• TABLE OF CONTENTS
HIDE
 Front Cover
 Copyright
 Table of Contents
 Introduction
 Acknowledgement
 Materials and methods
 Cranial osteology
 Fossil identification by cranial...
 Incisor enamel
 Auditory ossicles
 Postcranial osteology
 Systematics
 Origin of Erethizon
 Appenix A
 Appenix B
 Literature cited
 Back Cover






Group Title: Bulletin of the Florida State Museum
Title: A Revision of the fossil Erethizontidae of North America
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00095806/00001
 Material Information
Title: A Revision of the fossil Erethizontidae of North America
Series Title: Bulletin - Florida State Museum ; volume 27, number 1
Physical Description: 76 p. : ill. ; 23 cm.
Language: English
Creator: Frazier, Michael K.
Donor: unknown ( endowment )
Publisher: Florida State Museum, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1982
Copyright Date: 1982
 Subjects
Subject: Porcupines, Fossil   ( lcsh )
Paleontology -- North America   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: p. 71-76.
Statement of Responsibility: Michael K. Frazier.
 Record Information
Bibliographic ID: UF00095806
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 09643468
lccn - 82623268

Table of Contents
    Front Cover
        Front cover
    Copyright
        Page i
    Table of Contents
        Page 1
    Introduction
        Page 2
    Acknowledgement
        Page 3
    Materials and methods
        Page 4
        Page 5
        Page 6
        Page 7
    Cranial osteology
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
    Fossil identification by cranial measurements
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
    Incisor enamel
        Page 26
        Page 27
    Auditory ossicles
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
    Postcranial osteology
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
    Systematics
        Page 40
        Page 41
        Page 42
        Page 43
        Page 44
        Page 45
        Page 46
    Origin of Erethizon
        Page 47
        Page 48
        Page 49
        Page 50
    Appenix A
        Page 51
    Appenix B
        Page 52
        Page 53
        Page 54
        Page 55
        Page 56
        Page 57
        Page 58
        Page 59
        Page 60
        Page 61
        Page 62
        Page 63
        Page 64
        Page 65
        Page 66
        Page 67
        Page 68
        Page 69
        Page 70
    Literature cited
        Page 71
        Page 72
        Page 73
        Page 74
        Page 75
        Page 76
        Page 77
    Back Cover
        Page 78
Full Text









of the
FLORIDA STATE MUSEUM
Biological Sciences


1981


Number 1


A REVISION OF THE FOSSIL ERETHIZONTIDAE
OF NORTH AMERICA



MICHAEL K. FRAZIER


UNIVERSITY OF FLORIDA


Volume 27


GAINESVILLE







Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES,
are published at irregular intervals. Volumes contain varying numbers of pages and are not
necessarily completed in any one calendar year.








OLIVER L. AUSTIN, JR., Editor

RHODA J. BRYANT, Managing Editor

Consultants for this issue:

ROBERT WILSON

CHARLES A. WOODS
















Communications concerning purchase or exchange of the publications and all manuscripts
should be addressed to: Managing Editor, Bulletin; Florida State Museum; University of
Florida; Gainesville, Florida 32611.
Copyright 1982 by the Florida State Museum of the University of Florida

This public document was promulgated at an annual cost of $2028.00 or
$2.028 per copy. It makes available to libraries, scholars, and all interested
persons the results of researches in the natural sciences, emphasizing the
circum-Caribbean region.


Publication date: 5 January 1982


Price: $2.05








A REVISION OF THE FOSSIL ERETHIZONTIDAE OF
NORTH AMERICA


MICHAEL K. FRAZIER1


SYNOPSIS: The North American fossil record of porcupines (Erethizontidae) is reviewed. Com-
parisons of cranial osteology, incisor enamel, auditory ossicles, and post-cranial elements of
fossil samples with Recent Coendou and Erethizon samples indicate that all of the fossil
porcupines in North America are congeneric with Erethizon. The following four species of
Erethizon are recognized in the fossil record of North America: (1) E. bathygnathum appeared
approximately 2.5 million years ago and lived in western North America during late Blancan
and early Irvingtonian times. (2) E. cascoensis is recognized only in the early Irvingtonian El
Casco local fauna of California. (3) A new species of Erethizon is known only from the
early-middle Irvingtonian of Florida. (4) The extant E. dorsatum first appears in the United
States during middle Irvingtonian times, later dispersing throughout most of temperal North
America. Erethizon apparently differentiated from an ancestral form in South America prior to
its appearance in North America during late Pliocene times. Even the earliest fossils already
possessed the enlarged masticatory apparatus designed to consume the bark of temperate
conifers. Other functional specializations in the skeleton of Coendou and Erethizon are consid-
ered.




TABLE OF CONTENTS

INTRODUCTION ............. ................................................ 3
ACKNOW LEDGEMENTS .................................... .. ............. 5
MATERIALS AND METHODS ................................................. 6
NORTH AMERICAN DISTRIBUTION .. ........................................ 9
CRANIAL OSTEOLOGY ............... .................................... 13
FOSSIL IDENTIFICATION BY CRANIAL MEASUREMENTS ..................... 25
INCISOR ENAMEL ........................................................ .. 31
AUDITORY OSSICLES ........................................................ 35
POSTCRANIAL OSTEOLOGY ............... ............................... 40
SYSTEMATICS ............................................................. 45
ORIGIN OF ERETHIZON ...................................................... 53
APPENDIX A Pleistocene Faunas That Contain Porcupines .......................... 56
APPENDIX B Graphs of Cranial Measurements ................................ 58
LITERATURE CITED ......................................................... 77





'The author is Associate Curator of Paleontology at the Mississippi Museum of Natural Science, 111 North Jefferson Street,
Jackson 39202. This paper was submitted in partial fulfillment for the degree of Master of Science at the University of
Florida, Gainesville.


FRAZIER, MICHAEL K. 1981. A revision of the fossil Erethizontidae of North America.
Bull. Florida State Mus., Biol. Sci. 27(1):0-00.







BULLETIN FLORIDA STATE MUSEUM


INTRODUCTION

The oldest known erethizontids appear in the late Oligocene Deseaden
sediments of Patagonia and Bolivia, along with early records of other cavio-
morphs (Simpson 1950; Hoffstetter and Lavocat 1970). The origin of these
early South American caviomorphs has long been a topic of controversy
among rodent systematists. One school of thought favors an origin from an
African phiomorph, via rafting across a narrower Atlantic Ocean during the
Eocene (Hoffstetter and Lavocat 1970; Lavocat 1974; Raven and Axelrod
1975). Another school supports a North American ancestry from a primitive
Eocene hystricognathous rodent, possibly a franimorph (Wood and Patter-
son 1970; Wood 1974; Wahlert 1973). Presumably these ancestral popu-
lations arrived in South America via island hopping across the Eocene
Antillean Archipelago (Wood 1977). Wood (1975, 1977) has presented evi-
dence favoring the parallel evolution of the African and South American
hystricomorph rodents. Yet another school adds another dimension. Hus-
sain et al. (1978) suggests the caviomorphs are derived from the Chapat-
timyidae, a Southeast Asian late Eocene rodent group.
Numerous fossil erethizontid genera have been described from Tertiary
deposits in South America, all having the same basic tooth structure as
their living representatives (Patterson 1958; Fields 1957). The erethizon-
tids were restricted to the South American continent until their late
Pliocene dispersal across Middle America (Simpson 1950). At that time
they came north during the major faunal interchange between North and
South America (Webb 1976).
Fossil porcupines appear in North America during the late Pliocene, less
than three million years ago. There is little doubt that they are of South
American origin, where three of the four living genera still occur. This
study concentrates only on the two North American genera, Coendou and
Erethizon. Less is known about the other two genera, Chaetomys and
Echinoprocta, which occur exclusively in South America (Anderson and
Jones, 1967; Walker 1975). Chaetomys may even be an echymyid, based
upon the nature of the P4 (C.A. Woods, pers. comm.).
Erethizon lives exclusively in temperate and boreal North America, pre-
ferring wooded areas of conifers, junipers, and poplars, where it feeds
primarily upon the cambium layer of bark. It now ranges from northern
Mexico northward into the major portion of Canada and Alaska (Woods
1974). Erethizon is the only caviomorph rodent now occupying temperate
and boreal North America. Coendou inhabits lowland rain forests of the
Neotropical Realm from southern Mexico (20 latitude) to northern Argen-
tina, where it feeds upon leaves, tender stems, and fruit (Walker 1975).
The two genera are geographically separated by a distance of about 1000
km.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


Coendou and Erethizon show gross morphological differences. Coendou
has a quill-less, prehensile tail, a well developed prehallux in the hind foot,
and a masticatory apparatus designed for the consumption of soft tropical
vegetation. However some species of Coendou do not exhibit the pre-
hensile tail (Karl Koopman, pers. comm.). Erethizon, on the other hand,
has a short, stubby tail bearing quills, a vestigial prehallux, and a masti-
catory apparatus suited for the consumption of the much tougher tem-
perate vegetation, particularly the bark of conifers and hardwoods. In a
revision of the North American Erethizontidae, White (1970) presented
morphological evidence for referring the fossil porcupines from Grand
View (Coendou bathygnathum), Vallecito Creek (C. stirtoni), El Casco (C.
cascoensis), Cumberland Cave (C. cumberlandicus), and other localities to
the genus Coendou. He considered Coendou ancestral to Erethizon, with
the middle Irvingtonian Cumberland Cave sample from Maryland being
intermediate between the two genera. Discovery of an excellent sample of
Erethizon-like porcupines from the early Irvingtonian Inglis IA locality in
Florida prompted this further review of the fossil porcupines of North
America.
The principal goals of this study are (1) to decide whether the fossil
porcupines of North America represent Coendou, Erethizon, or both
genera, (2) to trace their stratigraphic and geographic distribution through
time, and (3) to outline any evolutionary trends that may be observed in
these North American erethizontids.


ACKNOWLEDGEMENTS

Specimens were made available for study by the following institutions: Academy of Natural
Sciences at Philadelphia (ANSP), American Museum of Natural History (AMNH or F:AM),
University of Arizona (UALP), Carnegie Museum of Natural History (CM), Florida State
Museum (UF), Imperial Valley College Museum (IVCM), University of Kansas Museum of
Natural History (KU), Los Angeles County Museum of Natural History (LACM), University of
Michigan Museum of Paleontology (UMMP), National Museum of Natural History, Smith-
sonian Institution (NMNH), University of Oregon Museum of Natural History (UO), and
Texas Memorial Museum, University of Texas (TMM). I am grateful to the people responsible
for the loan of these materials.
I express deep appreciation to S. David Webb, my supervisory chairman, for the pre-
liminary review of this thesis and his helpful comments and guidance through the course of
this work. I also wish to thank the other members of my supervisory committee, Pierce
Brodkorb, David Nicol, and Ronald G. Wolff, for helpful criticism of the manuscript and their
assistance during the last three years.
Valuable fossil specimens were donated by Allen Davis, Frank Garcia, Margaret C. Tho-
mas, and John S. Waldrop (Timberlane Research Organization). Mrs. Steven Kruger of Seff-
ner, Florida, loaned several porcupine mandibles from Apollo Beach and also allowed them to
be cast. Raymond Giron (Sant Fe Community College Zoo) provided a Coendou for wet
study. Jerry Duvendeck (Michigan Department of Natural Resources) and David Steadman
provided several specimens of Erethizon. Tim Breen and Robert Smidt provided valuable
assistance in the statistical analysis of the cranial measurements. John A. White and Charles







BULLETIN FLORIDA STATE MUSEUM


A. Woods helped with their discussions concerning the evolution and morphology of erethi-
zontids. Charles A. Repenning added valuable suggestions concerning Pleistocene strati-
graphy and zoogeography. Discussions with friends and fellow graduate students Jon Baskin,
David Frailey, Richard Franz, Andrew Homner, Jean Klein, John Meeder, Gary Morgan,
David Steadman, and John Waldrop added much to my thought processes. Additional thanks
go to Dr. Wolff and Robert Allen for their photographic assistance and to Chandra Aulsbrook
and Nancy Halliday for help with the figures. Special thanks go to Mr. and Mrs. Henry
Danker for the donation of the scanning electron microscope to the Department of Geology.
Robert Pierce provided assistance with the use of the microscope. Warm and special gratitude
goes to all the people who were not mentioned, but were helpful in many ways during the
course of this study.

MATERIALS AND METHODS

Cranial measurements of numerous Recent skulls and mandibles of Coendou and Erethizon
as well as all available fossil materials were taken in millimeters using a dial caliper to the
nearest tenth. On each Recent specimen 46 measurements were taken, of which the most
diagnostic are presented in Figures 1 and 2. On Recent specimens where a measurement
could not be taken, the sample mean of that measurement was substituted. The Recent
samples (over 100 skulls per genus) included males and females, adults and subadults, and
sampled the entire geographic range of each genus. Individuals with deciduous premolars
were considered subadults. The skulls and skeletons used were loaned from the mammalogy
collections of the American Museum of Natural History, National Museum of Natural History
(Smithsonian Institution), and the Florida State Museum.
A principal component analysis, analysis of variance, cluster analysis, and discriminant
analysis were performed upon the cranial measurements using the Statistical Analysis System
(SAS'76) (Barr et al. 1976). SAS'76 also produced the graphs of the cranial measurements of
Recent specimens (Appendix B).
Sagittal sections of lower incisor enamel of Coendou and Erethizon and fossil specimens
were studied using an ISI Super Mini-SEM II scanning electron microscope. The preparation
of the specimens followed the procedures described by Flynn and Wahlert (1978), Measure-
ments taken from the enamel photomicrographs follow Escala and GAllego (1977) and Wahlert
(1968). The preparations of Recent and fossil auditory bullae were accomplished using an S. S.
White Airbrasive jet machining unit.


NORTH AMERICAN DISTRIBUTION
The earliest porcupines in North America appear in late Pliocene (Blan-
can) and early Pleistocene (Irvingtonian) faunas from Grand View, Idaho,
Vallecito Creek and El Casco in southern California, and Wolf Ranch in the
San Pedro Valley of Arizona. These records all occur in the western United
States. Later records are known from other localities throughout North
America.
Stratigraphic correlation of these early records has been refined and
simplified by radiometric dating and magnetic polarity stratigraphy in the
Anza Borrego section of California (Opdyke et al. 1977), the San Pedro
Valley of Arizona (Johnson et al. 1975; Lindsay et al. 1975), the Pearlette
type ashes of the Midwest (Boellstorff 1973; Zakrzewski 1975a), and the
Grand View section of Idaho (Neville et al. 1979). Other correlations in this


Vol. 27 No. 1














\I 'I

3z


2

5r









SB C


FIGURE 1. Key to cranial measurements. (1) width of rostrum, (2) width of frontals, (3) width
of palate between P4 alveoli, (4) width of palate between M3 alveoli, (5) width at xygomatics,
(6) width across auditory meatuses, (7) width across occipital condyles, (8) medial width of U1
rostrum, and (9) width at infraorbital foramen.







BULLETIN FLORIDA STATE MUSEUM


A


FIGURE 2. Key to cranial measurements. (10) depth of rostrum, (11) length of auditory bulla,
(12) length of rostrum, (13) alveolar length of upper tooth row, (14) length from upper M3
alveolus to occipital condyle, (15) total skull length, (16) depth of mandible at lower P4, (17)
depth of mandible at lower M2, (18) depth from condlyle to angular process, (19) alveolar
length of lower tooth row, (20) length of diastema, and (21) length of symphysis.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


paper rely on biostratigraphic comparisons of diverse mammalian taxa
(Hibbard et al. 1965; Skinner and Hibbard 1972).
Harrison (1978) considered the Wolf Ranch local fauna to be late Blancan
in age, based upon the fossil mammals. Lindsay et al. (1975) established a
radiometric date for the Wolf Ranch sediments of approximately 2.5 my,
and their paleomagnetic studies establish the time of deposition at just
below the Gauss-Matuyama magnetic polarity boundary. These dates
establish the Wolf Ranch record as the oldest known porcupine in North
America.
The Grand View local fauna, as described by Shotwell (1970) and Wilson
(1933), is considered late Blancan in age by most paleontologists. A radio-
metric date of 1.36 my from the Bruneau Basalt, which is stratigraphically
above the Grand View deposits, provides a minimum age, and a date of 3.5
my for the Hagerman local fauna (Glenns Ferry Formation) below the
Grand View provides a maximum age for the fauna (Evernden et al. 1964).
Neville et al. (1979), using magnetic polarity stratigraphy, assign the Grand
View deposit to the earliest part of the Matuyama polarity epoch, between
1.8 and 2.5 my old. The above information and the occurrence of Boropha-
gus, Trigonictis, and Ondatra idahoensis support a late Blancan age for
the fauna. Thus, the holotype of Erethizon bathygnathum Wilson (1935) is
late Blancan.
The Anza Borrego faunas of southern California were originally studied
by Downs and White (1968), and recently the magnetic polarity stratig-
raphy was correlated with the mammal-bearing section (Opdyke et al.
1977). The stratigraphic occurrence of porcupine in the section coincides
with the appearance of Nothrotherium, Lepus, Microtus, Smilodon, and
?Euceratherium, which are considered early Irvingtonian indicators and
occur in the middle of the Matuyama polarity epoch in the Anza Borrego
section. The age of this record is approximately 1.9 or 1.6 my (Opdyke et
al. 1977).
The El Casco record occurs in a fauna collected in 1923 by Joseph Rak
and Childs Frick (White 1970; Henry Galiano, pers. comm.). As no stratig-
raphic data exist, correlation must rely on biostratigraphic methods. Canis
edwardi, Ondatra idahoensis (advanced form), Odocoileus, and Equus
(Equus) also occur in the El Casco local fauna.
Nelson and Semken (1970) have shown conclusively that a chronocline
exists in Ondatra during the Plio-Pleistocene, and its species are especially
useful biostratigraphic indicators. Ondatra idahoensis appears in the San
Pedro River Valley of Arizona approximately two million years ago (Lindsay
et al. 1975). It is known from the Borchers local fauna of Kansas, dated at
1.9 my (Zakrzewski 1975a), the Grand View local fauna (Wilson 1933; Shot-
well 1970), the White Rock local fauna (Eshleman 1975), the Mullen
assemblage of Nebraska (Martin 1972), and the California Wash and Gidley
Level local faunas from the San Pedro Valley (Johnson et al. 1975). All of







BULLETIN FLORIDA STATE MUSEUM


these records are considered late Blancan or early Irvingtonian.
The other taxa in the El Casco local fauna also indicate an early Pleis-
tocene age. Opdyke et al. (1977), in their study of the Anza Borrego sec-
tion, described the appearance of Odocoileus during the Matuyama polar-
ity epoch (approximately 2.1 my). Beryl Taylor (pers. comm.) considers the
wolf, Canis edwardi, to be an Irvingtonian species. Based upon the dental
development of the lower Ml in Ondatra idahoensis, the presence of the
post-Blancan Equus (Equus) sp. (Charles Repenning, pers. comm.), and
the occurrence of other early Pleistocene taxa, the El Casco local fauna is
considered early Irvingtonian in age (Frazier, MS).
The earliest procupines in the eastern United States occur in the Inglis
IA local fauna of Florida (Klein 1971; Webb 1974, 1976). This faunal
assemblage includes Smilodon gracilis, Chasmaporthetes, Glyptotherium
arizonae, Sigmodon curtisi, Canis edwardi, Platygonus bicalcaratus, Cap-
romeryx arizonensis, and an advanced Ondatra cf. idahoensis. These
mammalian taxa indicate an early Irvingtonian age. The Inglis IA local
fauna compares very closely with the early Irvingtonian Curtis Ranch fauna
as described by Gazin (1942). For example, G. arizonae, C. edwardi, Felis
lacustris, S. curtisi, Ondatra, Lepus, and C. arizonensis are shared by the
two faunas. Based upon these faunal similarities the Inglis IA local fauna
can be considered equivalent in age with the Curtis Ranch fauna, which
Johnson et al. (1975) assign to the Matuyama polarity epoch, approximately
1.9 my (near the Olduvai event).
The correlations of the major North American late Pliocene and early
Pleistocene faunas appear in Figure 3. Erethizontid occurrences are indi-
cated by asterisks.
As chronostratigraphic and paleomagnetic data are missing, the majority
of the middle Irvingtonian and Rancholabrean faunas that contain porcu-
pines must be correlated biostratigraphically. The Cumberland Cave, Port
Kennedy, and Trout Cave local faunas contain assemblages of middle
Irvingtonian arvicoline rodents (Hibbard 1955; Zakrzewski 1975b). The
majority of the Rancholabrean porcupine records were collected from cave
deposits throughout North America. Figure 4 presents the Recent ranges
of Coendou and Erethizon in North America along with selected late
Pliocene through late Pleistocene localities that have produced porcupine
fossils. A complete list of localities in North America is presented in
Appendix A.

CRANIAL OSTEOLOGY

Striking differences exist between the skulls of Erethizon and Coendou.
Most of them are presumably related to the profoundly different modes of
mastication in the two genera. Various cranial characters involved with the


Vol. 27 No. 1








FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


[ reversed normal erethizontid
polarity polarity
FIGURE 3. Correlation Chart of late Pliocene and early Pleistocene local faunas. Mammalian
faunal datum planes are geographically localized. 'Early Pleistocene Wellsch Valley local
fauna, Saskatchewan, Canada (Harington 1978). (Chart modified from Johnson et al. 1975;
Lindsay et al. 1975; Zakrzewski 1975a; Opdyke et al. 1977; Neville et al. 1979.)


masticatory apparatus have been proposed previously for generic deter-
mination of fossil porcupines in North America. White (1968, 1970) pre-
sented the following distinctions:
1) Upper P4 in Erethizon is generally larger than upper Ml, while in
Coendou the two teeth tend to be subequal in size.
2) The tooth rows are widely divergent in Erethizon, whereas they are
subparallel in Coendou.
3) "The incisors of Coendou are essentially orthodont, and the upper
and lower incisors occlude in such a manner as to provide an efficient
cutting mechanism. This is contrasted by the occlusion of the more
proodont incisors of Erethizon, which have less of a cutting and more
of a scraping function." (White 1970:12)
4) "In living Coendou the orbital width, or the greatest distance from
the supraorbital ridge to the zygomatic arch, is less than 75 percent of
the least interorbital constriction, whereas in Erethizon it is greater
than 80 percent." (White 1970:12)
5) "The angular process of the mandible in adults of Erethizon is in-
flected media and the posterolateral surface of the mandible is con-


1981








BULLETIN FLORIDA STATE MUSEUM


FIGURE 4. Erthizontid distribution in North America showing Recent ranges of Erethizon
(stippled) and Coendou (cross-hatched) (after Hall and Kelson 1959). Selected fossil porcupine
localities below correspond with numbers on map: (1) Vallecito Creek, (2) El Casco, (3) Grand
Vies, (4) Cumberland Cave, (5) Port Kennedy, (6) Conard Fissure, (7) Hartman's Cave, (8)
Merritt Island, (9) Inglis IA, (10) Coleman IIA, (11) Apollo Beach, (12) Haile XVI, (13)
Cedazo, (14) Cherokee Cave, (15) Durham's Cave, (16) San Josecito Cave, (17) Clamp Cave,
(18) Waccasassa River, (19) Wolf Ranch, (20) Trout Cave, (21) Port Charlotte.


vex, while in Coendou and in juvenile Erethizon the angular pro-
cesses are subparallel and the posterolateral surface of the mandible is
flattened." (White 1968:9)
6) "In adults of Erethizon the projection of a line superimposed upon
the longitudinal axis of the lower tooth row and bisecting the occlusal
surfaces of P4 and M3, passes media to the incisor, while in Coen-
dou it extends to the posteromedial surface or even laterad to the


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


incisor. This measurement makes it possible to determine if the man-
dibular cheek teeth converge as in adults of Erethizon, or are sub-
parallel as in Coendou and juvenile Erethizon." (White 1968:9)
7) "The fossa for the insertion of M. masseter medialis pars posterior on
the side of the mandible is deeper in Coendou than in adults of
Erethizon. This structure seems related to the degree of convexity
seen in the posterior part of the mandible." (White 1968:9)
8) "The ascending ramus of the mandible in Coendou slants posteriad to
a greater degree than in adults of Erethizon." (White 1968:9)
9) "The scratches on the enamel of the occlusal surfaces in both the
upper and lower cheek teeth are oriented anteromediad and form an
angle greater than 35 degrees with the longitudinal axis of the tooth
rows in Coendou." (Landry, 1957) "In Erethizon this angle is less than
30 degrees." (White 1968:9)
Martin (1974) added two other criteria to distinguish the genera:
1) The posterior border of palate at midline located opposite center or
posterior border of upper M3 in Coendou, while in Erethizon the
posterior border of palate is located opposite center or posterior bor-
der of upper M2.
2) Upper cheek tooth row in Coendou is less than one-fourth of skull
length (microdont), while in Erethizon the upper cheek tooth row is
greater than one-fourth the skull length (macrodont).

A thorough comparison of the cranial osteology of Recent Coendou and
Erethizon was deemed necessary to test the validity and variability of the
characters presented above and to search for other differences. A large
sample of skulls and mandibles of Coendou and Erethizon was measured
according to the previously described procedure (N greater than 100 for
each genus). The major cranial measurements of adults are presented in
Table 1, along with the corresponding statistical parameters. In almost
every dimension, living Coendou is much smaller than living Erethizon.
The largest differences in the cranial dimensions measured in the two
living genera occur in the following variables:
X1, X20 Alveolar length of upper and lower tooth rows
X2, X21 Width of upper and lower P4
X3, X22 Width of upper and lower M
X5 Width of palate between upper M3 alveoli
X6, X23 Width of upper and lower incisor
X7, X24 Anteroposterior length of upper and lower incisor
X8 Total skull length
X9 Length of rostrum
X27 Length of diastema
X10 Length of upper M3 to occipital condyle





TABLE 1.-Statistical values of major cranial measurements in adult Coendou and Erethizon.

Coendou Erethizon
Variable* N S SD MIN MAX N XSD MIN MAX


2.138
.615
.531
.916
1.146
.351
.483
9.345
4.178
4.026
2.439
3.096
4.882
5.322
4.053
2.378
5.735
3.700
2.330
2.259
.599
.494
.362
.486
2.146
1.452
2.678
3.725
3.111


21.8
6.5
5.9
6.3
9.2
3.8
5.3
100.4
36.0
47.3
28.6
40.7
56.9
48.9
28.7
21.0
40.9
33.7
28.4
23.7
6.0
5.6
3.9
4.8
22.0
14.1
19.8
30.7
30.8


26.6
8.7
7.1
3.8
11.0
4.3
4.9
108.8
36.9
44.9
24.9
45.6
71.2
50.4
23.0
18.4
31.3
29.7
25.1
29.1
7.5
6.4
4.5
4.8
25.1
16.0
22.8
36.5
36.6


1.346
.586
.450
.960
1.153
.366
.337
6.937
3.434
3.036
2.068
2.213
3.333
2.996
1.687
1.291
3.245
2.221
2.240
1.583
.540
.397
.333
.312
1.809
1.228
2.346
2.750
2.728


30.1
10.1
9.3
7.9
14.4
5.3
6.0
125.7
45.3
51.1
30.4
51.2
80.2
68.2
28.3
21.7
43.0
34.9
31.7
33.1
9.0
7.9
5.5
5.7
30.2
18.9
27.9
44.4
45.1


*Cranial measurements corresponding to variables are: SKULL-XI = alveolar length of upper tooth row, X2=width of upper P4, X3= width of upper M1, X4=width of palate between P4 alveoli,
X5 = width of palate between M1 alveoli, X6= width of upper incisor, X7 = anterior-posterior length of upper incisor, X8= total skull length. X9= rostrum length, X10=length from M3 alveolus to
occipital condyle, Xllwidth across occipital condyles X12width across auditory meatuses, X13= width at zygomatics, X14=width at infraorbital foramen, X15= width of rostrum, X16= medial
width of rostrum, X17= width at frontals, X18= depth of rostrum, X19=length of auditory bulla. MANDIBLE-X20= alveolar length of lower tooth row, X21= width of lower P4, X22= width of
lower Ml, X23= width of lower incisor, X24=anterior-posterior length of lower incisor, X25=depth of mandible at P4, X26=depth of mandible at M2, X27= length of diastema, X28= length of
symphysis, X29= depth from condyle to angular process.







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


A




























cm D



FICURE 5. Comparison of fossil and Recent porcupine skulls in lateral view. (A) Recent
Coendou, AMNH 190423; (B) Recent Erethizon dorsatum, NMNH 245525; (C) fossil E.
dorsatum, Cumberland Cave, Maryland, NMNH 7996 holotypee, C. cumberlandicus White
1970); (D) fossil E. cascoensis, El Casco, California, F:AM 17883-1 holotypee, C. cascoensis
White 1970).






BULLETIN FLORIDA STATE MUSEUM


I Cm


FIGURE 6. Comparison of fossil and Recent porcupine skulls in dorsal view. A D same as
figure 5.


Vol. 27 No. 1


1QI~I





FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


[i Icm / ' cm











r.. -.. ,..cm
.1cm





D


C
FIGURE 7. Comparison of fossil and Recent porcupine skulls in ventral view. A D same as
figure 5.







BULLETIN FLORIDA STATE MUSEUM


X12 Width of skull at auditory meatus
X13 Width of skull at zygomatics
X14 Width of skull at infraorbital foramen
X25 Depth of mandible at lower P4
X26 Depth of mandible at lower M2
X28 Length of symphysis
X29 Depth of mandible from condyle to angular process

In order to explore the range of variation in the overall cranial dimen-
sions of the two genera, a cluster analysis was performed. The computer
reviewed all of the data of Recent specimens and reclassified them into two
groups based upon all cranial variables. One group contained all Erethizon
and some large Coendou; the other contained the rest of the Coendou
sample and no Erethizon. This indicates a range of overlap in the cranial
measurements of large individuals of Coendou with those of adults and
subadults of Erethizon. The measurements in which there is no overlap in
adults are: width of upper P4, width of upper Ml, width of skull at audi-
tory meatus, width of skull at zygomatics, and alveolar length of upper and
lower tooth rows. Figures in Appendix B illustrate the relationships of
selected cranial measurements of subadults and adults of both genera.
These morphometric differences are evidently related directly to the dif-
ferent masticatory adaptations of the two genera. Other discriminatory
morphological characters in the crania are also apparent (Figs. 5-10).
The anterior end of the nasals is pointed in Coendou but indented in
Erethizon. This feature may relate to the fact that in Coendou the soft part
of the nose is much more massive than in Erethizon. In every Recent skull
examined, the above character clearly distinguishes these two genera.
Another character of the skull is the great inflation of the frontals in some
Coendou, as noted by Ellerman (1940). This inflation may be a nasal de-
velopment for more effective olfaction.
In diagnosing Coendou cumberlandicus from the Cumberland Cave in
Maryland, White (1970:7) stated: "Nasofrontal suture slightly curved pos-
teriad," as in Coendou, "not V-shaped" as in Erethizon. This character has
been shown to be quite variable in Erethizon (Anderson and Rand 1943).
Indeed Ahlberg (1969) used this feature to distinguish the eastern sub-
species of Erethizon, with a nearly straight transverse nasofrontal suture,
from the western subspecies with a V-shaped suture.
The degree of development of the sagittal crest, reflecting the size of the
temporal muscle that attaches to the parietals, is relatively greater in
Erethizon than Coendou. As the temporal muscles develop with age, juve-
nile Erethizon do not have such a well developed sagittal crest. The rela-
tive degree of divergence of the zygomatics in Erethizon is much greater
than in Coendou and the dorsoventral thickness of the zygomatic is less in


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


A B


cm








E F










G H

FIGURE 8. Comparison of fossil and Recent porcupine mandibles in lateral view. (A) Recent
Erethizon dorsatum, NMNH 245525; (B) fossil E. dorsatum, Haile XVI, Florida, UF 21490;
(C) Recent Coendou, AMNH 190423; (D) fossil E. dorsatum, Aguascalientes, Mexico, UMMP
V47106; (E) Erethizon kleini, new species, Inglis IA, Florida, UF 21474; (F) fossil E. bathy-
gnathum, Grand View, Idaho holotypee), NMNH 13684; (G) E. kleini, new species, Inglis
IA, Florida (holoype), UF 21473; (H) E. bathygnathum, Vallecito Creek, California, IVCM
309.







BULLETIN FLORIDA STATE MUSEUM


Coendou than in Erethizon. However, some large Coendou have relatively
thickened zygomatics, so that some overlap occurs in this feature. The
zygomatic processes of the squamosals extend ventrolaterally in Coendou,
while in Erethizon they extend laterally. All of these characters are a result
of much greater masseter muscle development in Erethizon.
The shape of the auditory bullae in Coendou is oval and elongate, while
in Erethizon the bullae are more rounded. Internal ear structures are
compared in a later section.
White (1970) showed that Erethizon skulls have a greater interorbital
width than those of Coendou, and suggested that this produces different
visual fields. The tangent to the line of optic projection in arboreal Coen-
dou makes an angle of about 650 with the horizontal plane; whereas in the
more terrestrial Erethizon, the angle is approximately 450, a more dorsal
view than in Coendou.
The posterior border of the palate at the skull midline was found to be
quite variable in both genera and cannot be used as a distinguishing char-
acteristic. Martin (1974) developed this criterion after comparing only six
skulls of Erethizon and three of Coendou. Martin's use of the ratio of the
upper tooth row length to the total skull length is also too variable to be
statistically reliable according to the larger samples studied here. The char-
acter holds for Coendou but tends to break down in the smaller Erethizon.
Thus the two characters he developed have proved to be quite variable.
As noted by White (1970), the incisors of Erethizon are more extended
anteriorly (proodont) than the essentially orthodont incisors of Coendou.
The lower incisors of Coendou occlude with the upper incisors at a more
oblique angle than Erethizon. The present study strongly supports this
distinction of White, and I agree with his suggestion that the proodont
incisors of Erethizon are better adapted for eating bark than the orthodont
incisors of Coendou.
The mandibular characters discussed by White (1968, 1970) are generally
distinctive but are somewhat variable. The most distinctive character is the
inflection of the angular process. As stated by White (1968), the angular
process is inflected sharply media and flattened on the ventral surface in
Erethizon, whereas the angle in Coendou exhibits a smaller degree of
inflection and has little or no flattening of the angular process.
White (1968, 1970) used the angle of divergence of the lower tooth row
from the midline to distinguish Coendou and Erethizon. This angle is
useful in separating the Recent genera in most cases but cannot be used
with total confidence on a small fossil sample. Erethizon exhibited a lateral
projection of the tooth row in relation to the lower incisor in only 2 out of
125 observations; Coendou had a medial projection in 6 of 53 cases. Among
other characters that have proven to be variable are the depth of the fossa
for the insertion of the M. masseter medialis pars posterior on the mandi-


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


A B


C









E


G H


FIGURE 9. Comparison of fossil and Recent porcupine mandibles in medial view. A through H
same as Figure 8.


1981







BULLETIN FLORIDA STATE MUSEUM


A B


C C
cm






E F









G H


FIGURE 10. Comparison of fossil and Recent porcupine mandibles in dorsal view. A through
H same as Figure 8.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


ble and the orientation of the ascending ramus.
The scratches on the enamel of the occlusal surface in both upper and
lower cheek teeth have also been used to distinguish Coendou from Erethi-
zon. Landry (1957:15-16) stated "In Coendou, ... the scratches on the
teeth indicate that the angle at which the lower teeth move across the
upper is about fifty degrees from transverse. White (1968:9) described this
angle as "greater than 35 degrees with the longitudinal axis of the tooth
rows in Coendou. In Erethizon this angle is less than 30 degrees." Generic
distinction based solely upon the orientation of tooth scars in a small sam-
ple of isolated teeth should be made with caution. Along with the other
morphological characters, the orientation of the tooth scratches on the
cheek teeth provides useful evidence of masticatory differences between
the genera.
Woods and Howland (1979) describes two types of mastication in hystri-
comorphous rodents. Orthognathous mastication is propalinal (anterior-
posterior) and bilateral, while plagiognathous is more oblique and unilater-
al. Woods (pers. comm.) considers Coendou to be more orthognathous
than the plagiognathous Erethizon. The major osteological characters of
Erethizon associated with plagiognathous mastication are the posterior di-
vergence of the cheek tooth rows from the midline, the increased width of
the skull and zygomatic development and the deepening of the mandible.
The increase in size of the upper P4 in relation to Ml and the enlarged
upper and lower incisors are all adaptations of Erethizon for consuming
coarse vegetation.

FOSSIL IDENTIFICATION BY CRANIAL MEASUREMENTS
Using the information supplied by statistical analysis of living Coendou
and Erethizon cranial dimensions, a discriminant analysis was performed
upon the cranial measurements of the fossil specimens from North Amer-
ica. As some fossils are fragmentary, the analysis assigned generic prob-
ability values for each specimen based only upon the available measure-
ments. These values are presented in Tables 2 and 3, together with the raw
measurements of the classified fossils.
The fossils from Grand View, Vallecito Creek, and El Casco were classi-
fied as Erethizon with a very high probability (greater than or equal to .99).
The palates of the Erethizon-like porcupine from Inglis IA and Merritt
Island in Florida were classified into both genera. In most cases the Inglis
IA and Merritt Island fossils tend to be intermediate in size between
Coendou and Erethizon. Based upon the mandibular measurements, the
Inglis IA porcupines were considered Erethizon with a probability of great-
er than or equal to .89. The Cumberland Cave sample was classified as
Erethizon with a probability of greater than or equal to .98. Figures in
Appendix B graphically present the measurements of most of the fossil
porcupines studied, together with the values of adults and subadults of
Coendou and Erethizon.









22 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1


TABLE 2.-Fossil cranial measurements and discriminant analysis classification (d = deciduous).




Locality v

Catalogue #


GRAND VIEW
UO F-16282
UO F-16284
UO F-16279
VALLECITO CREEK
LACM 17633 28.1 8.9 8.4 6.7
LACM 6136 26.4 --- --- 6.7
EL CASCO
F:AM 17883-1 25.0 8.7 --- 2.4*
F:AM 17882 26.2 7.5 6.7 4.0*
F:AM 17883-7
F:AM 18128 ----
INGLIS IA
UF 21468 ---- 6.8 6.0 3.2
UF 21469 21.2 6.7 6.2 4.7*
UF 21470 23.7 6.6 5.7
UF 21471 21.1 5.2 5.8
MERRITT ISLAND
UF 21465 ---- 7.5 6.1 3.7
UF 21466 ---- 6.4 5.5
CUMBERLAND CAVE
NMNH 7996 26.2 8.5 7.6 6.7
NMNH 7670 25.0 7.0 6.1 4.5
NMNH 25692 26.3 8.3 6.8 4.9
PORT KENNEDY
ANSP 122 ---- 7.3 6.8
COLEMAN IIA
UF 11774 24.7 6.2 6.3 3.4
CONARD FISSURE
AMNH 12422 29.1 9.4 8.1 5.0
HARTMAN'S CAVE
ANSP 658 23.5 7.7 6.5 2.7


*Specimens either reconstructed or damaged.








FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


SS ; Classification by
S Discrim. Analysis

< Coendou Erethizon


0.000
0.000
0.000

0.001
0.000


0.008
0.000
0.000

0.200
1.000
0.606
0.962

0.038
0.998

0.000
0.000
0.000

0.000

0.017

0.000

0.003


1.000
1.000
1.000

0.999
1.000


0.992
1.000
1.000

0.800
0.000
0.394
0.038d

0.962
0.002

1.000
1.000
1.000

1.000

0.983d

1.000


0.997


1981









BULLETIN FLORIDA STATE MUSEUM


TABLE 3.-Fossil mandibular measurements and discriminant analysis classification
(d = deciduous).





Locality

Catalogue #


GRAND VIEW
NMNH 13684
UO F-16271
UO F-16272
UO F-16275
VALLECITO CREEK
LACM 6136
LACM 61420
LACM 6210
IVCM 309
EL CASCO
F:AM 17883-2
F:AM 17883
F:AM 17883-4
F:AM 17883-5
F:AM 17883-6
INGLIS LA
UF 21473
UF 21474
UF 21475
UF 21480
CUMBERLAND CAVE
NMNH 8128
NMNH 25689
NMNH 7672
AGUASCALIENTES
UMMP V-47106
HAILE XVI
UF 21490
CHEROKEE CAVE
AMNH 45742
APOLLO BEACH
UF 24112
UF 24113
UF 24114
HARTMAN'S CAVE
ANSP 648


--- 5.9

i.8


7.0 6.8 4.8 5.5

7.1 6.7 5.5 5.7

7.5 6.1 5.5 5.1


29.8 7.0 6.0


Vol. 27 No. 1









FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


C a Classification by
Ss E- F-1 Discrim. Analysis
SErethizon
goen.-u ---------
SCoendou Erethizon


0.000
0.000

0.000


1.000
1.000
-----d
1.000


17.7
15.9
17.2
16.5


0.000

0.001
0.000

0.026
0.115

0.004


0.000


0.000

0.000

0.000


1.000

0.999
1.000

0.974
0.885

0.996


1.000


1.000

1.000

1.000


25.6 16.6


14.4







BULLETIN FLORIDA STATE MUSEUM


The mandibles from these same localities are wholly characteristic of
Erethizon rather than Coendou. The Inglis IA mandibles are referrable to
Erethizon in every feature; especially diagnostic are posterior depth of
mandible, ratio of mandibular depth at lower P4 and M2, and inflection of
angular process. A line down the tooth row in both Inglis IA mandibles
projects media to the incisor, a correlate of Erethizon's divergent tooth
rows (White 1968, 1970). The Grand View mandibles, although incom-
plete, look as though they had an angular process directed strongly
media, as in Erethizon. This character cannot be determined in the El
Casco and Vallecito Creek samples, but on all other features they were
classified as Erethizon (greater than or equal to .99) along with the Grand
View specimens. Figures 5 through 11 compare fossil cranial specimens
from various localities with Recent Erethizon and Coendou.
Thus all fossil porcupines in North America can be referred to Erethizon
using the cranial characters discussed above, although some Florida speci-
mens show some Coendou dimensions in their skulls. We turn next to
other sets of diagnostic characters revealed in the incisor enamel and audi-
tory ossicles.

INCISOR ENAMEL

The value of incisor enamel microstructure in rodent taxonomy was first
investigated by Tomes (1850) and later applied by Korvenkontio (1934).
More recent studies involving incisor enamel microstructure in rodents
include works by Wahlert (1968), Escala and Gfllego (1977), and War-
shawsky (1971). Wahlert (1968) compared the incisor enamel of various
fossil and Recent rodent groups as viewed in thin section. He examined the
degree of variability in the enamel and discussed the three basic types of
incisor enamel found in rodents. Escala and Gallego (1971), also using thin
sections, determined the variability and taxonomic usefulness of lower in-
cisor enamel in several genera of Muridae. Warshawsky (1971) examined in
detail the enamel microstructure of Rattus, using scanning electron micros-
copy.
In search of other taxonomic tools for the separation of Coendou and
Erethizon, I examined the sagittal sections of lower incisor enamel in both
of the living genera and in fossil specimens from various localities in North
America, using scanning electron microscopy. Both genera and the fossil
specimens have multiserial enamel as discussed by Wahlert (1968). Using
the measurement procedures presented by Wahlert (1968) and Escala and
Gfllego (1977), differences were found in the thickness of the inner enamel
and the total enamel in Coendou and Erethizon. Figure 12 pictures the
enamel of both genera in sagittal view and also shows the measurement
methods.


Vol. 27 No. 1






FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


B C


D E F




II
G H
Icm







I J K
FIGURE 11. Comparison of fossil and Recent porcupine palates in ventral view. (A) Recent
Coendou, AMNH 190423; (B) Recent Erethizon dorsatum, NMNH 245525; (C) fossil E.
dorsatum, Cumberland Cave, Maryland, NMNH 7996 holotypee, C. cumberlandicus White
1970); (D) E. kleini, new species, Inglis IA, Florida, UF 21468; (E) E. kleini, new species,
Inglis IA, Florida, UF 21469; (F) E. kleini, new species, Merritt Island, Florida, UF 21465;
(G) E. kleini, new species, Inglis IA, Florida, UF 21470; (H) E. kleini, new species, Inglis IA,
Florida, UF 21471; (I) E. bathygnathum, Vallecito Creek, California, LACM 17633 holotypee,
C. stirtoni White 1968); (J) E. cascoensis, El Casco, California, F:AM 17882; (K) E. cascoensis,
El Casco, California, F:AM 17883-1 holotypee, C. cascoensis White 1970).







BULLETIN FLORIDA STATE MUSEUM


Incisor enamel thickness in Erethizon is significantly greater than in
Coendou (Table 4). This is most likely related to the food habits of the
animals; Erethizon habitually consumes much tougher bark and other
temperate vegetation, while Coendou feeds upon generally softer tropical
foliage.
The fossils examined from the various early and late Pleistocene localities
exhibit values within the range of Erethizon (Table 5). From these data one
could hypothesize that the fossil erethizontids had developed the incisor
enamel thickness needed for the consumption of the bark of temperate
conifers prior to their dispersal into North America. Olson and McGrew
(1941) have found North American savannah mammals in Miocene sedi-
ments as far south as Honduras in Middle America. Southern Mexico sup-
ported such trees as fir, alder, beech, and elm as far back as Miocene time,
14 million years before the first record of porcupine in North America
(Raven and Axelrod 1975). It is thus probable that the first erethizontids to
enter North America immediately encountered many elements of more
temperate forests in Middle America.


AUDITORY OSSICLES

The auditory region is widely regarded as an excellent key to the sys-
tematics of various vertebrate groups. For example, Feduccia (1977) re-
cently developed a model for the evolution of perching birds based upon
the stapes morphology, and Hunt (1974) has used the auditory bulla in
discovering mammalian carnivore affinities. The most comprehensive re-
view of the auditory region in mammals is by Van der Klaauw (1931).
Recent studies of the auditory region in rodents by Segal (1971), Bugge
(1971, 1974), Lindsay (1972), and Wahlert (1974) have supplied vital
information concerning the interpretation of evolutionary relationships in
the order. Detailed studies of the auditory region in fossil hystricomorph
rodents have been carried out by Fields (1957) among dinomyids and by
Mones (1974) among hydrochoerids.
These many precedents encouraged me to examine the ear region in
North American porcupines in hopes of finding characters useful in the
classification of the early fossil forms. Comparison of the auditory regions
and particularly the ear ossicles of fossils from Vallecito Creek, El Casco,
and Inglis IA with those of Recent Erethizon and Coendou provides valu-
able systematic information. It is fortunate that these specimens exist, as
fossil ear ossicles are extremely rare.
No apparent differences were found in the general structure of the audi-
tory bullae of Coendou and Erethizon, although clear differences exist in
the morphology of the incus, malleus, and stapes. The ear ossicles from 12


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


OE



IE


C
FIGURE 12. Scanning electron microscope photomicrographs of porcupine lower incisor
enamel in sagittal section and measurement methods. (A) measurement methods: OE = outer
enamel thickness, IE = inner enamel thickness, TE = total enamel thickness, W = width of
enamel bands, a = angle of enamel bands in degrees; (B) Recent Coendou, AMNH 96326 (X
287); (C) Recent Erethizon dorsatum, UF 6349 (X 216).








BULLETIN FLORIDA STATE MUSEUM


TABLE 4.-Statistical values of lower incisor enamel parameters.


N = 12 Coendou Erethizon F-Values


Band Width
(W)

SD
MIN
MAX
Band Angle
(c)

SD
MIN
MAX
Outer Enamel
(OE)

SD
MIN
MAX
Inner Enamel
(IE)

SD
MIN
MAX
Total Enamel
(TE)

SD
MIN
MAX


.0236
.0033
.0191
.0280


39
6.10
29
46


.0288
.0077
.0179
.0426


.1795
.0388
.1172
.2580


.2099
.0398
.1459
.2960


.0273
.0045
.0206
.0368


44
6.17
32
52


.0307
.0076
.0108
.0433


.2403
.0374
.1879
.3053


.2710
.0374
.2241
.3509


4.3






.3614






15.28*


14.97*


*Indicates an F-ratio exceeding 14.4 (P < .001) (from Rohlf and Sokal 1969).


skulls of each living genus were examined. The auditory ossicle terminolo-
gy is presented in Figure 13.


INCUS-MALLEUS
The incus and malleus are fused in many rodent groups, including the
Erethizontidae (Wood 1974). Typical fused elements from Erethizon and
from Coendou are presented in Figure 14. The head of the malleus in
Coendou is bulbous, whereas in Erethizon it is elongate. The processus axialis
in Coendou is very deep and has a distinctive anterior projection, but in


Vol. 27 No. 1








FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


TABLE 5.-Incisor enamel values of fossil Erethizontidae.


Locality
Catalogue # W a OE IE TE U or L*


VALLECITO CREEK
LACM 6136-2
LACM 6136-3
LACM 23062
EL CASCO
F:AM 17883-5
F:AM 17883-6
F:AM 17883-7
GRAND VIEW
UO F-16275
UO F-16278
UO F-16281
INGLIS LA
UF 21480
UF 24134
COLEMAN IIA
UF 11794
UF 24135
WACCASASSA RIVER
UF 21487
SEMINOLE FIELD
AMNH 23114b
Steiromys
KU 11605


.0305
.030
.0325

.0250
.0250
.0263

.0325
.0250
.0325

.0315
.0263

.0263
.0263

.0263

.0222

.0277


.050 .3111 .3611
.070 .240 .310
.060 .310 .370

.0555 .3550 .4100
.0275 .2050 .2400
.0210 .2315 .2526


.0450 .2650
.0300 .2150
.0450 .1900


.3100
.2450
.2350


47 .0368 .2157 .2526
49 .0210 .2105 .2315

48 .0421 .2842 .3263
44 .0421 .2947 .3421

49 .0526 .3000 .3526

31 .0277 .2666 .2944

45 .0500 .2833 .3333


*U= Upper; L= Lower.


Erethizon it is shallow with no anterior projection. The manubrium is
relatively longer and thicker mediolaterally in Erethizon than in Coendou.
The heel of the manubrium in Coendou tends to be broad and rounded,
whereas in Erethizon the structure is relatively narrower.
In lateral view, the line of fusion between the incus and malleus is
marked by a trough on the dorsal surface in Coendou, whereas in Erethi-
zon that surface is flat. In Coendou the incus joins the malleus much closer
to the neck than in Erethizon. The dorsal surface of the incus in Coendou
slants more ventrally than in Erethizon.
The complete incus-malleus from Inglis IA shows the characters of
Erethizon rather than Coendou (Fig. 14). In silhouette the dorsal surface is
flat as in Erethizon. The processus axialis is shallow and no anterior projec-
tion is present. The heel of the manubrium is long and narrow. In each of








BULLETIN FLORIDA STATE MUSEUM


A B


INFERIOR SUPERIOR
CRURAL CRURAL
ARCH ARCH
ANTERIOR POSTERIOR
CRUS CRUS

FOOTPLATE



C
FIGURE 13. Terminology of hystricomorph auditory ossicles. (A) lateral or dorsal view of
incus-malleus; (B) medial or ventral view of incus-malleus; (C) lateral or dorsal view of stapes,
showing shape of footplate. (Modified from Fields 1957; Mones 1974.)


these characters the Inglis specimen closely resembles Recent Erethizon.
A partial incus-malleus from Vallecito Creek consists of the head of the
malleus and the upper portion of the incus with the processus brevis (Fig.
14). The dorsal surface of the prepared portion is flat and ungrooved as in
Erethizon. As the matrix that filled the auditory region is very hard, the
specimen was not prepared further, for fear of destroying it.

STAPES
Samples of 10 stapes from each living genus were examined. The stapes
of Erethizon is much larger in overall size than Coendou (Fig. 15). The
structure of the stapes head is quite variable in both genera. The anterior
and posterior crura are longer in Erethizon than in Coendou. The footplate
of the stapes, which rests in the fossa ovalis, has a long ovate outline in
Erethizon, whereas in Coendou it is circular.
The only known fossil erethizontid stapes from North America is that
from Inglis IA (Fig. 15). The footplate of the specimen is oval-elongate as in


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


h I(




A B

mm













C D
FIGURE 14. Lateral or dorsal view of fossil and Recent porcupine incus-mallei. (A) Recent
Erethizon dorsatum, NMNH 256661; (B) Recent Coendou, AMNH 190418; (C) E. bathy-
gnathum, Vallecito Creek, California, LACM 6136; (D) E. kleini, new species, Inglis IA,
Florida, UF 24176.

Erethizon. The anterior and posterior crura are not as long as in Recent
Erethizon specimens but are closer in length to Coendou specimens. The
shape of the fossa ovalis closely resembles the shape of the stapes footplate
(Fig. 15).
The shape of the fossa ovalis in the Inglis IA periotic is clearly oval as in
Erethizon. The fossa ovalis was prepared on the periotic of the El Casco
specimen, and its shape closely resembles that of the fossa in Recent
Erethizon. Although broken, enough of the fossa ovalis in the Vallecito






BULLETIN FLORIDA STATE MUSEUM


A :'^



mm










CI












FIGURE 15. Comparison of stapes and fossa ovalis in fossil and Recent porcupines. (A) Recent
Coendou, AMNH 186427; (B) Erethizon kleini, new species, Inglis IA, Florida, UF 24176; (C)
Recent E. dorsatum, NMNH 256661.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


Creek specimen is present to determine its shape as oval also. Thus, the
fossil porcupines from Vallecito Creek, El Casco, and Inglis IA can be
referred to Erethizon on the basis of several features of the auditory ossi-
cles and fossa ovalis.

POSTCRANIAL OSTEOLOGY

The gross osteology of Erethizon dorsatum was described by Swena and
Ashley (1956). Sutton (1972) made detailed observations of cranial and
postcranial variations in the skeleton of the species. Gupta (1966) made a
superficial comparison of the skeletons of Coendou and Erethizon. The
selected osteological comparisons presented here concern only the generic
assignment of the fossils from North America.
Ray (1958), in a study of the fusion of cervical vertebrae in Erethizontidae
and Dinomyidae, mentioned no characters to distinguish the cervical ver-
tebrae of Coendou and Erethizon. Likewise in this study no morphological
differences in the cervical vertebra or in those from other regions were
noticed.
Coendou has a long, prehensile tail, while the tail of Erethizon is short
and nonprehensile. Correspondingly Coendou has more caudal vertebrae
than Erethizon. Gupta (1966) reported 28 caudal vertebrae in Coendou and
18 in Erethizon. I found as many as 33 caudal vertebrae in Coendou.
Sutton (1972) reported 13 to 17 caudal vertebrae in Erethizon. Thus Coen-
dou has at least 10 more caudal vertebrae than Erethizon. Individual ver-
tebrae are not diagnostic and a complete caudal series is needed to assign
vertebrae to a particular genus. For this reason, the isolated caudal verte-
brae from Vallecito Creek and Inglis IA add little information about the
nature of the tail in these fossil forms.
No major distinctions were discovered in the forelimb. The humerus
(Fig. 16), ulna, and radius show only size-related differences.
Several major differences occur in the pes and hind limbs. The third
trochanter of the femur is relatively more developed in Erethizon than in
Coendou (Gupta 1966). The fossil femora from Vallecito Creek, Inglis IA,
Cumberland Cave, and Haile XVI exhibit a well developed third trochan-
ter as in Erethizon (Fig. 17).
At the distal end of the tibia in Coendou the medial malleolus extends
posteriorly, whereas in Erethizon this extension is reduced. This differ-
ence reflects the different modes of pedal articulation between Coendou
and Erethizon. Unfortunately, this character cannot be compared in the
fragmentary tibiae from Vallecito Creek, and no other fossil tibiae are
known.
The fibula in Coendou tends to be curved or bowed along its length,
while in Erethizon the fibula is straight. The fossil fibula from Inglis IA is








BULLETIN FLORIDA STATE MUSEUM


D E F
B C

FIGURE 16. Anterior view of fossil and Recent porcupine humeri. (A) Erethizon bathy-
gnathum, Vallecito Creek, California, LACM 4325; (B) E. bathygnathum, Vallecito Creek
California, LACM 6136; (C) Recent E. dorsatum, UF 7993; (D) E. kleini, new species, Inglis
IA, Florida, UF 24137; (E) E. kleini, new species, Inglis IA, Florida, UF 24138; (F) Recent
Coendou, AMNH 93040.





















C

FIGURE 17. Anterior view of fossil and Recent porcupine femora. (A) Recent Coendou,
AMNH 93040; (B) Erethizon kleini, new species, Inglis IA, Florida, UF 24155; (C) Recent E.
dorsatum, UF 7993; (D) fossil E. dorsatum, Cumberland Cave, Maryland, NMNH 214818;
(E) E. bathygnathum, Vallecito Creek, California, LACM 6136.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


straight as in Erethizon.
Several major differences in the pes distinguish Erethizon and Coendou.
Coendou, the more arboreal form, exhibits a highly specialized prehallux,
while the structure in Erethizon is almost vestigial (White 1968). The astra-
galus differs between Coendou and Erethizon in the following ways:

1) In ventral view the lateral articular surface in Erethizon exceeds the
medial articular surface area, whereas in Coendou these surfaces are
nearly equal in size.
2) In Erethizon the medial articular surface tends to be confluent with
the articular surface for the astragalus head; in Coendou the surfaces
tend to be separated.
3) In posterior view the lateral malleolar surface slopes more proximally
in Erethizon than in Coendou.
4) In dorsal view the astragalus of Erethizon exhibits an extended tuber-
osity just posterolateral to the medial malleolar surface, whereas in
Coendou this ossification does not exist.

Differences in the calcaneum that correspond with the above differences
in the astragalus are listed below:

1) In dorsal view the sustentaculum of the calcaneum in Erethizon slants
more anteriorly than in Coendou, extending beyond the cuboid ar-
ticular surface.
2) In dorsal view the posterior angle between the distomedial edges of
the sustentaculum and the lateral articular surface is more oblique in
Erethizon, whereas in Coendou the angle tends to be more perpen-
dicular.
3) In posterior view the angle between the medial articular surface on
the sustentaculum with the lateral articular surface is more oblique in
Coendou than in Erethizon.

These differences in the articular surfaces of the astragalus and calcaneum
of Coendou and Erethizon are probably related to their markedly different
modes of locomotion. The pes of Erethizon seems to have greater rotational
ability, ideal for its greater range of arboreal and terrestrial existence.
Astragali of the fossil specimens from Vallecito Creek and Inglis IA are
pictured in Figure 18 in comparison with Recent Coendou and Erethizon.
All of the fossil astragali exhibit the characters of Erethizon.
Fossil calcanea are known from Vallecito Creek, Inglis IA, and Haile
XVI. These specimens clearly have the articular surface characters of
Erethizon (Fig. 19). These postcranial comparisons suggest that the earliest
porcupines in North America should be referred to Erethizon.






BULLETIN FLORIDA STATE MUSEUM


cm


frm
~1 4p d


FIGURE 18. Fossil and Recent porcupine astragali in proximal (left) and distal (right) view. (A)
Recent Coendou, AMNH 134067; (B) Recent Erethizon dorsatum, UF 7993; (C) E. kleini,
new species, Inglis IA, Florida, UF 24161; (D) E. bathygnathum, Vallecito Creek, California,
LACM 4325.


Vol. 27 No. 1


Alth.






FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


cm


"Vp


FIGURE 19. Fossil and Recent porcupine calcanea in proximal (left) and medial (right) view.
(A) Recent Coendou, AMNH 134067; (B) Recent Erethizon dorsatum, UF 7993; (C) E. bathy-
gnathum, Vallecito Creek, California, LACM 4325; (D) E. kleini, new species, Inglis IA,
Florida, UF 24162.


I,
-I
y
-.i"L~uP_ -


ai4
p o;


'3


f! 3







BULLETIN FLORIDA STATE MUSEUM


SYSTEMATICS

ORDER RODENTIA
FAMILY ERETHIZONTIDAE THOMAS 1896
SUBFAMILY ERETHIZONTINAE THOMAS 1896
Erethizon F. CUVIER 1822

TYPE SPECIES. -Hystrix dorsata Linnaeus 1758 (monotypy).
KNOWN DISTRIBUTION. -Late Pliocene (Blancan) to Recent in temperate
North America.
EMENDED DIAGNOSIS.--A large erethizontid with a short, muscular non-
prehensile tail and a vestigal prehallux in the hind foot of the living spe-
cies. Astragalus has a tuberosity just posterolateral to the medial malleolar
surface. Upper and lower cheek tooth rows converge markedly anteriad,
more so in the living species than in extinct forms.

Erethizon bathygnathum WILSON 1935

E. bathygnathum Wilson 1935 (Type Locality: Jackass Butte [rather than
Castle Butte,; Hibbard 1959:32], Grand View, Owyhee County, Idaho;
Natl. Mus. Nat. Hist.).
E. bathygnathum: Shotwell 1970.
Coendou bathygnathum (Wilson) White 1970.
C. stirtoni White 1968 (Type locality: Vallecito Creek, Anza Borrego Des-
ert State Park, San Diego County, California; Los Angeles Co. Mus.
Nat. Hist.) NEW SYNONYMY
C. stirtoni: White 1970, Harrison 1978.
AGE.-Late Blancan to early Irvingtonian.
DISTRIBUTION.-Known only from the Great Basin and Pacific Coastal
regions of the western United States: Grand View, Idaho; Wolf Ranch,
Arizona; and Vallecito Creek, California.
EMENDED DIAGNOSIS.-A large porcupine, notably larger than the ex-
tant species, Erethizon dorsatum ff-irmaeus), with a much deeper and
more massive mandible. The cheek teeth and the anterior-posterior length
and width of the incisors tend to be larger than in E. dorsatum. Trochlear
groove of astragalus deeper and calcaneum more shallow and elongate than
in the living form.
HOLOTYPE.-NMNH 13684, fragmentary left mandible with broken in-
cisor and P4-M2.
REFERRED MATERIAL.-GRAND VIEW, IDAHO: UO F-16271, left
mandible with I, P4-M3; UO F-16272, left mandible with I, DP4-M2; UO
F-16275, left lower incisor; UO F-16278, left upper incisor; UO F-16279,
right upper incisor; UO F-16281, right upper incisor; UO F-16282, right


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


upper incisor; UO F-16289, left upper incisor. WOLF RANCH, ARIZO-
NA: UALP 4911, left upper incisor; UALP 4912. left upper M1. VALLE-
CITO CREEK, CALIFORNIA: LACM 17633, palate with left P4-M1 and
partial right P4-M2 holotypee, Coendou stirtoni); LACM 61420, edentulous
left mandible; LACM 6210, edentulous left mandible; LACM 23062, in-
cisors; LACM 4325, partial skeleton; LACM 6136, partial skeleton; IVCM
309, fragmentary left mandible with I, M1-M3; IVCM 125, cheek tooth and
incisor fragments.
DIscussIoN-This species of Erethizon is recognized from late Blan-
can-early Irvingtonian deposits in the western United States. No specific
differences can be found to distinguish the fossil form from Vallecito Creek
(Coendou stirtoni) from the Grand View specimens of E. bathygnathum.
Previously only mandibles were known from Grand View, and only crania
and mandibles without cheek teeth were available from Vallecito Creek,
thus making comparison difficult. Now a mandible from Vallecito Creek
(IVCM 309) permits direct comparison of the lower cheek teeth of C. stir-
toni with those of the type and topotypic mandibles of E. bathygnathum. No
specific differences separate the Vallecito Creek and Grand View samples,
assuming a range of variability comparable to that in Recent E. dorsatum.
All of the mandibles from the two localities have a short diastema, much
shorter than most E. dorsatum. The ratio of width of the deciduous P4 to
Ml in UO F-16272 from Grand View is outside the pattern found in E.
dorsatum.
Coendou stirtoni (Harrison 1978) from the Wolf Ranch local fauna, Ari-
zona, is referred to Erethizon bathygnathum, based upon the anterior-
posterior length (APL) and width of the upper incisor (UALP 4911) (Fig.
11; Appendix B) and the angle of the wear striations on the isolated upper
Ml (UALP 4912). These striations form an angle of approximately 35* with
the transverse line, and thus fall well within the range of Erethizon as
discussed previously (p.00). The APL and width of the upper incisor is
larger than in E. dorsatum and falls among the values of E. bathygnathum
(Fig. 11; Appendix B).
Erethizon bathygnathum is about 30% larger than the living porcupine,
E. dorsatum. This is evident from the cranial measurements (Table 2) and
the size of the limb bones (Figs. 16-19). The mandible of E. bathygnathum
is deeper relative to E. dorsatum (Figs. 8 and 9), although this feature
shows some overlap. Also the anterior-posterior length and width of the
incisors are generally larger in E. bathygnathum.
Erethizon bathygnathum can be distinguished from later species of
Erethizon by morphological differences in the astragalus and calcaneum.
The trochlear groove of the astragalus is deeper than in the living species.
The calcaneum is more elongate, but shallower than the calcaneum of E.
dorsatum (Fig. 19).


1981







BULLETIN FLORIDA STATE MUSEUM


It is interesting that the earliest Erethizon was apparently restricted to
the far west of North America. Blancan sites of appropriate ages in Texas,
Oklahoma, Kansas, and Nebraska are sufficiently well known to suggest
that this pattern is not likely an artifact of collecting. Apparently Erethizon
did not spread east of the Rocky Mountains until Irvingtonian times.

Erethizon cascoensis (WHITE)

Coendou cascoensis White 1970 (Type locality: El Casco, Riverside
County, California; Amer. Mus. Nat. Hist.).
AGE-Early Irvingtonian
DISTRIBUTION.-Known only from the El Casco local fauna, California.
EMENDED DIAGNOSIS.--A porcupine the size of the extant species,
Erethizon dorsatum (Linnaeus), with the internal reentrant (hypoflexus) of
upper P4 absent. Incisor anterior-posterior length and width relatively
large, as in E. bathygnathum. Incisor enamel thick as in other species of
Erethizon. Diastema relatively short as in E. bathygnathum. Sagittal crest
joined as in E. dorsatum. Internal ear structure comparable to Erethizon.
HOLOTYPE.-F:AM 17883-1, skull, rostrum missing (crushed).
REFERRED MATERIAL.-EL CASCO, CALIFORNIA: F:AM 17882,
crushed palate with right P4-M2 and left P4-M3; F:AM 17883, edentulous
left mandible with incisor; F:AM 17883-2,. crushed right mandible with
M1-M3; F:AM 17883-4, unworn right lower P4; F:AM 18128, left upper
incisor; F:AM 17883-5, right lower incisor; F:AM 17883-6, left lower in-
cisor; F:AM 17883-7, right upper incisor.
DIscussioN.--This species is recognized only from the El Casco local
fauna. Considering the variability in Erethizon dorsatum, the only charac-
ter that distinguishes E. cascoensis from E. bathygnathum is the absence of
the internal reentrant in the upper P4. The unworn right lower P4 (F:AM
17883-4) has the internal reentrant (hypoflexus). White (1970) states "...
seven out of 60 specimens of Coendou, .. ." have the hypoflexus missing
in the upper P4. I observed this absence in no Recent Erethizon. Based
upon these figures, this character is not acceptable as a generic determi-
nant but must be recognized as a specific differential. However, it is odd
that the lower P4 has the reentrant.
Distinct deformation and fractures are evident in the type skull of
Erethizon cascoensis (F:AM 17883-1) and the palate (F:AM 17882). The
parallel condition of the upper cheek tooth rows (White 1970), characteris-
tic of Coendou, is probably exaggerated by this distortion. In all other
features the El Casco porcupine sample is clearly characteristic of Erethi-
zon. The upper P4 is markedly larger than Ml. The fossa ovalis is oblong in
shape. The sagittal crests are joined; the results of enlarged masticatory
muscles. The zygomatic projection of the squamosals extends laterally,


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


typical of Erethizon. The incisor enamel is thick and the anterior-posterior
length and width of incisor are comparable with other species of Erethizon.
The diastema is short (F:AM 17883) and mandible deep (F:AM 17883-2) as
in E. bathygnathum. If the absence of the hypoflexus in upper P4 in the El
Casco specimens is a variability, E. cascoensis could be referred to E.
bathygnathum with a reasonable amount of assurance.
Erethizon kleini, new species

HOLOTYPE.--UF 21473, left mandible with incisor and P4-M3. Condyle
and angular process broken. Collected by Mr. Jean Klein in 1970.
TYPE LOCALITY.--Inglis IA, Citrus County, Florida. A sinkhole-cavefill
on the north side of the Cross-Florida Barge Canal approximately 3 miles
east of U.S. Highway 98 south of Inglis, Florida.
AGE. -Early Irvingtonian.
DISTRIBUTION. -Principal sample from Inglis IA in west-central Florida.
Possibly referrable material from Merritt Island, Brevard County, east-
central Florida.
DIAGNOSIs. -Notably smaller than any other species of Erethizon;
otherwise closely resembling E. dorsatum morphologically.
ETYMOLOGY.-Named in honor of Mr. Jean Klein for collecting this
specimen and for his contributions to Florida vertebrate paleontology.
REFERRED MATERIAL.-INGLIS IA, CITRUS COUNTY, FLORIDA:
UF 21474, left mandible with P4-M2; UF 21475, fragmentary left mandible
with M1-M3; UF 21469, partial palate with left P4-M2 and right P4-M3;
UF 21468, partial palate with right and left P4-M1; UF 21475, right palate
with P4-M3; UF 21471, right palate with DP4-M2; UF 21472, left partial
palate with DP4; UF 21476, left unerupted upper P4; UF 21478, right
upper M3; UF 21477, left unerupted upper P4; UF 21479, left upper M2;
UF 21481, left lower M2; UF 21482, right lower DP4; UF 21483, left
lower DP4; UF 21484, lower left incisor (immature); UF 21480, lower left
incisor; UF 24134, left upper incisor (fragmentary); UF 24176, left auditory
bulla with incus-malleus and stapes; UF 24167, nine isolated caudal ver-
tebrae; UF 24137, right humerus; UF 24138, right humerus; UF 24139,
fragmentary left humerus (immature); UF 24150, proximal left radius; UF
24151, proximal left radius; UF 24152, proximal left radius; UF 24146,
proximal left ulna; UF 24147, proximal right ulna; UF 24148, left ulna;
UF 24154 right femur; UF 24155, right femur; UF 24156, right femur; UF
24157, left femur; UF 24158, left femur; UF 24159, left femur; UF 24164,
right fibula; UF 24162, right calcaneum; UF 24160, left astragalus; UF
24161, right astragalus. MERRITT ISLAND, BREVARD COUNTY,
FLORIDA: UF 21465, partial palate with right and left P4-M2; UF 21466,
right palate with P4-M3. Possibly associated with a later Irvingtonian or
Rancholabrean fauna.


1981







BULLETIN FLORIDA STATE MUSEUM


DESCRIPTION AND DISCUSSION.--The type, a left mandible (UF 21473),
exhibits all the characteristics of Erethizon except for its small size (Figs.
8-10; Table 3). The mandibular depth at P4 and M2 falls clearly within the
range of Erethizon (Fig. 16; Appendix B). The angular process is inflected
media, as is typical of Erethizon. The lower incisor dimensions are clearly
within the range of Erethizon rather than Coendou (Fig. 12; Appendix B).
The cheek tooth size is intermediate between the two genera (Figs. 9 and
10; Appendix B). The cheek tooth rows also diverge posteriorly from the
midline as in Erethizon. The left mandible (UF 21474) also exhibits the
above characteristics (Fig. 10).
The palate and upper cheek teeth exhibit characters of Coendou and
Erethizon (Fig. 11). The P4 is markedly larger than Ml in UF 21468 and
UF 21465, while in UF 21469 and UF 21466 the P4 is more equal in size to
Ml. The tooth rows in UF 21468 and UF 21465 converge anteriorly as in
Erethizon. In UF 21469 the tooth rows appear more nearly parallel; but
this is questionable as the palate was reconstructed from isolated fragments
of the same individual, broken during screening for microfauna. Isolated
upper tooth rows clearly distinguish the species as Erethizon (Fig. 11). P4
in UF 21470 is markedly larger than Ml, and DP4 in UF 21471 is much
smaller than Ml. The above relationships are characteristic of Erethizon
and not Coendou.
The incus-malleus is preserved in UF 24176 and shows diagnostic fea-
tures of Erethizon (Fig. 14). The dorsal surfaces are smooth; the manu-
brium is long and thin; and the processus axialis is reduced. The shape of
the stapes and fossa ovalis is oval, also characteristic of Erethizon (Fig. 15).
The limb bones display the small size diagnostic of this species of Erethi-
zon. The humeri and femora of the fossil porcupines are compared in
Figures 16 and 17. The astragalus and calcaneum of E. kleini are similar in
size to Coendou but have the characteristics of Erethizon as discussed
earlier (p. 00-00). The astragali differ from E. bathygnathum in having a
more shallow trochlear groove, but the calcaneum is more similar in pro-
portions to E. bathygnathum than to E. dorsatum.
This fossil species, known only from Florida, is by far the smallest
Erethizon in North America, and it is also the earliest record of Erethizon
in eastern North America. The species probably dispersed northeastward
along the Gulf Coast from Central America about 2 mya, possibly becom-
ing endemic to the Gulf Coastal area of the Southeast.


Erethizon dorsatum (LINNAEUS)

Hystrix dorsata Linnaeus 1758.
Erethizon dorsatum (L.): F. Cuvier 1822.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


Erethizon cloacinum Cope 1871 (Type Locality: Port Kennedy Cave, Ches-
ter County, Pennsylvania; Amer. Mus. Nat. Hist.) NEW
SYNONYMY.
E. godfreyi Allen 1904 (Type Locality: White Mountain Apache Indian
Reservation, Navajo County, Arizona; Amer. Mus. Nat. Hist.) (= E.
dorsatum: White 1970).
Coendou cumberlandicus White 1970 (Type Locality: Cumberland Cave,
Allegany County, Maryland; Natl. Mus. Nat. Hist.) NEW
SYNONYMY.
Coendou sp.: White 1970:8-9.
AGE.--Middle Irvingtonian to Recent.
DISTRIBUTION. -North America (see Fig. 2 and Appendix A).
EMENDED DIAGNOSIS.--(same as genus). Anterior edge of nasals in-
dented. Calcaneum short and deepened. (See Woods 1974).
REFERRED MATERIAL. -PORT KENNEDY CAVE, CHESTER COUN-
TY, PENNSYLVANIA (Middle Irvingtonian): AMNH 8576, isolated right
upper M3; ANSP 15543, two associated upper incisors; ANSP 15542, three
isolated molars: ANSP 137, partial right mandible with incisor, M1-M2,
isolated right lower DP4 and left upper P4; ANSP 138, four isolated in-
cisors; ANSP 122, associated left upper P4-M2 and right upper P4.
CUMBERLAND CAVE, ALLEGANY COUNTY, MARYLAND (Middle
Irvingtonian): CM 12792, right astragalus; CM 20323, left lower M2; CM
20004, left upper M2; CM 34075, fragmentary left mandible with incisor,
P4-M1; NMNH 214775 and 214818, right and left femora, respectively;
NMNH 214816 proximal right humerus; (additional material listed by
White, 1970). CAVETOWN, WASHINGTON COUNTY, MARYLAND
(?Rancholabrean): NMNH 9208, left mandible with incisor, unerupted P4
and M1-M3. TROUT CAVE, PENDLETON COUNTY, WEST VIRGINIA
(Middle Irvingtonian): CM 12792, right upper P4. WINDY MOUTH
CAVE, GREENBRIAR COUNTY, WEST VIRGINIA (? Rancholabrean):
CM 24305, left humerus. CHEROKEE CAVE, ST. LOUIS COUNTY,
MISSOURI (Rancholabrean): AMNH 45742, associated right and left man-
dibles. CONARD FISSURE, NEWTON COUNTY, ARKANSAS (Middle
Irvingtonian): AMNH 12422, partial skull with palate and rostrum; AMNH
12423, three isolated cheek teeth. HAILE XVI, ALACHUA COUNTY,
FLORIDA (Middle Irvingtonian): UF 21490, left mandible with incisor,
P4-M1; UF 21492, fragmentary left mandible with M1-M2; UF 24166,
right palate with M2-M3; UF 24136, left upper M3; UF 21491, left lower
M3; UF 24140, 24141, 24142, 24143, 24144, and 24145, humeri; UF
24149, proximal right radius; UF 24150, proximal left femur; UF 24163,
right calcaneum. COLEMAN IIA, SUMTER COUNTY, FLORIDA (Late
Irvingtonian): UF 21494, left partial palate with DP4; UF 24135, right
lower incisor; UF 24176, left premaxilla; UF 24177, caudal vertebra.







BULLETIN FLORIDA STATE MUSEUM


APOLLO BEACH, HILLSBOROUGH COUNTY, FLORIDA (Rancho-
labrean, possibly Irvingtonian): UF 21467, left mandible with M2 and in-
cisor; UF 21489, right palate with DP4-M1; UF 24112 (cast), right mandi-
ble with P4-M2; UF 24113 (cast), left mandible with P4-M2; UF 24114
(cast), right mandible with P4-M3 and incisor. PORT CHARLOTTE,
CHARLOTTE COUNTY, FLORIDA (Irvingtonian): UF 21464, isolated
left upper M3. NEW PORT RICHEY, PASCO COUNTY, FLORIDA
(? Rancholabrean): UF 21493, right mandible with M2. WACCASASSA RIV-
ER, LEVY COUNTY, FLORIDA (Rancholabrean, possibly Irvingtonian):
Wac 7B-UF 21486, left mandible with DP4-M1; UF 14195, edentulous
left mandible: Wac 1A-UF 21485, left lower DP4; UF 21488, left upper
P4; UF 21487, left lower incisor. SEMINOLE FIELD, PINELLAS
COUNTY, FLORIDA (Rancholabrean): AMNH 23114a, b, c, isolated in-
cisors. CLAMP CAVE, McCOLLOCH COUNTY, TEXAS (Rancho-
labrean): TMM 1295-14, right mandible with incisor, P4-M2. SAN JOSE-
CITO CAVE, NUEVO LEON, MEXICO (Rancholabrean): LACM (CIT)
2968, immature right mandible with incisor, DP4-M2. CEDAZO, AGUAS-
CALIENTES, MEXICO (Rancholabrean): UMMP V47106, right mandible
with incisor, P4-M3.
DiscussioN.--All of the middle Irvingtonian through Rancholabrean fos-
sil porcupines are clearly referrable to the living species, Erethizon dorsa-
tum. The most difficult case, as it is the oldest sample, is the Cumberland
Cave sample of Coendou cumberlandicus (White 1970). Fortunately it is a
relatively large sample of well preserved specimens and is shown to fall
within the range of morphology observed in E. dorsatum. There are no
morphological structures in the type skull (NMNH 7996) or the complete
sample of skulls and mandibles from Cumberland Cave that substantiate
generic or specific distinction from the extant species (Figs. 5-7). The astra-
galus (CM 20322) is definitely referrable to E. dorsatum.
Cope (1871) based Erethizon cloacinum from the Port Kennedy Cave
upon an isolated cheek tooth (AMNH 8576). Later he (Cope 1899) ex-
pressed doubt as to the validity of the species. More specimens have been
found since, and they are clearly E. dorsatum. Based upon the age of Port
Kennedy, White (1970) referred this material to Coendou sp.
Hibbard and Mooser (1963) referred a right mandible (UMMP V47106)
from a Rancholabream site in Aguascalientes, Mexico, to Erethizon dorsa-
tum. White (1970) referred this specimen to Coendou sp. based upon the
degree of divergence in the tooth row and the scratches on the occlusal
surfaces of the cheek teeth. I found these characters to be quite variable in
Recent and fossil E. dorsatum and must agree with Hibbard and Mooser's
original assignment. In a restudy of this specimen, Mooser and Dalquest
(1975) also support this conclusion.


Vol. 27 No. 1







FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


The immature right mandible from San Josecito Cave, Mexico, cannot
be separated from Recent Erethizon dorsatum of similar age and develop-
ment, thus supporting the original assignment of Jakway (1958).
White (1970) assigned the type skull of Erethizon godfreyi (Allen 1904)
from a subrecent volcanic fissure on the White Mountain Apache Indian
Reservation in southern Arizona and the porcupine material from Conard
Fissure, Arkansas (Brown 1908), to E. dorsatum. I agree with his assign-
ments.
Ray et al. (1963) expressed doubt about the presence of Erethizon dorsa-
tum in the Seminole Field local fauna of Florida. At that time the incisors
described by Simpson (1929) were the only known record of Erethizon in
Florida. In a re-examination of the incisors, I found multiserial enamel as
in E. dorsatum. This occurrence now seems more probable, as many other
localities in Florida have produced more complete porcupine material dur-
ing the last 15 years.
Erethizon dorsatum appears to have its earliest representatives in the
middle Irvingtonian local faunas of Cumberland Cave, Port Kennedy,
Haile XVI, and other localities in the eastern United States (Appendix A).
It may also have existed in Mexico during the Irvingtonian, where it
appears relatively common in Rancholabrean sites. Jones and Genoways
(1968) discussed the Recent and fossil range of Erethizon in Mexico. To
judge from negative evidence, it did not spread to the Great Plains and the
western United States until the latest Pleistocene. It is now distributed
over temperate North America but has retreated from its former southern
distribution in Florida and Mexico. Ray and Lipps (1970) have correlated
this demise of Erethizon in the Southeast with the activity of man. Climatic
and floral changes during the Pleistocene may also provide relevant in-
formation. Parmalee (1965) states ". . it was apparently unable to survive
climatic/vegetation change or possibly hunting pressure by early man in
many areas of its former range in eastern United States."

ORIGIN OF ERETHIZON

As a wide ranging arboreal bark feeder, Erethizon dorsatum occupies an
adaptive zone for which it apparently has very little competition. Further-
more, Erethizon can function as a moderately large generalized herbivore
able to exist in a wide variety of habitats (Taylor 1935). In its fundamental
adaptations to this way of life, Erethizon has changed very little since its
late Pliocene appearance in North America. The fossil record shows no
major structural changes from the three early fossil forms to the living
species.
The geographic history of Erethizon forms a coherent pattern. In the late
Pliocene-early Pleistocene we find two large western species (E. bath-







BULLETIN FLORIDA STATE MUSEUM


ygnathum and E. cascoensis). Meanwhile in Florida, a small species (E.
kleini, n. sp.) appears in the early Pleistocene. By the middle Pleistocene
(during the last million years) the extant species (E. dorsatum) became
widespread throughout most of North America. This species retreated from
its former range in Mexico and the Southeast since the last glacial period.
The systematic relationships between these fossil forms will not be attemp-
ted until the earlier, possibly ancestral, South American forms have been
studied, in order to determine the primitive and derived conditions within
the Erethizontidae.
In many respects Erethizon has a more derived condition in its masti-
catory, locomotive, and optic functions than Coendou. On the other hand,
Coendou has specialized arboreal features of its own. For Coendou to have
given rise to Erethizon, a major genetic and morphologic change would
have had to occur. White (1970) hypothesized that Coendou is ancestral to
Erethizon. While this may be true, the fossil record offers no real inter-
mediate forms in support of it.
One basis for determining such relationships is karyotypic. George and
Weir (1974) present two phylogenies of select hystricomorph rodent based
upon chromosomes. One phylogeny favors White's hypothesis (Coendou
ancestral to Erethizon), while the other favors Erethizon as the ancestor to
Coendou. They prefer the change from Coendou to Erethizon, as more
likely genetically. It may be still more probable that the two genera differ-
entiated from a common ancestor sometime during the early Miocene or
the late Oligocene of South America.
Geographic considerations may also shed light on this problem. It is
almost certain that the North American erthizontids came from South Amer-
ica during Pliocene times. To reach temperate latitudes they had to pass
through what is now subtropical environments in Central America. As pre-
viously mentioned, a temperate forest has existed in southern Mexico since
Miocene times (Raven and Axelrod, 1975). Although not a preferred food of
Erethizon, pine has ranged as far south as Honduras and Nicaragua, presum-
ably since the late Cenozoic (Carr 1950; Parsons 1955). This presents a major
question: was Erethizon preadapted to a temperate forest vegetation prior to
its arrival in Central and North America?
The oldest record of Erethizon in North America is from the late Blancan
Wolf Ranch fauna of San Pedro Valley, Arizona. These fossil-bearing sedi-
ments have an estimated age of 2.5 million years using magnetic polarity
correlations (Harrison 1978). Shackleton and Opdyke (1977) record oxygen
isotopes and paleomagnetics from the equatorial Pacific that indicate the
first major oceanic cooling began about 3 mya and continued to the first
major continental glaciation about 2.3-2.4 mya. Boellstorff (1978) records a
glacial till in the Central U.S. older than 2.2 mya. With the lowering of sea
level in association with this first major Pleistocene glaciation, the exposed


Vol. 27 No. 1







FRAZIER. N.A. FOSSIL ERETHIZONTID REVISION


narrow land bridge in Central America allowed the erethizontids to enter
North America for the first time. The cool temperatures and possible
temperate-to-tropical environment in Central America apparently pre-
sented no problem to the adaptable porcupines as they dispersed north-
ward into a continent with an apparently open niche. It seems likely that
Erethizon was preadapted to the temperate forest environment prior to
this interchange between the two continents.
Assuming that Erethizon was preadapted prior to Pliocene times and
before arriving in North America, Erethizon and Coendou probably differ-
entiated in the Miocene or Oligocene of South America. On chronologic
and geographic grounds, an erethizontid in that area at that time might be
expected to approximate the common ancestry of the two taxa. That com-
mon ancestor may well be the fossil South American genus Steiromys. This
genus has been reported from several Miocene deposits in South America,
including the late Miocene La Venta beds of Columbia. It is broadly com-
parable to Erethizon in both size and tooth structure (Fields 1957; Scott
1905). Friant (1936) compared the dentitions of Erethizon, Coendou, and
Steiromys and concluded that Steiromys is morphologically more similar to
Erethizon than to Coendou. The examination of the lower incisor enamel
microstructure of one specimen of Steiromys from the Miocene of Patago-
nia (KU 11605) produced values corresponding to the dimensions found in
Erethizon (Table 5). Thus, Steiromys could have been the temperate South
American equivalent of Erethizon during the Miocene and Pliocene, or
even the direct ancestor of the North American genus. The most probable
hypothesis is that Steiromys was ancestral to Erethizon or differentiated
from the ancestor of Erethizon during the early Miocene and that a com-
mon ancestor with Coendou may be even older, possibly late Oligocene.
Further morphological and biochemical comparisons of the four living
genera (Echinoprocta, Chaetomys, Coendou, and Erethizon) along with a
more extensive survey of the fossil forms known from South America will
shed more light upon the systematic relationships within the family.


CONCLUSIONS

1) All of the late Blancan-early Irvingtonian fossil porcupines of North
America are Erethizon, not Coendou.
2) Four species of Erethizon are recognized in the fossil record of North
America: E. bathygnathum, a large extinct form from the late Blan-
can-early Irvingtonian of the western United States; E. cascoensis, an
extinct species known only from the early Irvingtonian El Casco Local
fauna of southern California; E. kleini n. sp., a small form from the
early Irvingtonian Inglis IA local fauna and later Irvingtonian Merritt







BULLETIN FLORIDA STATE MUSEUM


Island local fauna of Florida; and the extant species E. dorsatum, from
the remaining Irvingtonian-Rancholabrean localities, including Port
Kennedy and Cumberland Cave.
3) The erethizontids apparently dispersed to North America from South
America approximately 3 mya via the Central American land bridge
during the major faunal interchange between the two continents.
4) In the late Blancan-early Irvingtonian times (2.0-3.0 mya), three spe-
cies of porcupine lived in North America. Erethizon bathygnathum, a
large form, apparently followed the Cordillera northward. E. cascoen-
sis apparently was restricted to the Southwest as it is known only from
one locality in southern California. E. kleini, n. sp., a small Coendou-
sized porcupine, probably followed the Gulf Coast corridor into Flor-
ida as did many other members of the South American fauna. This
endemic smaller species probably became extinct during the Irving-
tonian land mammal age, possibly at the end of the Kansan glacial
period. E. dorsatum, appears in North America during the Irving-
tonian and had dispersed throughout the continent by the close of the
Pleistocene.
5) The genus Erethizon has changed very little ecologically or morpho-
logically since its appearance in North America about 2.5 mya.
6) Steiromys, a South American Miocene erethizontid, may be ancestral
to Erethizon or share a common ancestor with Erethizon. Erethizon
and Coendou probably differentiated in the late Oligocene of South
America.


Vol. 27 No. 1








FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION


APPENDIX A


PLEISTOCENE FAUNAS THAT CONTAIN PORCUPINES


Locality County State References


Wolf Ranch
El Casco
Vallecito Creek

Inglis IA
?Merritt Island
Grand View


Conard Fissure
Haile XVI
Port Charlotte
Apollo Beach
Coleman IIA
Cumberland Cave
Port Kennedy
Trout Cave


*Little Bear Creek
*Stanfield-Worley Site
Stuttgart
Wolcott #2
Waccasassa River
New Port Richey
Seminole Field
Spring Cave or
Harrisburg Cave
Schmitt Site
Jaguar Cave
American Falls
*Riverton Site
*Meyer Cave
Savage Cave
Welsh Cave
Cavetown
Brynjulfson Caves
Crankshaft Cave
*Tick Creek Cave
Cherokee Cave
Zoo Cave
Isleta Caves
*Hermit's Cave


LATE BLANCAN-EARLY IRVINGTONIAN
Cochise AZ Harrison 1978
Riverside CA White 1970
San Diego CA White 1968, 1970; Downs and White
1968
Citrus FL Klein 1971; Webb 1976
Brevard FL this paper
Owyhee ID Wilson 1935; Shotwell 1970


MIDDLE-LATE IRVINGTONIAN


Newton
Alachua
Charlotte
Hillsborough
Sumter
Allegany
Chester
Pendleton


RANCHOLABREAN*


Colbert
Colbert
Prairie
Pima
Levy
Pasco
Pinellas
Walker


Dubuque
Lemhi
Power
Crawford
Monroe
Logan
Woodford
Washington
Boone
Jefferson
Phelps
St. Louis
Taney
Bernalillo
Eddy


Brown 1908
Jackson 1978
this paper
this paper
Martin 1974
Gidley and Gazin 1938
Cope 1871, 1899
Zakrzewski 1975b; Frazier 1977


Barkalow 1961
Parmalee 1963
NMNH collections
Lindsay and Tessman 1974
Webb 1974; Jackson 1975
this paper
Simpson 1929
Ray and Lipps 1970

Eshelman 1972
Guilday and Adam 1967
Gazin 1935
Parmalee 1962
Parmalee 1967
Guilday, Hamilton and McCrady 1971
Guilday, Hamilton and McCrady 1971
Hay 1921; Franz and Slifer 1971
Parmalee 1971
Parmalee, Oesch and Guilday 1969
Parmalee 1965
Simpson 1949
Hood and Hawksley 1975
Harris and Findley 1964
Lloyd Tanner, pers. comm.








BULLETIN FLORIDA STATE MUSEUM


Locality County State References


New Paris No. 4
Durham's Cave
Hosterman's Pit
Sheep Rock Shelter
Hartman's Cave
*Vess Cave
Craig Quarry
Banshee Hold
Lookout Cave
*Bible Site
*Westmoreland-
Barber Site
Robinson Cave
Guy Wilson Cave
*Big Bone Cave
Clamp Cave
Silver Creek
Natural Chimneys
Clark's Cave
Ripplemead Fissures
Edinburg
Windy Mount Cave
Eagle Cave
Hoffman School Cave
Mandy Walters Cave
Cedazo

Cuatro Ciengas
San Josecito Cave


Bedford
Bucks
Centre
Huntingdon
Monroe
Anderson
Blount
Cumberland
Hamilton
Marion
Marion

Overton
Sullivan
Van Buren
McColloch
Summit
Augusta
Bath
Giles
Shenandoah
Greenbriar
Pendleton
Pendleton
Pendleton
Aguascalientes

Coahuila
Nuevo Leon


Guilday, Martin and McCrady 1964
Leidy 1887
Guilday 1967
Guilday and Parmalee 1965
Leidy 1887
Cahn 1939
Corgan 1976
Corgan 1976
Corgan 1976
Parmalee 1966
Guilday and Tanner 1966


TN Guilday, Hamilton and McCrady 1969
TN Corgan 1976
TN Mercer 1897
TX this paper
UT Miller 1976
VA Guilday 1962
VA Guilday, Parmalee and McCrady 1977
VA Weems and Higgins 1977
VA NMNH collections
WV this paper
WV Guilday and Hamilton 1973
WV Guilday and Hamilton 1978
WV Guilday and Hamilton 1978
Mexico Hibbard and Mooser 1963; Mooser and
Dalquest 1975
Mexico Gilmore 1947
Mexico Jakway 1958


*Indicates association with man.
APPENDIX B
GRAPHS OF CRANIAL MEASUREMENTS

RECENT


adult
subadult
adult
subadult

adult
subadult
adult
subadult
adult
subadult

KU 11605


FOSSIL


Erethizon dorsatum
E. dorsatum
Coendou
Coendou

Erethizon dorsatum
E. dorsatum
Erethizon bathygnathum
E. bathygnathum
Erethizon kleini
E. kleini
Erethizon cascoensis
Steiromys


Numbers in symbols correspond to locality numbers in Figure 2.


Vol. 27 No. 1









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FIGURE B-1


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65




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FIGURE B-12
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FIGURE B-15
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0
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0* * * *

+ * *



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1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 71

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76 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1

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and B. Patterson. 1970. Relationships among hystricognathous and hystricomor-
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Woods, C. A. 1974. Erethizon dorsatum. Mammalian Species 29:1-6.
and E. B. Howland. 1979. Adaptive radiation of capromyid rodents: Anatomy of
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Zakrzewski, R. J. 1975a. Pleistocene stratigraphy and paleontology in western Kansas: The
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1975b. The late Pleistocene arvicoline rodent Atopomys. Ann. Carnegie Mus.
45(12):255-261.








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