that its greyhoundike body
would make it a etter long distance runmer than letrans,
Fitey (Personal ‘communication, ‘September 1970) also. was
impressed by the long les of rufus and stated that it appeared
to him that the ved wolf was an animal adapted for coursing
In open county.
Steele (personal communication, October 1968) stated
that in east Texas most rufus old their tils down at a 45"
fanale when standing, but some animals hold it near back level
Nearly" ail carry it horizomally when. they run.” Duting
sreetings and courtship, they raise 7 high above their backs,
Dethaps to activate sent glands.
He also sated that rufus does not run Hike a dog, but
hhas a bounding motion, somewhat ke 2 rocking hore, pausing
‘when the shoulders are highest. Red wolves lavestigate sounds
nd noises by standing up on their hind lees, especially in
{all gras and weeds
Steele noted thet in the Texas Gulf Coast countos red
wolves are most active at night, generally at the same. time
Tabbite are Teeding. Sometimes ‘red. wolves bed down at
‘ight in the middle of a herd of atte, In daylight they rest
in‘ weedy fields or gras or broth pastes, rom April t0
‘mid-August red’ wolves reset their travels to ho pot that
track sigos all but disappear. Beginning in September, they
resume travel over & hurting range. Mated pairs, sometimes
with an extra tale, travel together. Pecks of from five to
Ti animale may get together temporaily, but break up into
family aroupe toon, afte exchanging. grecings, Pairs travel
round ‘range using established ranways. marked by scent
fs-and serach marke
Psled wolves have a long smooth howl that ends on a
slightly higher note. "They also have a wide variety of yodeling
fries that sound exactly Tike those of coyotes. Vernon Bailey
n'a 1908 special report tothe Ul. 8, Biological Survey wrote
Gf the wolves in the Big Thicket of Hardin County, Tex
“Their vice is 4 compromise between that of the coyote and
the Tobo (C. Iupush or rather a deeper vied ap yap and
howl of the coyote. I suggest the coyote much more than the
bo"
‘Stocle further noted that in the Texas Gulf Coast area
dens ure found in hollow logs, stumps road culverts sand
Knotts, and banks of canals, ditches, and rerervirs. They are
fneraly screened from view hy berry vines, wild rose, brash
Piles, reeset
GENETICS. Studies conducted at the M.D. Anderson
Hospital and Tumor Insite, University of Texas, Houston,
Ihave demonstrated thatthe diploid chromosome number of « red
‘wolf collected in 1966 in Jeflerson County, Texas, wat 72, and
that the X chromosome only is biarmed. The karyotype of this
specimen is thus indistinguishable from those of Zatrans, lupus,
land familiar’, and chromosomal factors ‘would not io
interbreeding. among. these species. (Frances E. Atrighi,
personal communication, August 1969; and Mamm
{Chromosomes Newsleter 21:15, July 1966).
‘Goldman (Young and Goldnian, 1944:480) first noted the
posbilty of hybridization between leirons and rufus. He
trrote: "Specimens collected in the vicinity of Llano, ‘Ts.
Jnclude typleal examples of both species and individuals not
Sharply distinctive of either. Cloce approach in eeeetial
details and the apparent absence of "any. invariable unit
character suggest the pousbiity of hybridism in some local
licen Texas
‘McCarley (1962) felt that hybridization with latrans was
possibly one of the factors that brought about the near
Patermipation of rufus. He further suggested that the entire
Subspecies C7, rifas might. be a population of natural
liybutds between C. fatrans and C. rufus gregory.
‘Lawrence and Bosser (1067), using a multiple character
analyeiy, found that a small sample of Canis from Fallvlle
‘Nevwton’ County, Arkansis, spanned the whole range of varia:
tion from coyote to wolf and felt that thie indleated possible
hybridization.
‘Paradiso (1968) examined a large serie of Canis fom east
‘Texas collected after'1060, and found that they also spanned
the whole range of variation from typical latrans to typical
rufus, with all intermediates represented. He concluded that
massive hybridization had occurred between the two spectes in
this region.Nowak (2970) reported that his studies with Paradiso at
the United States National Museum indicated that in most
fareas the red wolf died out as a result of heavy hunting and
teeming press ‘end Tasive evita changes that
were unfavorable to. the species ‘Specimens in the National
Collection indicated that in many areas rufus was replaced by
ppuroletrans and there, was no indication of ‘hybridization
Between the two. Specimens from the Edwards Plateaw of
central Tete, collected around the turn of the present century,
id. show intermediate characters between rufus and letras,
leading Nowak to postulate that a hybrid swarm formed here.
Ho further postulated that thie hybrid swarm migrated east
ward, occupying. territory from which Co rufus had been
extirpated, and today i occurs throughout ‘most of castern
‘Texas. and Louisana,, Specimens indicated, that the upper
Gulf Coast region of Texas and probably adjacent Louisiana
fare perhaps the only areas In which C. rufus continucs 10
Survive as 8 pare specs, Nowak reported that early specimens
ff C.F. rufus in the National collection led him to believe that
it was a valid subspecies of rufus and did not represent hybrids
as suggested by MeCarley (1962)
REMARKS. There are a number of differing opinions
regarding the taxonomic affinities of rufus. Goldman Young
‘and Goldman, 194) regarded rufus av a full species, distinct
from both Zettans and tapas. Paradiso. (1968), struck by what
he thought was, massive hybridization between rufus and
lairans in east ‘Texas, suggested that the two might be com
specie. "Lawrence and Bossert (1907:229) concluded. from
their uultiple character analysis that “easly populations
described. ae Canis niger [== C. rufus floridans) a
igregoryt (= C. rufus aregoryfl from the southeastern wooded
Fegions, east ofthe range of Canis latrans, are only a local form
of Canis fupus, not & dstint species of woll” Nowak (1970)
feported trenchant differences between rufus, Topus, and
lairans, and regarded rufus asa full species, Atkins and
Dillon (1971) alco presented evidence from brain morphology
that rufas should stand apart as a distinct species from other
North American Cants, ‘Studies currently ‘being conducted
by Nowak. at the University. of Kansas on Pleistocene and
ecent'Canis in North America, and serological studies by
Uljeres §. Seale of the University of Minnesota, may throw
additional light on the relationshtps of Canis rufus.
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‘America’ American Wildlife Insitute, Washington, D. C
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“Ainerican Wildlife Management Institute, Washington, D.
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‘The principal editor of thie account was SyoNEY ANMERSON.
oun L, Parsowso, Buneav oF Sront Fisttems axo WiLoure,
Bino. Axo “Mascara, Lavorarones, Nanionat Mustvst oF
Naturat. Histony, Wastunycron, D.C. 20560, axo RONALD
M. "Nowak, Muscost or Neonat Hisrony, Usivensiry oF
‘Kansas, LaWnincs, Kansas 65084.