American Society of Mammalogists

Phylogenetic Analysis of Recent Procyonid Genera

Author(s): Denise M. Decker and W. Chris Wozencraft
Source: Journal of Mammalogy, Vol. 72, No. 1 (Feb., 1991), pp. 42-55
Published by: American Society of Mammalogists
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 PHYLOGENETIC ANALYSISOF RECENT PROCYONID GENERA

DENISE M. DECKER AND W. CHRIS WOZENCRAFT

The VertebrateMuseum,ShippensburgUniversity,Shippensburg,
PA 17257
NationalMuseumof NaturalHistory,SmithsonianInstitution,Washington,DC 20560
Presentaddressof DMD:ZoologyDepartment,Universityof Tennessee,Knoxville,TN 37996

ABSTRACT.-A phylogeneticanalysisof all Recent procyonidgeneraand selectedmustelid

generawasperformedbasedon 69 cranial,28 dental,12 postcranial, and 20 soft morphological
characters.The mostparsimonious phylogenetictree (consistencyindex= 0.763)supportedtwo
monophyleticcladeswithinthe Procyonidae. Bassaricyonand Potoscomprisedthe firstbranch,
whereasBassariscus,Procyon,Nasua,and Nasuellaformedthe secondclade. Bassaricyonand
Potos were unitedby 10 synapomorphies features),amongtheseare:relatively
(shared-derived
uninflatedauditorybullae,reductionof dentalcusps,andfeaturesof the petrosalandearossicles.
Amongthe 14 synapomorphies thatuniteBassariscus,Procyon,Nasua,and Nasuellaarepalatine
inflationof the orbitalwall,presenceof a hypoconeon P4 and Ml, and an entoconidon m2.

The first widely accepted classification of the Procyonidae was that of Turner (1848) who
placed the raccoon (Procyon), coati (Nasua), ringtail (Bassariscus), and kinkajou (Cercoleptes
= Potos) in the family based on the absence of the alisphenoid canal and the presence of m
2/2. Turner (1848) stated that three closely related families could be characterized based on
their molar formulae: ursids, m 2/3; procyonids, m 2/2; and mustelids, m 1/2. Mivart (1885)
added to the classificationfeatures concerning modification of the carnassialshear and a relative
increase in the size of the molars. Lichtenstein (1827) originally described Bassariscus as a form
intermediate between Genetta (Viverridae) and Nasua, and Gray (1843) placed it within the
Viverridae. However, Turner (1848) and Flower (1869) argued, based primarily on basicranial
data, that Bassariscus be placed in the Procyonidae. Flower and Lydekker (1891) and Pocock
(1921a) further refined hypotheses regarding relationshipsat the generic level based on features
of the basicranial region and external anatomy; most authors have followed their classifications
(Honacki et al., 1982; Simpson, 1945; Wozencraft, 1989a).
Ailurus was placed in the family Procyonidae by Mivart (1885) based on presence of a simple
kidney (plesiomorphic character in carnivores)and absence of M3 (absent in all families except
canids and ursids);however, phylogenetic analyses using biochemical (Sarich, 1976; Todd and
Pressman, 1968) and morphological evidence (Ginsburg, 1982; Wozencraft, 1989b) place the
red panda (Ailurus) within or near the Ursidae.
There has been only one phylogenetic study (sensu Hennig, 1966) of relationshipswithin the
Procyonidae (Baskin, 1982). Baskin (1982, 1989) tentatively united Bassaricyon with the Nasua-
Procyon clade and placed Bassariscusas the firstbranch within the Recent procyonids(Bassaricy-
on and Potos were not discussed by Baskin, 1989). We have reservations in accepting this
hypothesis because: Bassariscus was considered part of the outgroup, therefore was precluded
from being placed anywhere except as the sister taxon of the remaining members (rectified in
Baskin, 1989); 76%of Baskin's(1982) charactersthat unite Bassaricyon with Nasua and Procyon
concern the morphology of P4/ml (modificationof the carnassialshear);polarityof the remaining
characters that unite Bassaricyon with Procyon and Nasua must be reversed in Bassaricyon to
keep it within this clade; many of Baskin's (1982, 1989) characters concern relative size and
were not scaled; some charactersare repeated within a transformationseries (e.g., enlarged M2);
and Potos (Baskin, 1982, 1989) or Bassaricyon (Baskin, 1989) was not included in the analysis.
Several early workers proposed, based primarily on dental similarities, that ursids were the
sister group of procyonids (Flower, 1869; Mivart, 1885; Turner, 1848) because both groups have
additional cusps on the molars. More recent phylogenetic analyses suggest that ursids are an
outgroup to the procyonid-mustelid-phocid clade and that mustelids are the sister group to
procyonids (Baskin, 1989; Bugge, 1978; Ginsburg, 1982; Tedford, 1976; Wozencraft, 1989b).

J. Mamm., 72(1):42-55, 1991 42

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February 1991 DECKER AND WOZENCRAFT-PHYLOGENY OF PROCYONIDS 43

Many workers have used the hypocone of P4 on some procyonids to characterize the family
Procyonidae (Baskin, 1989; Flower and Lydekker, 1891; Lankester, 1901; Mivart, 1885). Baskin
(1989) united New World procyonids based on presence of a well-developed parastyle on P4
(relative to mustelids) and presence of a new posteriorly placed hypocone on M1. Wozencraft
(1989a) united procyonids based on presence of a uniquely derived suprameatal fossa and an
anteriorly bilobed baculum. Bassaricyon and Potos lack a distinguishable hypocone on M1, and
other characters suggest that its absence represents a secondary loss in these taxa.
Herein, we examine genealogical relationships among Recent procyonid genera. We exclude
fossil taxa to yield a more complete data set (fossil material of New World Tertiary procyonids
is restricted largely to dentition-Baskin, 1982, 1989; Linares, 1982), and to avoid certain prob-
lems that arise when combining fossil and Recent classifications(Wiley, 1981). We do not consider
the red panda a member of the Procyonidae, therefore exclude it from our analysis. The rela-
tionship of the red panda to canoid carnivores is discussed elsewhere (Wozencraft, 1989b). The
hypothesis we propose here is the most parsimonious arrangement for the Recent Procyonidae
and employs a classification scheme that conveys the most information (Wiley, 1979).

METHODS
Standardphylogenetic techniques were employed using comparative morphology (Wiley, 1981). Character
polaritieswere establishedbased on the outgroup-comparison method and independentpolarizationof
transformation seriesof coded features(Watrousand Wheeler,1981).To avoid problemsresultingfrom
genus-levelparaphyly,the operationaltaxonomicunitsfor our analysiswerespecies.Characterswith more
thantwo statesweredividedintomultistatecharactersfollowingthe additivebinary-coding method(Brooks
et al., 1984).Characters wereanalyzedby useof a computeralgorithm,PAUP(PhylogeneticAnalysisUsing
Parsimony-Swofford,1984), with the global-branch-swapping and multiple-parsimonyoptions(Appen-
dix I).
Characters exceptfor 10soft-anatomical
initiallyobtainedfromthe literaturewereverifiedby observation,
featuresfor whichwe reliedon publisheddescriptions(Bugge,1978;Davis,1964;Story,1951;Windleand
Parsons,1897,1898).We examinedcranialandpostcranial materialfromadultandjuvenilemaleandfemale
specimensfor each taxon(AppendixII). All procyonidspeciesrecognizedby Honackiet al. (1982)were
examined,exceptforBassaricyonlasiusandB. beddardi,speciesof dubiousvalidity(Ewer,1973;Poglayen-
Neuwalland Poglayen-Neuwall, 1965;Wozencraft,1989b).
We used the Mustelidae as the outgroup; this family is widely accepted as the sister group of the Recent
Procyonidae (Baskin, 1989; Ginsburg, 1982; Schmidt-Kittler, 1981; Tedford, 1976; Wozencraft, 1989b).
Representatives of each of the following subfamilies were examined for outgroup comparison (Appendix I):
Lutrinae (Lutra lutra), Mephitinae (Spilogale putorius), Melinae (Melogale personata), and Mustelinae
(Martes americana and Eira barbara). Character transformationsthat were ambiguous based on outgroup
comparisonwere treated as unordered. The branching pattern within the Procyonidae was the same regardless
wasusedas the outgroup.Therefore,we choseto present
of whichsubfamily(orcombinationof subfamilies)
our resultswith Eira and Martesas the outgroupsbecausethey are similarto some procyonidsin both
morphology and ecology. Martes represents one of the more primitive members of the Mustelidae and Eira
showsmanystrikingsimilaritiesto Potos.
Bassariscustraditionallywas recognizedas the mostprimitiveRecentprocyonidbecauseof the number
of primitivedentalfeaturesit shareswith othercarnivores(Baskin,1982, 1989;Mivart,1885).However,
similarityof primitivefeaturesdoes not indicaterelationship(Hennig,1966;Wiley, 1981).Also,Potos has
been consideredeithera primitivemusteloid(Baskin,1982;Segall,1943)or the mostdivergentprocyonid
(Pocock, 1929). We tested the hypothesis that Bassariscus was the most primitive procyonid by placing it
as the firstbranchto the othermembers(Fig. la). Anotherusertreewasconstructedto examinethe question
of the relationshipbetweenBassaricyonand Potos (monophylyor diphyly);this tree placedPotos as the
first,Bassaricyonas the second,and Bassariscusas the thirdbranch(Fig. lb).
We compared our results with those of Baskin (1982, 1989). Because Baskin was not concerned solely with
the relationshipsamong Recenttaxa, he selectedcharacters(primarilydentition)that best distinguished
amongfossilremains.However,we restructured his datato conformto ourformatusinghis interpretations
and transformationsas a guide. We coulddiscern52 characters,basedon his coding,of which 42 (82%)
weredentalcharacters.Ourreanalysisof Baskin's(1982)dataexcludedfossilprocyonids(whichshouldnot
affectbranchingpattern),includedPotos,and usedMustelidaeas the outgroup.

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 44 JOURNALOF MAMMALOGY Vol. 72, No. 1

Martes Martes BassaricYon

a Eira
b Eira
C
Potos Potos Nasua

Bassaricyon Bassaricyon

Procyon Bassariscus Nasuella

Nasua Procyon
Procyon
Nasuella Nasua

Bassariscus Nasuea Bassascu

Bassartscus

FIG.1.-a, Testof specifiedtree withBassariscus

as the firstcladewithinthe Procyonidae. Reanalysisof
Baskin's(1982)dataproducedan identicaltree;b, test of specifiedtree with Potosas the firstcladeto the
c, tree proposedby Baskin(1982).
restof the procyonids;

RESULTS

The most parsimonious cladogram (Fig. 2) has a length of 135 and a consistency index of
0.763. This cladogram contains two monophyletic clades within the Procyonidae. The first clade
(Potosinae) consists of Bassaricyon and Potos, and the other clade (Procyoninae) consists of
Bassariscus, Procyon, Nasua, and Nasuella. The relative merits of this arrangement can be
evaluated based on the number of synapomorphiesat each branching point. The Bassaricyon-
Potos clade is supported by 10 synapomorphies (no reversals), and the Bassariscus-Procyon-
Nasua-Nasuella clade is supported by nine synapomorphies (two reversals).
The Potosinae share modifications in dentition such as rounded molars, reduction or loss of
cusps, loss of the carnassialshear (related to broadening of the lingual cingulum and reduction
of the metastyle), and grooved canines. Other cranial features of the Potosinae are a relatively
small entotympanic bone, internal auditory meatus situated on the ventromedial edge of the
petrosal, shortened rostrum, laterally bowed zygomatic arch, and long supraorbitalprocesses.
The ventral borders of the anterior opening of the infraorbital canal are positioned distinctly
posteriorto the dorsalborders.Steno'sforamen is located medially, between the incisive foramina.
In the postcranial skeleton, the acromion process on the scapular spine is sharply angled.
The kinkajou (Potos) has the most autapomorphies of any procyonid. The skull is shortened
and rounded. The mandible is modified by a triangular-shapedcoronoid process (which it shares
with Martes), a reduced angular process, and a posteriorly located mandibular foramen. Ossi-
fication in the lacrimal region of the orbit is incomplete, with no distinct naso-lacrimal canal.
Rather than a long narrow mastoid process, more typical of procyonines, there is a wide flat
ventral surface separating the small paroccipital process from the bullae. The prehensile tail and
the ventral cutaneous glands (also autapomorphiccharacters)were discussed in detail by Pocock
(1921a). Bassaricyon has two autapomorphies that distinguish it from other procyonids. The
posterior border of the nasals is U-shaped and the lateral edges are parallel. Also, Bassaricyon
is the only procyonid with a poorly developed trochanteric fossa on the femur.
The Procyoninae are distinguished by presence of a hypocone on P4 and M1, an entoconid
on m2, a laterally expanded palatine giving the orbital wall an inflated appearance, medial or
posterior attachment of the vomer to the palatine, and an annulated tail.
Bassariscus has primitive carnivore dentition, with only a slight reduction in carnassialshear
(Baskin, 1982); it is the most carnivorousof the procyonids (Walker et al., 1968). The suprameatal
fossa of Bassariscus has a deep posterolateral orientation, whereas in all other procyonids the
fossa has a posteromedial orientation. The mesopterygoid region of the palate is absent in

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February 1991 DECKER AND WOZENCRAFT-PHYLOGENY OF PROCYONIDS 45

Bassaricyon

Potos

Bassa risc us

SNasua

Nasu ella

Procyon

Martes

Eira

FIG.2.-The mostparsimoniousphylogenetictree of the RecentProcyonidaewith Martesand Eira as

the outgroup.The numberin parentheseson each stem refersto the corresponding
numbersin Appen-
dix I.

Bassariscus and, as a reflection of this, the palatine is much shorter than the maxilla. Although
Bassariscus is not the most primitive clade on the tree, this taxon has the fewest number of
apomorphies as measured from the base of the tree.
The clade containing Procyon, Nasua, and Nasuella is distinguished from Bassariscus by a
large parastyle on P4, the metastylar blade is reduced, and the carnassial shear is functionally
lost. In Procyon, Nasua, and Nasuella the mesopterygoid region is depressed dorsally along the
midline, the deep fossa for insertion of the tensor tympani in the petrosal is partially covered
by a thin bony lamina, and the floccular fossa fills in with age.
Procyon has four autapomorphic traits: the maxilla-palatine suture is truncate; the external
auditory meatal tube is long and extends to the lateral edge of the squamosal; the black facial
mask is complete; and interdigital webbing is absent. The coatis (Nasua and Nasuella) are

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46 JOURNALOF MAMMALOGY Vol. 72, No. 1

distinguishedby their rostralmorphology:a long Steno'sforamen between the incisive foramina;

upturned nasals;flexible snouts;and a deep nasolabialisfossa. Shared dental charactersinclude:
reduced 13 (equal in size to I1 and 12) displaced laterally on the premaxilla, procumbent il-3,
and laterally flared upper canines. Nasua can be distinguished from other procyonids by a deep
zygomaticus muscular fossa in the orbital wall, a distinctly enlarged basal whorl of the promo-
torium in the petrosal, and a round window considerably larger than the oval window. Nasuella
exhibits progressive trends in several characters such as reduced dentition and absence of su-
praorbital processes.
Figure la differs from our most parsimonioustree (Fig. 2) only in the placement of Bassariscus
as the first branch of the procyonids. The tree in Fig. la (length, 140; consistency index, 0.736)
is longer than our final tree, and requires 26 character reversals. The tree in Fig. lb is even
longer (length, 143; consistency index, 0.720) and requires 34 character reversals. Careful con-
sideration must be given to alternative hypotheses concerning the evolution of this group. This
must be more than a simple consideration of tree length, because the characters involved may
form character complexes or they may be related in a manner that we do not understand fully.
Differences between the trees in Fig. la and Fig. 2 include features of the nasolabialisfossa,
relative size of the palatine, vomer attachment to the maxilla, inflation of the orbital wall of the
palatine, and the position of Steno'sforamen. Several features of the basioccipitalregion, features
of the rostralprocess of the maxilla, and the position of Steno's foramen and the external pinnae
were reversed in Fig. lb. These features are relatively conservative in the evolution of the
Carnivora(Flynn and Galiano, 1982; Flynn et al., 1988; Hunt, 1974, 1987; Turner, 1848; Wozen-
craft, 1984, 1989a; Wyss, 1987).
In our final tree (Fig. 2), characterreversalsnot sharedwith Figs. la and lb involved postcranial
characters that show a high amount of homoplasy across the Carnivora. Analogous conditions
can be found within the Viverridae and Mustelidae. Some charactersreversed in all three trees
were related to the carnassialshear, one of the main synapomorphiesthat defines the Carnivora.
Procyonids, like viverrids, have invaded the frugivorus niche and have corresponding modifi-
cations in dentition.
Our final tree (Fig. 2) was chosen not only because of less homoplasy and fewer character
reversals, but also because of the nature of the character distribution. Alternative topologies
involved reversalsin charactersthat show little homoplasy in other groups of carnivores,whereas
our most parsimonioustree only showed homoplasy in characters that appear to be susceptible
to parallel and convergent evolution when compared to other carnivoran families.
The most parsimonioustree produced from our analysis of Baskin's(1982) data differed from
our final phylogeny (Fig. 2) only in placing Bassariscus as the first branch of the procyonids (no
reversals;length, 37; consistency index, 0.532). It agreed with our tree in uniting Bassaricyon
and Potos (Fig. la). The remaining procyonids (excluding Bassariscus) are united by four
synapomorphies involving characters dealing with relative size of cusps. Baskin (1982) did not
intend to resolve relationships among Recent procyonids; he selected characters based on their
ability to resolve relationships among fossil taxa. Nevertheless, we were encouraged by how
closely our independent analyses agreed.
DISCUSSION
Our most parsimonious tree (Fig. 2) suggests major changes in the traditional view of rela-
tionships among procyonid genera. Bassariscus is not the most primitive clade, but is the taxon
with the fewest number of derived states. Bassaricyon and Potos form a highly autapomorphic
group (Potosinae);Bassaricyon is essentiallya primitive versionof Potos. The Recent Procyoninae
consists of Bassariscus, Procyon, Nasua, and Nasuella, with the latter three taxa sharing a suite
of morphological features reflecting their omnivorous and cursorial habits.
We did not attempt to determine what possiblefeatures united the Procyonidaeto the exclusion
of other carnivores. However, key innovations that unite the Procyonidae and are supported by
our tree include the posterior orientation of the suprameatal fossa, the presence of a squamosal

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February1991 DECKERAND WOZENCRAFT-PHYLOGENY
OF PROCYONIDS 47

epitympanic sinus, and a distinctly bilobed baculum. The suprameatal fossa was recognized as
the most distinctive characteristic of the Procyonidae (Hunt, 1974; Schmidt-Kittler, 1981; Ted-
ford, 1976; Wozencraft, 1989b). Schmidt-Kittler (1981) recognized a more dorsally placed ho-
mologous fossa in the mustelids that connects to the epitympanic recess, but Wyss (1987) ques-
tioned this interpretation.Ursidsand Ailurus have a shallow, dish-shaped depressionin the dorsal
wall, unlike the posteriorly projecting deep fossa of the procyonids. Segall (1943) described the
presence of an epitympanic sinus in the squamosal as a character distinguishing procyonids from
other caniforms,but stated that Potos lacked such a sinus. However, our examinationof specimens
indicates that all procyonids, including Potos, have squamosal epitympanic sinuses, absent in
Ailurus.
The relative inflation of the ecto- and entotympanic is also an important feature of procyonids.
The condition varies in mustelids: Martes has an inflated entotympanic and Spilogale has a
relatively uninflated entotympanic. Pocock (1921b) suggested that because dissimilargenera such
as Meles, Gulo, and Martes all have inflated bullae, this condition probably is primitive in the
Mustelidae. If so, Bassaricyon and Potos would have the derived condition (uninflated entotym-
panic) and it would be another synapomorphy to support the monophyly of the potosines.
However, if the uninflated entotympanic is considered primitive, the inflated condition would
unite the Procyoninae, and the potosines clade would be the first outgroup to the procyonines.
On the basis of ontogeny, this character might be considered plesiomorphic (primitive-Hum-
phries and Funk, 1984). Regardless of whether we consider the inflated or the uninflated ento-
tympanic as primitive, our arrangementof taxa is supported.Furthermore,the degree of inflation,
or the percentage of the bullar floor composed of the entotympanic is a relative categorization.
Strict outgroup comparison among carnivore families (Baskin, 1982; Pocock, 1921a; Wozencraft,
1989b) indicates that the uninflated entotympanic is the derived state, and our data support this
hypothesis.
The dental characters used by Mivart (1885) and Turner (1848) to unite the Procyonidae are
not supported at the base of our tree. If the hypocone on P4 were used to unite the Procyonidae,
two characterchanges would be required;using the hypocone on P4 to unite only the Procyoninae
would require one. Bassaricyon and Potos have a posterolingual widening of P4 that might
suggest secondary loss of the cusp, and its variable presence in Bassariscus sumichrasti suggests
that it could have been lost more than once.
Of major characters that unite the Procyonidae, homology of the suprameatal fossa appears
to be most open to question. Morphological features as diverse as the shallow dish-shaped fossa
in ursids, the deep posterolateral tapering pocket in Bassariscus, and the lateral pocket off of
the epitympanic cavity of the middle ear in some mustelids have been referred to as suprameatal
fossas. It is difficult to determine whether all of these features are homologous, especially when
the functional significance of the fossa is unknown. However, resolving the homology and function
of this feature will not affect the topology of our most parsimonious tree, but may affect the
phylogenetic placement of various extinct European carnivores (Broiliana, Stromeriella, and
Zodiolestes) considered procyonids primarily based on the presence of this feature.
The common ancestorof living potosinesmost likely was arborealand frugivorous,and potosine
synapomorphies are, to a great extent, a reflection of these adaptations. Forward placement of
the orbits may allow for greater stereoscopic vision, an adaptation for arboreality. Bunodont
dentition and grooved canines can be attributed to frugivorous habits (Eisenberg, 1981). The
frugivorous small-toothed palm civet (Arctogalidia trivirgata) of southeastern Asia also shows
reduction and simplification of tooth cusps, long postorbital processes, and grooved canines
demonstrating parallel evolution of adaptations for frugivory. Story (1951) illustrated features
in potosines that also were related to adaptations for arboreal frugivory: development of the
masticatory muscle (temporal, zygomatico-mandibular, masseter, and pterygoids) is greater in
the horizontal plane than in the sagittal plane; one main artery supplies the cranium whereas
forms with longer crania tend to have two well-developed meningeal arteries; and the orbital
region, but not the nasal region, is well supplied with vessels.

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48 JOURNALOF MAMMALOGY Vol. 72, No. 1

Potos has the most autapomorphiesof any procyonid. Simpson (1945) postulated that it must
have diverged from all other procyonids early in the Miocene. However, uniqueness does not
convey information regarding the relationship of Potos to other taxa. We are hesitant to place
Bassaricyon and Potos in the same genus because of the uncertain status of several proposed
Bassaricyon species. Should future research support suggestions by Ewer (1973) and Poglayen-
Neuwall and Poglayen-Neuwall (1965) that all Bassaricyon species are conspecific, then consid-
eration should be given to placing B. gabbii in the genus Potos. This produces the simplest
classification with the most information content (the uniqueness of Potos would be recognized
and the sister group relationship of Potos and Bassaricyon conveyed).
The pattern of cranial blood supply in the procyonines Bassariscus, Procyon, Nasua, and
Nasuella is similar to the pattern seen in the Mustelidae, Canidae, and Ursidae (Story, 1951).
This suggeststhat procyoninespossessthe more primitive condition of blood supply to the cranium
and the potosines have the derived condition. Variability in the arterial pattern in the face of
procyonids may be related to functional necessities accompanying changes in facial proportions
(Story, 1951). Procyon, Nasua, and Nasuella have evolved a longer rostrum and smaller orbits,
whereas Bassaricyon and Potos have shorterrostraand enlarged orbits.Bassariscusis a functional
intermediate, with enlarged orbits, but without a reduced rostrum (Story, 1951).
Baskin (1982:92) interpreted the fourth cusp in the posterolingual position on Ml as a meta-
conule, based on its connection to the protocone by a short postprotocrista (plagiocrista of
Hershkovitz, 1971). This requires development of the metaconule, its enlargement to replace
functionally the hypocone in Nasua and Procyon lotor, its subsequent reduction and movement
to a more internal position on the postprotocrista,and development of a large hypocone in
Procyon cancrivorus. In Nasua examined, the cusp may or may not be connected with the
plagiocrista. If this cusp is interpreted as a metaconule, it must have moved off the crista onto
the posterolingual cingulum in a manner similar to the hypocone. Thus, the most parsimonious
interpretation is that this cusp is homologous to the hypocone (Butler, 1941; Hershkovitz, 1971).
The long Steno'sforamen in Nasua and in Nasuella is associated with the rich supply of blood
vessels to the nasal region (Story, 1951) and the placement of the Jacobson'sorgan. The "nose-
up" posture that coatis display in intense threatening situations(Kaufmann, 1962) and the general
flexibility of the snout is reflected in the upturned nasalsand large muscle fossas.The maxillonaso-
labialis muscle that attaches to the nasolabialisfossa is developed extensively and serves as the
primary mechanism for rhinarial mobility. This morphology may be associated with grooming
behavior in coatis, that frequently use their incisorsas pincers, nipping the skin lightly in a rapid
series of bites (Kaufmann, 1962). In addition, dental synapomorphies shared by Nasua and
Nasuella may be related to their insectivoroushabits.
Based on our tree (Fig. 2), we propose the following phylogenetic classification of Recent
procyonid genera and related taxa used in this study.

SYSTEMATICS
Class Mammalia
Infraclass Eutheria
Order Carnivora Bowdich, 1821
Suborder Caniformia Kretzoi, 1945
Family Procyonidae Gray, 1825
Subfamily Potosinae Trouessart, 1904
Potos E. Geoffroy and G. Cuvier, 1795
Bassaricyon Allen, 1876
Subfamily Procyoninae Gray, 1825
Bassariscus Coues, 1887
Procyon Storr, 1780
Nasua Storr, 1780
Nasuella Hollister, 1915

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February 1991 DECKER AND WOZENCRAFT-PHYLOGENY OF PROCYONIDS 49

Family Mustelidae Fischer de Waldheim, 1817

Subfamily Mustelinae Gill, 1872
Martes Pinel, 1792
Eira H. Smith, 1842
Subfamily Mephitinae Gill, 1872
Spilogale Gray, 1865
Subfamily Lutrinae Baird, 1857
Lutra Schreber, 1776

ACKNOWLEDGMENTS

We express our appreciation to the following persons for their careful review of and helpful comments
on this manuscript: J. A. Baskin, A. Berta, D. Frost, J. L. Gittleman, R. S. Hoffmann, T. Holmes, R. Kay,
M. L. Kennedy, K. Koopman, R. D. Owen, M. Ritke, M. E. Taylor, R. W. Thorington, R. H. Tedford, and
A. R. Wyss. R. W. Thorington and L. Gordon (National Museum of Natural History, SmithsonianInstitution),
G. Musser (American Museum of Natural History), M. E. Rutzmoser (Museum of Comparative Zoology,
Harvard), R. M. Timm (Museum of Natural History, University of Kansas),B. D. Patterson (Field Museum
of Natural History), G. L. Kirkland (Vertebrate Museum, Shippensburg University), and M. L. Kennedy
(Museum of Zoology, Memphis State University) kindly offered use of their respective collections. Funds
and support for this study were provided by The Vertebrate Museum, ShippensburgUniversity, the Museum
of Zoology, Memphis State University, the Office of the Assistant Secretary for Research, Smithsonian
Institution, and the National Museum of Natural History, Smithsonian Institution.

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 APPENDIXI
TABLEA.-Autapomorphic (unique-derived) and synapomorphic (shared-derived) characters for the phylogeny
numbers in Fig. 2). Primitive and derived conditions are listed for each feature.
Feature Primitive condition

(1) Procyonidae
Basioccipital rugosity Absent
Epitympanic sinuses Absent
Epitympanic recess Small
Suprameatal fossa Absent or vestigial
Baculum Long, stylized
(2) Potosinae
Steno's foramen On posterior edge of incisive foramina
Infraorbital canal, ventral border Ventral and dorsal borders in line
Supraorbitalprocess Small or absent
Malleus Long and slender
Number of cusps on molars >3
Internal auditory meatus Middle of petrosal
Entotympanic Well inflated
Acromion process Straight
External pinnae Long, pointed
(3) Potos
Trigonid ml Present
P3 One or two roots
Spinous process of axis Prominent
Prehensile tail Absent
Ventral cutaneous glands Absent
Hamular process of pterygoids Extends below horizontal plane of post-glenoid pr
Naso-lacrimal foramen Present
Lambdoidal ridge Extends to mastoid
Entotympanic inflation Present
Mandibularsymphysis Weak
Mandibular foramen Posterioventral to anterior edge of coronoid proces
Angular process Pronounced
(4) Bassaricyon
Posterior edge of nasals V-shaped
Trochanteric fossa on femur Well developed

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 APPENDIXI
TABLEA.-Continued.
Feature Primitive condition

(5) Procyoninae
Nasolabialis fossa Absent
Midline length of palatine excluding mesopterygoid Less than or equal to length of maxilla
Vomer Attached to anterior part of palate
Orbital wall of palatine Diverging, not inflated
Hypocone on P4 Absent
Hypocone on MI Absent
Entoconid on m2 Absent
Interosseuscrest of ulna Absent or small
Banded tail rings Absent
(6) Bassariscus
Midline length of palatine plus mesopterygoid Greater than or equal to length of maxilla
Mesopterygoid region Present
Carnassialshear on P4 Trenchant
M. sartorius,caudal head Single
(7)
Mesopterygoid region Flat or absent
Ethmoid foramen Small
Petrosal, floccular fossa Present, deep
Insertion of tensor tympani Deep, exposed fossa
Parastyle on P4 Absent or vestigial
(8)
Steno's foramen Lost or vestigial
Lateral profile of nasals Straight
Length of rostrum <3 times width of nasals
Rostral process of lacrimal Absent
Antorbital fossa Absent or vestigial
Pterygoid medialis Absent

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 I
APPENDIX
TABLEA.-Continued.
Feature Primitive condition

Diastema-I2 and I3 Absent

Diastema-C and P1 Absent
Diastema-PI and P2 Absent
13 Larger than Ii and I2
Lower incisors Vertical
Upper canines Directed ventrally
Scapular spine Straight
(9) Nasua
Basal whorl of cochlea Not enlarged
Fenestra rotunda Equal in size to fenestra vestibuli
Deep zygomaticus muscular fossa in orbital wall Absent
(10) Nasuella
Supraorbitalprocess Present
Sagittal crest (males) Present
(11) Procyon
Maxilla-palatine suture U-shaped
External auditory meatal tube Short
Black face mask Absent
Interdigital webbing Present

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54 JOURNAL OF MAMMALOGY Vol. 72, No. 1

APPENDIXII

Specimens Examined
Holotypes, indicated by an asterisk, are followed by the original name assigned to the specimen (in
parentheses) if different from the current name. Museum acronyms follow Yates et al. (1987).
Bassaricyon alleni.-USNM 194315; FMNH 41501, 34693, 34717, 41502, 55429, 62079, 65787-65789,
65805, 86908, 86909, 104935.
Bassaricyon gabbii.-AMNH 28486*; ANSP 18852; KU 104639; USNM 14214/12237* (B. richardsoni),
79053, 171138, 179157* (B. g. orinomus), 194315, 194316, 281484, 284773, 284903, 284933-284935, 307035,
307037, 305748-305753, 307036, 310666, 316230, 324293, 324296, 335767, 335768, 337632, 338348, 363343,
363344, 516945, 516946.
Bassaricyon pauli.-ANSP 17911*, 18851.
Bassariscus astutus.-ANSP 970, 971, 972* (B. a. flavus), 1614* (B. flavus oregonus), 3969, 5245, 6326,
6483, 8655, 51864, 82260, 108703, 125714, 126162* (B. a. consitus), 126181, 126199, 126235, 126237-
126239, 187950-187953, 187955; KU 4866, 68328, 91367, 119392, 125713; USNM 976/97* (B. raptor), 1619,
24023, 30619, 31157, 32246, 44715, 44717, 44829, 49734, 79031* (B. saxicolus), 79034* (B. a. insulicola),
126352, 108702, 118988, 144528, 174760* (B. a. nevadensis), 205388* (B. a. arizonensis), 146192* (B. A.
palmarius).
Bassariscus sumichrasti.-ANSP 18896; USNM 3273, 8977/9422, 13849, 51507, 108290, 108291* (Jen-
tinkia sumichrasti campechensis), 181254-181257, 292254, 340735-340740.
Eira barbara.-USNM 3839, 38481, 61203, 104546, 104547, 240886, 241383, 244901, 251153, 255132,
259003, 281466-281475, 281478, 281480, 290885, 305755, 307039, 307040, 338978, 339676, 361036, 362120,
362244, 398512, 411757.
Lutra lutra.-USNM 23217, 132224, 133686, 188626, 197708, 484480, 484805, 484826, 484842, 484843.
Martes americana.-USNM 3285, 3286, 23215, 116168, 134021, 136121, 148569, 148570, 188218, 209945,
209947, 209981, 210014, 210018, 210032, 210033, 210035, 210038-210040, 210045, 221877, 287847, 505316,
546163, 546165-546170.
Melogale personata.-USNM 239586, 253410, 254588, 254918, 255528, 258014, 258015, 260614, 292887,
297171, 330698, 356597-356600, 356604, 356605, 356828, 356931, 357537, 358072-358077, 358570, 505242,
543122.
Nasua narica.-ANSP 13631, 17903; AMNH 7125* (N. n. pallida), 18901* (N. n. panamensis), 19210*
(N. n. bullata); USNM 12037/10468* (N. n. yucatanica), 21983, 22810, 22811, 32630/44553* (N. n. molaris),
37287, 38421, 44554, 49644, 51331* (N. n. richmondi), 63112, 74681* (N. n. isthmica), 108311-108313,
108315, 108519, 108537, 116754* (N. n. tamaulipensis), 244896, 244900, 257672, 258606, 307020, 314547,
316212, 324286-324288, 338347, 520614, 520615, 541322.
Nasua nasua.-AMNH 131166* (N. candace dichromatica); KU 876, 11989, 11990, 26954, 29985, 29986,
63118, 63119, 65928; USNM 1661, 172991, 172993, 241335-241338, 270365, 281513, 283471, 362119, 374838,
388236, 388316, 398713-398719, 544417-544419, 546101.
Nasua nelsoni.-AMNH 17557* (N. phaeocephala); KU 92570; USNM 108520*-108524, 108615, 239962.
Nasuella olivacea.-AMNH 33045* (N. o. lagunetae); USNM 143658, 240034, 251906, 372854, 372855.
Potos flavus.-AMNH 1412/353* (P. f. chapadensis), 14186* (P. f. caucensis), 18926* (P. f. chiriquensis),
32722* (P. f. tolimensis); KU 19266, 65933, 68057, 68058, 83588, 93822-93824, 93826, 96901, 100052,
115610, 115611, 115613, 115614; USNM 13055, 74683* (P. f. guerrerensis), 179042* (P. f. isthmicus),
181266* (P. f. campechensis), 194317, 241181, 249829, 250319, 250320* (P. f. arborensis), 255123, 257315,
257352, 281522, 310664, 338858, 372732, 372734, 372857, 372858, 406525, 406827, 406829, 406832, 406833,
406836, 442896, 442892-442894, 442896, 442898, 443270, 443273, 443277, 443461, 443465, 503823, 503825,
503826.
Procyon cancrivorous.-AMNH 23492* (P. proteus), 36458* (P. [Euprocyon]aequatorialis); ANSP 4677;
USNM 6949, 49718, 123351, 171229, 171669* (Euprocyoncancrivoruspanamensis), 172733, 172987, 179049,
281516, 281518, 281519, 283470, 291139, 305747, 310656, 372851, 388315, 442891, 443521, 517720, 517721.
Procyon gloveralleni.-MCZ 18591*; USNM 267380, 267381.
Procyon insularis.-USNM 88978", 88982* (P. insularis vicinus).
Procyon lotor.-KU 14604, 72004, 72005, 80046; MCZ 3502* (P. 1. elucus); USNM 1386, 2018/1051* (P.
1. mexicanus), 2450* (P. 1. litoreus), 12191/14191* (P. 1. crassidens), 12796* (P. 1. dickeyi), 35090, 35136,
35185, 36517, 46344, 46392, 47130, 59900* (P.1. ochraceus), 60675* (P.1. californicus), 63055* (P.1. fuscipes),
72190, 91427-91429, 93137* (P. psora), 99272* (P. pallidus), 135457* (P. 1.vancouverensis), 135455, 135456,
135460-135462, 140568, 144069, 147181* (P. 1. grinnelli), 158246* (P. 1. varius), 171983* (P. pumilus),

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February 1991 DECKER AND WOZENCRAFT-PHYLOGENY OF PROCYONIDS 55

177546* (P. 1. shufeldti), 236214* (P. 1. excelsus), 187926* (P. 1. hirtus), 254989* (P. 1. marinus), 255037*
(P. 1. inesperatus), 255060* (P. 1. incautus), 255080* (P. 1. auspicatus), 256027* (P. 1. solutus), 256057,
259131, 261298, 271883, 275290* (P. 1. maritimus), 284886, 285118-285123, 532412.
Procyon maynardi.-MCZ 7750*.
Procyon minor.-USNM 38417/15481*.
Procyon pygmaeus.-KU 92565; USNM 108503, 108510, 108511*, 108512.
Spilogale putorious.-USNM 57586, 58700, 64468, 65632, 73522, 76232, 91430, 99580, 109074, 115947,
126326, 126327, 126339, 126340, 126370, 126643, 126644, 126726, 126727, 135473-135476.

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