Journal of Mammalian Evolution, Vol. 8, No.

4, December 2001 ( 2002)

Comments on the Systematics and Classification of the

Beavers (Rodentia, Castoridae)
William W. Korth1

The systematics of the beavers (Castoridae) are reviewed and definitions are presented for
each subfamilial group. Four subfamilies are recognized: primitive Agnotocastorinae (divided
into two tribes, Agnotocastorini and Anchitheriomyini); burrowing beavers, Palaeocastorinae;
giant beavers, Castoroidinae (containing two tribes, Castoroidini and Trogontheriini); and the
Castorinae. The agnotocastorines are viewed as the ancestral group for all later subfamilies. The
Palaeocastorinae is viewed as a side-branch, not ancestral or closely related to any of the later
subfamilies. The Castorinae and Castoroidinae may represent a distinct clade united by dental
features.

KEY WORDS: Phylogeny, Systematics, Classification, Castoridae, Beaver.

INTRODUCTION
The beavers (family Castoridae) are represented by only two existing species of the same
genus ranging throughout North America and Eurasia. However, they have an extensive
fossil record from North America, Europe and Asia, ranging from the latest Eocene or
early Oligocene. The beavers are a unique family and do not appear closely related to
other Recent groups of rodents (see Korth, 1994). A thorough review of the systematics of
fossil castorids has not been undertaken since Stirtons (1935) work over sixty years ago.
Stirton did not designate subfamilies, but he believed that the genera of castorids were
separable into two groups based on the cross-sectional shape of the incisors: the ante-
rior enamel surface was either rounded or flattened (or semi-flattened). Classifications of
castorids have been presented by a number of authors since that time, usually with lit-
tle explanation (Table I). Simpson (1945), the first to present formally a classification of
the family, identified two subfamilies, Castorinae and Castoroidinae. Martin (1987) pre-
sented a review of the burrowing beavers from the Arikareean of North America, refer-
ring them to a new subfamily, the Palaeocastorinae. Korth (1994), in his survey of North

1 Rochester Institute of Vertebrate Paleontology, 265 Carling Road, Rochester, New York 14610, and Depart-
ment of Earth Science and Science Education, Buffalo State College, Buffalo, New York 14222. e-mail:
korthww@buffalostate.edu
2 Correspondence should be sent to the following address: William W. Korth, Rochester Institute of Vertebrate
Paleontology, 265 Carling Road, Rochester, NY 14610.

279

1064-7554/ 01/ 1200-0279/ 0 2002 Plenum Publishing Corporation

 280
Table I. Previous Classifications of Castoridae
Korth, 1994 McKenna and Bell, 1997
Stirton, 1935 Simpson, 1945 (North America only) Xu, 1995 This Paper
Semi-flattened incisors Castorinae Uncertain subfamily Agnotocastorinae Castorinae Agnotocastorinae
Agnotocastor Agnotocastor Agnotocastor Agnotocastor Agnotocastor Agnotocastorini
Palaeocastor Palaeocastor Hystricops Castorinae Capacikalini Agnotocastor
Euhapsis Euhapsis Castorinae Palaeocastor Capacikala Neatocastor
Palaeomys Palaeomys Castor Capacikala Pseudopalaeocastor Hystricops
Castor Sinocastor Palaeocastorinae Steneofiber Castorini Anchitheriomyini
Convex incisors Castor Palaeocastorini Hystricops Euhapsina Anchitheriomys
Steneofiber Castoroidinae Palaeocastor Castor Euhapsis Propalaeocastor
Monosaulax Steneofiber Capitanka Fossocastorinae Fossorcastor Oligotheriomys
Eucastor Monosaulax Capacikalini Euhapsis Castorina Palaeocastorinae
Dipoides Eucastor Capacikala Capatanka Propalaeocastor Palaeocastor
Castoroides Dipoides Pseudopalaeocastor Nannasfiber Palaeomys Capacikala
Amblycastor Procastoroides Euhapsini Fossorcastor Palaeocastor Pseudopalaeocastor
Hystricops Castoroides Euhapsis Asiacastorinae Hystricops Euhapsis
Trogontherium Trogontherium Fossorcastor Asiacastor Castor Fossocastor
Conodontes Amblycastor Castoroidinae Anchitherioimys Castoroidinae Castoroidinae
Monosaulax Youngofiber Steneofiber Castoroidini
Eucastor Trogontherium Neatocastor Monosaulax
Dipoides Castoroidinae Asiacastor Eucastor
Procastoroides Eucastor Youngofiber Dipoides Schlosser
Castoroides Dipoides Trogontherium Castoroides
Procastoroides Eucastor Paracastoroides
Castoroides Schreuderia Trogontheriini
Dipoides Trogontherium
Boreofiber Boreofiber
Romanocastor Euroxenomys
Zamolxfiber Youngofiber
Procastoroides Asiacastor
Castoroides Castorinae
Paradipoides Castor
Palaeomys
(=Chalicomys)
Steneofiber
Zamolxifiber

Korth
Romanofiber
Schreuderia
Systematics of the Beavers 281

American rodents, provided a historical review of the classification of beavers and rec-
ognized three subfamilies (Castorinae, Castoroidinae, and Palaeocastorinae), along with
the primitive and problematical genus Agnotocastor, which was not assigned to any of
the three recognized subfamilies.
A more recent attempt at a classification of beavers was in an unpublished disser-
tation by Xu (1995), where craniometric parameters were used to define the separate
subfamilies of the Castoridae. Xu (1995) recognized five subfamilies: Simpsons (1945)
Castorinae and Castoroidinae along with three new subfamilies (Agnotocastorinae, Fos-
sorcastorinae, Asiacastorinae), whereas he abandoned the concept of the Palaeocastori-
nae. In a classification for all mammals, McKenna and Bell (1997) generally followed
Xu (1995) but included all genera in the two subfamilies (Castorinae and Castoroidi-
nae) recognized originally by Simpson (1945). McKenna and Bell (1997) also included
the Rhizospalacidae as possible Eurasian castoroids. The most recent discussion of the
classification of beavers was the naming of a new subfamily, Agnotocastorinae, for the
nominal genus and similar primitive genera by Korth and Emry (1997).
The purpose of this work is to present diagnoses of the subfamilial groupings of the
Castoridae, which will help to better distinguish different lineages or clades within the
family and to suggest a possible classification for the group.

MATERIALS AND METHODS

Cranial and dental characteristics of beavers were examined and a cladistic analysis
of these features is applied here to the systematic classification of the Castoridae. Direct
observations on castorids from North America were made on specimens predominantly
from the collections of the American Museum of Natural History, Carnegie Museum of
Natural History, and the University of Nebraska State Museum. A few species of Euro-
pean and Asian beavers available at the American Museum were also studied directly.
Observations on species and genera of castorids not directly studied (mainly Eurasian
taxa) are based on literature cited throughout this paper. Dental terminology for castorids
follows that presented by Stirton (1935: also see Fig. 1), and identification of cranial
foramina follows Wahlert (1974, 1977: also see Fig 2).

RESULTS
Supergeneric Characters
The Castoridae is a distinct family of rodents originating in the latest Eocene of
North America. Definitions or characterizations of the family have been presented by a
number of authors based on dental and cranial features (Stirton, 1935; Wahlert, 1977;
Korth, 1994; Xu, 1995; Hugueney, 1999). The rodent family believed to be the clos-
est sister-group to the castorids is the Eutypomyidae [see Wahlert (1977) for a review
of eutypomyid systematics]. The genera of castorids, as viewed here, are divisible into
four distinct subfamilies. The subfamily Agnotocastorinae retains a number of primi-
tive characteristics of the dentition and skull that are shared with the castoroid family
Eutypomyidae: 1) complex occlusal pattern of cheek teeth; 2) upper tooth rows paral-
lel; 3) premolars subequal in size to molars; 4) retained stapedial foramen; 5) posterior
282 Korth

Fig. 1. Schematic drawings of castorid cheek teeth, demonstrating the specialized dental nomenclature.

palatine foramen within palatine bone; 6) elongated rostrum; 7) P3 or dP3 retained; and
8) palatal surface smooth (Korth, 1996; Korth and Emry, 1997). The remainder of the
beaver genera have lost the primitive eutypomyid features of the agnotocastorines and
are characterized by: 1) decreased complexity of the occlusal pattern of cheek teeth (with
smooth-sided re-entrant valleys); 2) posteriorly divergent upper tooth rows; 3) loss of P3
and dP3; 4) enlargement of last premolars (relative to molars); 5) loss of the stapedial
foramen; 6) parallel grooves on the palatal surface; 7) shortening of the incisive foramina;
and 8) anterior position of the posterior palatine foramina (within the palatine-maxillary
suture).
Systematics of the Beavers 283

Fig. 2. Specialized cranial features used in text to define subfamilies or tribes of castorids. Above: anterior
extent of jugal bone on zygoma (lateral view of skulls). A, late Miocene castoroidine Dipoides (note anterior
extent of jugal just anterior to middle of zygoma). B, Recent castorine Castor (note jugal extends anteriorly to
contact lacrimal bone). Below: cranial foramina of castorids (ventral view of skulls). C, Oligocene Steneofiber
(note lack of stapedial foramen [st] and occurrence of posterior palatine foramen [ppl] within the palatine-
maxillary suture). D, Oligocene Agnotocastor (stapedial foramen present in bulla and posterior palatine foramen
entirely within palatine bone). Figures not to scale. Figure A modified from Wagner (1983); B and C modified
from Hugueney (1999); and D modified from Korth (2001b).

The non-agnotocastorine castorids are readily separable into three groups based on
the morphologies of their respective postcranial skeletons. Members of the Palaeocastori-
nae are small with fossorial adaptations (enlarged forelimbs, low, broad skull, shortened
tail), the Castoroidinae are characterized by their much larger size and general massive-
ness of the skeleton, and the Castorinae have distinct aquatic adaptations (broadening of
limb bones, flattening of caudal vertebrae). However, these postcranial differences are
often observable only in the most derived species and cannot be recognized in the ear-
284 Korth

Table II. Derived Characters in Different Tribes and Subfamilies of Castoridsa

Ag An Pal Cod Tro Cas
P3/dP3 lost + + + +
stapedial foramen (artery) lost + + + +
posterior palatine foramina in
palatine-maxillary suture + + + +
premolars larger than molars + + + +
upper tooth rows strongly divergent posteriorly + + + +
no accessory crenulations on cheek teeth + + + +
grooved palate + + + +
incisive foramen less than 30%
length of diastema +/ + + + +
rostrum broad and shortened +
upper incisor flattened +
deep (dorsoventrally) zygomata + + + +
no contact between jugal and lacrimal bones + +
alternating alignment of mandibular
condyles (zig-zag) +/ +/ + +
increased hypsodonty of cheek teeth + + +
ridged incisors +/ +/ +/
M3 elongated anteroposteriorly +
very large body size +/ + +
a+, indicates presence of character; lack of character; +/ variable occurrence. Abbreviations: Ag,
Agnotocastorini; An, Anchitheriomyini; Pal, Paleocastorinae; Cod, Castoroidini; Tro, Trogontheriini; Cas,
Castorinae.

lier species due to their primitive morphology or to the lack of postcranial elements in
the fossil record. Therefore, the characters used here to distinguish between subfamilies
are limited to those of the skull and dentition (Table II).
Xu (1995) recognized two additional cranial characters that he used to separate the
beavers into subfamilial groups. The first is the morphology of the jugal bone on the
zygoma (Fig. 2). In primitive rodents such as paramyines (Wood, 1962) and sciuravids
(Dawson, 1961) the jugal contacts the lacrimal. Other sciuromorphous rodents, such as
sciurids, also share this feature. However, hystricomorphous and myomorphous rodents
(and geomyoids) do not have a contact between the lacrimal and jugal [see examples in
Ellerman (1940)]. The primitive ctenodactyloid Cocomys also lacks this contact (Li et al.,
1988). Clearly, this is a primitive feature only for protrogomorphous and sciuromorphous
rodents, including the castorids. In the derived condition (castsoroidines and trogontheri-
ines among the beavers), the maxillary bone separates the jugal from the lacrimal and
there is no contact between the latter.
The second feature utilized by Xu (1995) is the alignment of the mandibular pro-
cesses. Primitively in all rodents the angle, condyle, and coronoid process of the mandible
form a straight line when the mandible is viewed from the posterior (Fig. 3). This is the
condition in nearly all sciurognathous rodent families. The derived condition is one where
the condyle is medial to the other two processes and the angle is flattened and projected
medially, giving the posterior alignment of the processes a zig-zag pattern.
The following classification is based on the use of the above listed characters for
the subfamilies and tribes of beavers.
Systematics of the Beavers 285

Fig. 3. Alignment of mandibular condyles in castorids. All right mandibles, viewed from posterior. A, Recent
castorine Castor. B, Middle Miocene castoroidine Eucastor. C, Middle Miocene anchitheriomyine Anchithe-
riomys. D-F, latest Oligocene to early Miocene palaeocastorine beavers showing progressive development of
zig-zag pattern of processes. D, Capacikala. E, Palaeocastor. F, Pseudopalaeocastor. Figures not to scale.
Figures A, B, D-F modified from Xu (1995); C modified from Korth (2001a).

Classification and Definitions

Family CASTORIDAE Hemprich, 1820
Definition: Cranial featuresskull generally low, broad, and robust with broad
zygomata; postorbital process lacking; sciuromorphous zygomasseteric structure, with
infraorbital foramen low on rostrum; sciurognathy; digastric (symphyseal) eminence on
mandible; interorbital foramen posterior to optic foramen. Dental featuresincisor micro-
structure uniserial; dental formula I11 C00 P(2)11 M33; cheek teeth lophate, forming series
286 Korth

of transverse or oblique re-entrant valleys (cflexi) or elongated fossettes of enamel; cheek

teeth at least mesodont in crown height, commonly attaining hypsodonty; crowns of
molars taper towards base of the crown (shorten anteroposteriorly with wear). Postcranial
featuresintermediate to large rodents; skeletons generally robust to massive, developing
fossorial or semiaquatic adaptations in some lineages.

Subfamily AGNOTOCASTORINAE Korth and Emry, 1997

Definition: Retained primitive features (shared with Eutypomyidae)Complex
occlusal pattern of cheek teeth; upper tooth rows parallel; premolars subequal in size
to molars; retained stapedial foramen; posterior palatine foramen within palatine bone;
length of incisive foramen more than 30% total length of upper diastema; P3 or dP3
retained; and palatal surface smooth. Derived castorid featurescheek teeth crowns taper
toward base (shorten with wear) and are at least mesodont in crown height; promi-
nent digastric process on mandible. Probable autapomorphic featureselongated rostrum;
procumbant incisors.

Tribe AGNOTOCASTORINI Korth and Emry, 1997

Definition: Agnotocastorines with simpler occlusal pattern on cheek teeth than in
Anchitheriomyini (fewer crenulations); smooth enamel surface on incisors.
Included genera: Hystricops Leidy, 1856; Agnotocastor Stirton, 1935; Neatocastor
Korth, 1996.
Range: Latest Eocene (Chadronian) to late Miocene (Hemphillian) of North Amer-
ica, and middle Oligocene of Asia.

Tribe ANCHITHERIOMYINI Korth, 2001a

Definition: Agnotocastorines with beginnings of posterior divergence of upper tooth
rows; shortening of nasal bones; greater complexity of occlusal pattern of cheek teeth than
in Agnotocastorini (greater number of crenulations or fossettes); loss of P3 except in most
primitive species, but retention of dP3 (lost in adult specimens); posterior maxillary notch
rather than foramen; shortening of incisive foramen (as short as 12% of total length of
upper diastema); ridged anterior surface of incisors on all but most primitive genera; most
advanced genera with zig-zag arrangement of mandibular condyles.
Included genera: Anchitheriomys Roger, 1898; Propalaeocastor Borisoglebskaya,
1967; Oligotheriomys Korth, 1998b.
Range: Oligocene (Orellan) and early to middle Miocene (Hemingfordian to Barsto-
vian) of North America, middle to late Oligocene and early late Miocene of Asia, and
middle Miocene (MN 5-8) of Europe.
Discussion: The earliest occurring and most primitive group of beavers (Castoridae)
are the agnotocastorines, which can be readily derived from a Eutypomys-like ancestor
(Korth, 1996, 2000). The remaining three subfamilies of castorids are united by a num-
ber of features that appear to be derived over the condition in agnotocastorines, which is
generally shared with eutypomyids (Korth and Emry, 1997). None of the currently known
agnotocastorines is morphologically transitional between eutypomyids and the other cas-
torids; however, it is likely that an early agnotocastorine was ancestral to later castorids.
Within the agnotocastorines there are two separable groups of genera. Agnotocas-
tor, Neatocastor, and Hystricops have generally similar skulls and less complex occlusal
pattern of the cheek teeth (Stirton, 1935; Wahlert, 1977; Korth, 1996, 1998a). These
Systematics of the Beavers 287

genera are referred to here as the Tribe Agnotocastorini. Oligotheriomys and the Asian
Propalaeocastor are similar to Anchitheriomys in the more complex occlusal patterns of
the cheek teeth (Borisoglebskaya, 1967; Lyschev, 1970; Hugueney, 1975; Korth, 1998b).
These latter genera also differ from the other agnotocastorines in the presence of a poste-
rior maxillary notch instead of a foramen as in other castorids, and the upper tooth rows
diverge slightly posteriorly (Korth and Emry, 1997; Korth, 1998b, 2001a). The other cra-
nial features that separate Anchitheriomys from the agnotocastorines cannot be determined
on the basis of known material of either Oligotheriomys or Propalaeocastor. However,
it appears that these three genera represent a distinct clade within the Agnotocastorinae,
recognized here as the Tribe Anchitheriomyini. Species of Anchitheriomys also share the
condition of the zig-zag alignment of the mandibular condyles with some palaeocas-
torines and castoroidines.

Subfamily PALAEOCASTORINAE Martin, 1987

Definition: Synapomorphic features in common with castoroidines and castorines
incisive foramina less than 30% total length of upper diastema; upper tooth rows strongly
divergent posteriorly; loss of stapedial branch of carotid artery (loss of stapedial foramen);
loss of dP3 and P3; P4/ p4 largest of cheek teeth; nasal bones extend posteriorly on dor-
sal rostrum to same point as premaxillaries; posterior palatine foramina within palatine-
maxillary suture; palatal surface grooved. Plesiomorphic featurescheek teeth primitive
(mesodont, rooted, occlusal surface with fossettes rather than flexi); jugal extends ante-
riorly to meet with lacrimal bone. Autapomorphic featuresfossorial adaptations (skull
with shorter rostrum and broader postorbital constriction and neurocranium than other
castorids; forelimbs more robust; tails shortened); anterior enamel surface of incisors
flattened; ascending ramus of mandible more anteriorly placed than in other beavers;
advanced genera develop more medial position of condyle on mandible.
Included genera: Palaeocastor Leidy, 1869; Euhapsis Douglass, 1905; Capacikala
Macdonald, 1963; Fossorcastor Martin, 1987; Pseudopalaeocastor Martin, 1987.
Range: Latest Oligocene (Whitneyan) to earliest Miocene (Arikareean) of North
America.
Discussion: The Whitneyan is the earliest occurrence of a palaeocastorine in the fos-
sil record (Martin, 1987; Korth, 2001b). The entire range of palaeocastorines is from the
Whitneyan through the Arikareean only. During this short time they diversified rapidly,
developing many fossorial adaptations. The palaeocastorines are easily separable from
other beavers based on dental and cranial features. All palaeocastorines have a distinc-
tively flattened anterior surface of the incisor and nearly all of the flexi of the cheek teeth
are closed-off at the margin of the tooth, producing elongated fossettes after only moder-
ate wear, indicating relatively short striae. In all other castorids, including the Castorinae,
the anterior surface of the incisors is always anteriorly convex.
Although some of the palaeocastorines have been included within the Castorinae
(Xu, 1995, 1996; McKenna and Bell, 1997), this does not appear likely. The Castorinae
is clearly an Old World radiation beginning with Steneofiber in the early Oligocene. The
development of derived fossorial adaptations of the skull and flattening of the incisors
of the earliest palaeocastorines clearly eliminate them as possible ancestors of the Cas-
torinae. Early castorines such as Steneofiber have cheek teeth that are similar to those of
palaeocastorines with the formation of fossettes from the flexi in relatively early stages
288 Korth

of wear (short striae). However, in later castorines such as Palaeomys and Castor, the
lingual flexi on the lower cheek teeth and buccal flexi on the upper cheek teeth remain
open to the base of the crown, indicating that these striae run the full height of the tooth.
The skull and skeleton of palaeocastorines are clearly fossorially adapted. In general
appearance, the skull of palaeocastorines is short, low, and posteriorly broadened, typical
of many burrowing rodents such as mylagaulids and bathyergids. The forelimb of the
burrowing beavers is also broadened and robust, another character of other burrowing
rodents, and the tail is often shortened. In Castor both the forelimb and hindlimb are
broadened, most likely as an adaptation for swimming. The tail of Castor consists of
caudal vertebra that are dorsoventrally flattened to form the paddle tail. Hugueney and
Escuillie (1995) demonstrated that the semi-aquatic adaptations of castorines were already
beginning in early Miocene species of the Eurasian Steneofiber.
Xu (1996) argued that the more primitive palaeocastorines (Palaeocastor and
Capacikala) were castorines and the more derived genera (Fossorcastor and Euhapsis)
were closer to the castoroidines. However, this was all based on the development of the
zig-zag pattern of the processes of the mandible and position of the infraorbital fora-
men on the rostrum. Clearly, the early stages of the medial positioning of the mandibular
condyle are present in the mandibles of Palaeocastor and Capacikala (Xu, 1995:figs. 5-
1, 5-2), simply indicating that the specialized condition of the mandibular processes was
achieved within the palaeocastorines, beginning with the primitive condition (see Martin,
1987:fig. 12). Similarly, the primitive position of the infraorbital foramen in beavers is
low on the side of the rostrum as in Castor and primitive palaeocastorines. In derived
palaeocastorines and some castoroidines, it is much higher on the side of the rostrum.
Clearly, this derived condition is attained separately within the palaeocastorines and cas-
toroidines. Palaeocastor and Capacikala are united with the other palaeocastorines based
on the specializations of the skull and skeleton (fossorial changes) and incisors (flattened
anterior surface), whereas they retain the earlier, more primitive condition of the mandible
and infraorbital foramen.
The less derived palaeocastorines are neither ancestral to, nor should be included
in, the Castorinae, because the two cranial features supposedly uniting them (pattern of
mandibular processes and position of the infraorbital foramen) are primitive conditions
for all castorids.
Castoroidines and palaeocastorines share the zig-zag pattern of mandibular pro-
cesses. Xu (1995) noted that there were significant differences in the development of this
alignment between the condition in palaeocastorines and castoroidines, although he later
(Xu, 1996) suggested that the more derived palaeocastorines were closely related to the
castoroidines. The use of alignment of mandibular processes to unite the palaeocastorines
and castoroidines is not substantiated. It appears that this specialization arose three times
within the Castoridae (Castoroidinae, Palaeocastorinae, Anchitheriomys). Since there are
no other shared-derived features that would unite the palaeocastorines with the cas-
toroidines, it is likely that this character was achieved in parallel in more than one sub-
family.
Martin (1987) divided the palaeocastorines into three tribes. This was reflected in
Korths (1994) classification of the North American beavers (Table I). No attempt to
justify separate tribes of beavers is offered here; Martins (1987) definitions of the tribes
will be followed without examination.
Systematics of the Beavers 289

Xu (1996) named a new genus and species of palaeocastorine, Nannasfiber ostella-

tus. However, it was based on a previously named species, Steneofiber barbouri Peterson
(1905), which was listed as a synonym and not recognized elsewhere in his review of
the Oligocene beavers. Although Xu (1996) selected a different holotype, the trivial name
he assigned is invalid and Petersons barbouri has priority. Martin (1987) named a new
genus Pseudopalaeocastor to include only P. barbouri based on Petersons (1905) type
material. Therefore, Pseudopalaeocastor has priority over Nannasfiber because the latter
is based on the same material as the previously named genus, and Nannasfiber ostellatus
is a junior synonym of Pseudopalaeocastor barbouri. The additional species of Nannas-
fiber described by Xu (1996), N. osmagnus, should be transferred to Pseudopalaeocastor.
Subfamily CASTOROIDINAE ALLEN, 1877
Definition: Synapomorphic features in common with castorines and palaeocasto-
rinesincisive foramina less than 30% total length of upper diastema; upper tooth rows
strongly divergent posteriorly; loss of stapedial branch of carotid artery (loss of stape-
dial foramen); loss of dP3 and P3; premolars largest of cheek teeth; nasal bones extend
posteriorly on dorsal rostrum to same point as premaxillaries; posterior palatine foram-
ina within palatine-maxillary suture; palatal surface grooved. Synapomorphic feature in
common with castorinescheek teeth progressively hypsodont (loss of roots). Plesiomor-
phic featuresskeleton massive but unspecialized (no aquatic or fossorial adaptations).
Autapomorphic featuresjugal does not contact lacrimal bone; condyle of mandible more
medially placed than coronoid or angle (zig-zag pattern); marked increase in size (largest
of castorids).

Tribe CASTOROIDINI Allen, 1877

Definition: S-pattern of molars (elongation of one lingual and one buccal flexus)
ultimately resulting in the separation of molars into three enamel plates.
Included genera: Monosaulax Stirton, 1935; Eucastor Leidy, 1858; Dipoides
Schlosser, 1902; Castoroides Foster, 1838; Procastoroides Barbour and Schultz, 1937.
Range: Early Miocene (Hemingfordian) to Pleistocene of North America, late
Miocene (MN 11) to Pliocene of Europe, and Pliocene of Asia.
Tribe TROGONTHERIINI Lyschev, 1973
Definition: Castoroidines with elongation of M3 (anteroposteriorly longer than ante-
rior molars); cheek teeth retain minute fossettes along with longer buccal and lingual flexi
(no S-pattern); elongated flexi anteriorly concave.
Included genera: Trogontherium Fischer de Waldheim, 1809; Boreofiber Radulesco
and Samson, 1972; Euroxenomys Samson and Radulesco, 1973; Youngofiber Chow and
Li, 1978; Asiacastor Lyschev, 1971.
Range: Early to middle Miocene (Hemingfordian to early Barstovian) of North
America, early to late Miocene (MN 4 to 13) of Europe, and late Oligocene to Pleis-
tocene of Asia.
Discussion: The earliest castoroidine, Monosaulax, is known from the Hemingfor-
dian and Barstovian of North America [see Korth (1999) for review of species]. In North
America, the sequence of Monosaulax-Eucastor-Dipoides-Castoroides (from earliest and
most primitive to latest and most derived), recognized by Stirton (1935), still appears rel-
evant. This lineage would also include the migration of Dipoides to Eurasia in the later
290 Korth

Tertiary. In this clade there is an increase in size and development of the S-pattern on
the cheek teeth through time.
The lineage of Eurasian castoroidines, ending in Trogontherium, also appears to have
originated in North America. The earliest member of this clade is Euroxenomys. It was
first named for Trogontherium minutus from the Miocene of Europe (Samson and Rad-
ulesco, 1973). Many later authors have not recognized this genus and have maintained
it in Trogontherium (Savage and Russell, 1983; McKenna and Bell, 1997). However, the
size, relative development of M3, lack of ridges on the incisors, and shorter flexi on the
cheek teeth of T. minutus (Stromer, 1928:pl.2; Engesser, 1972:abb. 64) are clearly more
primitive than the conditions in Plio-Pleistocene specimens of Trogontherium. Therefore,
the Miocene species that appear ancestral to later Trogontherium are referable to a dis-
tinct genus. Stefen (1997) has even suggested that the European species of Euroxenomys
be retained as Steneofiber minutus because of its great similarity to other species of the
latter genus. Most recently, Hugueney (1999) has included Euroxenomys as a subgenus
of Trogontherium.
In North America, there are two species of Euroxenomys from the Hemingfordian
and early Barstovian, respectively, that are generically separable from the contempora-
neous North American species of Monosaulax (Sutton and Korth, 1995). However, these
differences (lengths of the striae on P4, relative length of M3, crown height) in the Eurox-
enomys dental pattern are as easily derived from that of a primitive Monosaulax as from
Steneofiber, suggesting the possibility of a North American origin for the trogontheri-
ine castoroids. In Asia, Youngofiber occurs in the early part of the Miocene and is more
primitive than later trogontheriines (prominent roots and shorter flexi on cheek teeth).
It is likely that the later Miocene Asiacastor is directly derived from Youngofiber and
represents a separate clade of trogontheriines.
The genera that have the castoroidine-like alignment of the mandibular processes
include those North American genera traditionally included in the Castoroidinae (Mono-
saulax, Eucastor, Dipoides, Castoroides) but also the large Eurasian beavers such as Tro-
gontherium, Youngofiber, and Asiacastor. All of these genera have the derived morphol-
ogy of the jugal and of the mandibular processes. North American castoroidines develop
a distinctive S-pattern on the occlusal surface of the molars that is produced by the elon-
gation of two flexi (one buccal, one lingual) and the elimination of all others. Ultimately,
these flexi completely cross the surface of the tooth, dividing it into three separate sec-
tions that are flattened enamel rings surrounding dentine with layers of cement in between
the enamel rings. The Eurasian castoroidines similarly elongate two flexi on each molar,
but retain minute fossettes in all but the most derived species, and the flexi are anteri-
orly concave. They do not develop the separate enamel plates seen in the molars of North
American castoroidines. M3 of Trogontherium and the other Eurasian castoroidines is also
anteroposteriorly elongated, having a greater length than the anterior molars. Clearly the
Eurasian and North American castoroidines can be divided on the basis of these dental
characters.
Some authors have argued that the primitive Hemingfordian to Barstovian castoroi-
dine Monosaulax was synonymous with the Barstovian to Clarendonian Eucastor (Stout,
in Skinner and Taylor, 1967; Xu, 1994, 1995; McKenna and Bell, 1997), but these genera
are clearly separable based on dental and cranial criteria (Korth, 1999). In their recent
classification, McKenna and Bell (1997) listed two questionable Pliocene genera, Eocas-
Systematics of the Beavers 291

toroides Hibbard (1937) and Paradipoides Rinker and Hibbard (1952). The former was
listed as a castorine and the latter as a castoroidine. However, both of these genera appear
to be junior synonyms of previously named castoroidines. Kurten and Anderson (1980)
list the type (and only) species of Eocastoroides as a junior synonym of the type species
of Procastoroides, P. sweeti Barbour and Schultz (1937). Martin (1969) and Zakrzweski
(1969) both have questioned the validity of Paradipoides. Based on the arguments of the
latter authors, it appears that Paradipoides should be included as a junior synonym of
Dipoides.

Subfamily CASTORINAE Hemprich, 1820

Definition: Synapomorphic features in common with castoroidines and palaeocasto-
rinesincisive foramina less than 30% total length of upper diastema; upper tooth rows
strongly divergent posteriorly; loss of stapedial branch of carotid artery (loss of stapedial
foramen); loss of dP3 and P3; posterior premolars largest of cheek teeth; nasal bones
extend posteriorly on dorsal rostrum to same point as premaxillaries; posterior palatine
foramina within palatine-maxillary suture; palatal surface grooved. Synapomorphic fea-
ture in common with castoroidinescheek teeth progressively hypsodont (loss of roots).
Plesiomorphic featuresmandibular processes (coronoid, condyle, angle) form straight
line; jugal extends anteriorly to meet the lacrimal bone; rounded to semi-flattened anterior
surface of incisors. Autapomorphic featuresflexi on cheek teeth progressively remain
open to base of crown (three lingual flexids remain open on lower cheek teeth, three
buccal flexi on upper cheek teeth) indicating longer striiae, but extend just slightly more
than half the width of the occlusal surface of the tooth; advanced species show special-
ization of the postcranial skeleton for swimming (flattened caudal vertebrae, broadened
limb bones and digits).
Included genera: Castor Linnaeus, 1758; Palaeomys Kaup, 1832 or Chalicomys
Kaup, 1832; Steneofiber Geoffroy Saint-Hilaire, 1833; Zamolxifiber Radulesco and Sam-
son, 1967; Romanofiber Radulesco and Samson, 1967; Schreuderia Aldana Carrasco,
1992.
Range: Oligocene to Recent of Europe, late Miocene (Hemphillian) to Recent of
North America, and latest Miocene to Recent of Asia.
Discussion: The earliest castorine is Steneofiber from the Oligocene of Europe. The
cheek teeth of this earliest genus are not specialized (flexi close to form fossettes), and
the skull shows the primitive condition of the jugal and alignment of the mandibular
processes. Progressively, in the Steneofiber-Palaeomys-Castor clade, the flexi (lingual on
lower cheek teeth, buccal on upper cheek teeth) remain open through the entire height
of the cheek teeth. This same progression of genera was recognized by Stirton (1935).
The remaining later Tertiary genera of European castorines (Schreuderia, Romanofiber,
Zamolxifiber) appear to be late aberrant side branches of the Castorinae.
There appears to be some discrepancy in the use of the genus name Chalicomys Kaup
(1832) and Palaeomys Kaup (1832). The majority of European authors use the former (see
Hugueney, 1999), whereas North American authors use Palaeomys (McKenna and Bell,
1997). Any reference to Palaeomys in this text recognizes its synonymy with Chalicomys
and is not necessarily a judgment of the validity of using one genus name over the other.
Schreuderia Aldana Carrasco (1992) has the occlusal morphology of the cheek teeth
of a castorine (three lingual flexi remain open and do not exceed half the occlusal width of
292 Korth

the tooth). The depth of the lingual striids on the cheek teeth of Schreuderia is greater than
those of Steneofiber, but they do not reach the base of the crowns as in Castor; similar to
that of Palaeomys. The dental morphology makes Schreuderia clearly a castorine rather
than a castoroidine. Hugueney (1999) listed Schreuderia as a subgenus of ?Eucastor.
However, the figures provided by Hugueney (1999:fig. 28.8) show a primitive castorine
pattern rather than that of a castoroidine. It is more likely that Schreuderia is a subgenus
of Palaeomys (or Chalicomys) rather than Eucastor.
Two additional genera of Pliocene beavers from Europe, Romanofiber and Zamolxi-
fiber (Radulesco and Samson, 1967) were listed as members of the Castoroidinae by
McKenna and Bell (1997). They are characterized by a simple Steneofiber-like castorine
dentition, although they occur at a much later time than species of the latter. Romanofiber
and Zamolxifiber are nearly identical in dental morphology, having very short lingual stri-
ids on p4 that are eliminated early in wear, giving a primitive Steneofiber-like occlusal
morphology, and differing from the latter only in having higher crowned cheek teeth.
The two respective type species are nearly identical except for a slight difference in size
and depth of the mesostriid on p4. It is likely that these two genera may represent a sin-
gle genus and ultimately may be recognized as synonyms. Further examination of the
mandibular morphology of referred specimens will also assist in the subfamilial assign-
ment of these genera.

DISCUSSION
Phylogeny and Biogeography
The suggested phylogenetic relationships of the Castoridae, based on the cranial,
dental, and skeletal characters discussed above, are presented in Fig. 4. The most prim-
itive group, Agnotocastorinae, retains many primitive eutypomyine characters that sep-
arate them from the remainder of the beavers. Within the agnotocastorines, two groups
are recognizable, the earliest and most primitive Agnotocastorini and the larger, later, and
more specialized Anchitheriomyini that develop convergently some of the characters of
the more advanced beavers (divergence of upper tooth rows, shortening of incisive foram-
ina, enlargement of premolars). It is likely that the later castorids were derived from an
agnotocastorine. If this is the case, this ancestor would more likely be from the more
generalized Agnotocastorini rather than an anchitheriomyine.
Agnotocastorines appear nearly simultaneously in both North America and Asia
(Savage and Russell, 1983; Korth, 1994; Xu, 1994). It appears likely that the Anchitheri-
omyini first developed in North America, were introduced into Eurasia, then reintroduced
into North America in the Miocene (Korth, 1998b). The more continuous record of Agno-
tocastorini in North America suggests that the possible ancestor of later subfamilies was
also North American.
The suite of derived characters that separates the remainder of the castorids from
the agnotocastorines (Fig. 4, node 5) is consistent with the remaining three recognized
subfamilies, Palaeocastorinae, Castorinae, and Castoroidinae. Here, the latter two sub-
families are grouped together (node 7) because of the generally more hypsodont cheek
teeth. However, this may be a parallel development in these two groups, thus making
the separation of the Palaeocastorinae somewhat artificial. It may be possible to consider
Systematics of the Beavers 293

Fig. 4. Phylogenetic relationships of the Castoridae above the genus level. Explanation of nodes: 1)
Castoroideasciuromorphous zygomasseteric structure; postorbital process lacking; sciurognathy; interor-
bital foramen posterior to optic foramen; dorsal palatine foramen separate from sphenopalatine foramen and
within maxilla or maxillary-suture; incisor microstructure uniserial; dental formula I11 C00 P(2)11 M33. 2)
Castoridaeskull generally low, broad, and robust with broad zygomata; infraorbital foramen low on rostrum;
digastric (symphyseal) eminence on mandible; less complex occlusal pattern of cheek teeth, forming series of
transverse or oblique fossettes of enamel; cheek teeth at least mesodont in crown height, commonly attain-
ing hypsodonty; crowns of molars taper towards base of the crown (shorten anteroposteriorly with wear). 3)
Agnotocastorinaecomplex occlusal pattern of cheek teeth; upper tooth rows parallel; premolars subequal in
size to molars; retained stapedial foramen; posterior palatine foramen within palatine bone; elongated rostrum;
P3 or dP3 retained; and palatal surface smooth. 4) Anchitheriomyinibeginnings of posterior divergence of
upper tooth rows; shortening of nasal bones; greater complexity of occlusal pattern of cheek teeth; loss of
P3; posterior maxillary notch rather than foramen; shortening of incisive foramen; ridged anterior surface of
incisors. 5) unnamed nodedecreased complexity of the occlusal pattern of cheek teeth (with smooth-sided
re-entrant valleys and fossettes); posteriorly divergent upper tooth rows; loss of P3 and dP3; enlargement of last
premolars (relative to molars); loss of the stapedial foramen; parallel grooves on the palatal surface; shortening
of the incisive foramina (less than 30% length of upper diastema); anterior position of the posterior palatine
foramina (within the palatine-maxillary suture). 6) Palaeocastorinaefossorial adaptations of skull and skeleton
(low, broad skull, hypertrophied forelimbs, shortened tail); flattened anterior surface of incisors; development
of zig-zag alignment of mandibular processes. 7) unnamed nodeincreased hypsodonty of cheek teeth (greater
than mesodont). 8) Castoroidinaejugal does not contact lacrimal; zig-zag alignment of mandibular processes;
largest species of the family; 9) Trogontheriinielongation of M3; reentrants on cheek teeth lengthen, but do
not attain S-pattern. 10) CastoroidiniS-pattern on occlusal surface of molars (two deep re-entrants crossing
at least half the width of the tooth, ultimately completely cross molars). 11) Castorinaeprogressively, all re-
entrants on cheek teeth remain open until late in wear; postcranial skeleton adapted for aquatic locomotion
(broadened limb bones; elongated tail with flattened caudal vertebrae).
294 Korth

this node as a tricotomy. Further work, possibly on postcranial material, should allow for
a better examination of the relationships of the palaeocastorines with the castorines and
castoroidines.
The Palaeocastorinae is a short-lived group that is limited geographically to North
America. It appears to be an aberrant side-branch of the castorids, not ancestral to either
the castorines or castoroidines.
The most diverse group of castorids is the Castoroidinae (Fig. 4, node 8). There
are two groups of giant beavers recognizable in this clade, the predominantly North
American Castoroidini (Fig. 4, node 10) and the predominantly Eurasian Trogontheriini
(Fig. 4, node 9). Both groups appear to have originated in North America at or near the
beginning of the Miocene (Hemingfordian). The trogontheriines migrated almost immedi-
ately to Eurasia and continued to diversify there (Xu, 1994). The Castoroidini diversified
throughout the remainder of the Tertiary in North America with Dipoides emigrating to
Eurasia in the later Miocene.
The large size of the individuals of both tribes of castoroidines appears to have
been attained separately on different continents. The earliest and most primitive (small-
est) species of each lineage are clearly distinguishable from one another at their first
occurrence (Sutton and Korth, 1995), suggesting this parallel development in size.
The dental pattern of the castorines is distinctive and can be traced from the earliest
species to Recent genera (increased length of striae on cheek teeth). However, the cranial
morphology of this subfamily is persistently primitive. The first castorine to appear in
North America is Castor in the late Miocene [Hemphillian (Savage and Russell, 1983)].

ACKNOWLEDGMENTS
The author thanks J. H. Wahlert of the American Museum of Natural History and N.
Rybczynski of Duke University for helpful discussions about castorid morphology and
relationships. This paper was critically reviewed by the editors of the journal and two
anonymous reviewers.

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