Hind Limb Myology of The Ringtail (Bassariscus Astutus) and The Myology of Hind Foot Reversal

Journal of Mammalogy, 97(1):211–233, 2016
DOI:10.1093/jmammal/gyv171
Published online October 27, 2015
Hind limb myology of the ringtail (Bassariscus astutus) and the

myology of hind foot reversal
Margaret Liu, Shawn P. Zack, Lynn Lucas, Darien Allen, and Rebecca E. Fisher*
School of Life Sciences, Arizona State University, 427 East Tyler Mall, Tempe, Arizona 85287, USA (ML, REF)
Department of Basic Medical Sciences, University of Arizona College of Medicine Phoenix, 435 North 5th Street, Phoenix,
Arizona 85004, USA (SPZ, LL, REF)
School of Human Evolution and Social Change, Arizona State University, 900 Cady Mall, Tempe, Arizona 85287, USA (SPZ, DA)
Margaret Liu and Shawn P. Zack are joint first-authors and contributed equally.
* Correspondent: rfisher@email.arizona.edu
The ringtail (Bassariscus astutus) is a widely distributed small carnivorous mammal (Procyonidae) in Mexico
and the southwestern United States. As in other procyonids, the ringtail is capable of rotating its hind foot to allow
headfirst descent of vertical substrates. The osteological correlates of this process, termed hind foot reversal, are
well documented, but potential myological correlates have never been investigated. We present the 1st detailed
study of the muscular anatomy of the hind limb of B. astutus, including the 1st muscle maps of the pelvis and pes
of any procyonid. Comparison of the hind limb myology of the ringtail with other arctoid carnivorans, including
taxa incapable of hind foot reversal, indicates that the muscles responsible for the action of reversal do not differ
significantly between nonreversing forms and taxa capable of partial or full reversal. This suggests that specific
myological adaptations are not necessary to achieve hind foot reversal. However, increased development of the
digital flexors, which maintain a grip while body mass is supported by the hind limb, may characterize taxa that
make use of reversed postures. The hind limb myology of members of Procyonidae does not strongly support
either morphological or molecular hypotheses of relationship, in part because relatively few differences among
members of the family can be documented.
Key words: hind foot reversal, hind limb, myology, Procyonidae, ringtail
© 2015 American Society of Mammalogists, www.mammalogy.org
The ringtail (Bassariscus astutus) is a small scansorial pro- 1984). Trapp (1972) suggested, based on postmortem manipu-
cyonid carnivoran native to Mexico and the southwestern lations of B. astutus, that this is aided by more proximal rota-
United States. The ringtail inhabits areas containing oak, pin- tion at the hip, but this is not supported by in vivo data.
yon pine, juniper, conifer, and chaparral (Taylor 1954; Bradley Hind foot reversal occurs in a diversity of arboreal and
and Hansen 1965; Grinnell et al. 1987; Poglayen-Neuwall scansorial mammals, including some didelphimorphians,
and Toweill 1988) and the ringtail diet is variable but broadly scandentians, sciurid rodents, and strepsirhine and platyrhine
omnivorous, including fruit as well as invertebrates and small primates (Trapp 1972; Jenkins and McClearn 1984; Cartmill
vertebrates (Taylor 1954; Toweill and Teer 1977; Trapp 1978; 1985; Fleagle and Meldrum 1988). Among carnivorans, hind
Alexander et al. 1994; Harrison 2012). As a scansorial mam- foot reversal has been documented in nandiniids (Nandinia
mal, the ringtail uses a variety of climbing techniques to move binotata—Taylor 1970), herpestids (Herpestes ichneumon—
through an uneven, irregular environment, including trees and Haines 1958), felids (Leopardus wiedii—Leyhausen 1963),
rock faces (Trapp 1972). and mustelids (Martes—Haines 1958; Heptner and Naumov
Central to the broad locomotor repertoire of B. astutus is the 1967). Additionally, except for the poorly known mountain
presence of substantial mobility in the limbs, including hind coati (Nasuella spp.), partial or complete capacity for hind
foot reversal, the ability to rotate the hind foot 180° while foot reversal is documented in all members of Procyonidae
descending a vertical support. In placental mammals, in vivo (Kaufmann 1962; Poglayen-Neuwall and Poglayen-Neuwall
radiographs indicate that hind foot reversal occurs via a combi- 1965; Trapp 1972; Jenkins and McClearn 1984; McClearn
nation of plantarflexion at the crurotalar joint and inversion at 1992). As in Bassariscus, the kinkajou (Potos flavus) exhib-
the subtalar and transverse tarsal joints (Jenkins and McClearn its complete hind foot reversal (Trapp 1972; Jenkins and
211
Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

by guest
on 28 August 2018
212 JOURNAL OF MAMMALOGY
McClearn 1984). The more terrestrial genera Nasua (white- Materials and Methods
nosed and South American coatis) and Procyon (raccoons) are
A total of 4 hind limbs, comprising the left and right hind limbs
only capable of partial reversal, rotating the foot 90° rather than
of 2 captive ringtails (B. astutus), were dissected at Arizona
180° from its normal anatomical position in which the distal
State University. The 1st individual, a fresh frozen specimen,
phalanges face cranially (McClearn 1992).
was an adult male of unknown age from the Phoenix Zoo. The
The osteological correlates of hind foot reversal have been
2nd individual, preserved in 70% ethanol, was an adult female
investigated by Jenkins and McClearn (1984) and Meldrum
of unknown age, originally from the Chicago Zoological
et al. (1997). In placental mammals, these include a reduced
Society and currently housed in the Field Museum of Natural
medial malleolus of the tibia, an arc of the tibial facet of the
History (FMNH 135423). During each stage of dissection, mus-
astragalus exceeding 180°, a medially angled astragalar neck,
cles were identified, labeled, and digitally photographed using
an expanded astragalar sustentacular facet continuous with the
a Canon T31 18MP CMOS Digital SLR Camera. Digital pho-
navicular facet, a bulbous and mediolaterally broad astragalar
tographs were also taken of a male hind limb skeleton (FMNH
head, a smoothly concave astragalar ectal facet, an inclined long
53761) from the Field Museum for muscle mapping. An addi-
axis of the calcaneal ectal facet, a concave calcaneal cuboid
tional hind limb skeleton (FMNH 53869) was utilized as a ref-
facet, and a calcaneal tuber that is deflected plantad (Jenkins
erence during dissection. Both of the latter specimens were also
and McClearn 1984; Meldrum et al. 1997). Most of these fea-
adult, captive individuals from the Chicago Zoological Society.
tures reflect modifications of the joint surfaces of the ankle to
During dissection, muscle origins and insertions were
facilitate inversion and plantarflexion. Only the last mentioned
recorded on transparency paper overlying the digital photo-
may reflect muscular adaptation, as Sarmiento (1983) has
graphs, creating muscle maps. Muscle weights were recorded
shown that this morphology increases the leverage of m. flexor
for each hind limb muscle in the preserved right limb and fresh
digitorum brevis, which is an important digital flexor during
frozen left and right limbs using a Scout Pro 400 g portable
hind limb suspension. Osteological features of the tarsus have
balance (Ohaus Corporation, Pine Brook, New Jersey). Each
been used to infer the capacity for hind foot reversal in extinct
muscle was removed at its origin and insertion points and mea-
mammals without close modern relatives such as multitubercu-
sured inclusive of tendinous fibers, unless otherwise noted.
lates and hyaenodontids (Jenkins and Krause 1983; Gingerich
The muscles of the preserved left limb were not weighed, as
and Deutsch 1989).
this limb was utilized to collect moment arm measurements.
In contrast, the myology of hind foot reversal is not well
Unfortunately, these measurements were difficult to replicate,
studied. In part, this reflects the fact that no modern study has
so we decided not to continue this set of measurements and
presented a focused examination of the relevant musculature in
they are not reported here. Muscle origins and insertions were
a mammal capable of hind foot reversal. Based on the motions
described based on dissection; muscle actions were described
that occur during the process of reversal, plantarflexion and
based on the way(s) in which the muscle crosses the joint(s)
inversion, it can be hypothesized that modifications should
and the actions of the myology of the red panda hind limb,
be sought in pedal plantarflexors and inverters. Relative to
which represent the consensus of muscle manipulations in
the same muscles in nonreversers, these muscles should show
multiple limbs during dissection (Fisher et al. 2008). As a rule,
enlargement and/or a proximal shift in their origin (to increase
the muscle maps and illustrations in this manuscript represent
leverage). Pierotti (in Jenkins and McClearn 1984) has sup-
the morphology present in the majority of the dissected limbs,
ported the role of m. tibialis cranialis as the prime mover dur-
unless otherwise noted in the text. In rare cases, when variation
ing reversal indicating that this muscle in particular may show
is high and a consensus morphology is not present, in-text cita-
modification in taxa capable of reversal.
tions will designate which variant is represented in an accom-
To investigate the potential myological correlates of hind
panying figure(s).
foot reversal, this study presents the 1st description of the
In order to test whether or not there are specific myologi-
hind limb myology of B. astutus. To date, information on
cal adaptations related to hind limb reversal, the myology of
Bassariscus myology has been limited to a few observations
the ringtail was compared to accounts from the literature of the
by Davis (1949, 1964). Most of these observations pertained
myology of a diversity of other arctoid carnivorans (Table 1).
to forelimb musculature and were not accompanied by either
Among non-procyonids, an emphasis was placed on the com-
detailed descriptions or illustrations. Description of the hind
parisons to the lesser grison (Galictis cuja) and the red panda
limb myology of B. astutus fills a significant gap in knowl-
(Ailurus fulgens). Both belong to families (Mustelidae and
edge of the anatomy of this animal. In addition, this study
Ailuridae, respectively) that are closely related to Procyonidae
presents an opportunity to produce muscle maps of all major
and the hind limb myology of both taxa has recently been
hind limb elements of B. astutus. Muscle maps document the
revised in detailed, extensively illustrated accounts that facili-
bony origins and insertions of muscles and facilitate compar-
tate comparisons (Fisher et al. 2008; Ercoli et al. 2013). Of the
isons between taxa and identification of osteological corre-
2, the lesser grison is terrestrial and typically does not climb,
lates of myological features. To date, hind limb muscle maps
whereas the red panda is arboreal. There is no indication in
of members of Procyonidae are restricted to the femur, tibia,
the literature that the red panda is or is not capable of hind
and fibula of the common raccoon (Procyon lotor—Gowell
foot reversal. However, based on observations of video of cap-
1897).
tive A. fulgens, we believe that partial or complete reversal is

by guest
on 28 August 2018
LIU ET AL.—HIND LIMB MYOLOGY OF THE RINGTAIL 213
Table 1.—Sources of data used for comparisons to Bassariscus. Superscripts indicate number of specimens examined, if known. Locomotor
(Loc.) data from MacLeod and Rose (1993), Nowak (1999), and Egi (2001). A, arboreal; Aq, semiaquatic; F, semifossorial; S, scansorial; T,
generalized terrestrial.
Taxon Loc. Sources

Ursidae
Ailuropoda S Davis (1964)1
Helarctos S Ray (1949)1
Ursus S/T Cuvier and Laurillard (1849); Haughton (1867b)1; Shepherd (1883)1;
Kelley (1888)1; Windle and Parsons (1898)1; Davis (1964)3
Mephitidae
Mephitis F Hall (1926)1
Spilogale T Hall (1926)1
Ailuridae
Ailurusa S Carlsson (1925)1; Fisher et al. (2008)4
Mustelidae
Eira S Macalister (1873a)1
Enhydra Aq Howard (1975)1
Galictis T Windle and Parsons (1898)1; Ercoli et al. (2013)3
Ictonyx F Windle and Parsons (1898)2
Lontra Aq Fisher (1942)3; Williams (1955)
Lutra Aq Cuvier and Laurillard (1849); Haughton (1867c)1; Cunningham (1879);
Windle and Parsons (1898)1
Lutrinaeb Macalister (1873b)1
Martesa S Cuvier and Laurillard (1849); Mackintosh (1875–1877)2; Hall (1926)1
Meles F Cuvier and Laurillard (1849); Haughton (1867a)2; Windle and Parsons
(1898)1; Scherling (1989)6
Melogale F Beddard (1905)1
Mustela T Alix (1876); Barone and Deutsch (1953)
Neovison Aq Williams (1955)
Pekania S Feeney (1999)4
Taxidea F Hall (1927)1
Procyonidae
Bassaricyona A Beddard (1900)1; Davis (1964)
Bassariscusa S Davis (1964)
Nasuaa S Cuvier and Laurillard (1849); Mackintosh (1875–1877)2; McClearn (1985)2
Potosa A Beswick-Perrin (1871)1; Windle and Parsons (1898)2; Julitz (1909)1
Procyona S Allen (1882)2; Windle (1888)1; Gowell (1897)3; Windle and Parsons (1898)1;
McClearn (1985)3; Feeney (1999)2
Taxon capable of partial or full hind foot reversal. The status of some other taxa is unclear (e.g., Eira).
a
b
Identified as Aonyx leptonyx (=Aonyx cinereus) by Macalister (1875). However, Macalister (1873b) had earlier stated that the specimen did not match the
morphology of A. leptonyx, and his later reference to this as a “large Indian Otter” is inconsistent with A. cinereus, the smallest living otter. This may represent
Lutrogale perspicillata, the largest otter recorded in India, but this cannot be determined with certainty.
likely in this species (Wildscreen Arkive 2015). For this reason, of arboreal habitats without being capable of hind foot reversal.
G. cuja was used as the primary comparative exemplar of a Unfortunately, all of these taxa are too distantly related to per-
nonreversing carnivoran. mit meaningful comparisons of details of muscular anatomy.
Ideally, to isolate the potential myological correlates of Distinctions reflecting differences in positional behavior would
hind foot reversal, comparisons would be made to a closely be difficult to isolate from phylogenetic effects.
related arboreal/scansorial taxon that is not capable of hind The muscle terminology used in this manuscript conforms
foot reversal. Unfortunately, no taxon that is adequately docu- to the standards of the Nomina Anatomica Veterinaria (Waibl
mented myologically fits these criteria. A few scansorial/arbo- et al. 2005) as modified by Fisher et al. (2008). An additional
real musteloids (e.g., Eira) may conceivably be incapable of divergence from Waibl et al. (2005) concerns the muscle they
hind foot reversal, but both positional behavior and myology refer to as m. gluteofemoralis. The more widely used m. femo-
are inadequately documented for detailed comparisons. Ursids rococcygeus is employed in this work.
are somewhat more distantly related but are broadly scansorial.
However, extant bears are much larger than Bassariscus, and,
while they are capable climbers, are better classified as scanso- Results
rial/terrestrial (e.g., MacLeod and Rose 1993). Table 2 presents the hind limb muscle weights of B. astutus for
More distantly related taxa (e.g., cercopithecoid pri- the preserved right limb and fresh frozen left and right limbs.
mates) would better fit the desired positional criteria. Within M. gluteus superficialis.—M. gluteus superficialis originates
Carnivora, some felids (e.g., L. wiedii) also make extensive use from the iliac crest, the fascia of mm. intertransversarii dorsales

by guest
on 28 August 2018
Table 2.—Hind limb muscle weights for Bassariscus astutus. Absolute weights are given in grams Followed by percentage weights in paren-
theses. Values preceded by the “>” sign were damaged and the reported weight is an underestimate. A dash “-” indicates muscles that were too
damaged to meaningfully weigh. Weights listed as “< 0.01” did not register on the scale, which was accurate to 0.01 g.
Muscle Specimen
FMNH 135423 Frozen specimen
R L
M. gluteus superficialis 1.09 (2.6) 1.45 (3.0) 1.37 (2.7)
M. femorococcygeus 1.02 (2.4) 0.90 (1.8) 0.94 (1.9)
M. gluteus medius 1.69 (4.1) 2.30 (4.7) 2.27 (4.5)
M. piriformis 0.10 (0.2) 0.03 (0.1) -
Mm. gluteus profundus + gemellus superioris 0.32 (0.8) 0.33 (0.7) 0.40 (0.8)
M. gemellus inferioris 0.09 (0.2) 0.06 (0.1) 0.07 (0.1)
M. obturator internus - - 0.87 (1.7)
M. quadratus femoris 0.58 (1.4) 0.64 (1.3) 0.62 (1.2)
M. biceps femoris 3.98 (9.5) 4.31 (8.8) 3.51 (6.9)
M. abductor cruris caudalis 0.26 (0.6) 0.07 (0.1) 0.07 (0.1)
M. semitendinosus 1.77 (4.2) 2.26 (4.6) 2.41 (4.7)
M. semimembranosus 5.10 (12.2) 5.52 (11.3) 6.01 (11.8)
M. iliopsoas > 1.30 (3.1) > 2.65 (5.4) 2.83 (5.6)
M. sartorius 1.93 (4.6) 2.12 (4.3) 2.21 (4.4)
M. tensor fasciae latae > 0.66 (1.6) 1.00 (2.0) > 0.89 (1.8)
Mm. quadriceps femorisa 5.51 (13.2) 8.20 (16.7) 9.83 (19.4)
M. rectus femoris 1.74 (4.2) 2.59 (5.3) 3.27 (6.4)
M. vastus lateralis 2.44 (5.8) 3.65 (7.4) 4.50 (8.9)
M. vastus intermedius 0.23 (0.6) 0.34 (0.7) 0.42 (0.8)
M. vastus medialis 0.90 (2.2) 1.12 (2.3) 1.26 (2.5)
M. gracilis 1.82 (4.4) 2.35 (4.8) > 1.05 (2.1)
M. pectineus 0.42 (1.0) 0.48 (1.0) 0.51 (1.0)
M. adductor longus 0.45 (1.1) > 3.40 (6.9) > 4.10 (8.1)
M. adductor magnus 2.93 (7.0)
M. adductor brevis 0.12 (0.3) 0.11 (0.2) 0.11 (0.2)
M. obturator externus 0.71 (1.7) 0.75 (1.5) 0.74 (1.5)
Mm. gastrocnemius + flexor digitorum superficialis 3.46 (8.3) 3.67 (7.5) 3.85 (7.6)
M. soleus 0.67 (1.6) 0.87 (1.8) 0.78 (1.5)
M. popliteus 0.24 (0.6) 0.28 (0.6) 0.28 (0.6)
Long digital flexors
M. flexor digitorum lateralis 0.84 (2.0) 0.73 (1.5) 0.63 (1.2)
M. flexor digitorum medialis 0.39 (0.9) 0.39 (0.8) 0.32 (0.6)
Combined tendons 1.41 (3.4) 1.34 (2.7) 1.21 (2.4)
M. tibialis caudalis 0.17 (0.4) 0.22 (0.4) 0.17 (0.3)
M. fibularis longus 0.18 (0.4) 0.33 (0.7) 0.28 (0.6)
M. fibularis brevis > 0.34 (0.8) 0.33 (0.7) 0.30 (0.6)
M. extensor digitorum lateralis 0.08 (0.2) 0.05 (0.1) 0.04 (0.1)
M. tibialis cranialis 1.30 (3.1) 1.31 (2.7) 1.39 (2.7)
M. extensor digiti I longus 0.10 (0.2) 0.09 (0.2)
M. extensor digitorum longus 0.26 (0.6) 0.30 (2.7) 0.32 (0.6)
M. extensor digitorum brevis 0.11 (0.3) 0.13 (0.3) 0.10 (0.2)
M. abductor digiti V < 0.01 0.03 (0.1) 0.02 (0.0)
M. flexor digitorum brevis 0.14 (0.3) 0.13 (0.3) 0.15 (0.3)
M. interflexores 0.03 (0.1) < 0.01 -
M. quadratus plantae 0.04 (0.1) 0.02 (0.0) 0.02 (0.0)
Mm. lumbricales < 0.01 0.02 (0.0) -
Mm. adductores digitorum 0.02 (0.0) 0.02 (0.0) < 0.01
Mm. flexores breves profundi 0.46 (2.4) 0.31 (2.0) 0.36 (2.0)
M. flexor brevis profundus digit I 0.03 (0.1) < 0.01 0.03 (0.1)
M. flexor brevis profundus digit II 0.08 (0.2) 0.06 (0.1) 0.08 (0.2)
M. flexor brevis profundus digit III 0.07 (0.2) 0.10 (0.2) 0.08 (0.2)
M. flexor brevis profundus digit IV 0.14 (0.3) 0.07 (0.1) 0.11 (0.2)
M. flexor brevis profundus digit V 0.07 (0.2) 0.07 (0.1) 0.08 (0.2)
M. flexor hallucis profundus < 0.01 < 0.01 -
a
Including quadriceps tendon and patella.

by guest
on 28 August 2018
caudale, and the dorsal aspect of the transverse process of the was fused with the caudal head of m. semitendinosus. In both
1st caudal vertebra (Figs. 1A and 1C). This broad muscle lies fresh frozen limbs, the caudal fibers of the muscle were fused
superficial to mm. gluteus medius and gluteus profundus and with the proximal fibers of m. femorococcygeus. The muscle
becomes narrower as it dives deep to mm. femorococcygeus inserts along the inferior border of the greater trochanter and
and biceps femoris (Fig. 2). There was a variable amount of the lateral femoral shaft (Figs. 3B and 3D). The cranial fibers
fusion in the 4 limbs. In the preserved left limb, the cranial of m. gluteus superficialis flex and weakly abduct the hip joint,
fibers shared an attachment with m. tensor fasciae latae, while and the caudal fibers weakly extend and abduct the hip joint.
the caudal fibers fused with the cranial fibers of m. semimem- M. femorococcygeus (m. gluteofemoralis, m. abductor
branosus; however, in the fresh frozen right limb, the muscle cruris cranialis).—M. femorococcygeus originates from the
A Gluteus superficialis
B
Iliacus Sartorius
Gluteus medius
Tensor fascia latae
Rectus femoris
Psoas minor
Gluteus profundus
+
Gemellus superioris
Pectineus
Adductor brevis
Adductor longus
Obturator internus
Gemellus inferioris
Gracilis Quadratus femoris
Biceps femoris
Semitendiosus,
Adductor magnus ischial head
Biceps femoris Obturator externus Semimembranosus
Semitendinosus,
ischial head
C Gluteus superficialis
Gluteus medius
Sartorius
Gluteus profundus
+
Tensor fasciae latae
Gemellus superioris
Gemellus Quadratus
Rectus femoris inferioris femoris
Biceps femoris
Semitendinosus,
ischial head
Pectineus Obturator
externus
Adductor brevis
Adductor longus
Semimembranosus
Adductor magnus
Gracilis
Fig. 1.—Pelvis muscle maps for Bassariscus astutus (left side): A) dorsal; B) ventral; and C) lateral views.

by guest
on 28 August 2018
Gluteus
superficialis (cut)
Intertransversarius
dorsalis caudalis
Gluteus
medius (cut)
Gluteus
profundus Femorococcygeus
Sartorius Semitendinosus
Abductor cruris
caudalis (dotted)
Tensor
fasciae
latae Biceps femoris
Fig. 2.—Lateral view of the gluteal region and left thigh in Bassariscus astutus (dotted lines indicate deep muscles).
transverse processes of the 1st and 2nd caudal vertebrae and greater trochanter (Figs. 3D and 4). In the fresh frozen left
the caudal fibers of m. gluteus superficialis; the origin also lies limb, the muscle belly was partially removed during the dissec-
deep to mm. intertransversarii dorsales caudale (Fig. 2). The tion of m. gluteus medius. M. piriformis extends and abducts
proximal third of the muscle belly is distinct, then fuses with the hip joint.
m. biceps femoris, and becomes distinct again at the distal fifth M. gluteus profundus.—M. gluteus profundus is located deep
of the muscle belly. In the preserved left limb, the proximal to m. gluteus medius. The muscle originates from the ilium,
muscle fibers had additional fusion with the proximal fibers of just caudal to the origin of m. gluteus medius (Figs. 1A, 1C, 2,
m. abductor cruris caudalis. However, in the preserved right and 4). The cranial and ventral fibers share a border with the
limb, there was no fusion with the aforementioned muscles. origin of m. tensor fasciae latae, while the caudal fibers origi-
The muscle inserts just cranial to a ridge along the lateral fem- nate from a rugosity along the dorsal aspect of the acetabulum,
oral shaft, immediately caudal to m. vastus lateralis (Figs. 3B confluent with the origin of m. gemellus superioris (Fig. 1C).
and 3D). M. femorococcygeus abducts and weakly extends the The muscle gives rise to a tendon just prior to its insertion on
hip joint. the greater trochanter (Figs. 3B and 3D). M. gluteus profundus
M. gluteus medius.—M. gluteus medius lies deep to m. glu- extends and abducts the hip joint.
teus superficialis and originates from the dorsal ilium, iliac Mm. gemelli.—M. gemellus superioris originates from a
fossa, the fascia of mm. intertransversarii dorsales caudale, rugosity along the dorsal aspect of the acetabulum, confluent
and the ventral aspect of the sacrum (Figs. 1A, 1C, 2, and 4). with the caudal end of the origin of m. gluteus profundus (Figs.
The sacral origin is separable from the remainder of the muscle 1A and 1C). It passes distally, giving rise to a tendon just prior
proximally, but they fuse distally to share a common insertion. to its insertion along the proximal aspect of the trochanteric
The main belly of the muscle lies within the iliac fossa and fossa (Fig. 3A). M. gemellus inferioris originates from dorso-
gives rise to a short, robust tendon that joins with the tendon caudal ischium (Figs. 1B and 1C). In the preserved left limb,
from the sacral portion to insert on the dorsal aspect of the the tendon of m. obturator internus divided the muscle into
greater trochanter (Figs. 3A, 3B, and 3D). In the fresh frozen superficial and deep bellies. In the other 3 limbs, the tendon
left limb, the sacral origin was separable into 2 distinct por- of m. obturator internus was deep to and fused with m. gemel-
tions and in the preserved right limb, the insertion of m. glu- lus inferioris at its insertion in the trochanteric fossa (Figs. 3A
teus medius extended farther down the greater trochanter and and 4). Mm. gemelli laterally rotate the hip joint; m. gemellus
partially onto the insertion of m. gluteus profundus. M. gluteus superioris also weakly abducts the hip joint.
medius extends and abducts the hip joint. M. obturator internus.—M. obturator internus originates
M. piriformis.—M. piriformis originates from the dorsal from the obturator membrane and the medial aspect of the
aspect of the 3rd sacral and 1st caudal vertebrae and extends ischium and pubis (Fig. 1A). As the muscle exits the pelvis, it
distally and laterally to insert on the proximal aspect of the gives rise to a tendon that merges with m. gemellus inferioris at

by guest
on 28 August 2018
A Gluteus medius
B
Gluteus medius
Gemellus superioris
Obturator Gluteus profundus
Obturator internus + externus
Gemellus inferioris Vastus lateralis
Iliopsoas
Iliopsoas
Quadratus femoris
Adductor brevis, Gluteus superficialis
dorsal belly
Adductor brevis,
ventral belly
Vastus medialis
Femorococcygeus
Pectineus
Adductor magnus,
caudal belly Adductor longus
Adductor magnus,
cranial belly
Vastus intermedius
Gastrocnemius, Semimembranosus
lateral head + Soleus
Gastrocnemius,
medial head
Extensor digitorum
longus
C D
Gluteus medius
Gluteus profundus
Piriformis
Iliopsoas
Gluteus superficialis
Vastus medialis
Adductor brevis, Vastus lateralis
dorsal belly Adductor magnus,
caudal belly
Pectineus
Femorococcygeus
Adductor longus
Vastus intermedius
Adductor magnus,
cranial belly
Adductor magnus,
caudal belly
Gastrocnemius, lateral head +

Soleus
Semimembranosus
Gastrocnemius,
medial head
Medial collateral Popliteus
ligament Extensor digitorum Lateral collateral ligament
longus
Fig. 3.—Femur muscle maps for Bassariscus astutus (left side): A) caudal; B) cranial; C) medial; and D) lateral views.
its insertion point on the trochanteric fossa (Fig. 3A). M. obtu- ischium (Figs. 1B and 1C). In 3 limbs, there were dorsal and
rator internus laterally rotates and weakly abducts the hip joint. ventral muscle bellies (Fig. 4), but in the fresh frozen right
M. quadratus femoris.—M. quadratus femoris originates limb, there was only one muscle belly. The muscle insertion
deep to mm. biceps femoris and gemellus inferioris from the is on the caudal aspect of the lesser trochanter, extending to a

by guest
on 28 August 2018
shallow depression on the caudal aspect of the proximal femur with one another. M. semitendinosus extends the hip joint and
inferior to the greater trochanter (Fig. 3A). M. quadratus femo- flexes the knee joint.
ris extends and laterally rotates the hip joint. M. semimembranosus.—M. semimembranosus (Fig. 5) has
M. biceps femoris.—M. biceps femoris covers the majority of superficial and deep heads that are only distinct medially. Both
the lateral thigh and lies superficial to m. abductor cruris cauda- heads originate from the caudal aspect of the ischium (Figs.
lis (Fig. 2). It originates from the ischial tuberosity, deep to the 1B and 1C), deep to mm. semitendinosus and biceps femoris.
ischial origin of m. semitendinosus (Figs. 1A–C). The muscle The muscle inserts on the caudomedial distal femur (Figs. 3A
inserts on the lateral aspect of the knee and the crural fascia of and 3C), a distinct rugosity on the proximal, medial tibia (Figs.
the proximal lateral leg (Fig. 2). In both fresh frozen limbs, the 6B and 6C), and knee joint capsule as it fuses with the fascia
muscle was fused at its insertion point with m. abductor cru- of m. vastus medialis. In the fresh frozen left limb, the 2 heads
ris caudalis. The muscle also fuses with the tendon of m. ten- shared an origin but not an insertion: the deep head inserted
sor fasciae latae in the middle of the thigh. M. biceps femoris on the distal femur, while the superficial head inserted on the
extends the hip joint and flexes the knee joint. distinct rugosity on the proximal, medial tibia, and the crural
M. abductor cruris caudalis (m. tenuissimus).—M. abductor fascia. In the fresh frozen right limb, part of the deep head
cruris caudalis is a long, thin muscle that originates from the fused with the middle portion of m. adductor magnus. M. semi-
transverse process of the 1st caudal vertebra (Fig. 2). In the membranosus extends and adducts the hip joint and flexes the
preserved left limb, the proximal fibers were deep to and fused knee joint.
with the proximal fibers of m. femorococcygeus. Although there M. psoas minor.—M. psoas minor originates from the bodies
was no fusion in the fresh frozen limbs, the muscle was closely of the cranial lumbar vertebrae and inserts on the ventral ilium
associated with m. femorococcygeus. The muscle travels deep (Fig. 1B). M. psoas minor flexes the vertebral column.
to the caudal border of m. biceps femoris to insert on the cru- M. iliopsoas.—M. iliopsoas comprises mm. psoas major
ral fascia in the middle of the lateral leg (Fig. 2). M. abductor and iliacus. M. psoas major originates from the transverse pro-
cruris caudalis extends the hip joint and flexes the knee joint. cesses and bodies of the lumbar vertebrae, beginning with L3.
M. semitendinosus.—M. semitendinosus (Fig. 2) has a larger M. iliacus fuses with the ventral aspect of m. psoas major in the
ischial head and a smaller caudal head that are distinct proxi- iliac fossa. However, in the fresh frozen right limb, m. iliacus
mally but fuse from a third of the way down the muscle belly was removed prior to muscle identification. M. iliopsoas inserts
to its insertion point. The ischial head originates from the dor- on the medial aspect of the lesser trochanter, just cranial to the
sal aspect of the ischial tuberosity, superficial to the origin of insertion of m. pectineus (Figs. 3A–C, and 5). M. iliopsoas
m. biceps femoris (Figs. 1A–C), while the caudal head origi- flexes the hip joint.
nates from the transverse process of the 2nd caudal vertebra, M. articularis coxae.—M. articularis coxae is absent in the
just superficial to the ischial head. The muscle courses to the ringtail.
medial thigh, where it fuses distally with m. gracilis and inserts M. sartorius.—M. sartorius originates from the ventral and
on the proximal crural fascia and (with m. gracilis) onto the cranial aspect of the iliac crest (Figs. 1B and 1C) and courses
proximal tibia. In the fresh frozen left limb, there was no fusion over the cranial and medial thigh (Figs. 2 and 5). The muscle
with m. gracilis, but the 2 muscles were very closely associated fuses with the tendon of m. tensor fasciae latae at the lateral
Gluteus medius
Piriformis
Gluteus
profundus
Gemellus
inferioris
Quadratus
Rectus femoris:
femoris dorsal belly,
ventral belly
Vastus
lateralis
Fig. 4.—Deep view of the gluteal region of Bassariscus astutus (left side).

by guest
on 28 August 2018
Iliopsoas
Rectus femoris
Vastus medialis
Sartorius
Pectineus
Adductor Adductor
brevis Adductor
magnus
(dotted) longus Semimembranosus
Gracilis (cut)
Fig. 5.—Medial view of the left thigh of Bassariscus astutus.
knee and, in both preserved limbs, is also fused distally with fibers are fused with the distal fibers of m. vastus medialis.
m. gracilis near the medial knee. Although there was no fusion M. vastus medialis originates from the craniomedial femoral
with m. gracilis in the fresh frozen limbs, mm. sartorius and shaft (Figs. 3B and 3C). Mm. vasti lateralis, intermedius, and
gracilis were closely associated. The muscle inserts along the medialis extend the knee joint.
crural fascia of the medial knee and leg and the medial aspect M. gracilis.—M. gracilis originates just dorsal to a ridge on
of the tibial crest (Figs. 6B and 6C). M. sartorius flexes the hip the ventral pubis (Figs. 1B and 1C) and courses down the medial
and knee joints and weakly adducts the hip joint. thigh, with the cranial fibers fusing distally with m. sartorius,
M. tensor fasciae latae.—M. tensor fasciae latae originates and the caudal fibers fusing distally with m. semitendinosus to
from the ventral aspect of the iliac crest, deep to m. sartorius insert onto the crural fascia of the medial knee and leg (Fig. 5).
(Figs. 1B, 1C, and 2). It gives rise to a broad tendon starting in Additionally, some cranial fibers insert onto the proximal tibia,
the middle of the thigh, which fuses with the tendons of mm. just distal to the insertion of m. sartorius (Figs. 6B and 6C). In
sartorius and biceps femoris before inserting onto a ridge on the fresh frozen left limb, only the distal portion of the mus-
the lateral femoral shaft (Figs. 3B and 3D) and along the lat- cle was intact due to previous damage, but that aspect was not
eral knee. Additionally, the distal tendinous fibers insert with fused with either mm. sartorius or semitendinosus. M. gracilis
mm. quadriceps femoris along the cranial aspect of the knee. In adducts the hip and knee joints and flexes the knee joint.
the preserved right limb, the tendinous fibers also inserted onto M. pectineus.—M. pectineus originates from the cranial
the muscle belly of m. vastus medialis. M. tensor fasciae latae pubic ramus on the iliopectineal eminence (Figs. 1B, 1C, and
flexes and weakly abducts the hip joint. 5). In the preserved left limb, the muscle belly was fused with
Mm. quadriceps femoris.—Mm. quadriceps femoris consists the distal end of m. adductor longus. The muscle inserts along
of mm. rectus femoris, vastus lateralis, vastus intermedius, and the caudomedial aspect of the proximal femoral shaft, just infe-
vastus medialis. M. rectus femoris originates from a rugosity rior to the lesser trochanter, and continues halfway down the
on the distal ilium (Figs. 1A and 1C), deep to the origin of femoral shaft (Figs. 3A and 3C). M. pectineus adducts the hip
m. gluteus profundus (Fig. 4). The muscle courses along the joint.
cranial thigh, fusing distally with mm. vasti to form the quad- M. adductor longus.—M. adductor longus is caudal to mm.
riceps tendon that inserts directly onto the patella and, via the adductor brevis and pectineus (Fig. 5) and originates from the
patellar ligament, onto the tibial tuberosity. The muscle fuses cranial pubic ramus, deep to m. gracilis (Figs. 1B and 1C). In
with m. vastus lateralis halfway down the thigh. M. rectus fem- both fresh frozen limbs, the muscle was completely fused with
oris flexes the hip joint and extends the knee joint. M. vastus m. adductor magnus. The muscle inserts on the caudal femo-
lateralis lies deep to mm. tensor fasciae latae and biceps femoral shaft (Figs. 3A and 3C). M. adductor longus adducts and
ris (Fig. 4). The muscle originates from the cranial aspect of the weakly extends the hip joint.
greater trochanter, continuing down the craniolateral femoral M. adductor magnus.—M. adductor magnus has caudal and
shaft (Figs. 3B and 3D). M. vastus intermedius begins as a thin cranial bellies that originate deep to m. gracilis from the ven-
belly and thickens distally. The muscle lies deep to mm. rec- tral aspect of the pubis (Figs. 1B and 1C) and run toward the
tus femoris and vastus lateralis and originates from the cranial distal femur (Fig. 5). There were variable insertion sites in the
aspect of the femoral shaft (Figs. 3B and 3D). The proximal 4 limbs. In the preserved left limb, both bellies inserted along

by guest
on 28 August 2018
A B
Patellar ligament
Lateral collateral Semimembranosus Tibialis cranialis
ligament Lateral collateral ligament
Soleus Fibularis longus
Medial collateral ligament
Flexor digitorum Sartorius
Fibularis longus medialis
Gracilis
+
Extensor digiti I longus
Tibialis caudalis Semitendinosus
Fibularis brevis
Popliteus
Popliteus
Flexor digitorum
lateralis
Fibularis brevis
C D
Semimembranosus
Lateral collateral ligament
Patellar ligament
Patellar ligament Soleus
Soleus
Medial collateral Flexor digitorum lateralis
ligament Fibularis longus
Flexor digitorum
Sartorius
medialis Tibialis cranialis
Gracilis
Tibialis caudalis +
Semitendinosus Extensor digiti I longus
Popliteus
Fibularis brevis
Flexor digitorum
medialis
Fig. 6.—Tibia and fibula muscle maps for Bassariscus astutus (left side): A) caudal; B) cranial; C) medial; and D) lateral views.
a ridge on the caudal femoral shaft: the caudal belly insertion defines the medial border of the muscle. In the preserved right
was longer and more laterally positioned, began inferior to the limb, the caudal and cranial bellies were fused at their origin
greater trochanter and continued along the entire femoral shaft and fused with the cranial fibers of m. adductor longus. The
(Figs. 3A, 3C, and 3D), while the cranial belly insertion was caudal belly insertion began distal to the greater trochanter and
smaller, more distal, and medially positioned (Figs. 3A and ended two-thirds of the way down the femur, while the cranial
3C). Additionally, there is a ridge on the medial femur that belly inserted halfway down the caudomedial femoral shaft. In

by guest
on 28 August 2018
both fresh frozen limbs, the bellies were fused with m. adductor medial supracondylar tuberosity (Figs. 3A and 3C) and the
longus. In the fresh frozen left limb, the caudal belly inserted caudal aspect of the knee joint capsule, while the lateral head
just proximal to the lateral femoral condyle, while the cranial originates from the lateral supracondylar tuberosity (Figs. 3A
belly inserted just caudal to the insertion of m. adductor brevis and 3D) and the caudolateral aspect of the knee joint capsule.
and continued down the entire caudomedial femoral shaft. In Fabellae are embedded within both heads. The lateral head
the fresh frozen right limb, the distal portion of the caudal head and part of the medial head are fused on their deep aspect with
had been previously damaged, so it was difficult to discern the m. flexor digitorum superficialis (Fig. 7A). The medial and
insertion; in this limb, there was extensive fusion between the lateral heads fuse distally, and along with m. flexor digitorum
cranial and caudal bellies. M. adductor magnus adducts and superficialis, give rise to the calcaneal tendon, which inserts on
weakly extends the hip joint. the calcaneal tuberosity (Figs. 7A, 8A, and 8B). M. gastrocne-
M. adductor brevis.—M. adductor brevis has dorsal and mius flexes the knee joint and plantarflexes the ankle joint.
ventral bellies that are deep to m. pectineus (Fig. 5) and origi- M. soleus.—M. soleus lies deep to m. flexor digitorum super-
nate from the cranial pubic ramus, just ventral to the origin of ficialis and the lateral head of m. gastrocnemius (Fig. 7A). The
m. pectineus (Figs. 1B and 1C). The dorsal belly inserts just muscle originates from a bony protuberance on the caudal
distal to the lesser trochanter (Figs. 3A and 3C), while the ven- aspect of the fibular head (Figs. 6A, 6C, and 6D). The mus-
tral belly inserts on the caudal aspect of the proximal femo- cle gives rise to a tendon that fuses with the calcaneal tendon
ral shaft, between the insertions of mm. adductor magnus and to insert on the calcaneal tuberosity (Figs. 7A, 8A, and 8B).
pectineus (Fig. 3A). However, in the preserved right and fresh M. soleus plantarflexes the ankle joint.
frozen left limbs, there was only one muscle belly. In the fresh M. flexor digitorum superficialis (m. plantaris).—M. flexor
frozen left limb, the muscle had a small insertion just distal to digitorum superficialis is fused with and originates from the lat-
the greater trochanter on the caudal femoral shaft. M. adductor eral head and part of the medial head of m. gastrocnemius (Figs.
brevis adducts and weakly extends the hip joint. 3A, 3D, and 7A). However, the medial border of the muscle is
M. obturator externus.—M. obturator externus originates separable from m. gastocnemius. The muscle gives rise to the
from the obturator membrane and the lateral aspect of the shared calcaneal tendon with m. gastrocnemius to insert on the
ischium and pubis (Figs. 1B and 1C). The muscle has a robust calcaneal tuberosity (Figs. 7A, 8A, and 8B). M. flexor digito-
tendon that inserts in the trochanteric fossa (Fig. 3A). M. obtu- rum superficialis plantarflexes the ankle joint.
rator externus laterally rotates and weakly adducts the hip joint. M. popliteus.—M. popliteus lies deep to m. gastrocnemius
M. gastrocnemius.—M. gastrocnemius has medial and lat- (Fig. 7B). The muscle originates from the lateral femoral con-
eral heads (Fig. 7A). The medial head originates from the dyle and courses toward the medial tibial shaft (Figs. 3D and
A B
Gastrocnemius:
medial head,
lateral head
Popliteus
Flexor
digitorum
superficialis
(dotted)
Flexor
digitorum
Fibularis medialis
longus Tibialis
caudalis
Extensor
Flexor digitorum Flexor
Soleus
digitorum lateralis digitorum
medialis (dotted) lateralis
Fibularis
Tibialis
Fibularis brevis
caudalis
longus
Fibularis Flexor
brevis digitorum
lateralis
Fig. 7.—Caudal view of the left leg of Bassariscus astutus: A) superficial and B) deep dissections.

by guest
on 28 August 2018
A B
Flexores breves profundi: Tendo Extensor expansions:
a. Digit I medial belly Tendo Digit I: extensor digiti I longus
calcaneus
b. Digit I lateral belly calcaneus + extensor digitorum brevis digiti I
Abductor digiti V
c. Digit II medial belly Digits II-IV: extensor digitorum brevis digiti II-IV
d. Digit II lateral belly + extensor digitorum longus
e. Digit III medial belly + lumbricales digiti II-IV + flexores breves
f. Digit III lateral belly Quadratus profundi digiti II-IV
g. Digit IV medial belly plantae Digit V: extensor digitorum longus
h. Digit IV lateral belly Extensor + extensor digitorum lateralis
i. Digit V medial belly digitorum + lumbricalis digiti V + flexores breves
j. Digit V lateral belly Tibialis caudalis brevis profundi digiti V
Flexor hallucis profundus
Fibularis brevis Fibularis
Abductor digiti V Tibialis cranialis brevis
j
i
h c
g d
f e
Fibularis longus
Adductores digitorum a
b
i+j c
d+c
g Flexor digitorum lateralis +

+ Flexor digitorum medialis
I
h f
e I
Flexor digitorum brevis +
g +
Interflexores
f Extensor expansions
Flexor digitorum brevis
Flexor digitorum lateralis +
Flexor digitorum medialis
II
V
II
V
III IV
III
IV
Fig. 8.—Pes muscle maps for Bassariscus astutus (left side): A) plantar and B) dorsal views.
7B). The muscle inserts a third of the way down the medial caudomedial aspect of the fibular head, the interosseous mem-
tibial shaft (Figs. 6A–C). M. popliteus flexes the knee joint and brane, and the caudomedial tibial shaft (Figs. 6A and 6C). The
medially rotates the tibia. tendon travels caudal to the medial malleolus into the plantar
pes, where it fuses with the tendon of m. flexor digitorum late-
Mm. flexores digitorum profundi ralis to form a conjoined tendinous plate in the plantar pes.
M. flexor digitorum lateralis (m. flexor digiti I longus, m. flexor M. flexor digitorum medialis plantarflexes the ankle joint and
fibularis).—M. flexor digitorum lateralis (Figs. 7A and 7B) origi- flexes the metatarsophalangeal and the proximal and distal
nates from the fascial intermuscular septum that separates the lat- interphalangeal joints of digits I through V.
eral and caudal compartments of the leg, the caudal aspect of the M. tibialis caudalis.—M. tibialis caudalis originates from
fibular head and shaft, the interosseous membrane, and the cau- the caudal fibular head, the interosseous membrane, and the
dolateral aspect of the tibial shaft (Figs. 6A and 6D). In all limbs proximal tibial shaft (Figs. 6A and 6C). It gives rise to a ten-
except the preserved right limb, the muscle belly was fused with don halfway down the leg, travels caudal to the medial mal-
m. tibialis caudalis. In the preserved right limb, the muscle was leolus and into the plantar pes to insert on the plantarmedial
fused proximally with mm. fibularis brevis and extensor digito- aspect of the navicular and medial cuneiform (Figs. 7A, 7B,
rum lateralis. In the fresh frozen left limb, the lateral portion of and 8A). M. tibialis caudalis plantaflexes the ankle joint and
the muscle was fused proximally with m. fibularis brevis. The inverts the pes.
tendon fuses with the tendon of m. flexor digitorum medialis to M. fibularis longus.—M. fibularis longus (Figs. 7A, 7B, 10,
form a conjoined tendinous plate in the plantar pes (Figs. 9B and and 11) typically originates from a bony protuberance on the
9C). The common tendon plate gives rise to 5 tendons that insert caudolateral aspect of the fibular head (Figs. 6A, 6B, and 6D).
on the distal phalanges of digits I through V (Figs. 8A, 9B, and However, in the preserved right limb, the origin was from the
9C). M. flexor digitorum lateralis plantarflexes the ankle joint cranial aspect of the fibular head. In all limbs except the fresh
and flexes the metatarsophalangeal and the proximal and distal frozen right limb, the muscle belly was fused with the proxi-
interphalangeal joints of digits I through V. mal portion of m. tibialis cranialis. In the fresh frozen right
M. flexor digitorum medialis (m. flexor digitorum longus, limb, the proximal fibers were fused with the proximal fibers of
m. flexor tibialis).—M. flexor digitorum medialis lies just dis- m. fibularis brevis. The muscle belly gives rise to a tendon that
tal to m. popliteus (Fig. 7B). The muscle originates from the travels caudal to the lateral malleolus and deep to the fibular

by guest
on 28 August 2018
Fig. 9.—Plantar views of the left pes of Bassariscus astutus: A) superficial layer; B) layer 2; C) layer 3; and D) deep layer.
retinaculum that also encompasses the tendons of mm. fibularis base of the 1st metatarsal (Fig. 8A). M. fibularis longus everts
brevis and extensor digitorum lateralis (Figs. 10 and 11). The the pes and plantarflexes the ankle joint.
tendon travels deep to the long plantar ligament and through M. fibularis brevis.—M. fibularis brevis (Figs. 7A, 7B, and
an osseofibrous tunnel, which is bordered by the tarsals proxi- 10) lies deep to m. fibularis longus and originates from the cra-
mally and bases of metatarsals III through V distally, into the niolateral aspect of the fibular shaft (Figs. 6A, 6B, and 6D).
plantar pes. The insertion is on the plantarlateral aspect of the There was variable fusion in the 4 limbs. In the preserved

by guest
on 28 August 2018
Extensor
digitorum
longus
Tibialis
cranialis
Fibularis
longus
Extensor
digiti I longus
Extensor
digitorum
brevis
Tibialis
cranialis Extensor
digitorum
lateralis
Fibularis longus
Extensor digitorum
longus
Fibularis brevis
Extensor digiti I
longus
Fig. 10.—Craniolateral view of the left leg of Bassariscus astutus.
left limb, the muscle fused distally with the medial portion of
m. extensor digitorum lateralis near the lateral malleolus. In the
preserved right limb, only the proximal fibers were fused with Fig. 11.—Dorsal view of the left pes of Bassariscus astutus.
m. extensor digitorum lateralis; in this same limb, the proxi-
mal fibers of the muscle acted as the origin site for m. extensor malleolus. In the other 3 limbs, the muscle was fused proxi-
digitorum lateralis. In the fresh frozen left limb, there was no mally at its origin site with m. fibularis brevis until halfway
fusion with m. fibularis longus, but the proximal portion of the down the fibula; however, there was no association with m. tibi-
muscle belly also acted as the origin site for m. extensor digi- alis caudalis. In the fresh frozen right limb, there was additional
torum lateralis. M. fibularis brevis gives rise to a robust tendon fusion with m. fibularis longus. M. extensor digitorum latera-
halfway down the leg, which becomes fully tendinous at the lis gives rise to a tendon just proximal to the lateral malleolus
level of the lateral malleolus. The tendon courses deep to the and becomes fully tendinous once past the lateral malleolus. It
fibular retinaculum, alongside mm. fibularis longus and exten- travels alongside the tendons of mm. fibularis longus and fibu-
sor digitorum lateralis (Fig. 10), to insert on the tuberosity at laris brevis, deep to the fibular retinaculum. The insertion is on
the base of the 5th metatarsal (Figs. 8A and 8B). M. fibularis the distal phalanx of digit V, forming the extensor expansion
brevis plantarflexes the ankle joint and everts the pes. of digit V with the tendons of mm. extensor digitorum longus,
M. extensor digitorum lateralis.—M. extensor digitorum lumbricalis digiti V, and flexores breves profundi digiti V (Figs.
lateralis lies deep to m. fibularis brevis and is closely associ- 8B and 11). M. extensor digitorum lateralis plantarflexes the
ated with mm. fibularis longus and fibularis brevis (Fig. 7B). ankle joint, everts the pes, and extends and abducts the tarso-
In the preserved left limb, the muscle originated from the lat- metatarsal joint of digit V.
eral portion of the proximal belly of m. fibularis longus. In this M. tibialis cranialis.—M. tibialis cranialis covers the major-
same limb, the medial portion was fused with the lateral portion ity of the cranial leg (Fig. 10). It originates from a fossa formed
of m. tibialis caudalis. The muscle then joined mm. fibularis by the tibial tuberosity and the lateral tibial condyle, the proxi-
longus and fibularis brevis distally at the level of the lateral mal third of the tibial shaft, and the proximal portion of the

by guest
on 28 August 2018
interosseous membrane (Figs. 6B and 6D). The muscle belly of the calcaneus, just distal to the calcaneal tendon (Figs. 8A
is superficial to m. extensor digiti I longus (Figs. 10 and 11). and 9A). However, in the preserved left limb, the muscle origi-
M. tibialis cranialis gives rise to a tendon just proximal to the nated from the plantarlateral aspect of the calcaneus. It gives
medial malleolus, courses deep to the extensor retinaculum, and rise to a short tendon prior to inserting on the plantar surface of
travels medial to the tendon of m. extensor digiti I longus (Figs. the tubercle on the 5th metatarsal (Fig. 8A). M. abductor digiti
10 and 11). The tendon inserts on the plantarmedial aspect of V abducts the tarsometatarsal joint of digit V.
the base of the 1st metatarsal (Fig. 8A). M. tibialis cranialis M. flexor digitorum brevis.—M. flexor digitorum brevis orig-
dorsiflexes the ankle joint and inverts the pes. inates from the plantar surface of the calcaneal tendon (likely
M. extensor digiti I longus.—M. extensor digiti I longus lies the portion derived from the tendon of m. flexor digitorum
deep to m. tibialis cranialis (Figs. 10 and 11). It originates from superficialis) and has 4 muscle bellies that give rise to tendons
the craniomedial aspect of the proximal half of the fibular shaft traveling to digits II through V (Figs. 8A and 9A). The lateral
and is separated from the origin of m. fibularis brevis by a ridge 3 tendons fuse with the tendons of mm. interflexores going to
on the fibula (Figs. 6B and 6D). The muscle belly follows the digits III through V (Figs. 8A and 9A). All 4 tendons bifurcate
path of m. tibialis cranialis and gives rise to a tendon at the at the metatarsophalangeal joints, where the conjoined tendons
extensor retinaculum or at the level of the metatarsals on the of mm. flexor digitorum lateralis and flexor digitorum medialis
dorsal pes. After traveling through the retinaculum, it joins the pass through to the distal phalanges. After bifurcation, the short
tendon of m. extensor digitorum brevis to form the extensor flexor tendons insert together, deep to the tendons of mm. flexor
expansion of digit I, inserting on the distal phalanx (Figs. 8B digitorum lateralis and flexor digitorum medialis, on the middle
and 11). M. extensor digiti I longus dorsiflexes the ankle joint phalanges of digits II through V (Figs. 8A and 9A). M. flexor
and extends the metatarsophalangeal and the proximal and dis- digitorum brevis flexes the metatarsophalangeal and proximal
tal interphalangeal joints of digit I. interphalangeal joints of digits II through V.
M. extensor digitorum longus.—M. extensor digitorum lon- Mm. interflexores.—Mm. interflexores consist of 3 mus-
gus lies deep to m. tibialis cranialis (Fig. 10) and originates via cle bellies that originate from the common tendon plate of
a long tendon from the lateral femoral condyle, just cranial and mm. flexor digitorum lateralis and flexor digitorum medialis
slightly distal to the lateral collateral ligament (Figs. 3B and (Fig. 9B). They give rise to tendons that fuse with the tendons
3D). The muscle belly gives rise to a tendon at the level of the of m. flexor digitorum brevis going to digits III through V (Figs.
lateral malleolus and travels deep to the fibular retinaculum, 8A, 9A, and 9B). These conjoined short flexor tendons bifur-
where it splits into 4 tendons that help form the extensor expan- cate at the metatarsophalangeal joints, thus permitting the long
sions of digits II through V, inserting on the distal phalanges flexor tendons to pass through. After bifurcation, these tendons
(Figs. 8B and 11). M. extensor digitorum longus dorsiflexes the insert together on the middle phalanges of digits III through
ankle joint and extends the metatarsophalangeal and the proxi- V (Fig. 8A). In the fresh frozen left limb, the origin and inser-
mal and distal interphalangeal joints of digits II through V. tion sites of mm. interflexores were damaged during dissection
M. extensor digitorum brevis.—M. extensor digitorum bre- and could not be confirmed. M. interflexores flex the metatar-
vis originates from the dorsal calcaneus (Fig. 8B) and has 4 sophalangeal and proximal interphalangeal joints of digits III
or 5 muscle bellies that give rise to corresponding tendons at through V.
the midshafts of the metatarsals of digits I through IV. These M. quadratus plantae.—M. quadratus plantae originates
tendons join the tendons of mm. extensor digiti I longus and from the plantarlateral aspect of the calcaneus in a fossa created
extensor digitorum longus to help form the extensor expan- by the peroneal tubercle (Fig. 8A). The muscle typically lies
sions of digits I through IV (Figs. 8B and 11). In the preserved deep and distal to the origin of m. abductor digiti V (Figs. 8A
limbs, single muscle bellies served digit I and digit II, respec- and 9B). However, in the preserved right limb, the origin was
tively. In the preserved left limb, digit III had 2 muscle bellies: level with the origin of m. abductor digit V. M. quadratus plan-
a smaller medial belly and a larger lateral belly. However, in tae gives rise to a tendon that travels medially and passes over
the preserved right limb, there was only one muscle belly for the tendon of m. flexor digitorum lateralis before inserting onto
digit III. The muscle belly for digit IV gave rise to a tendon just the tendinous plate of mm. flexor digitorum lateralis and flexor
proximal to the base of the metatarsals, but the muscle belly digitorum medialis (Fig. 9B). M. quadratus plantae assists
continued on the plantar aspect of the tendon until halfway mm. flexor digitorum lateralis and flexor digitorum medialis
down the distal phalanx. In both fresh frozen limbs, there were in flexing the metatarsophalangeal, proximal, and distal inter-
4 muscle bellies. Digit II had 2 muscle bellies, with a larger phalangeal joints of digits I through V. At least in humans,
medial belly and a smaller lateral belly. The muscle bellies for m. quadratus plantae is the primary operator of the long flexor
digits III and IV were fused but gave rise to distinct tendons. tendons during voluntary flexion of the digits (Sooriakumaran
M. extensor digitorum brevis dorsiflexes the ankle joint and and Sivananthan 2005).
extends the metatarsophalangeal and interphalangeal joints of Mm. lumbricales.—Mm. lumbricales consist of 4 muscle bel-
digits I through IV. lies that originate from the common tendon plate of mm. flexor
M. abductor digiti V.—M. abductor digiti V originates via a digitorum lateralis and flexor digitorum medialis (Fig. 9C).
stout tendon, partially deep to m. flexor digitorum brevis and However, in the fresh frozen left limb, the muscle belly travel-
superficial to m. quadratus plantae, on the plantarmedial surface ing to digit II could not be confirmed due to damage during

by guest
on 28 August 2018
dissection. The muscle bellies are located between the long belly of m. flexor brevis profundus digiti II on the cartilaginous
flexor tendons and insert on the medial aspect of the extensor condensation in the plantar ligaments at the base of the 3rd
expansions for digits II through V (Figs. 8B, 9B, and 9C). Mm. metatarsal (Fig. 8A). The muscle inserts on the joint capsule of
lumbricales flex the metatarsophalangeal joints and extend the the metatarsophalangeal joint, including the medial sesamoid
proximal and distal interphalangeal joints of digits II through V. for digit III. The lateral belly originates from a cartilaginous
Mm. adductores digitorum.—Mm. adductores digitorum condensation of the plantar ligaments on the distal aspect of
consist of 3 muscle bellies that travel to digits I, II, and V the bases of the 3rd and 4th metatarsals (Fig. 8A). The lateral
(Fig. 9D). However, in the fresh frozen left limb, there was belly splits into 2 heads distally to insert on both the medial and
no muscle belly or tendon traveling to digit I. The muscle bel- lateral joint capsules of the metatarsophalangeal joint, includ-
lies are located superficial to mm. flexores breves profundi ing the sesamoids of digit III (Fig. 8A). The medial and lateral
(Fig. 9D). The 3 muscle bellies are fused at their origin from heads also give off thin tendons that contribute to the extensor
the plantar calcaneonavicular ligament. The muscle bellies for expansion of digit III on the dorsal pes (Fig. 8B). In the pre-
digits I and II insert on the lateral aspects of the proximal pha- served right limb, the medial belly originated from the carti-
langes of the respective digit (Fig. 8A). The muscle belly for laginous condensation at the base of the 3rd metatarsal (sharing
digit V gives rise to a large, flat tendon that dives deep to mm. an origin with the lateral belly of digit II), and the lateral belly
flexores breves profundi to insert along the medial aspect of the originated from the condensation at the base of only the 4th
distal metatarsal of digit V and the proximal phalanx (Fig. 8A). metatarsal. In the same limb, the medial head gave off a slip
M. adductores digitorum adduct and weakly flex the metatarso- of muscle that inserted with the lateral head on the lateral joint
phalangeal joints of digits I, II, and V. capsule. In the fresh frozen right limb, the medial and lateral
Mm. flexores breves profundi.—Mm. flexor breves profundi bellies were fused, and the medial belly split distally to insert
travel to digits I through V, with medial and lateral bellies for on the medial and lateral joint capsules at the metatarsopha-
each digit (Fig. 9D). The bellies of digits II through V give off langeal joints. The lateral belly then sent a slip of muscle that
small tendinous slips that insert on the extensor expansions of distally fused with the medial belly of digit IV.
digits II through V (Fig. 8B). Mm. flexores breves profundi flex Mm. flexores breves profundi digiti IV. The medial belly
the metatarsophalangeal joints, as well as abduct or adduct the originates from a cartilaginous condensation of the plantar liga-
metatarsophalangeal joints, according to their respective loca- ments at the base of the 4th metatarsal (Fig. 8A), the long plan-
tions on the pes. Additionally, they extend the proximal and tar ligament, and the plantar cuneometatarsal ligament. The
distal interphalangeal joints of digits II through V. medial belly splits into 2 heads distally to insert on both the me-
Mm. flexores breves profundi digiti I. The medial and lat- dial and lateral joint capsules of the metatarsophalangeal joint,
eral bellies originate via a thin tendon from the plantar calca- including the sesamoids of digit IV (Fig. 8A). The lateral belly
neonavicular ligament. The medial belly inserts on the plantar originates from the sesamoid on the base of the 5th metatarsal
aspect of the head of the 1st metatarsal and the joint capsule of (Fig. 8A), the plantar ligament, and the plantar cuneometatarsal
the metatarsophalangeal joint of digit I, including the medial ligament. This belly inserts on the joint capsule of the metatar-
sesamoid (Fig. 8A). The lateral belly inserts on the joint cap- sophalangeal joint, including the lateral sesamoid of digit IV.
sule of the metatarsophalangeal joint of digit I, including the The medial and lateral heads also give off thin tendons that con-
lateral sesamoid (Fig. 8A). tribute to the extensor expansion of digit IV on the dorsal pes
Mm. flexores breves profundi digiti II. The medial belly (Fig. 8B). In the preserved left limb, the lateral belly split into 2
originates from a cartilaginous condensation of the plantar liga- heads distally to insert on the medial and lateral joint capsules
ments at the lateral aspect of the base of the 1st metatarsal and and sesamoids of the metatarsophalangeal joint.
the medial aspect of the base of the 2nd metatarsal (Fig. 8A). Mm. flexores breves profundi digiti V. The medial belly
The medial belly inserts on the joint capsule of the metatarso- originates from the medial aspect of the sesamoid at the base of
phalangeal joint, including the medial aspect of the head of the the 5th metatarsal (Fig. 8A). It inserts on the joint capsule of the
2nd metatarsal and the medial sesamoid (Fig. 8A). The medial metatarsophalangeal joint, including the medial aspect of the
belly also gives off a slip of muscle to insert with the lateral belly medial sesamoid of digit V; this insertion is not shown on the
(Fig. 8A). The lateral belly originates from a cartilaginous con- muscle map because the proximal phalanx of digit V was tilted
densation of the plantar ligaments at the base of the 3rd metatar- medially on the articulated pes. The medial belly also gives off
sal and inserts on the joint capsule of the metatarsophalangeal a slip of muscle that inserts with the lateral belly (Fig. 8A). The
joint, including the head of the 2nd metatarsal and the lateral lateral belly originates from the lateral aspect of the sesamoid
sesamoid (Fig. 8A). Both heads also give off thin tendons that at the base of the 5th metatarsal and inserts on the joint capsule
contribute to the extensor expansion of digit II on the dorsal pes of the metatarsophalangeal joint, including the lateral sesa-
(Fig. 8B). In the preserved right limb, the medial head did not moid of digit V (Fig. 8A). The medial and lateral bellies also
originate from a cartilaginous condensation associated with the have a tendinous insertion on the extensor expansion of digit V
1st metatarsal, and the lateral belly originated from the base of (Fig. 8B). In the preserved right limb, the medial belly origi-
the 2nd metatarsal and the plantar calcaneonavicular ligament. nated from the long plantar ligament, as well as a sesamoid at
Mm. flexores breves profundi digiti III. The medial belly the base of the 5th metatarsal, while the lateral belly originated
originates slightly distal and lateral to the origin of the lateral solely on the long plantar ligament. In the fresh frozen limbs,

by guest
on 28 August 2018
the medial and lateral heads inserted solely on the medial and only 3 procyonid genera (Bassariscus—this study; Potos—
lateral joint capsules. Julitz 1909; Procyon—Gowell 1897). The myology of Nasua
M. flexor hallucis profundus.—M. flexor hallucis profun- nasua is extensively illustrated by Cuvier and Laurillard (1849),
dus originates from the navicular and lies deep to the ligament but a thorough description is lacking for this genus. Muscle
spanning the navicular and medial cuneiform (Fig. 8A). The maps are only available for Bassariscus and, in part, Procyon
muscle inserts on the plantar aspect of the base of the 1st meta- and muscle weights only for Bassariscus itself, which makes it
tarsal (Fig. 8A). M. flexor hallucis profundus weakly flexes the impossible to compare proportions across taxa. It is not surpris-
tarsometatarsal joint of digit I. ing that many of the features that show clear variation involve
Intraspecific variation in Bassariscus astutus.—Muscle vertebral attachments, as, due to the segmental nature of the
weights varied between the preserved female and fresh frozen vertebral column, differences in vertebral attachments can be
male, with typically lower weights in the female (Table 2). assessed from the literature with minimal ambiguity.
There was an ample amount of intraspecific variation in the Proximally, m. gluteus superficialis takes partial origin from
ringtail hind limbs regarding muscle fusion. The sample size the transverse processes of the caudal vertebrae in all pro-
is small, so it is unclear whether this is due to sexual dimor- cyonids. Most taxa resemble B. astutus in having an origin
phism or whether there is general variation in fusion within the restricted to Ca1 (Mackintosh 1875–1877; Allen 1882; Feeney
population. For example, m. gluteus superficialis was fused 1999), but in P. flavus, its origin extends to Ca3 (Julitz 1909) or
with m. semimembranosus in a preserved limb, m. femorococ- Ca4 (Beswick-Perrin 1871).
cygeus in a fresh frozen limb, and m. semitendinosus in a 2nd There is considerable variation in the presence and attach-
fresh frozen limb. Additional examples include fusion between ments of m. femorococcygeus. In addition to B. astutus, this
mm. femorococcygeus and biceps femoris in a preserved limb, muscle is present in Bassaricyon alleni, Nasua narica, and
as well as m. gracilis with mm. sartorius and semitendinosus P. flavus, although it is variously misidentified as m. agitator
in a fresh frozen limb. Another intraspecific difference relates caudae (Mackintosh 1875–1877), a head of m. biceps femo-
to the varying number of muscle bellies. Examples include ris (Beddard 1900), m. caudofemoralis (Julitz 1909), and a
m. piriformis, which possessed 2 bellies in a fresh frozen limb; portion of the caudal head of m. semitendinosus (Beswick-
m. gemellus inferioris, which contained 2 bellies in a preserved Perrin 1871). However, m. femorococcygeus is either lost
limb; and m. quadratus femoris, which had 2 bellies in 3 limbs. or inseparably fused with adjacent m. gluteus superficialis
The most variable features of the ringtail hind limbs occurred in Procyon spp. (Allen 1882; Windle 1888; Gowell 1897;
in the deep caudal leg. Differences between the 4 limbs were Feeney 1999). When the muscle is present, its origin is
observed in the origins of mm. flexor digitorum medialis and variable. N. narica has an origin from Ca1 (Mackintosh
fibularis longus, fusion between mm. tibialis caudalis and flexor 1875–1877), similar to the origin from Ca1–2 in B. astu-
digitorum lateralis, fusion at the origin of m. extensor digitorum tus. P. flavus has a more distal origin from Ca3 (Julitz 1909)
lateralis, and the variable number of muscle bellies of m. exten- or Ca3–4 (Beswick-Perrin 1871). The femoral insertion is
sor digitorum brevis. The majority of the variation occurred at variable, ranging from the proximal third in B. astutus and
the site of origin, while the insertions remained fairly constant. B. alleni (Beddard 1900) to the distal two-thirds in Nasua
Analysis of a greater number of specimens would help deter- spp. (Mackintosh 1875–1877), to a very distal insertion just
mine if this variability is completely representative of this spe- above the lateral femoral condyle in P. flavus (Beswick-
cies or if there is additional variability. Perrin 1871; Julitz 1909).
Variation within Procyonidae.—While some variation in The vertebral level from which m. abductor cruris caudalis
procyonid hind limb myology is apparent, it is relatively lim- takes origin is only specified in a few taxa. In both Nasua spp.
ited (Table 3). Additional myological differences may exist but (Mackintosh 1875–1877: m. biceps accessorius) and B. astu-
these cannot be detected due to limitations of the comparative tus, this occurs at the level of Ca1. In P. flavus, in contrast,
literature. Detailed, well-illustrated accounts are available for the muscle takes origin from the level of Ca2 (Beswick-Perrin
Table 3.—Distribution of hind limb features in procyonids and related taxa. Data are derived from the references listed in Table 1. A question
mark “?” indicates that the relevant morphology is not documented in the literature on a taxon. A dash “-” indicates that the character is inappli-
cable for that taxon due to absence of the structure whose variation is being described.
Character Galictis Ailurus Potos Bassaricyon Nasua Procyon Bassariscus

M. gluteus superficialis caudal origin Ca1–2 Ca1 Ca3 or Ca4 ? Ca1 Ca1 Ca1
M. femorococcygeus Absent Present Present Present Present Absent Present
M. femorococcygeus origin - Ca1–3 Ca3–4 ? Ca1 - Ca1–2
M. femorococcygeus insertion - Femoral midshaft Distal femur Proximal femur Femoral - Proximal femur
midshaft
M. abductor cruris caudalis origin Ca1 Ca2 Ca2 or Ca3 ? Ca1 ? Ca1
M. semitendinosus caudal head Present Absent Present Absent Present Present Present
M. semitendinosus caudal head origin Ca1–3 - Ca3–4 - Ca1 Ca1–2 Ca2
Fusion of m. presemimebranosus with Fused Separate or fused Separate ? Fused Fused Fused
m. semimembranosus

by guest
on 28 August 2018
1871: 2nd head of m. biceps femoris) or Ca3 (Julitz 1909: 2nd at the tibiotarsal joint and inversion at the subastragalar and
head of m. biceps femoris). transverse tarsal joints (Jenkins and McClearn 1984). Trapp’s
There is possible variation in the presence of a caudal head (1972) claim that rotation at the hip contributes to reversal of
of m. semitendinosus. This head is present in most procyonids the foot in the ringtail has not been substantiated, although in
(Cuvier and Laurillard 1849; Beswick-Perrin 1871; Mackintosh vivo radiographic data are unavailable for B. astutus. Potential
1875–1877; Windle 1888; Gowell 1897; Julitz 1909; this study) movements of the hip that would achieve reversal of the pes
but is stated to be absent in B. alleni (Beddard 1900). Among would include extension, abduction, and external rotation.
taxa that have a caudal head, there is variation in the level of There is modest evidence of hypertrophy of the hip extensors
origin. Nasua spp. has a proximal origin from Ca1 (Mackintosh and abductors in B. astutus, as compared with the more terres-
1875–1877), whereas B. astutus has a slightly more distal origin trial G. cuja (data from Ercoli et al. 2013). Extensors constitute
from Ca2. The origin in P. lotor spans Ca1–2 (Gowell 1897). In between 52% and 59% of the total mass of the musculature
P. flavus, the origin is more distal, from Ca3–4 (Beswick-Perrin acting on the hip in B. astutus, compared to 52% in G. cuja.
1871; Julitz 1909). For abductors, the percentages are 15–17% in B. astutus ver-
A femoral head of m. semimembranosus is present in B. astu- sus 14% in G. cuja. The difference in the external rotators is
tus, Nasua spp., and Procyon spp. (Cuvier and Laurillard 1849; larger. External rotators constitute between 4% and 6% of total
Mackintosh 1875–1877; Windle 1888; Gowell 1897; this study) muscle mass in B. astutus compared with 3% in G. cuja. The
but is not noted in P. flavus (Beswick-Perrin 1871; Windle and relative hypertrophy of the external rotators in B. astutus sug-
Parsons 1898; Julitz 1909). Beddard’s (1900) description of gests that if any movement at the hip is associated with reversal
mm. adductores matches the morphology of m. semimem- of the hind foot, it is likely to be rotation rather than exten-
branosus in the first 3 taxa, suggesting that B. alleni also has sion or abduction. The muscle responsible for the differential
a femoral head of the muscle. However, this would leave the development of the external rotators can be readily identified.
identity of his m. semimembranosus (described only as lacking M. quadratus femoris has approximately twice the relative
notable features) unclear. In taxa in which it is clearly present, mass in Bassariscus as compared with Galictis. It has a promi-
the femoral head likely corresponds to m. presemimembrano- nent insertion on the caudal surface of the lesser trochanter and
sus of Windle and Parsons (1898). Carlsson (1920) noted that intertrochanteric region, contrasting with the smaller insertion
a muscle described and illustrated in P. flavus as m. adductor in G. cuja.
magnus by Julitz (1909) likely corresponds, at least in part, The strong development of m. quadratus femoris in B. astu-
to m. presemimembranosus. Beswick-Perrin (1871) describes tus is potentially consistent with a link between movements at
m. adductor magnus as being pierced by the femoral artery in the hip and reversal of the hind foot. However, a large m. qua-
his individual of P. flavus. Given that Windle and Parsons (1898) dratus femoris is widely distributed among arboreal mammals
indicate that the femoral artery separates m. adductor magnus in general. Among other taxa, it characterizes dermopterans
from m. presemimembranosus, this suggests that m. presemi- and strepsirhine primates and, based on the distribution of an
membranosus was also fused with m. adductor magnus in this enlarged surface on the lesser trochanter, most early euarchon-
individual. If these inferences are correct, P. flavus does retain tans (Beard 1989). Within Carnivora, more arboreal taxa such
m. presemimembranosus, but it is not closely associated with as A. fulgens, Ailuropoda melanoleuca, and Genetta spp. (Davis
m. semimembranosus. 1964; Taylor 1976; Fisher et al. 2008) have relatively larger sur-
Finally, the presence of an undivided m. sartorius in B. astu- faces on the caudal femur for attachment of m. quadratus femo-
tus agrees with the morphology present in B. alleni, P. flavus, ris than do more terrestrial taxa including herpestids, felids, and
Procyon spp., and the specimen of N. nasua illustrated by canids (Langguth 1969; Taylor 1976; Evans 1993; Reighard
Cuvier and Laurillard (1849). This conflicts with Davis (1964, and Jennings 1901; Carlon and Hubbard 2012). Beard (1989)
table 16) who indicated that Bassariscus has 2 heads of sarto- suggested that hypertrophy of m. quadratus femoris reflects the
rius, a claim not substantiated by descriptions or illustrations. fact that arboreal mammals habitually use externally rotated
Mackintosh (1875–1877) appears to describe a double-headed postures when grasping branches. Sargis (2002) supported this
sartorius in Nasua spp. However, this author merely notes the hypothesis, noting that the femoral surface for attachment of
presence of an insertion onto both the patella and tibia and never m. quadratus femoris is much larger in the arboreal scandentian
clearly indicates the presence of 2 distinct bellies. This descrip- Ptilocercus than in terrestrial Tupaia. At the same time, these
tion is consistent with the presence of a continuous sheet-like functions are not mutually exclusive, as many taxa capable of
attachment spanning both bones, as occurs in B. astutus and hind foot reversal also habitually grasp branches.
all other adequately illustrated procyonids, including N. nasua Hind foot reversal and hind limb suspension.—More distally,
(Cuvier and Laurillard 1849). few differences exist between the plantarflexors or invertors of
the feet of B. astutus and P. flavus, which is also capable of full
hind foot reversal. The same is true for species of Nasua and
Discussion Procyon which, unlike B. astutus, are only capable of partial
Adaptations at the hip to hind foot reversal.—As is dis- reversal. Comparisons with Bassaricyon are limited due to the
cussed in the introduction, in vivo radiographic data indicate very brief account of this taxon, but no differences are evident.
that hind foot reversal involves a combination of plantarflexion More surprisingly, these muscles do not differ meaningfully

by guest
on 28 August 2018
from other musteloid taxa that exhibit little or no capacity for including the European otter (Lutra lutra: 14%), American
hind foot reversal (e.g., Galictis). This includes m. tibialis cra- black bear (Ursus americanus: 18%), and cheetah (Acinonyx
nialis, which Pierotti (in Jenkins and McClearn 1984) regards jubatus: 18%—Haughton 1867b, 1867c; Hudson et al. 2011).
as the prime mover in the action of hind foot reversal, at least in The long digital flexors comprise a greater percentage of mus-
sciurid rodents. The lack of meaningful myological distinctions cle mass in B. astutus than in any comparative taxon. Although
between Bassariscus and taxa that lack the capacity for hind data from additional species would be preferable, this suggests
foot reversal indicates that the ability to reverse the hind foot that the long digital flexors are relatively more developed in
does not have significant muscular correlates. Bassariscus than in a diversity of carnivorans that do not use
The apparent lack of myological features associated with hind limb suspension. This is consistent with the hypothesis that
hind foot reversal suggests that osteology, particularly the these muscles are used to maintain digital flexion for extended
shape, position, and orientation of joint surfaces, is likely to periods of time, an action that is not necessary in other taxa.
be the primary determinant of whether and to what degree the Although their very small size makes quantitative compari-
hind foot can be reversed. When joint surfaces permit reversal, sons difficult, the development of some intrinsic muscles of the
m. tibialis cranialis and other involved muscles can achieve hind plantar pes also supports the hypothesis that the musculature of
foot reversal without significant change to their morphology or taxa capable of hind limb reversal and suspension is adapted for
attachments. This is consistent with studies of the osteology of maintenance of digital flexion. Relative to the terrestrial G. cuja,
hind foot reversal (Jenkins and McClearn 1984; Meldrum et al. mm. interflexores are much larger in B. astutus. A. fulgens,
1997) which have primarily identified features associated with which also appears to use reversed, suspended postures, resem-
joint surfaces, rather than muscle attachments, as characteriz- bles B. astutus in this regard (Fisher et al. 2008). Mm. inter-
ing those taxa capable of reversal. flexores aids in flexion the digits at the metatarsophalangeal and
In fact, direct myological adaptations to hind foot reversal proximal interphalangeal joints. The presence of larger, more
may not be selectively beneficial. Mammals that are capable powerful bellies would be beneficial to an animal that habitually
of reversal generally make use of this capability to position the needs to maintain digital flexion against resistance.
foot for either headfirst descent of a vertical substrate or hind Another intrinsic pedal muscle that may show specializa-
limb suspension of the body from a horizontal substrate. In tions related to hind foot reversal and hind limb suspension is
these situations, the hind foot is unlikely to be reversed against m. quadratus plantae. As noted above, m. quadratus plantae is
resistance nor is speed likely to be critical. Once the foot is active during voluntary flexion of the digits in humans where,
reversed, gravity will maintain the position, eliminating the by virtue of its insertion on the long flexor tendons, it can act as
need for the muscles involved in reversal to remain active. All a flexor of the metatarsophalangeal and interphalangeal joints
of this argues that, so long as one or more muscles is positioned (Sooriakumaran and Sivananthan 2005). M. quadratus plantae
such that it can achieve reversal, no other specialization of the takes origin from the plantar surface of the peroneal tubercle of
musculature is likely to be necessary. the calcaneum. It has been noted that the peroneal tubercle tends
In contrast, adaptations to hind limb suspension are more to be larger and more proximally positioned in arboreal mam-
likely to result in myological correlates. In most situations in mals (e.g., Gunnell 1989; Rose and Chinnery 2004). When an
which the hind foot is reversed, most or all of the body weight explanation for this relationship has been proposed, it has typi-
is being supported by the hind limb against the pull of gravity. cally focused on the effect on m. fibularis longus, which passes
Unlike the action of hind foot reversal, keeping the body in a directly caudal to the tubercle (Gunnell 1989). However, a
suspended posture is likely to require sustained muscular activ- larger, more proximally positioned tubercle would also increase
ity. In particular, the capacity to maintain a grip on a substrate the area of origin and leverage of m. quadratus plantae, increas-
with the hind foot for extended periods is likely to be critical ing its utility as a digital flexor during hind limb suspension. The
in taxa that make extensive use of hind limb suspension. This broad overlap between arboreal mammals and taxa capable of at
suggests that the digital flexors should be strongly developed in least some degree of hind foot reversal suggests that adaptation
taxa that habitually engage in hind limb suspension (and hind to hind limb suspension may account for the apparent associa-
foot reversal). tion between a proximal peroneal tubercle and arboreality.
One feature of the digital flexors of B. astutus may represent M. flexor hallucis profundus.—Fisher et al. (2008) identified a
an adaptation to habitual use of hind limb suspension, particu- new deep flexor of the 1st digit, m. flexor hallucis profundus, in
larly when climbing headfirst. The ringtail shows hypertrophy their account of the myology of the red panda, A. fulgens. They
of mm. flexores digitorum lateralis and medialis, the most pow- also reported that a dissection of the sun bear, Helarctos malaya-
erful and best-leveraged digital flexors. In comparison with nus, revealed this muscle but were unable to confidently identify
Galictis, for which detailed muscle weights have also been any records of m. flexor hallucis profundus in the literature. The
published (Ercoli et al. 2013), these muscles and their tendons identification of m. flexor hallucis profundus in B. astutus consti-
comprise 21–23% of the total muscle mass of muscles that tutes the 3rd record of this muscle in carnivorans, and the first in
are primarily or completely distal to the knee versus 13% in procyonids. The morphology of the muscle matches that of the sun
Galictis. When only muscles originating proximal to the ankle bear (H. malayanus) but differs somewhat from that of Ailurus in
are considered, these values are 22–24% and 14%, respectively. having an insertion near the base of the 1st metatarsal, rather than
The latter values can be compared to a few other carnivorans, more distally, near the metatarsophalangeal joint of the 1st digit.

by guest
on 28 August 2018
Further examination of the procyonid literature reveals 2 Because these aspects of procyonid morphology are develop-
additional records of m. flexor hallucis profundus and sug- mentally and functionally integrated, it is not unexpected that
gests that the presence of this muscle may be characteristic they might respond in the same way to common selective pres-
of Procyonidae. Beswick-Perrin (1871) described a muscle, sures in different taxa. As a more distant and functionally dis-
m. obliquus tarsi in the kinkajou (P. flavus), that is identi- tinct aspect of procyonid anatomy, hind limb myology might
cal in its position and attachments to the ringtail m. flexor show a different pattern of variation than craniodental osteol-
hallucis profundus. Similarly, a muscle illustrated as ogy, potentially more in agreement with molecular data.
m. scaphocuneiformis in P. lotor by Jenkins and McClearn The available data indicate that the morphology of Potos
(1984) closely matches the morphology of m. flexor hal- (Table 3) contrasts with that of other procyonids (primarily
lucis profundus. Both taxa exhibit the more proximal inser- Bassariscus, Nasua, and Procyon). Mm. gluteus superficialis,
tion of B. astutus rather than the more distal insertion of femorococcygeus, and abductor cruris caudalis as well as the
Ailurus, consistent with this being the typical morphology caudal head of m. semitendinosus in the kinkajou are character-
of procyonids. ized by a more distal origin from the base of the tail, arising from
While the distribution of m. flexor hallucis profundus the 2nd through 4th caudal vertebrae in Potos (Beswick-Perrin
remains incompletely known, a tentative pattern is emerging. 1871; Julitz 1909), compared to more proximal levels in other
All 5 taxa in which the muscle has been recorded are scanso- procyonids (Nasua—Mackintosh 1875–1877; Procyon—Allen
rial or arboreal. No terrestrial taxon has been documented to 1882; Gowell 1897; Feeney 1999; Bassariscus—this study).
possess this muscle, including members of other arctoid fami- Additional features that differentiate Potos from other procyo-
lies. A possible explanation for this pattern is that the presence nids are a more distal insertion of m. femorococcygeus and sepa-
of m. flexor hallucis profundus increases the independence ration of m. presemimembranosus from m. semimembranosus
of the 1st digit relative to the lateral digits, providing some (Potos—Beswick-Perrin 1871; Julitz 1909; Nasua—Mackintosh
capacity to grip supports. If this is the case, it predicts that 1875–1877; Procyon—Allen 1882; Gowell 1897; Feeney 1999;
further study of other arboreal carnivorans will reveal further Bassaricyon—Beddard 1900; Bassariscus—this study). Fewer
records of this muscle. Members of Miacidae, the paraphy- features differentiate other procyonid taxa. Most notably, m. fem-
letic stem group of Carnivora (Wesley-Hunt and Flynn 2005; orococcygeus is absent or indistinguishably fused with m. glu-
Spaulding and Flynn 2012), show substantial arboreal adapta- teus superficialis in Procyon (Allen 1882; Windle 1888; Gowell
tions (Heinrich and Rose 1995, 1997; Spaulding and Flynn 1897; Feeney 1999), a morphology that also occurs in some
2009; Spaulding et al. 2010), implying an arboreal ancestry other musteloids (e.g., Galictis—Ercoli et al. 2013). A caudal
for Carnivora (Rose 2006). This suggests that it is plausible head of m. semitendinosus is described as absent in Bassaricyon,
that the presence of m. flexor hallucis profundus may be prim- although confirmation of the latter would be preferable given the
itive for the order. brevity and lack of illustrations in Beddard’s (1900) account.
Phylogenetic implications.—Molecular phylogenetic studies This would be unique among procyonids, although other mus-
(Fulton and Strobeck 2006; Koepfli et al. 2007; Wolsan and teloids (e.g., Ailurus—Fisher et al. 2008) also lack this head.
Sato 2010) recover a well-resolved, strongly supported topol- The dichotomy between Potos and other procyonids is con-
ogy for Procyonidae at the generic level. Potos is identified as sistent with molecular support for a basal position of the kin-
the most basal extant procyonid. Remaining genera are divided kajou with respect to other procyonids. However, comparisons
into 2 clades, one containing Bassaricyon, Nasua, and Nasuella with other musteloids (e.g., Ailurus, Galictis) indicate that, for
(nested within a paraphyletic Nasua) and the other containing most features in which there is a dichotomy between Potos and
Bassariscus and Procyon. This topology is in broad conflict other procyonids, the morphology of Potos is likely derived.
with results of rigorous morphological analyses of procyo- For these features other procyonids more closely approximate
nid interrelationships (Decker and Wozencraft 1991; Baskin the plesiomorphic morphology, which would render these char-
2004; Ahrens 2012). The latter have consistently supported acteristics parsimony uninformative. Additionally, there are no
a sister taxon relationship between Procyon and Nasua plus features to support a special relationship between Bassaricyon
Nasuella as well as a sister taxon relationship between Potos and Potos, although few of the distinctive features in Potos can
and Bassaricyon. The position of Bassariscus in morphologi- be assessed from Beddard’s (1900) account.
cal studies has been more variable, but the genus has emerged Finally, it is also noteworthy that no hind limb myological
as either the basally diverging living procyonid (Ahrens 2012) feature supports a placement of Bassariscus basal to all other
or as the sister taxon to the Procyon/Nasua clade (Decker and procyonids. For all variable features, the morphology present in
Wozencraft 1991; Baskin 2004). Bassariscus is shared by at least one other member of the fam-
The lack of agreement between molecular and morphologi- ily such that there is no myological feature that plausibly unites
cal studies has been ascribed by Koepfli et al. (2007) to a com- all other procyonids to the exclusion of Bassariscus.
bination of morphological convergence between ecologically
similar taxa and a failure to account for unrecognized non-
independence of characters that share common genetic and/ Acknowledgments
or developmental underpinnings. All morphological analyses The authors thank W. Stanley and J. D. Phelps, Jr. from the
published to date have relied heavily on craniodental osteology. Field Museum for the loan of FMNH 135423, FMNH 53761,

by guest
on 28 August 2018
and FMNH 53869, K. Reed from Arizona State University for Cunningham, D. J. 1879. The intrinsic muscles of the mammalian
the loan of the Phoenix Zoo specimen, and C. Johnston from foot. Journal of Anatomy and Physiology 13:1–16.
the Natural History Collections at Arizona State University for Cuvier, G., and M. Laurillard. 1849. Anatomie comparée. Recueil
the loan of a suitable storage container. The authors are also de planches de myologie dessinées par Georges Cuvier ou exécu-
tées sous ses yeux par M. Laurillard. Imprimerie d’E. Duverger,
grateful for K. Whitfield’s assistance in translating Carlsson
Paris.
(1925). D. Kang produced Figs. 1–11, and H. Sydiongco edited
Davis, D. D. 1949. The shoulder architecture of bears and other carni-
Figs. 2, 4, 5, 7, 10, and 11. The authors also thank the review- vores. Fieldiana: Zoology 31:285–305.
ers, including M. D. Ercoli and 1 anonymous colleague, for Davis, D. D. 1964. The giant panda: a morphological study of evolu-
providing comments that greatly improved an earlier version tionary mechanisms. Fieldiana: Zoology, Memoirs 3:1–339.
of this manuscript. ML thanks the members of her honors the- Decker, D. M., and W. C. Wozencraft. 1991. Phylogenetic
sis committee, Drs. R. Fisher, R. Hinrichs, and K. Kusumi for analysis of recent procyonid genera. Journal of Mammalogy
their feedback and support. Funding for this project was pro- 72:42–55.
vided to ML from Barrett, The Honors College, Arizona State Egi, N. 2001. Body mass estimates in extinct mammals from limb
University. bone dimensions: the case of North American hyaenodontids.
Palaeontology 44:497–528.
Ercoli, M. D., S. Echarri, F. Busker, A. Álvarez, M. M. Morales,
Literature Cited and G. F. Turazzini. 2013. The functional and phylogenetic impli-
Ahrens, H. E. 2012. Craniodental characters and the relationships cations of the myology of the lumbar region, tail, and hind limbs of
of Procyonidae (Mammalia: Carnivora). Zoological Journal of the the lesser grison (Galictis cuja). Journal of Mammalian Evolution
Linnean Society 164:669–713. 20:309–336.
Alexander, L. F., B. J. Verts, and T. P. Farrell. 1994. Diet of ring- Evans, H. E. 1993. Miller’s anatomy of the dog. Saunders, an imprint
tails (Bassariscus astutus) in Oregon. Northwestern Naturalist of Elsevier, Philadelphia, Pennsylvania.
75:97–101. Feeney, S. 1999. Comparative osteology, myology, and locomotor
Alix, E. 1876. Memoire sur la myologie du putois (Putorius commu- specializations of the fore and hind limbs of the North American
nis, Cuv.). Journal de Zoologie 5:152–188. foxes Vulpes vulpes and Urocyon cinereoargenteus. Ph.D. disserta-
Allen, H. 1882. The muscles of the limbs of the raccoon (Procyon tion, University of Massachusetts, Amherst.
lotor). Proceedings of the Academy of Natural Sciences of Fisher, E. M. 1942. The osteology and myology of the California
Philadelphia 34:115–144. River Otter. Stanford University Press, Stanford University,
Barone, R., and H. Deutsch. 1953. Myologie des membres du furet Stanford, California.
(Putorius furo). Bulletin de la Société des Sciences Vétérinaires de Fisher, R. E., B. Adrian, C. Elrod, and M. Hicks. 2008. The phylog-
Lyon 55:445–454. eny of the red panda (Ailurus fulgens): evidence from the hindlimb.
Baskin, J. A. 2004. Bassariscus and Probassariscus (Mammalia, Journal of Anatomy 213:607–628.
Carnivora, Procyonidae) from the early Barstovian (middle Fleagle, J. G., and D. J. Meldrum. 1988. Locomotor behavior and
Miocene). Journal of Vertebrate Paleontology 24:709–720. skeletal morphology of two sympatric pitheciine monkeys, Pithecia
Beard, K. C. 1989. Postcranial anatomy, locomotor adaptations, and pithecia and Chiropotes satanas. American Journal of Primatology
paleoecology of Early Cenozoic Plesiadapidae, Paromomyidae, 16:227–249.
and Micromomyidae (Eutheria, Dermoptera). Ph.D. dissertation, Fulton, T. L., and C. Strobeck. 2006. Molecular phylogeny of
The Johns Hopkins University School of Medicine, Baltimore, the Arctoidea (Carnivora): effect of missing data on supertree
Maryland. and supermatrix analyses of multiple gene data sets. Molecular
B eddard , F. E. 1900. On the anatomy of Bassaricyon Phylogenetics and Evolution 41:165–181.
alleni. Proceedings of the Zoological Society of London Gingerich, P. D., and H. A. Deutsch. 1989. Systematics and evolu-
1900:661–675. tion of early Eocene Hyaenodontidae (Mammalia, Creodonta) in
Beddard, F. E. 1905. Some notes upon the anatomy of the ferret- the Clarks Fork Basin, Wyoming. Contributions from the Museum
badger. Proceedings of the Zoological Society of London 2:21–29. of Paleontology, The University of Michigan 27:327–391.
Beswick-Perrin, J. 1871. On the myology of the limbs of the Kinkajou Gowell, R. C. 1897. Myology of the hind limb of the raccoon. Kansas
(Cercoleptes caudivolvulus). Proceedings of the Zoological Society University Quarterly 6:121–126.
of London 1871:547–559. Grinnell, J., J. S. Dixon, and J. M. Linsdale. 1937. Fur-bearing
Bradley, W. G., and C. G. Hansen. 1965. Observations on the dis- mammals of California. University of California Press, Berkeley.
tribution of the ring-tailed cat in southern Nevada. Southwestern Gunnell, G. F. 1989. Evolutionary history of Microsyopoidea
Naturalist 10:310–311. (Mammalia, ?Primates) and the relationship between
Carlon, B., and C. Hubbard . 2012. Hip and thigh anatomy of Plesiadapiformes and Primates. University of Michigan Papers on
the clouded leopard (Neofelis nebulosa) with comparisons Paleontology 27:1–157.
to the domestic cat (Felis catus). The Anatomical Record Haines, R. W. 1958. Arboreal or terrestrial ancestry of placental
295:577–589. mammals. The Quarterly Review of Biology 33:1–23.
Carlsson, A. 1920. Über Arctictis binturong. Acta Zoologica Hall, E. R. 1926. The muscular anatomy of three mustelid mam-
1:337–380. mals, Mephitis, Spilogale, and Martes. University of California
Carlsson, A. 1925. Über Ailurus fulgens. Acta Zoologica 6:269–305. Publications in Zoology 30:7–38.
Cartmill, M. 1985. Climbing. Pp. 73–88 in Functional vertebrate Hall, E. R. 1927. The muscular anatomy of the American badger
morphology (M. Hildebrand, D. M. Bramble, K. F. Liem, and D. (Taxidea taxus). University of California Publications in Zoology
B. Wake, eds.). The Belknap Press, Cambridge, Massachusetts. 30:205–219.

by guest
on 28 August 2018
Harrison, R. L. 2012. Ringtail (Bassariscus astutus) ecology and foina). Proceedings of the Royal Irish Academy of Sciences, Series
behavior in central New Mexico, USA. Western North American 2 2:48–55.
Naturalist 72:495–506. MacLeod, N., and K. D. Rose. 1993. Inferring locomotor behavior in
Haughton, S. 1867a. On the muscular anatomy of the badger. Paleogene mammals via Eigenshape analysis. American Journal of
Proceedings of the Royal Irish Academy of Sciences, Series 2 Science 293-A:300–355.
ix:507–508. McClearn, D. 1985. Anatomy of raccoon (Procyon lotor) and Coati
Haughton, S. 1867b. On the muscles of the Virginian bear. Proceedings (Nasua narica and N. nasua) forearm and leg muscles: relations
of the Royal Irish Academy of Sciences, Series 2 ix:508–511. between fiber length, moment-arm length, and joint-angle excur-
Haughton, S. 1867c. On the muscular anatomy of the otter (Lutra sion. Journal of Morphology 183:87–115.
vulgaris). Proceedings of the Royal Irish Academy of Sciences, McClearn, D. 1992. Locomotion, posture, and feeding behav-
Series 2 ix:511–515. ior of kinkajous, coatis, and raccoons. Journal of Mammalogy
Heinrich, R. E., and K. D. Rose. 1995. Partial skeleton of the primi- 73:245–261.
tive carnivoran Miacis petilus from the early Eocene of Wyoming. Meldrum, D. J., M. Dagosto, and J. L. White. 1997. Hindlimb
Journal of Mammalogy 76:148–162. suspension and hind foot reversal in Varecia variegata and other
Heinrich, R. E., and K. D. Rose. 1997. Postcranial morphology and arboreal mammals. American Journal of Physical Anthropology
locomotor behaviour of two early Eocene miacoid carnivorans, 103:85–102.
Vulpavus and Didymictis. Palaeontology 40:279–305. Nowak, R. M. 1999. Walker’s mammals of the world. 6th ed. The
Heptner, V. G., and N. P. Naumov. 1967. Mammals of the Sovietic Johns Hopkins University Press, Baltimore, Maryland.
Union, Vol. II, Part 1. Vysshaya Shkola Publishers, Moscow. Poglayen-Neuwall, I., and I. Poglayen-Neuwall. 1965.
Howard, L. D. 1975. Muscular anatomy of the hind limb of the sea Gefangenschaftsbeobachtungen an makibären (Bassaricyon Allen
otter (Enhydra lutris). Proceedings of the California Academy of 1876). Zeitschrift für Säugertierkunde 30:321–366.
Sciences, Fourth Series 40:335–416. Poglayen-Neuwall, I., and D. E. Toweill. 1988. Bassariscus astutus.
Hudson, P. E., S. A. Corr, R. C. Payne-Davis, S. N. Clancy, E. Lane, Mammalian Species 327:1–8.
and A. M. Wilson. 2011. Functional anatomy of the cheetah Ray, L. J. 1949. The myology of the inferior extremity of the Malay
(Acinonyx jubatus) hindlimb. Journal of Anatomy 218:363–374. Bear, Ursus malayanus. Proceedings of the Zoological Society of
Jenkins, F. A., and D. W. Krause. 1983. Adaptations for climb- London 119:121–132.
ing in North American multituberculates (Mammalia). Science Reighard, J. E., and H. S. Jennings. 1901. Anatomy of the cat. 2nd
220:712–715. ed. H. Holt, New York.
Jenkins, F. A., and D. McClearn. 1984. Mechanisms of hind Rose, K. D. 2006. The beginning of the age of mammals. The Johns
foot reversal in climbing mammals. Journal of Morphology Hopkins University Press, Baltimore, Maryland.
182:197–219. Rose, K. D., and B. J. Chinnery. 2004. The postcranial skeleton of
Julitz, C. 1909. Osteologie und Myologie der Extremitäten und des early Eocene rodents. Bulletin of Carnegie Museum of Natural
Wickelschwanzes vom Wickelbären, Cercoleptes caudivolvulus, History 36:211–244.
mit besonder Berücksichtigung der Anpassungserscheinungen an Sargis, E. J. 2002. The postcranial morphology of Ptilocercus
das Baumleben. Archiv für Naturgeschichte Berlin 75:143–188. lowii (Scandentia, Tupaiidae): an analysis of primatomorphan
Kaufmann, J. H. 1962. Ecology and social behavior of the coati, and volitantian characters. Journal of Mammalian Evolution
Nasua narica, on Barro Colorado Island, Panama. University of 9:137–160.
California Publications in Zoology 60:95–222. Sarmiento, E. E. 1983. The significance of the heel process in anthro-
Kelley, E. A. 1888. Notes on the myology of Ursus maritimus. poids. International Journal of Primatology 4:127–152.
Proceedings of the Academy of Natural Sciences of Philadelphia Scherling, A. 1989. Zur Anatomie des Bewegungsapparates des
1888:141–154. Dachses (Meles meles L.). Ph.D. dissertation, Universität Gießen,
Koepfli, K.-P., et al. 2007. Phylogeny of the Procyonidae Gießen, Germany.
(Mammalia: Carnivora): molecules, morphology and the Great Shepherd, F. J. 1883. Short notes on the myology of the American
American Interchange. Molecular Phylogenetics and Evolution black bear (Ursus americanus). Journal of Anatomy and Physiology
43:1076–1095. 18:103–117.
Langguth, A. 1969. Die südamerikanischen Canidae unter beson- Sooriakumaran, P., and S. Sivananthan. 2005. Why does man have a
derer Berücksichtigung des Mähnenwolfes Chrysocyon brachy- quadratus plantae? A review of its comparative anatomy. Croatian
urus Illiger (Morphologische, systematische und phylogenetische Medical Journal 46:30–35.
Untersuchungen). Zeitschrift für wissenschaftliche Zoologie Spaulding, M., and J. J. Flynn. 2009. Anatomy of the postcranial
179:1–188. skeleton of “Miacis” uintensis (Mammalia: Carnivoramorpha).
Leyhausen, P. 1963. Über südamerikanische Pardelkatzen. Zeitschrift Journal of Vertebrate Paleontology 29:1212–1223.
für Tierpsychologie 20:627–640. Spaulding, M., and J. J. Flynn. 2012. Phylogeny of the
Macalister, A. 1873a. The muscular anatomy of the civet and tyra. Carnivoramorpha: the impact of postcranial characters. Journal of
Proceedings of the Royal Irish Academy of Sciences, Series 2 Systematic Palaeontology 10:653–677.
1:506–513. Spaulding, M., J. J. Flynn, and R. K. Stucky. 2010. A new basal
Macalister, A. 1873b. On the anatomy of Aonyx. Proceedings of the carnivoramorphan (Mammalia) from the ‘Bridger B’ (Black’s Fork
Royal Irish Academy of Sciences, Series 2 1:539–547. Member, Bridger Formation, Bridgerian NALMA, middle Eocene)
Macalister, A. 1875. On two new species of Pentastomida. of Wyoming, USA. Palaeontology 53:815–832.
Proceedings of the Royal Irish Academy of Sciences, Series 2 Taylor, W. P. 1954. Food habits and notes on life history of the ring-
2:62–66. tailed cat in Texas. Journal of Mammalogy 35:55–63.
Mackintosh, H. W. 1875–1877. Notes on the myology of the coati- Taylor, M. E. 1970. Locomotion in some East African viverrids.
mondi (Nasua narica and N. fusca) and common marten (Martes Journal of Mammalogy 51:42–51.

by guest
on 28 August 2018
Taylor, M. E. 1976. The functional anatomy of the hindlimb of some Wildscreen Arkive. 2015. Red panda (Ailurus fulgens) videos. http://
African Viverridae (Carnivora). Journal of Morphology 148:227–254. www.arkive.org/red- panda/ailurus-fulgens/videos.html. Accessed
Toweill, D. E., and J. G. Teer. 1977. Food habits of ringtails in 1 March 2015.
the Edwards Plateau region of Texas. Journal of Mammalogy Williams, R. C. 1955. The osteology and myology of the ranch mink
58:660–663. (Mustela vison). M.S. thesis, Cornell University, Ithaca, New York.
Trapp, G. R. 1972. Some anatomical and behavioral adapta- Windle, B. C. A. 1888. Notes on the limb myology of Procyon cancriv-
tions of ringtails, Bassariscus astutus. Journal of Mammalogy orus and of the Ursidae. Journal of Anatomy and Physiology 23:81–89.
53:549–557. Windle, B. C. A., and F. G. Parsons. 1898. The myology of the ter-
Trapp, G. R. 1978. Comparative behavioral ecology of the ringtail restrial carnivora. Part II. Proceedings of the Zoological Society of
and gray fox in southwestern Utah. Carnivore 1:3–32. London 1898:152–186.
Waibl, H., et al. 2005. Nomina anatomica veterinaria. 5th ed. International Wolsan, M., and J. J. Sato. 2010. Effects of data incompleteness on
Committee on Veterinary Gross Anatomical Nomenclature, World the relative performance of parsimony and Bayesian approaches
Association of Veterinary Anatomists, Hanover, Germany. in a supermatrix phylogenetic reconstruction of Mustelidae and
Wesley-Hunt, G. D., and J. J. Flynn. 2005. Phylogeny of the Procyonidae (Carnivora). Cladistics 26:168–194.
Carnivora: basal relationships among the carnivoramorphans, and
assessment of the position of ‘Miacoidea’ relative to Carnivora. Submitted 18 March 2015. Accepted 10 October 2015.
Journal of Systematic Palaeontology 3:1–28.
Associate Editor was John S. Scheibe.

by guest
on 28 August 2018

Hind Limb Myology of The Ringtail (Bassariscus Astutus) and The Myology of Hind Foot Reversal

Uploaded by

Copyright:

Available Formats

You might also like

Hind Limb Myology of The Ringtail (Bassariscus Astutus) and The Myology of Hind Foot Reversal

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Hind Limb Myology of The Ringtail (Bassariscus Astutus) and The Myology of Hind Foot Reversal

Uploaded by

Copyright:

Available Formats

Journal of Mammalogy, 97(1):211–233, 2016

Hind limb myology of the ringtail (Bassariscus astutus) and the

© 2015 American Society of Mammalogists, www.mammalogy.org

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Taxon Loc. Sources

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Tensor fascia latae

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Gastrocnemius, lateral head +

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Fig. 4.—Deep view of the gluteal region of Bassariscus astutus (left side).

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Fig. 5.—Medial view of the left thigh of Bassariscus astutus.

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

g Flexor digitorum lateralis +

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Fig. 10.—Craniolateral view of the left leg of Bassariscus astutus.

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Character Galictis Ailurus Potos Bassaricyon Nasua Procyon Bassariscus

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

Downloaded from https://academic.oup.com/jmammal/article-abstract/97/1/211/2459704

You might also like